Land use, habitat integrity, and aquatic insect assemblages in Central Amazonian streams

23
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Transcript of Land use, habitat integrity, and aquatic insect assemblages in Central Amazonian streams

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bull If we do not receive your corrections within 48 hours we will send you a reminder

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Fax to +44 207 806 8278 or +44 870 762 8807 (UK)or +91 44 4208 9499 (INDIA)To Springer Correction Team

6amp7 5th Street Radhakrishnan Salai Chennai Tamil Nadu India ndash 600004Re Hydrobiologia DOI101007s10750-008-9441-x

Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsAuthors JorgeL Nessimian middot EduardoM Venticinque middot Jansen Zuanon middot Paulo Marco middot Marcelo

Gordo middot Luana Fidelis middot Joana Drsquoarc Batista middot Leandro Juen

Permission to publishI have checked the proofs of my article andq I have no corrections The article is ready to be published without changes

q I have a few corrections I am enclosing the following pagesq I have made many corrections Enclosed is the complete article

Date signature ______________________________________________________________________________

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Metadata of the article that will be visualized in OnlineFirst

ArticleTitle Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsArticle Sub-Title

Article CopyRight - Year Springer Science+Business Media BV 2008(This will be the copyright line in the final PDF)

Journal Name Hydrobiologia

Corresponding Author Family Name NessimianParticle

Given Name Jorge LSuffix

Division Departamento de Zoologia IB

Organization Universidade Federal do Rio de Janeiro (UFRJ)

Address CP 68044 21944-970 Rio de Janeiro RJ Brazil

Email nessimiaacdufrjbr

Author Family Name VenticinqueParticle

Given Name Eduardo MSuffix

Division

Organization Wildlife Conservation Society (WCS) Andes Amazon Conservation Program

Address Rua dos Jatobaacutes 274 CEP 69085-380 Manaus AM Brazil

Division

Organization INPA (National Institute of Amazonian Research) PDBFF

Address Manaus AM Brazil

Division Departamentos de Ecologia e Silvicultura

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address Manaus AM Brazil

Email

Author Family Name ZuanonParticle

Given Name JansenSuffix

Division CPBA

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name MarcoParticle DeGiven Name PauloSuffix Jr

Division Departamento de Ecologia Geral

Organization Universidade Federal de Goiaacutes (UFG)

Address 74001-970 Goiania GO Brazil

Email

Author Family Name GordoParticle

Given Name MarceloSuffix

Division Instituto de Ciecircncias Bioloacutegicas

Organization Universidade Federal do Amazonas (UFAM)

Address Manaus AM Brazil

Email

Author Family Name FidelisParticle

Given Name LuanaSuffix

Division DCEN

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name Drsquoarc BatistaParticle

Given Name JoanaSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Author Family Name JuenParticle

Given Name LeandroSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Schedule

Received 26 March 2007

Revised 15 May 2008

Accepted 26 May 2008

Abstract The distribution and composition of aquatic insect communities in streams at a local scale are considered tobe primarily determined by environmental factors and interactive relationships within the system Here weevaluated the effects of forest fragmentation and forest cover changes on habitat characteristics of streamlets(igarapeacutes) in Amazonian forests and on the aquatic insect communities found there We also developed ahabitat integrity index (HII) based on Petersenrsquos protocol (1992) to evaluate physical integrity of thesestreamlets and to determine its efficiency to interpret the environmental impacts on this system We studied20 small streams at the Biological Dynamics of Forest Fragments Project (BDFFP INPASI) study areasCentral Amazonia 80 km north of Manaus Amazonas State Brazil The vegetation cover was estimated byusing LANDSAT images and classified in the following categories exposed soil pastures secondary forests(capoeiras) and primary forests Stream habitat features were evaluated by using a HII based on visualassessment of local characteristics Aquatic insects were sampled in four major stream substrates litter

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

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Sectionparagraph Details required Authorrsquos response

Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

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Habitat integrity and faunal metrics

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Please provide citation for Reference Resh and Jackson (1993)

Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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Hydrobiologia

DOI 101007s10750-008-9441-x

Au

tho

r P

ro

of

UNCORRECTEDPROOF

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

123

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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UNCORRECTEDPROOF

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

123

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UNCORRECTEDPROOF

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

Fax to +44 207 806 8278 or +44 870 762 8807 (UK)or +91 44 4208 9499 (INDIA)To Springer Correction Team

6amp7 5th Street Radhakrishnan Salai Chennai Tamil Nadu India ndash 600004Re Hydrobiologia DOI101007s10750-008-9441-x

Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsAuthors JorgeL Nessimian middot EduardoM Venticinque middot Jansen Zuanon middot Paulo Marco middot Marcelo

Gordo middot Luana Fidelis middot Joana Drsquoarc Batista middot Leandro Juen

Permission to publishI have checked the proofs of my article andq I have no corrections The article is ready to be published without changes

q I have a few corrections I am enclosing the following pagesq I have made many corrections Enclosed is the complete article

Date signature ______________________________________________________________________________

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Metadata of the article that will be visualized in OnlineFirst

ArticleTitle Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsArticle Sub-Title

Article CopyRight - Year Springer Science+Business Media BV 2008(This will be the copyright line in the final PDF)

Journal Name Hydrobiologia

Corresponding Author Family Name NessimianParticle

Given Name Jorge LSuffix

Division Departamento de Zoologia IB

Organization Universidade Federal do Rio de Janeiro (UFRJ)

Address CP 68044 21944-970 Rio de Janeiro RJ Brazil

Email nessimiaacdufrjbr

Author Family Name VenticinqueParticle

Given Name Eduardo MSuffix

Division

Organization Wildlife Conservation Society (WCS) Andes Amazon Conservation Program

Address Rua dos Jatobaacutes 274 CEP 69085-380 Manaus AM Brazil

Division

Organization INPA (National Institute of Amazonian Research) PDBFF

Address Manaus AM Brazil

Division Departamentos de Ecologia e Silvicultura

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address Manaus AM Brazil

Email

Author Family Name ZuanonParticle

Given Name JansenSuffix

Division CPBA

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name MarcoParticle DeGiven Name PauloSuffix Jr

Division Departamento de Ecologia Geral

Organization Universidade Federal de Goiaacutes (UFG)

Address 74001-970 Goiania GO Brazil

Email

Author Family Name GordoParticle

Given Name MarceloSuffix

Division Instituto de Ciecircncias Bioloacutegicas

Organization Universidade Federal do Amazonas (UFAM)

Address Manaus AM Brazil

Email

Author Family Name FidelisParticle

Given Name LuanaSuffix

Division DCEN

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name Drsquoarc BatistaParticle

Given Name JoanaSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Author Family Name JuenParticle

Given Name LeandroSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Schedule

Received 26 March 2007

Revised 15 May 2008

Accepted 26 May 2008

Abstract The distribution and composition of aquatic insect communities in streams at a local scale are considered tobe primarily determined by environmental factors and interactive relationships within the system Here weevaluated the effects of forest fragmentation and forest cover changes on habitat characteristics of streamlets(igarapeacutes) in Amazonian forests and on the aquatic insect communities found there We also developed ahabitat integrity index (HII) based on Petersenrsquos protocol (1992) to evaluate physical integrity of thesestreamlets and to determine its efficiency to interpret the environmental impacts on this system We studied20 small streams at the Biological Dynamics of Forest Fragments Project (BDFFP INPASI) study areasCentral Amazonia 80 km north of Manaus Amazonas State Brazil The vegetation cover was estimated byusing LANDSAT images and classified in the following categories exposed soil pastures secondary forests(capoeiras) and primary forests Stream habitat features were evaluated by using a HII based on visualassessment of local characteristics Aquatic insects were sampled in four major stream substrates litter

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

Author Query Form

Please ensure you fill out your response to the queries raised below

and return this form along with your corrections

Dear Author

During the preparation of your manuscript for typesetting some questions have arisen These are listed below Please check your typeset proof carefully and mark any corrections in the margin of the proof or compile them as a separate list This form should then be returned with your marked prooflist of corrections to spr_corrections1spscoin Disk use In some instances we may be unable to process the electronic file of your article andor artwork In that case we have for

efficiency reasons proceeded by using the hard copy of your manuscript If this is the case the reasons are indicated below Disk damaged Incompatible file format LaTeX file for non-LaTeX journal Virus infected Discrepancies between electronic file and (peer-reviewed therefore definitive) hard copy Other We have proceeded as follows Manuscript scanned Manuscript keyed in Artwork scanned Files only partly used (parts processed differently helliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphellip) Bibliography If discrepancies were noted between the literature list and the text references the following may apply The references listed below were noted in the text but appear to be missing from your literature list Please complete the list or

remove the references from the text Uncited references This section comprises references that occur in the reference list but not in the body of the text

Please position each reference in the text or delete it Any reference not dealt with will be retained in this section Queries andor remarks

Sectionparagraph Details required Authorrsquos response

Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

Please provide details for Reference Laurence (2001) which is cited in text but is not present in the reference list

Reference Merritt and Cummins (1984) has been changed to Merritt and Cummins (1996) as per the reference list Please check

Habitat integrity and faunal metrics

The variable ldquop = 0590rdquo has been changed to ldquor = 0590rdquo Please check

Please provide citation for Reference Resh and Jackson (1993)

Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Hydrobiologia

DOI 101007s10750-008-9441-x

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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Metadata of the article that will be visualized in OnlineFirst

ArticleTitle Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsArticle Sub-Title

Article CopyRight - Year Springer Science+Business Media BV 2008(This will be the copyright line in the final PDF)

Journal Name Hydrobiologia

Corresponding Author Family Name NessimianParticle

Given Name Jorge LSuffix

Division Departamento de Zoologia IB

Organization Universidade Federal do Rio de Janeiro (UFRJ)

Address CP 68044 21944-970 Rio de Janeiro RJ Brazil

Email nessimiaacdufrjbr

Author Family Name VenticinqueParticle

Given Name Eduardo MSuffix

Division

Organization Wildlife Conservation Society (WCS) Andes Amazon Conservation Program

