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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
Author Family Name ZuanonParticle
Given Name JansenSuffix
Division CPBA
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name GordoParticle
Given Name MarceloSuffix
Division Instituto de Ciecircncias Bioloacutegicas
Organization Universidade Federal do Amazonas (UFAM)
Address Manaus AM Brazil
Author Family Name FidelisParticle
Given Name LuanaSuffix
Division DCEN
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name JuenParticle
Given Name LeandroSuffix
Division Departamento de Biologia Geral
Organization Universidade Federal de Viccedilosa (UFV)
Address 36571-000 Vicosa MG Brazil
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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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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Journal HYDR-10750 Article 9441
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Please provide details for lsquoboldrsquo emphasis present in Table 3
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
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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
Hydrobiologia
123
Journal Medium 10750 Dispatch 3-6-2008 Pages 15
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UNCORRECTEDPROOF
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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of
UNCORRECTEDPROOF
UNCORRECTEDPROOF
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
Journal Medium 10750 Dispatch 3-6-2008 Pages 15
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UNCORRECTEDPROOF
UNCORRECTEDPROOF
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
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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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
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
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
Author Family Name ZuanonParticle
Given Name JansenSuffix
Division CPBA
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name GordoParticle
Given Name MarceloSuffix
Division Instituto de Ciecircncias Bioloacutegicas
Organization Universidade Federal do Amazonas (UFAM)
Address Manaus AM Brazil
Author Family Name FidelisParticle
Given Name LuanaSuffix
Division DCEN
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name JuenParticle
Given Name LeandroSuffix
Division Departamento de Biologia Geral
Organization Universidade Federal de Viccedilosa (UFV)
Address 36571-000 Vicosa MG Brazil
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
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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
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Please provide citation for Reference Resh and Jackson (1993)
Journal HYDR-10750 Article 9441
Please provide accessed date for Reference StatSoft Inc (2001)
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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
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
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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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Article No 9441 h LE h TYPESET
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of
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
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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
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of
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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
Author Family Name ZuanonParticle
Given Name JansenSuffix
Division CPBA
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name GordoParticle
Given Name MarceloSuffix
Division Instituto de Ciecircncias Bioloacutegicas
Organization Universidade Federal do Amazonas (UFAM)
Address Manaus AM Brazil
Author Family Name FidelisParticle
Given Name LuanaSuffix
Division DCEN
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name JuenParticle
Given Name LeandroSuffix
Division Departamento de Biologia Geral
Organization Universidade Federal de Viccedilosa (UFV)
Address 36571-000 Vicosa MG Brazil
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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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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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
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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
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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
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
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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
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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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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tho
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of
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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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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
Author Family Name ZuanonParticle
Given Name JansenSuffix
Division CPBA
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name GordoParticle
Given Name MarceloSuffix
Division Instituto de Ciecircncias Bioloacutegicas
Organization Universidade Federal do Amazonas (UFAM)
Address Manaus AM Brazil
Author Family Name FidelisParticle
Given Name LuanaSuffix
Division DCEN
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name JuenParticle
Given Name LeandroSuffix
Division Departamento de Biologia Geral
Organization Universidade Federal de Viccedilosa (UFV)
Address 36571-000 Vicosa MG Brazil
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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Journal HYDR-10750 Article 9441
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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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DOI 101007s10750-008-9441-x
Au
tho
r P
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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
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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
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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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
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Article No 9441 h LE h TYPESET
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tho
r P
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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
123
Journal Medium 10750 Dispatch 3-6-2008 Pages 15
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tho
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Author Family Name GordoParticle
Given Name MarceloSuffix
Division Instituto de Ciecircncias Bioloacutegicas
Organization Universidade Federal do Amazonas (UFAM)
Address Manaus AM Brazil
Author Family Name FidelisParticle
Given Name LuanaSuffix
Division DCEN
Organization Instituto Nacional de Pesquisas da Amazocircnia (INPA)
Address CP 478 69011970 Manaus AM Brazil
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
Author Family Name JuenParticle
Given Name LeandroSuffix
Division Departamento de Biologia Geral
Organization Universidade Federal de Viccedilosa (UFV)
Address 36571-000 Vicosa MG Brazil
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
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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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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
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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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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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
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Article No 9441 h LE h TYPESET
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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
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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
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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
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
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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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Article No 9441 h LE h TYPESET
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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
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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
123
Journal Medium 10750 Dispatch 3-6-2008 Pages 15
Article No 9441 h LE h TYPESET
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tho
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ro
of
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Please check the publication numbers in Article note and contribution numbers in Acknowledgments
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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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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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UNCORRECTEDPROOF
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
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
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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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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
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Article No 9441 h LE h TYPESET
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tho
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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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Article No 9441 h LE h TYPESET
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Please provide accessed date for Reference StatSoft Inc (2001)
Please provide details for lsquoboldrsquo emphasis present in Table 3
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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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
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
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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
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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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
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Article No 9441 h LE h TYPESET
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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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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
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
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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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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
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Article No 9441 h LE h TYPESET
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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
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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
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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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UNCORRECTEDPROOF
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
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
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tho
r P
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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
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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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
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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
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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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Article No 9441 h LE h TYPESET
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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
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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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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
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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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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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
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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UNCORRECTEDPROOF
UNCORRECTEDPROOF
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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Article No 9441 h LE h TYPESET
MS Code HYDR2696 h CP h DISK4 4
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r P
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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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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
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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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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
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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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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
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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
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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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Article No 9441 h LE h TYPESET
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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
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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
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
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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
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tho
r P
ro
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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
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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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
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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
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
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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UNCORRECTEDPROOF
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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UNCORRECTEDPROOF
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
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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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r P
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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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Article No 9441 h LE h TYPESET
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tho
r P
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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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Article No 9441 h LE h TYPESET
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tho
r P
ro
of
UNCORRECTEDPROOF
UNCORRECTEDPROOF
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
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
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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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
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
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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
tho
r P
ro
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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
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
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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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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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
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
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