Address Rua dos Jatobaacutes 274 CEP 69085-380 Manaus AM Brazil

Division

Organization INPA (National Institute of Amazonian Research) PDBFF

Address Manaus AM Brazil

Division Departamentos de Ecologia e Silvicultura

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address Manaus AM Brazil

Email

Author Family Name ZuanonParticle

Given Name JansenSuffix

Division CPBA

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name MarcoParticle DeGiven Name PauloSuffix Jr

Division Departamento de Ecologia Geral

Organization Universidade Federal de Goiaacutes (UFG)

Address 74001-970 Goiania GO Brazil

Email

Author Family Name GordoParticle

Given Name MarceloSuffix

Division Instituto de Ciecircncias Bioloacutegicas

Organization Universidade Federal do Amazonas (UFAM)

Address Manaus AM Brazil

Email

Author Family Name FidelisParticle

Given Name LuanaSuffix

Division DCEN

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name Drsquoarc BatistaParticle

Given Name JoanaSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Author Family Name JuenParticle

Given Name LeandroSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Schedule

Received 26 March 2007

Revised 15 May 2008

Accepted 26 May 2008

Abstract The distribution and composition of aquatic insect communities in streams at a local scale are considered tobe primarily determined by environmental factors and interactive relationships within the system Here weevaluated the effects of forest fragmentation and forest cover changes on habitat characteristics of streamlets(igarapeacutes) in Amazonian forests and on the aquatic insect communities found there We also developed ahabitat integrity index (HII) based on Petersenrsquos protocol (1992) to evaluate physical integrity of thesestreamlets and to determine its efficiency to interpret the environmental impacts on this system We studied20 small streams at the Biological Dynamics of Forest Fragments Project (BDFFP INPASI) study areasCentral Amazonia 80 km north of Manaus Amazonas State Brazil The vegetation cover was estimated byusing LANDSAT images and classified in the following categories exposed soil pastures secondary forests(capoeiras) and primary forests Stream habitat features were evaluated by using a HII based on visualassessment of local characteristics Aquatic insects were sampled in four major stream substrates litter

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

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Habitat integrity and faunal metrics

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Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
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Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
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UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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Hydrobiologia

DOI 101007s10750-008-9441-x

Au

tho

r P

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of

UNCORRECTEDPROOF

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

123

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

123

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

Hydrobiologia

123

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

Hydrobiologia

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UNCORRECTEDPROOF

318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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tho

r P

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of

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

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Metadata of the article that will be visualized in OnlineFirst

ArticleTitle Land use habitat integrity and aquatic insect assemblages in Central Amazonian streamsArticle Sub-Title

Article CopyRight - Year Springer Science+Business Media BV 2008(This will be the copyright line in the final PDF)

Journal Name Hydrobiologia

Corresponding Author Family Name NessimianParticle

Given Name Jorge LSuffix

Division Departamento de Zoologia IB

Organization Universidade Federal do Rio de Janeiro (UFRJ)

Address CP 68044 21944-970 Rio de Janeiro RJ Brazil

Email nessimiaacdufrjbr

Author Family Name VenticinqueParticle

Given Name Eduardo MSuffix

Division

Organization Wildlife Conservation Society (WCS) Andes Amazon Conservation Program

Address Rua dos Jatobaacutes 274 CEP 69085-380 Manaus AM Brazil

Division

Organization INPA (National Institute of Amazonian Research) PDBFF

Address Manaus AM Brazil

Division Departamentos de Ecologia e Silvicultura

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address Manaus AM Brazil

Email

Author Family Name ZuanonParticle

Given Name JansenSuffix

Division CPBA

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name MarcoParticle DeGiven Name PauloSuffix Jr

Division Departamento de Ecologia Geral

Organization Universidade Federal de Goiaacutes (UFG)

Address 74001-970 Goiania GO Brazil

Email

Author Family Name GordoParticle

Given Name MarceloSuffix

Division Instituto de Ciecircncias Bioloacutegicas

Organization Universidade Federal do Amazonas (UFAM)

Address Manaus AM Brazil

Email

Author Family Name FidelisParticle

Given Name LuanaSuffix

Division DCEN

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name Drsquoarc BatistaParticle

Given Name JoanaSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Author Family Name JuenParticle

Given Name LeandroSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Schedule

Received 26 March 2007

Revised 15 May 2008

Accepted 26 May 2008

Abstract The distribution and composition of aquatic insect communities in streams at a local scale are considered tobe primarily determined by environmental factors and interactive relationships within the system Here weevaluated the effects of forest fragmentation and forest cover changes on habitat characteristics of streamlets(igarapeacutes) in Amazonian forests and on the aquatic insect communities found there We also developed ahabitat integrity index (HII) based on Petersenrsquos protocol (1992) to evaluate physical integrity of thesestreamlets and to determine its efficiency to interpret the environmental impacts on this system We studied20 small streams at the Biological Dynamics of Forest Fragments Project (BDFFP INPASI) study areasCentral Amazonia 80 km north of Manaus Amazonas State Brazil The vegetation cover was estimated byusing LANDSAT images and classified in the following categories exposed soil pastures secondary forests(capoeiras) and primary forests Stream habitat features were evaluated by using a HII based on visualassessment of local characteristics Aquatic insects were sampled in four major stream substrates litter

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

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remove the references from the text Uncited references This section comprises references that occur in the reference list but not in the body of the text

Please position each reference in the text or delete it Any reference not dealt with will be retained in this section Queries andor remarks

Sectionparagraph Details required Authorrsquos response

Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

Please provide details for Reference Laurence (2001) which is cited in text but is not present in the reference list

Reference Merritt and Cummins (1984) has been changed to Merritt and Cummins (1996) as per the reference list Please check

Habitat integrity and faunal metrics

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Please provide citation for Reference Resh and Jackson (1993)

Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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Hydrobiologia

DOI 101007s10750-008-9441-x

Au

tho

r P

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

123

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

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tho

r P

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of

UNCORRECTEDPROOF

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

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Au

tho

r P

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of

Email

Author Family Name GordoParticle

Given Name MarceloSuffix

Division Instituto de Ciecircncias Bioloacutegicas

Organization Universidade Federal do Amazonas (UFAM)

Address Manaus AM Brazil

Email

Author Family Name FidelisParticle

Given Name LuanaSuffix

Division DCEN

Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)

Address CP 478 69011970 Manaus AM Brazil

Email

Author Family Name Drsquoarc BatistaParticle

Given Name JoanaSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Author Family Name JuenParticle

Given Name LeandroSuffix

Division Departamento de Biologia Geral

Organization Universidade Federal de Viccedilosa (UFV)

Address 36571-000 Vicosa MG Brazil

Email

Schedule

Received 26 March 2007

Revised 15 May 2008

Accepted 26 May 2008

Abstract The distribution and composition of aquatic insect communities in streams at a local scale are considered tobe primarily determined by environmental factors and interactive relationships within the system Here weevaluated the effects of forest fragmentation and forest cover changes on habitat characteristics of streamlets(igarapeacutes) in Amazonian forests and on the aquatic insect communities found there We also developed ahabitat integrity index (HII) based on Petersenrsquos protocol (1992) to evaluate physical integrity of thesestreamlets and to determine its efficiency to interpret the environmental impacts on this system We studied20 small streams at the Biological Dynamics of Forest Fragments Project (BDFFP INPASI) study areasCentral Amazonia 80 km north of Manaus Amazonas State Brazil The vegetation cover was estimated byusing LANDSAT images and classified in the following categories exposed soil pastures secondary forests(capoeiras) and primary forests Stream habitat features were evaluated by using a HII based on visualassessment of local characteristics Aquatic insects were sampled in four major stream substrates litter

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

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efficiency reasons proceeded by using the hard copy of your manuscript If this is the case the reasons are indicated below Disk damaged Incompatible file format LaTeX file for non-LaTeX journal Virus infected Discrepancies between electronic file and (peer-reviewed therefore definitive) hard copy Other We have proceeded as follows Manuscript scanned Manuscript keyed in Artwork scanned Files only partly used (parts processed differently helliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphelliphellip) Bibliography If discrepancies were noted between the literature list and the text references the following may apply The references listed below were noted in the text but appear to be missing from your literature list Please complete the list or

remove the references from the text Uncited references This section comprises references that occur in the reference list but not in the body of the text

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Sectionparagraph Details required Authorrsquos response

Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

Please provide details for Reference Laurence (2001) which is cited in text but is not present in the reference list

Reference Merritt and Cummins (1984) has been changed to Merritt and Cummins (1996) as per the reference list Please check

Habitat integrity and faunal metrics

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Please provide citation for Reference Resh and Jackson (1993)

Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Hydrobiologia

DOI 101007s10750-008-9441-x

Au

tho

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

123

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

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tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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ro

of

deposited in pools or backwaters litter retained in riffles sand and marginal banks Stream habitatcharacteristics were significantly correlated to land use and riparian forest condition Overall aquatic insectrichness and Ephemeroptera Plecoptera and Trichoptera (EPT) richness were significantly lower in pasturestreams and their taxonomic composition differed significantly from streams in forested areas Howeverthese metrics were not significantly correlated to the stream HII Taxonomic composition of bank insectassemblages changed significantly between streams with low and high values of HII There was no significantrelationship between the proportion of primary forest cover and the faunal metrics Only drastic changes inthe vegetal cover seem to induce significant changes in the aquatic insect community Matrix habitatheterogeneity distance to forest fragments the presence of areas of secondary forest and the intrinsic capacityto disperse in many of the insect groups may have contributed to attenuate the effects of habitat disturbanceon aquatic insect assemblages in streamlets

Keywords (separated by -) Habitat integrity - Streams - Forest cover - Forest fragments - Aquatic insects - Amazonia

Footnote Information Publication number 00 of the PDBFF Technical Series PDBFFmdashINPASI and number 00 of the IgarapeacutesProjectHandling editor J TrexlerElectronic supplementary material The online version of this article (doi101007s10750-008-9441-x)contains supplementary material which is available to authorized users

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remove the references from the text Uncited references This section comprises references that occur in the reference list but not in the body of the text

Please position each reference in the text or delete it Any reference not dealt with will be retained in this section Queries andor remarks

Sectionparagraph Details required Authorrsquos response

Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

Please provide details for Reference Laurence (2001) which is cited in text but is not present in the reference list

Reference Merritt and Cummins (1984) has been changed to Merritt and Cummins (1996) as per the reference list Please check

Habitat integrity and faunal metrics

The variable ldquop = 0590rdquo has been changed to ldquor = 0590rdquo Please check

Please provide citation for Reference Resh and Jackson (1993)

Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
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We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

UNCORRECTEDPROOF

UNCORRECTEDPROOF

PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Hydrobiologia

DOI 101007s10750-008-9441-x

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

123

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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r P

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of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

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of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

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of

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Please check affiliation of ldquoEduardo M Venticinquerdquo

Please check the publication numbers in Article note and contribution numbers in Acknowledgments

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Journal HYDR-10750 Article 9441

Jorge Nessimian
Note
Lawrence WF 2001 The Hyper-Diverse Flora of the Central Amazon An Overwiew pp 47-53 In Bierregaard R O Jr Gascon C Lovejoy T E amp Mesquita R (eds) Lessons from Amazonia the ecology and conservation of a fragmented forest Yale University Press New Haven and London
Jorge Nessimian
Note
Please remove the reference at lines 916-92013
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Checked
Jorge Nessimian
Note
Publication number 515 of the PDBFF Technical Series13PDBFF-INPASTRI (Biological Dynamics of Forest Fragments Project Instituto Nacional de Pesquisas da Amazonia and Smithsonian Tropical Research Institute) and number 09 of the Igarapeacutes Project13

Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

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PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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DOI 101007s10750-008-9441-x

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

Hydrobiologia

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

Hydrobiologia

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

123

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

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Please provide accessed date for Reference StatSoft Inc (2001)

Please provide details for lsquoboldrsquo emphasis present in Table 3

Jorge Nessimian
Note
Please add to caption Significative values of p are in bold
Jorge Nessimian
Note
We did not accessed the page Please exclude (wwwstatsoftcom) from the reference at line 598

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PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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Hydrobiologia

DOI 101007s10750-008-9441-x

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

123

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

Hydrobiologia

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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PRIMARY RESEARCH PAPER1

2 Land use habitat integrity and aquatic insect assemblages

3 in Central Amazonian streams

4 Jorge L Nessimian Eduardo M Venticinque

5 Jansen Zuanon Paulo De Marco Jr Marcelo Gordo

6 Luana Fidelis Joana Drsquoarc Batista Leandro Juen

7 Received 26 March 2007 Revised 15 May 2008 Accepted 26 May 20088 Springer Science+Business Media BV 2008

9 Abstract The distribution and composition of aqua-

10 tic insect communities in streams at a local scale are

11 considered to be primarily determined by environ-

12 mental factors and interactive relationships within the

13 system Here we evaluated the effects of forest

14 fragmentation and forest cover changes on habitat

15characteristics of streamlets (igarapes) in Amazonian

16forests and on the aquatic insect communities found

17thereWe also developed a habitat integrity index (HII)

18based on Petersenrsquos protocol (1992) to evaluate

19physical integrity of these streamlets and to determine

20its efficiency to interpret the environmental impacts on

J Zuanon

CPBA Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

P De Marco Jr

Departamento de Ecologia Geral Universidade Federal de

Goias (UFG) 74001-970 Goiania GO Brazil

M Gordo

Instituto de Ciencias Biologicas Universidade Federal do

Amazonas (UFAM) Manaus AM Brazil

L Fidelis

DCEN Instituto Nacional de Pesquisas da Amazonia

(INPA) CP 478 69011970 Manaus AM Brazil

J Drsquoarc Batista L JuenDepartamento de Biologia Geral Universidade Federal de

Vicosa (UFV) 36571-000 Vicosa MG Brazil

A1 Publication number 00 of the PDBFF Technical Series

A2 PDBFFmdashINPASI and number 00 of the Igarapes Project

A3 Handling editor J Trexler

A4 Electronic supplementary material The online version ofA5 this article (doi101007s10750-008-9441-x) containsA6 supplementary material which is available to authorized users

A7 J L Nessimian (amp)

A8 Departamento de Zoologia IB Universidade Federal do

A9 Rio de Janeiro (UFRJ) CP 68044 21944-970

A10 Rio de Janeiro RJ Brazil

A11 e-mail nessimiaacdufrjbr

A12 E M Venticinque

A13 Wildlife Conservation Society (WCS) Andes Amazon

A14 Conservation Program Rua dos Jatobas 274

A15 CEP 69085-380 Manaus AM Brazil

A16 E M Venticinque

A17 INPA (National Institute of Amazonian Research)

A18 PDBFF Manaus AM Brazil

A19 E M Venticinque

A20 Departamentos de Ecologia e Silvicultura Instituto

A21 Nacional de Pesquisas da Amazonia (INPA) Manaus

A22 AM Brazil

123

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Hydrobiologia

DOI 101007s10750-008-9441-x

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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tho

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UNCORRECTEDPROOF

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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tho

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UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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262626262626 this system We studied 20 small streams at the

27 Biological Dynamics of Forest Fragments Project

28 (BDFFP INPASI) study areas Central Amazonia

29 80 km north of Manaus Amazonas State Brazil The

30 vegetation cover was estimated by using LANDSAT

31 images and classified in the following categories

32 exposed soil pastures secondary forests (capoeiras)

33 and primary forests Stream habitat features were

34 evaluated by using a HII based on visual assessment of

35 local characteristics Aquatic insects were sampled in

36 four major stream substrates litter deposited in pools

37 or backwaters litter retained in riffles sand and

38 marginal banks Stream habitat characteristics were

39 significantly correlated to land use and riparian forest

40 conditionOverall aquatic insect richness andEpheme-

41 roptera Plecoptera and Trichoptera (EPT) richness

42 were significantly lower in pasture streams and their

43 taxonomic composition differed significantly from

44 streams in forested areas However these metrics were

45 not significantly correlated to the stream HII Taxo-

46 nomic composition of bank insect assemblages

47 changed significantly between streams with low and

48 high values of HII There was no significant relation-

49 ship between the proportion of primary forest cover

50 and the faunal metrics Only drastic changes in the

51 vegetal cover seem to induce significant changes in the

52 aquatic insect community Matrix habitat heterogene-

53 ity distance to forest fragments the presence of areas

54 of secondary forest and the intrinsic capacity to

55 disperse in many of the insect groups may have

56 contributed to attenuate the effects of habitat distur-

57 bance on aquatic insect assemblages in streamlets

58 Keywords Habitat integrity Streams

59 Forest cover Forest fragments Aquatic insects

60 Amazonia

61

62 Introduction

63 Human occupation in Amazonia and the associated

64 loss of forest cover have resulted in the degradation

65 of rivers and streams Major impacts on Amazonian

66 rivers include sediment filling and removal of

67 substrate material in river beds water draining

68 modification of shore areas dam and reservoir

69 construction raw domestic sewage inputs as well

70 as agricultural cattle mining and industrial effluents

71(McClain amp Elsenbeer 2001 Davidson et al 2004

72Melo et al 2005) In addition removal or substitu-

73tion of riparian vegetation has a direct negative effect

74on the input of organic matter that constitutes the

75primary energy source of rivers trophic chains (De

76Long amp Brusven 1994 Pozo et al 1997) These

77have resulted in changes in physical habitat hydrol-

78ogy and water quality in streams and rivers All these

79factors lead to drastic changes in aquatic biota

80causing loss of diversity in the system The effects of

81human activity especially deforestation in Central

82Amazonia affect directly and negatively the small

83watercourses Since there is a large amount of forest

84cover still intact the impacts of forest loss and

85fragmentation are not readily perceived in higher

86order stretches (Smith et al 1995 Davidson et al

872004) Besides that it is not always possible to detect

88the effects of these negative environmental impacts

89on the aquatic biota present in streamlets in a simple

90and fast way because there is no protocol of

91environmental evaluation adapted to the Amazonian

92conditions

93Riparian forests are essential for the protection of

94fluvial systems as they prevent erosion loss of nutrients

95intake of sediments and other pollutants and they also

96contribute to the maintenance of the biota (Zweig amp

97Rabeni 2001 Sparovek et al 2002) Modifications

98such as fragmentation or changes in the forest cover

99lead to alterations in the habitat structure including

100litter fall (Sizer 1992) and changes in the composition

101of allochthonous material carried to the streams which

102determines changes in their structure and function

103(Benstead et al 2003 Benstead amp Pringle 2004) In

104Brazil Amazonian deforestation has happened at a fast

105rate despite the effortsmade by governmental and non-

106governmental agencies for the last years In Manaus

107area central Amazonia studies about the effects of

108forest fragmentation on biota have been performed for

109more than 25 years (Bierregaard et al 2001 Gascon

110et al 2001) One of the central objectives of the

111Biological Dynamics of Forest Fragments Project

112(BDFFP) is to study the ecological effects of forest

113fragmentation on Tropical Forest areas (Lovejoy et al

1141983 Gascon et al 2001) Nevertheless almost all

115results obtained so far correspond to terrestrial systems

116As part of theBDFFP the Igarapes Project is devoted to

117the study of effects of forest fragmentation and changes

118of vegetation cover on the integrity of structure and

119function in small forest streams This study aimed to

Hydrobiologia

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

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UNCORRECTEDPROOF

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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120 evaluate the effects of landscape changes on the

121 structure and functioning of small forest streams in

122 the Brazilian central Amazon as perceived by changes

123 in aquatic insect communities Furthermore to accom-

124 plish this objective we employed an index of habitat

125 integrity adapted to Amazonian environmental condi-

126 tions and compared the results with faunal

127 characteristics of insect communities

128 Study area

129 The BDFFP study site is composed of replicated

130 series of forest preserves of 1 10 and 100 ha areas

131 experimentally isolated from the surrounding contin-

132 uous primary forest matrix (for details on isolation

133 procedures and characteristics of forest fragments

134 see Gascon amp Bierregaard 2001) It is situated 60ndash

135 90 km north from the city of Manaus (Amazonas

136 State Brazil) The forest cover of BDFFP area was

137 partially removed 30 years ago for establishing cattle

138 farms Some forest fragments were maintained some

139 of the cleared areas were abandoned and the process

140 of regeneration was naturally established All the area

141 is surrounded by primary forest which extends

142 unbroken for hundreds of kilometers to the north

143 east and west (Gascon amp Bierregaard 2001 Gascon

144 et al 2001) (Fig 1)

145The study area is classified as tropical moist forest

146with a mean annual rainfall of about 2200 mm

147ranging 1900ndash2500 mm There is a pronounced dry

148season from June to October with less than 100 mm of

149monthly rainfall The forest canopy reaches up to 30ndash

15037 m tall with emergent trees as high as 55 m This is

151one of the most diverse forest communities in the

152world with at least 280 tree speciesha (Oliveira amp

153Mori 1999) About 1300 tree species occur in the

154project area (Laurence 2001) The landscape is located

155in Pleistocene terraces of interglacial origin (RADAM

156Brasil 1978) at 80ndash100 m above sea level and

157altitudinal differences of 40ndash50 m between plateaus

158and stream valleys (Gascon amp Bierregaard 2001)

159The first to third order streamlets (150000 scale)

160we studied belong to catchments of three different

161rivers (Urubu Cuieiras and Preto da Eva Rivers) with

162similar general characteristics such as geomorphology

163and distance from the Negro and Amazonas Rivers

164The streams have black acidic waters (pH 45ndash54)

165with low conductivity (79ndash167 lS cm-1) and the

166mean water temperature is of approximately 24C

167(Mortati 2004) Their streambeds are typically com-

168prised of sand patches and litter

169Forest streams under natural conditions (primary

170forest areas) are highly shaded due to the reduced

171canopy openness and they present distinct channel

172morphology depending on the terrain characteristics

173In wide flat-bottomed stream valleys (lsquolsquobaixiosrsquorsquo)

174there is more connectivity between the shallow river-

175bed and the marginal ponds or flooded adjacent areas

176Banks when formed are firmly sustained by roots and

177vegetation with cuttings only under roots and espe-

178cially in narrow-angled meanders The height of banks

179depends on the terrain declivity stream valley width

180and stream size Banks are sometimes incipient in

181small order streams There is a large amount of leaf and

182wood detritus deposited in pools as well as in

183meanders and logs and twigs constitute the main

184structures that retain this material These obstacles to

185the stream flow produce pools and small riffles that

186increase the habitat heterogeneity in the system In

187streams that cross open areas such as pastures light

188incidence is very high the streams get progressively

189shaded in old secondary forests where herbaceous

190plants grow on the margins or on the streambed

191Although qualitative changes are expected in the

192allochthonous matter deposited in the streams there

193are no differences in litter availability in secondaryFig 1 Sampling sites in the BDFFP areas Amazonas state

Brazil

Hydrobiologia

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

Hydrobiologia

123

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

Hydrobiologia

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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194 forest streams when compared to primary forests

195 (Webster et al 1990Mortati 2004) In pastures there

196 are no retention mechanisms (such as logs and twigs)

197 on the streambeds which are frequently silted and the

198 water spreads widely over the ground Stands of the

199 riparian vegetation (herbs shrubs and trees) may

200 wither away and eventually die Furthermore the

201 marginal banks are fragile and may crumble as they

202 are protected only by grass

203 Methods

204 Experimental design

205 We established 150 m reaches of 20 streams for

206 sampling (Fig 1) The sampling design intended to

207 produce independent samples and to maximize their

208 distribution into the BDFFP area We had two sample

209 reaches of one stream but these had different landscape

210conditions (a stream that drains a 10-ha forest fragment

211and subsequently crosses a secondary forest area)

212Catchments exhibited a range in land use from

213primary forest to secondary forest or pasture (Table 1)

214The secondary forest landscape included three vege-

215tation types (1) areas dominated by Vismia Vand

2161788 (Clusiaceae) (2) areas dominated by Cecropia

217Loefl 1758 (Cecropiaceae) or (3) mixed conditions

218where both species were present These landscape

219differences result from the managing practices

220employed during the process of forest fragment

221isolation Vismia secondary forests grew where the

222original vegetation was burned whereas Cecropia

223dominated where the original vegetation was only

224logged (Williamson et al 1998) According to More-

225ira (2002) the ages of secondary forests in the study

226area varied between 2 and 20 years old and were

227distributed in three age classes (Table 1) Older

228secondary forests may grow up to 25 m tall but with

229lower tree density than in primary forests The studied

Table 1 Sampling sites of aquatic insects in the areas of the Biological Dynamics of Forest Fragments Project (BDFFP-INPA)

Site Predominant cover Basin Latitude Longitude PFC150 Order Curr

(cm s-1)

Disch

(m3 min-1)

Depth

(cm)

Width

(cm)

1 Vismia secondary forest Preto da Eva River 022425 595351 061 1 301 79 155 1883

2 Primary forest Preto da Eva River 022444 595447 097 1 132 25 185 1733

3 Primary forest Preto da Eva River 022408 595415 100 2 336 73 198 2100

4 Cecropia secondary

forest

Urubu River 022355 595242 036 1 40 32 148 1750

5 Primary forest Urubu River 022366 595134 079 1 64 04 114 1250

6 10 ha Forest fragment Urubu River 022435 595203 027 1 114 03 48 883

7 Primary forest Urubu River 022603 595103 099 1 139 08 80 883

8 Primary forest Urubu River 022624 594642 100 1 290 07 47 983

9 Primary forest Urubu River 022643 594648 099 1 183 19 98 1767

10 Primary forest Urubu River 022698 594629 100 1 150 14 92 1667

11 Pasture Urubu River 022111 595905 028 1 286 26 176 1283

12 Pasture Urubu River 022155 595922 027 1 185 18 182 850

13 100 ha Forest fragment Urubu River 022174 595827 095 1 279 30 122 2267

14 Primary forest Urubu River 022605 595427 076 3 374 648 467 5833

15 Vismia secondary forest Cuieiras River 021967 600466 045 3 466 449 422 4367

16 10 ha Forest fragment Cuieiras River 022018 600679 084 1 118 11 105 1383

17 Mixed secondary forest Cuieiras River 022036 600681 025 1 18 04 173 1110

18 Primary forest Cuieiras River 022100 600582 080 2 236 128 285 3033

19 Mixed secondary forest Cuieiras River 022098 600549 036 1 61 09 114 2033

20 100 ha Forest fragment Cuieiras River 022075 600556 084 1 188 10 76 1500

21 Pasture Urubu River 022501 595215 0 1 ndash ndash 80 960

PFC150 percentage of forest cover in a 150-m width linear buffer zone Curr current in cm s-1 Disch discharge in m3 min-1

Hydrobiologia

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

Hydrobiologia

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

Hydrobiologia

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

Hydrobiologia

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

Hydrobiologia

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

Hydrobiologia

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

123

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

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tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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230 areas in primary forest landscapes have different sizes

231 (10 100 ha and continuous forest areas) The maxi-

232 mum distance between the sampling sites and the

233 nearby continuous forest was of 1 km (Fig 1)

234 Landscape analysis

235 We used Landsat TM 5 (Thematic Mapper) images

236 (path 232 row 62mdash2001) RGB bands 3 (063ndash

237 069 lm) 4 (076ndash090 lm) 5 (155ndash175 lm) with

238 30 m resolution for the landscape analysis in this

239 study We produced a classified image by using

240 Maximum Likelihood algorithm in supervised clas-

241 sification with ERDAS 87 Software We generated a

242 linear buffer zone 150 m wide around each stream

243 stretch by using ArcView 32 For each buffer zone

244 we calculated the proportion of the area covered by

245 primary forest secondary forest pasture and exposed

246 soil Some studies researched the classification of

247 these categories and mapped the extent and temporal

248 dynamics of secondary vegetation at local level in

249 Amazon (eg Adams et al 1995 Alves amp Skole

250 1996 Steininger 1996)

251 Stream habitat evaluation

252 We measured 12 habitat characteristics (items) to

253 describe the environmental conditions in the studied

254 reaches based on the protocols of Petersen (1992) for

255 visual assessment relative to land use riparian zone

256 streambed characteristics and stream channel mor-

257 phology to produce a habitat integrity index (HII)

258 Each item was composed of four to six alternatives

259 ordered in relation to perceived aspects of habitat

260 integrity To assure that each item had the same

261 weight in the analysis the observed values (ao) were

262 standardized in relation to the maximum value for

263 each item (am Eq 1) The final index is the mean

264 value for the total sampled habitat characteristics (n

265 Eq 2) These transformations produce an index that

266 vary between 0 and 1 and that is directly related to

267 the integrity of habitat conditions (Table 2)

pi frac14ao

ameth1THORN

269269

HII frac14

Pn

ifrac141

Pi

neth2THORN

271271

272Biological variables

273We took one subsample of macroinvertebrates com-

274posed by three sweeps from each of the four main

275substrates randomly spread in a stretch of 50 m at

276each stream with an aquatic sweep net of 1 mm mesh

277size and 30 cm of diameter The substrates sampled

278were leaf litter in pool and backwater areas leaf litter

279in riffle areas sand patches and rootvegetation at

280stream banks Subsamples from the two sampling

281events (March and October 2001) were combined

282into one sample from each stream The total sampled

283area for each stream was 17 m2 The samples were

284washed and preliminarily sorted in the field and fixed

285at 80 ethanol Later we sorted the specimens into

286morphospecies and identified them up to species or

287higher taxonomic level using identification keys (eg

288Belle 1992 Angrisano 1995 Merritt amp Cummins

2891996 Wiggins 1996 Nieser amp Melo 1997 Carvalho

290amp Calil 2000 Da Silva et al 2003 Olifiers et al

2912004 Manzo 2005 Pes et al 2005) and aid of

292specialists All insects in the samples were enumer-

293ated and identified Three different metrics were used

294to represent aquatic insect communities taxonomic

295richness Ephemeroptera Plecoptera and Trichoptera

296(EPT) richness and aquatic insect taxonomic com-

297position (Benstead et al 2003 Benstead amp Pringle

2982004) These metrics are considered sensitive to

299habitat changes in the aquatic environment (Barbour

300et al 1996 Silveira et al 2005)

301Data analysis

302We performed Spearmanrsquos rank correlation tests

303among the HII values and the values of each

304measured protocol variable and vegetation cover as

305well as with insect community metrics For calcula-

306tions taxa composition of each stream reach area

307(total or by substrate type) was represented by the

308coordinates of the first axis of a NMDS analysis

309based on species presencendashabsence data (one dimen-

310sion distance measure Euclidian distance) The

311percentage of variation of the Euclidian distance

312matrix captured by the first axis is expressed by r2

313The ANOSIM was used to compare and determine

314differences between stream communities of primary

315forest forest fragments secondary forest and pas-

316ture Taxa richness comparisons between streams

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

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UNCORRECTEDPROOF

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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Table 2 Habitat characteristics used in evaluation of sampling sites for HII calculations

Characteristic Condition Score

F1 Land use pattern beyond the

riparian zone

Primary continue forest100 ha fragment10 ha fragment 6

Cecropia secondary forestmixed secondary forest 5

Vismia secondary forest 4

Pasture 3

Perennial crops 2

Short-cycle cropsexposed soil 1

F2 Width of riparian forest Continuous forest 6

Forest width between 30 and 100 m 5

Forest width between 5 and 30 m 4

Forest width between 1 and 5 m 3

Riparian forest absent but some shrub species and pioneer trees 2

Riparian forest and shrub vegetation absent 1

F3 Completeness of riparian forest Riparian forest intact without breaks in vegetation 4

Breaks occurring at intervals of[50 m 3

Breaks frequent with gullies and scars at every 50 m 2

Deeply scarred with gullies all along its length 1

F4 Vegetation of riparian

zone within 10 m of channel

More than 90 plant density by non-pioneer trees or shrubs 4

Mixed pioneer species and mature trees 3

Mixed grasses and sparse pioneer trees and shrubs 2

Grasses and few tree shrubs 1

F5 Retention devices Channel with rocks andor old logs firmly set in place 4

Rocks andor logs present but backfilled with sediment 3

Retention devices loose moving with floods 2

Channel of loose sandy silt few channel obstructions 1

F6 Channel sediments Little or no channel enlargement resulting from sediment accumulation 4

Some gravel bars of coarse stones and little silt 3

Sediment bars of rocks sand and silt common 2

Channel divided into braids or stream channel corrected 1

F7 Bank structure Banks inconspicuous 5

Banks stable with rock and soil held firmly by grasses shrubs or tree roots 4

Banks firm but loosely held by grasses and shrubs 3

Banks of loose soil held by a sparse layer of grass and shrubs 2

Banks unstable easily disturbed with loose soil or sand 1

F8 Bank undercutting Little not evident or restricted to areas with tree root support 4

Cutting only on curves and at constrictions 3

Cutting frequent undercutting of banks and roots 2

Severe cutting along channel banks falling in 1

F9 Stream bottom Stone bottom of several sizes packed together interstices obvious 4

Stone bottom easily moved with little silt 3

Bottom of silt gravel and sand stable in some places 2

Uniform bottom of sand and silt loosely held together stony substrate absent 1

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

Hydrobiologia

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

Hydrobiologia

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

123

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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Article No 9441 h LE h TYPESET

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tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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318318 were made using a rarefaction method (Gotelli amp

319 Colwell 2001) We performed individual-based rar-

320 efactions because we only had one composite sample

321 per stream The ANOVA and the Tukey HSD test

322 were used to compare and to determine differences

323 between the resultant richness values of stream

324 communities of primary forest forest fragments

325 secondary forest and pasture

326 Due to the use of multiple Spearmanrsquos correlation

327 tests we performed the false discovery rate (FDR)

328 approach (Benjamini amp Hochberg 1995) to control

329 for type I errors (n = 223 a = 005) In our analysis

330 after application of FDR we accepted P-values

331 0006 For details and discussion on this method

332 see Garcıa (2004) The species indicator analysis

333 (Dufrene amp Legendre 1997) was used to relate taxa

334 and landscapes The statistical programs Past 14

335 (Hammer et al 2001) PC-ORD 4 (McCune amp

336 Mefford 1999) and STATISTICA 60 (StatSoft

337 2001) were used For all analyses taxa with only one

338 specimen were not considered We adopted a signif-

339 icance level of a = 005 in all tests

340 Results

341 The aquatic insect fauna

342 We observed 151 taxa distributed in 10 orders in the

343 samples which held 5746 individuals The numbers

344 of taxa recorded in each stream reach area ranged

34525ndash70 (Appendix in Electronic supplementary mate-

346rial) The average numbers of insect taxa in stream

347reaches of primary and secondary forests were very

348similar to one another (518 plusmn 95 and 500 plusmn 56

349respectively) and larger than in pasture areas

350(34 plusmn 102) EPT was represented by 68 taxa of

351which 5ndash35 were collected from each stream reach

352area Trichoptera was composed of 44 taxa Epheme-

353roptera of 20 and Plecoptera of only 4 The average

354numbers of EPT taxa in stream reaches of primary

355forest secondary forest and pasture were 282 plusmn 51

356262 plusmn 28 and 153 plusmn 100 respectively

357Regarding to secondary forests older forests

358([14 years old) showed higher number of taxa than

3592ndash7 year-old forests

360Comparing rarefaction-based species richness pas-

361ture streams showed lower values of insect taxa

362richness (F = 10217 P = 0000) and EPT richness

363(F = 6934 P = 0003) (Figs 2 and 3) than streams

364in primary forests forest fragments and secondary

365forests Regarding substrate diversity pasture streams

366showed lower insect taxa richness in sand (F = 7052

367P = 0003) and riffle litter (F = 16558 P = 0000)

368and lower values of EPT richness in riffle litter

369(F = 12080 P = 0000) and pool litter (F = 4770

370P = 00137) Banks showed no differences between

371vegetation covers

372The results of ANOSIM showed that pasture

373streams have different communities from primary

374and secondary forest streams but not of forest

375fragment streams (Table 3) The first axis of a NMDS

Table 2 continued

Characteristic Condition Score

F10 Riffles and pools or meanders Distinct occurring at intervals of 5ndash79 the stream width 4

Irregularly spaced 3

Long pools separating short riffles meanders absent 2

Meanders and rifflepools absent or stream corrected 1

F11 Aquatic vegetation When present consists of moss and patches of algae 4

Algae dominant in pools vascular plants along edge 3

Algal mats present some vascular plants few mosses 2

Algal mats cover bottom vascular plants dominate channel 1

F12 Detritus Mainly consisting of leaves and wood without sediment 5

Mainly consisting of leaves and wood with sediment 4

Few leaves and wood fine organic debris with sediment 3

No leaves or woody debris coarse and fine organic matter with sediment 2

Fine anaerobic sediment no coarse debris 1

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

Hydrobiologia

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

123

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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MS Code HYDR2696 h CP h DISK4 4

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of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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376(final stress = 3606 r2 = 046) also provided a

377contrast of pasture secondary forest and primary

378forest areas (Fig 4) Primary forest streams had more

379exclusive taxa (14 insect taxa 5 EPT) whereas

380secondary forest and pasture streams presented seven

381exclusive taxa each (1 EPT) Comparing primary

382forest plus secondary forest streams with secondary

383forest plus pasture streams more taxa were exclusive

384of the first group (46 insect taxa 24 EPT) than of the

385second one (6 3) Continuous forest fragment (both

386primary forests) and secondary forest streams

387showed very similar fauna but few species were

388considered characteristic of primary forest streams by

389the indicator species analysis and none for secondary

390forest streams (Fig 4 and Appendix in Electronic

391supplementary material)

392Forest cover relationships

393Stream reaches with larger percentages of canopy

394cover showedhigherHII values (r = 077P 0001)

395Among the 12measured habitat variables present inHII

396composition the following seven showed significant

397correlations with vegetation cover (1) land use pattern

398beyond the riparian zone (r = 0760 P 0001) (2)

399width of riparian forest (r = 0606 P = 0004) (3)

400completeness of riparian forest (r = 0596

401P = 0004) (4) vegetation of riparian zone within

40210 m of channel (r = 0762 P 0001) (5) retention

403devices (r = 0713 P 0001) (6) channel sediments

404(r = 0708 P 0001) and (7) aquatic vegetation

405(r = 0662 P 0001) These results indicate a closeFig 3 Median values of EPT taxa richness of streams under

different vegetation covertures

Table 3 Pairwise comparisons of habitats sampled

Primary

forest

Forest

fragment

Secondary

forest

Pasture

Primary

forest

01947 04113 00043

Forest

fragment

01947 03461 0054

Secondary

forest

04113 03461 00367

Results from ANOSIM based on Euclidian distances between

streams in primary forest forest fragments secondary forests

and pastures The omnibus test indicated significant difference

among habitats (number of permutations = 1000 r = 0282

P = 0015)

Fig 4 Median values of NMDS analysis first axis scores of

streams under different vegetation covertures based on taxa

presenceabsence

Fig 2 Median values of insect taxa richness of streams under

different vegetation covertures

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

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UNCORRECTEDPROOF

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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406 relationship between the physical structure of the

407 streams and the integrity of the riparian forest (Fig 5)

408 However there was no significant correlation between

409 forest cover and the faunalmetrics insect richness EPT

410 richness and insect taxa composition (except for the

411 stream bank fauna)

412 Habitat integrity and faunal metrics

413 There was not significant correlation between HII

414 values and aquatic insect metrics (total insect richness

415r = 0223 P = 0330 EPT richness r = 0252

416P = 0290 insect taxa composition r = -0552

417P = 0010) None of the habitat variables was signif-

418icantly correlated with total insect richness and only

419one aquatic vegetation was correlated with EPT

420richness (r = 0620 P = 0003) Six variables were

421significantly correlated to insect taxa composition (1)

422land use pattern beyond the riparian zone (r = 0590

423P = 0005) (2) width of riparian forest (r = 0800

424P 0001) (3) completeness of riparian forest

425(r = 0675 P = 0001) (4) retention devices

426(r = 0607 P = 0004) (5) channel sediments

427(r = 0702 P = 0001) and (6) aquatic vegetation

428(r = 0810 P 0001) The variables vegetation of

429riparian zone within 10 m of channel bank structure

430bank undercutting stream bottom and pool-riffle or

431meander distances and detritus showed no significant

432relationships with the faunal metrics These contrast-

433ing results ie the strong correlation between forest

434cover and streamhabitat integrity theweak correlation

435between forest cover and the faunal metrics and the

436significant relation between some stream habitat

437variables and faunal metrics point out an indirect

438relation between forest cover and faunal variables

439(Fig 6andashc)

440Some taxa were positively correlated with HII

441values Calcopteryx (Polythoridae) Anacroneuria

Fig 5 Relationship between percentage of primary forest in a

150-m linear buffer zone and HII values in 21 sampled sites

Fig 6 Relationship

between HII and faunal

metrics in 21 sampled sites

(andashc) all substrates (d) taxa

composition in marginal

banks

Hydrobiologia

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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442 (Perlidae) Protosialis (Sialidae)GyrelmisHeterelmis

443 Macrelmis (Elmidae)Helicopsyche (Helicopsychidae)

444 andMacrostemum (Hydropsychidae) ConverselyCal-

445 libaetis (Baetidae) Belostoma (Belostomatidae)

446 Tenagobia (Corixidae) Smicridea (Hydropsychidae)

447 and Pyralidae were negatively correlated and occurred

448 only in deforested areas

449 Separate analyses by substrate type showed sig-

450 nificant correlation only for insects dwelling in banks

451 and HII concerning taxa composition (r = -0591

452 P = 0005) (Fig 6d) Regarding habitat variables

453 taxa composition showed higher values and more

454 significant correlations than others (1) land use

455 pattern beyond the riparian zone (r = -0614

456 P = 0003 for banks) (2) width of riparian forest

457 (r = -0612 P 0003 for riffle litter) (3) com-

458 pleteness of riparian forest (r = 0628 P = 0002 for

459 sand substrate) (4) vegetation of riparian zone within

460 10 m of channel (r = -0594 P = 0005 for banks)

461 (5) retention devices (r = -0690 P = 0001 for

462 banks) and (6) aquatic vegetation (r = 0685

463 P 0001 for riffle litter) The latter variable was

464 the only significantly correlated with other metrics

465 (r = 0678 P = 0001 for insect richness in pool

466 litter and r = 0580 P = 0006 for insect richness in

467 riffle litter)

468 Discussion

469 Forest cover condition

470 Stream habitat attributes closely matched the vege-

471 tation cover condition as initially expected

472 However there were no significant relationships

473 between modifications in the vegetation cover and

474 variables such as bank structure and undercutting

475 stream substrate and pool-riffle or meander distances

476 Some of these variables are also dependent on the

477 stream size slope and geological features of the

478 sampling areas (Gordon et al 1992 Wood amp

479 Armitage 1997 Church 2002) Moreover depend-

480 ing on vegetation cover type or land use (eg cattle

481 raising different agricultural practices forestry)

482 changes in vegetation cover may lead to few or

483 discrete physical changes in the streams (Allan et al

484 1997 Harding et al 1999 Price amp Leigh 2006) The

485 environmental quality and proportion of the second-

486 ary forests at the buffer zone around the sampled

487stream reaches may have influenced our results The

488streams included in the present study are character-

489istic of the region they have no rocky substrates and

490streambeds are mainly constituted of sand and

491detritus (Zuanon amp Sazima 2004 Mendonca et al

4922005) In addition few differences were indicated in

493HII between substrates with different proportions of

494clay and silt although significant increase was

495expected in sediment input (Allan et al 1997 Wood

496amp Armitage 1997 Nerbonne amp Vondracek 2001

497Church 2002) Sediment inputs induce siltation

498within and on the surface of the substrate leading

499to habitat modifications disturbance of trophic

500resources and in feeding mechanisms of stream fauna

501(Fossati et al 2001 Mol amp Ouboter 2004) In the

502present study only two pasture streams showed this

503condition

504There was no significant relationship between the

505amount of detritus (litter) and vegetation cover

506Davies et al (2005) compared streams with different

507logging intensity in Tasmania and showed a

508decrease in the input and retention of organic

509matter De Long amp Brusven (1994) suggested that

510lower rates of allochthonous input may result in a

511system with detrital dynamics which bears macro-

512invertebrate communities different from those found

513in comparable undisturbed streams According to

514Webster et al (1990) differences in litter input

515between disturbed and undisturbed catchments seem

516to be due to the replacement of the original mature

517vegetation by successional species Successional

518forests which consist of herbaceous species and

519small shrubs typically contribute less litter to a

520stream than mature forests Furthermore the absence

521of large limbs and fallen trees reduces stream

522capacity to retain and process organic material after

523entering the system (Webster et al 1990) Besides

524sedimentation may diminish the availability of

525detritus by burying leaf packs (Fossati et al 2001

526Mol amp Ouboter 2004) Nevertheless Mortati (2004)

527did not find significant differences in detritus

528availability in the streams included in the present

529study although a reduction of detritus was expected

530in open areas The 20-year-old second growth forest

531that comprises the riparian vegetation along one of

532these streams reaches up to 20 m tall and shows a

533complex highly structured environment that may

534have contributed to restore and maintain a detritus

535source and processing

Hydrobiologia

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

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tho

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

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857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

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536 Faunal metrics

537 Despite the fact that streamlets in primary forest areas

538 had largest number of insect taxa there was no

539 significant relationship between the proportion of

540 primary forest around the stream reaches and insect

541 richness Fidelis da Silva (2006) showed that the

542 decrease of macroinvertebrate taxa richness in these

543 same streams was related to proportion of pasture and

544 gaps in canopy cover Roque amp Trivinho-Strixino

545 (2000) and Roque et al (2003) compared forested

546 and non-forested sites in Atlantic Forest streams in

547 the State of Sao Paulo Brazil and found highest

548 values for taxonomic richness in stream sections with

549 intact riparian zones On the other hand in a study of

550 Atlantic Forest streams in the State of Rio de Janeiro

551 Brazil Egler (2002) found significant differences

552 between species richness from forested and cultivated

553 areas but not between forested and deforested or

554 second growth areas

555 Although pasture streams showed significantly

556 lower values of richness none of the habitat

557 variables nor HII or percentage of forest cover were

558 good predictors to taxa richness We expected some

559 significant relationships because the width and

560 structure of riparian forest and aquatic vegetation

561 are associated features related to environmental

562 integrity in regard to canopy openness and light

563 (Petersen 1992) As mentioned above the lack of

564 riparian forest directly contributes to an increase of

565 sediment and decrease of organic matter inputs

566 influencing the structure of the macroinvertebrate

567 community by reducing habitat availability or by

568 making the habitat unsuitable for survival (Vannote

569 et al 1980 Sizer 1992 Wood amp Armitage 1997

570 Nerbone amp Vondraek 2001 Zweig amp Rabeni 2001

571 Sparovek et al 2002 Davies et al 2005) However

572 relationships between aquatic insect and EPT rich-

573 ness with sediment and organic matter inputs were

574 not significant in this study These two insect metrics

575 only showed differences across sites in relation to HII

576 values and percentage of vegetation cover Aquatic

577 insect assemblages were strongly altered and impov-

578 erished in highly disturbed sites with very low

579 vegetation cover

580 The results related to the taxonomic composition

581 suggest a replacement of faunal components associ-

582 ated with characteristics of the physical environment

583 and habitat integrity but a single marginally

584significant relationship was obtained with HII Some

585factors seem to have stronger impact on the taxo-

586nomic changes in aquatic insect communities such as

587riparian forest width and structure canopy openness

588retention devices aquatic vegetation and sediments

589Retention devices are important in keeping high

590habitat heterogeneity (Barbour et al 1999) Many

591insect taxa were absent or represented by smaller

592numbers of individuals under low HII values while

593others such as grazers and algal piercers showed

594increasing number of individuals under the same

595conditions These results corroborate several studies

596that pointed out changes in insect taxonomic compo-

597sition and function related to deforestation (eg De

598Long amp Brusven 1994 Barbour et al 1996 Naiman

599amp Decamps 1997 Benstead et al 2003 Benstead amp

600Pringle 2004 Silveira et al 2005)

601The absence of significant correlations between

602habitat variables HII scores and insect taxonomic

603composition in pool and riffle litter (except width of

604riparian forest and aquatic vegetation for riffle litter)

605suggests that assemblage composition is more homo-

606geneous in these substrates Alternatively these

607assemblages may only be affected in terms of

608composition by impacts that drastically modify the

609habitat However there were significant relationships

610between habitat integrity and insect composition in

611banks Similar results were found by Roy et al

612(2003) and were attributed to banks acting as refuges

613for the fauna from other substrates in streams under

614perturbation

615We observed conspicuous gaps in the distribution

616of values for biological variables and HII values

617between areas with no forest and those presenting

618small percentages of vegetation cover Even limited

619riparian vegetation seems to maintain the distinctive

620habitat characteristics in streams that are essential for

621the occurrence of many insect taxa

622Our results highlight the importance of riparian

623vegetation in the maintenance of the biota of streams

624and its function as ecological corridors (eg Naiman

625amp Decamps 1997 De Lima amp Gascon 1999

626Anbumozhi et al 2005 Nakamura amp Yamada

6272005) Four factors related to the present study are

628worth further discussion the importance of secondary

629forests the extensive area of primary forest around

630the pasture matrix of the study area that some

631streamlets cross areas with different vegetation and

632the potential capacity of dispersion in aquatic insects

Hydrobiologia

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UNCORRECTEDPROOF

633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

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of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

633 Both forest fragments and secondary forests pres-

634 ent faunal characteristics similar to the continuous

635 forest areas in most cases Older secondary forests

636 behave like forests with reduced light incidence

637 recovery of retention devices on streambeds and less

638 loss of sediments to streamlets Younger secondary

639 forests are more open dominated by shrubs herbs

640 and grasses They present higher light incidence and

641 less capacity to keep sediments from entering the

642 streamlets Abandoned pasture areas may present

643 variable growth of secondary forests In some

644 settings a field abandoned for 2 years can have 4-

645 m-tall woody vegetation while elsewhere a similarly

646 aged plot could still be dominated by grasses and

647 sedges (Walker et al 1999) Thus the habitat matrix

648 presents great heterogeneity and the secondary forest

649 may play an important role in the connectivity among

650 the forest areas which facilitates the dispersal of

651 aquatic insects as well as the colonization of streams

652 Depending on the area or their position streams in

653 forest fragments suffer more or less edge effects

654 which allows the entrance of habitat matrix speci-

655 mens On the other hand parts of streams that cross

656 pasture areas or secondary forests are influenced by

657 upstream forests The isolation of forest fragments

658 depends on the distance of continuous forest areas or

659 other fragments and on the connectivity with sec-

660 ondary forests In the study area the distance from

661 fragments and secondary forests to primary forest

662 areas is not longer than 1 km Another feature to be

663 taken into account is the occurrence of at least one

664 narrow riparian corridor in most studied streamlets

665 The dispersal of aquatic insects may occur longi-

666 tudinally (in the same stream) or transversally among

667 watersheds (Malmqvist 2002 Elliott 2003 Macne-

668 ale et al 2005) Some species have great capacity of

669 dispersal (further than 1 km) as in Ephemeroptera

670 Odonata and Coleoptera (Bilton et al 2001 Peter-

671 sen et al 2004 Macneale et al 2005) Since one of

672 the main determinant colonization factors is adequate

673 substrate (Sanderson et al 2005) its occurrence

674 even in small proportion may favor the presence of a

675 determined taxon Distance is an important factor for

676 dispersal Petersen et al (2004) during studies in the

677 UK observed that the number of Plecoptera

678 Ephemeroptera and Trichoptera adults captured

679 decreases as the distance of the river increases

680 However they did not find differences between

681 forests and deforestation areas in relation to the

682occurrence of dispersal Sanderson et al (2005)

683compared macroinvertebrate assemblages at 188

684running-water sites in the catchment of the River

685Rede northeast England and concluded that there

686was a significant influence of the species composition

687of neighboring sites on determining local species

688assemblages

689These previously published results support our

690findings in some way They might explain that the

691low faunal metrics response in relation to changes in

692vegetation cover and to fragmentation can be due to

693the effect of the heterogeneous matrix as well as the

694presence and proximity of a wide area of surrounding

695forests in the present study

696Acknowledgments This work was supported by the BDFFP697FAPEAMPIPT 2004ndash2006 Fundacao OBoticario de Protecao a698Natureza (0630_20041) and CNPq (Edital Universal 2005ndash6992007) We thank Ocırio Pereira and Jose Ribamar Marques de700Oliveira for invaluable field assistance Ana Maria Oliveira Pes701(DCEN INPA) Alcimar do Lago Carvalho (Museu Nacional702UFRJ) Elidiomar Ribeiro da Silva (UNI-RIO) and Maria Ines703da Silva dos Passos (IB UFRJ) helped with the identification of704the insects Darcılio Fernandes Baptista (FIOCRUZ) provided705valuable comments and suggestions on this manuscript Daniela706Maeda Takiya (UFPR) revised the final English text The707manuscript was greatly improved by comments and critical708reviews fromDr Joel Trexler and two anonymous referees This709is contribution number 00 of Projeto Igarapes and contribution710number 000 of the BDFFP Technical Series

711References

712Adams J B D E Sabol V Kapos D A Roberts M O713Smith amp A R Gillespie 1995 Classification of multi-714spectral images based on fractions of end members715Application to land-cover change in the Brazilian Ama-716zon Remote Sensing of Environment 52 137ndash154717Allan D D L Erickson amp J Fay 1997 The influence of718catchment land use on stream integrity across multiple719spatial scales Freshwater Biology 37 149ndash161720Alves S D amp D Skole 1996 Characterizing land cover721dynamics using multi-temporal imagery International722Journal of Remote Sensing 17 835ndash839723Anbumozhi V J Radhakrishnan amp E Yamaji 2005 Impact724of riparian buffer zones on water quality and associated725management considerations Ecological Engineering 24726517ndash523727Angrisano E B 1995 Insecta Trichoptera In Lopretto E C728amp G Tell (eds) Ecosistemas de Aguas Continentales Ed729SUR La Plata 1199ndash1238730Barbour M T J Gerritsen G E Griffith R Frydenborg731E McCarron J S White amp M L Bastian 1996 A732framework for biological criteria for Florida streams733using benthic macroinvertebrates Journal of the North734American Benthological Society 15 185ndash211

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

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979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

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735 Barbour M T J Gerritsen B D Snyder amp J B Stribling736 1999 Rapid Bioassessment Protocols for Use in Streams737 amp Wadeable Rivers Periphyton Benthic Macroinverte-738 brates amp Fish 2nd edn United States Environmental739 Protection Agency EPA 841-B-99-002 Washington DC740 Belle J 1992 Studies on ultimate instar larvae of Neotropical741 Gomphidae with description of Tibiagomphus gen nov742 (Anisoptera) Odonatologica 2 1ndash25743 Benstead J P amp C M Pringle 2004 Deforestation alters the744 resource base and biomass of endemic stream insects in745 eastern Madagascar Freshwater Biology 49 490ndash501746 Benstead J P M M Douglas amp C M Pringle 2003 Rela-747 tionships of stream invertebrate communities to748 deforestation in eastern Madagascar Ecological Appli-749 cations 13 1473ndash1490750 Benjamini Y amp Y Hochberg 1995 Controlling the false751 discovery rate A practical and powerful approach to752 multiple testing Journal of the Royal Statistical Society B753 57 289ndash300754 Bierregaard R O Jr C Gascon T E Lovejoy amp755 R Mesquita (eds) 2001 Lessons from Amazonia The756 Ecology and Conservation of a Fragmented Forest Yale757 University Press New Haven758 Bilton D T J R Freeland amp B Okamura 2001 Dispersal in759 freshwater invertebrates Annual Review of Ecology and760 Systematics 32 159ndash181761 Carvalho A L amp E R Calil 2000 Chaves de identificacao762 para as famılias de Odonata (Insecta) ocorrentes no Brasil763 adultos e larvas Papeis Avulsos de Zoologia 41 223ndash241764 Church M 2002 Geomorphic thresholds in riverine land-765 scapes Freshwater Biology 47 541ndash557766 Gotelli N amp R K Colwell 2001 Quantifying biodiversity767 Procedures and pitfalls in the measurement and compar-768 ison of species richness Ecology Letters 4 379ndash391769 Da Silva E R F F Salles J L Nessimian amp L B N Coelho770 2003 A identificacao das famılias de Ephemeroptera771 (Insecta) ocorrentes no Estado do Rio de Janeiro Chave772 pictoricapara as ninfasBoletimdoMuseuNacional 508 1ndash6773 Davidson E A C Neill A V Krusche V V R Ballester774 D Markewitz amp R O Figueiredo 2004 Loss of nutri-775 ents from terrestrial ecosystems to streams and the776 atmosphere following land use change in Amazonia In777 Ecosystems and Land Use Change Geophysical Mono-778 graph Series 147ndash158779 Davies P E L S J Cook P D McIntosh amp S A Munks780 2005 Changes in stream biota along a gradient of logging781 disturbance 15 years after logging at Ben Nevis Tas-782 mania Forest Ecology and Management 219 132ndash148783 De Long M D amp M A Brusven 1994 Allochthonous imput784 of organic matter from different riparian habitats of an785 agriculturally impacted stream Environmental Manage-786 ment 18 59ndash71787 Egler M 2002 Utilizando a comunidade de macroinverte-788 brados bentonicos na avaliacao da degradacao de789 ecossistemas de rios em areas agrıcolas Masterrsquos Thesis790 ENSP FIOCRUZ Rio de Janeiro791 Elliott J M 2003 A comparative study of the dispersal of 10792 species of stream invertebrates Freshwater Biology 48793 1652ndash1668794 Fossati O J G Wasson C Hery R Marin amp G Salinas795 2001 Impact of sediment releases on water chemistry and

796macroinvertebrate communities in clear water Andean797streams (Bolivia) Archiv fur Hydrobiologie 151 33ndash50798De Lima M G amp C Gascon 1999 The conservation value of799linear forest remnants in Central Amazonia Biological800Conservation 91 241ndash247801Dufrene M amp P Legendre 1997 Species assemblages and802indicator species The need for a flexible asymmetrical803approach Ecological Monographs 67 345ndash366804Fidelis da Silva L 2006 Estrutura da comunidade de insetos805aquaticos em igarapes na Amazonia Central com difer-806entes graus de preservacao da cobertura vegetal e807apresentacao de chave de identificacao para generos de808larvas da ordem Odonata Masterrsquos Thesis UFAMINPA809Manaus810Garcıa L V 2004 Escaping the Bonferroni iron claw in811ecological studies Oikos 105 657ndash663812Gascon C amp R O Bierregaard Jr 2001 The Biological813dynamics of forest fragments projectmdashThe study site814experimental design and research activity In Bierregaard815R O Jr C Gascon T E Lovejoy amp R Mesquita (eds)816Lessons from Amazonia The Ecology and Conservation817of a Fragmented Forest Yale University Press New818Haven 31ndash45819Gascon C W F Lawrence amp T E Lovejoy 2001 Frag-820mentacao florestal e biodiversidade na Amazonia Central821In Garay I amp B F S Dias (orgs) Conservacao da bio-822diversidade em ecossistemas tropicais avancos823conceituais e revisao de novas metotologias de avaliacao e824monitoramento Editora Vozes Petropolis 112ndash127825Gordon N D T A McMahon amp B L Finlayson 1992826Stream hydrology An introduction for ecologists John827Wiley amp Sons Chichester828Hammer O D A T Harper amp P D Ryan 2001 Past829Paleontological statistic software for education and data830analysis Paleontologia Eletronica 4(1) 9 pp831Harding J S R G Young J W Hayes K A Shearer amp J D832Stark 1999 Changes in agricultural intensity and river833health along a river continuum Freshwater Biology 42834345ndash357835Lovejoy T E R O Bierregaard J M Rankin amp H O R836Schubart 1983 Ecological dynamics of tropical forest837fragments In Sutton S L T C Whitmore amp A C Chad-838wick (eds) Tropical Rain Forest Ecology and Management839Blackwell Scientific Publication Oxford 377ndash384840Macneale K H B L Peckarsky amp G E Likens 2005 Stable841isotopes identify dispersal patterns of stonefly populations842living along stream corridors Freshwater Biology 508431117ndash1130844Malmqvist B 2002 Aquatic invertebrates in riverine land-845scapes Freshwater Biology 47 679ndash694846Manzo V 2005 Key to the South America of Elmidae847(Insecta Coleoptera) with distributional data Studies of848Neotropical Fauna and Environment 40 201ndash208849McClain M E amp H Elsenbeer 2001 Terrestrial inputs to850Amazon streams and internal biogeochemical processing851In McClain M E E Victoria amp J Rishey (eds) The852Biogeochemistry of the Amazon Basin Oxford University853Press Oxford 185ndash207854McCune B amp M J Mefford 1999 Multivariate Analysis of855Ecological Data Version 414 MjM Software Gleneden856Beach Oregon

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

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UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

857 Melo E G F M S R Silva amp S A F Miranda 2005858 Influencia antropica sobre aguas de igarapes na cidade de859 Manaus ndash Amazonas Caminhos de Geografia 5 40ndash47860 Mendonca F P W E Magnusson amp J Zuanon 2005861 Relationships between habitat characteristics and fish862 assemblages in small streams of Central Amazonia Co-863 peia 4 750ndash763864 Merritt R W amp K W Cummins (eds) 1996 An Introduction865 to the Aquatic Insects of North America KendallHunt866 Publishing Company Dubuque867 Mol J H amp P E Ouboter 2004 Downstream effects of868 erosion from small-scale gold mining on the instream869 habitat and fish community of a small neotropical rain-870 forest stream Conservation Biology 18 201ndash214871 Moreira M P 2002 O uso de sensoriamento remoto para872 avaliar a dinamica de sucessao secundaria na Amazonia873 Central Masterrsquos Thesis INPAUFAM Manaus874 Mortati A F 2004 Colonizacao por peixes no folhico875 submerso implicacoes das mudancas na cobertura flor-876 estal sobre a dinamica da ictiofauna de igarapes na877 Amazonia Central Masterrsquos Thesis INPAUFAM878 Manaus879 Naiman R J amp H Decamps 1997 The ecology of interfaces880 Riparian zones Annual Review of Ecology and System-881 atics 28 621ndash658882 Nakamura F amp H Yamada 2005 Effects of pasture devel-883 opment on the ecological functions of riparian forests in884 Hokkaido in northern Japan Ecological Engineering 24885 539ndash550886 Nerbone B A amp B Vondracek 2001 Effects of local land use887 on physical habitat benthic macoinvertebrates and fish in888 the Whitewater River Minesota USA Environmental889 Management 28 87ndash99890 Nieser N amp A L Melo 1997 Os heteropteros aquaticos de891 Minas Gerais Editora UFMG Belo Horizonte892 Olifiers M H L F M Dorville J L Nessimian amp893 N Hamada 2004 A key to Brazilian genera of Ple-894 coptera (Insecta) based on nymphs Zootaxa 651 1ndash15895 Oliveira A A amp S Mori 1999 A central Amazonian terra896 firme forest I High tree species richness on poor soils897 Biodiversity and Conservation 8 1219ndash1244898 Pes A M O N Hamada amp J L Nessimian 2005 Chaves de899 identificacao de larvas para famılias e generos de Tri-900 choptera (Insecta) da Amazonia Central Brasil Revista901 Brasileira de Entomologia 49 181ndash204902 Petersen R C Jr 1992 The RCE A riparian channel and903 environmental inventory for small streams in agricultural904 landscape Freshwater Biology 27 295ndash306905 Petersen I Z Masters A G Hildrew amp S J Ormerod 2004906 Dispersal of adult aquatic insects in catchments of dif-907 fering land use Journal of Applied Ecology 41 934ndash950908 Price P amp D S Leigh 2006 Morphological and sedimento-909 logical responses of streams to human impact in the910 southern Blue Ridge Mountains USA Geomorphology911 78 142ndash160912 Pozo J E Gonzalez J R Dıez J Molinero amp A Elosegui913 1997 Inputs of particulate organic matter to streams with914 different riparian vegetation Journal of the North Amer-915 ican Benthological Society 16 602ndash611916 Resh V H amp J K Jackson 1993 Rapid assessment917 approaches to biomonitoring using macroinvertebrates In

918Rosemberg D M amp V H Resh (eds) Freshwater Bio-919monitoring and Benthic macroinvertebrates Chapman amp920Hall New York 195ndash233921Roque F O amp S Trivinho-Strixino 2000 Fragmentacao de922habitats nos corregos do Parque Estadual do Jaragua (SP)923Possıveis impactos na riqueza de macroinvertebrados e924consideracoes para conservacao in situ In Anais do II925Congresso Brasileiro de Unidades de Conservacao926Campo Grande 752ndash760927Roque F O S Trivinho-Strixino G Strixino RC Agostinho928amp J C Fogo 2003 Benthic macroinvertebrates in streams929of Jaragua State Park (Southeast of Brazil) considering930multiple spatial scale Journal of Insect Conservation 793163ndash72932Roy A H A D Rosemond DS Leigh M J Paul amp933B Wallace 2003 Habitat-specific responses of stream934insects to land cover disturbance Biological conse-935quences and monitoring implications Journal of North936American Benthological Society 22 292ndash307937Sanderson R A M D Eyre amp S P Rushton 2005 The938influence of stream invertebrate composition at neigh-939bouring sites on local assemblage composition940Freshwater Biology 50 221ndash231941Silveira M P D F Baptista D F Buss J L Nessimian amp942M Egler 2005 Application of biological measures for943stream integrity assessment in south-east Brazil Envi-944ronmental Monitoring and Assessment 101 117ndash128945Sizer N C 1992 The impact of edge formation on regener-946ation and litterfall in a tropical rain forest fragment in947Amazonia PhD Thesis University of Cambridge948Cambridge949Sparovek G S B L Ranieri A Gassner I C De Maria950E Schnug R F Santos amp A Joubert 2002 A conceptual951framework for definition of the optimal width of riparian952forests Agriculture Ecosystems and Environment 90953169ndash175954Smith N J H E A S Serrao P T Alvim amp I C Falesi9551995 Amazonia Resiliency and dynamism of the land956and its people United Nations University Press Tokyo957StatSoft Inc (2001) STATISTICA (data analysis software958system) version 6 wwwstatsoftcom959Steininger M K 1996 Tropical secondary forest regrowth in960the Amazon Age area and change estimation with961Thematic Mapper data International Journal of Remote962Sensing 17 9ndash27963Vannote R L G W Minshall KW Cummins J R Sedell amp964C Cushing 1980 The river continuum concept Canadian965Journal of Fisheries and Aquatic Sciences 37 130ndash137966Walker R W Salas G Urquhart M Keller D Skole amp M967Pedlowski (orgs) 1999 Secondary vegetation ecological968social and remote sensing issues Report on the work-969shop lsquolsquoMeasurement and Modeling of the Inter-Annual970Dynamics of Deforestation and Regrowth in the Brazilian971Amazonrsquorsquo Florida State University972Webster J R S W Golladay E F Benfield D J DrsquoAngelo amp973G T Peters 1990 Effects of forest disturbance on partic-974ulate organic matter budgets of small streams Journal of975North American Benthological Society 9 120ndash140976Wiggins G B 1996 Larvae of North American caddisfly977genera (Trichoptera) 2nd edn University of Toronto978Press Toronto

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

997

Hydrobiologia

123

Journal Medium 10750 Dispatch 3-6-2008 Pages 15

Article No 9441 h LE h TYPESET

MS Code HYDR2696 h CP h DISK4 4

Au

tho

r P

ro

of

UNCORRECTEDPROOF

UNCORRECTEDPROOF

979 Williamson G B R C G Mesquita K Ickes amp G Ganade980 1998 Estrategias de arvores pioneiras nos Neotropicos In981 Gascon C amp P Moutinho (eds) Floresta Amazonica982 Dinamica Regeneracao e Manejo Instituto Nacional de983 Pesquisas da Amazonia (INPA) Manaus Amazonas984 Brazil 131ndash144985 Wood P J amp P D Armitage 1997 Biological effects of fine986 sediment in the lotic environment Environmental Man-987 agement 21 203ndash207

988Zuanon J amp I Sazima 2004 Natural history of Staurogl-

989anis gouldingi (Siluriformes Trichomycteridae) a990miniature sand-dwelling candiru from central Amazonia991streamlets Ichthyological Exploration of Freshwaters99215 201ndash208993Zweig L D amp C H Rabeni 2001 Biomonitoring for994deposited sediment using benthic invertebrates A test on9954 Missouri streams Journal of the North American Ben-996thological Society 20 643ndash657

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Hydrobiologia

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