STRUCTURE AND DIVERSITY OF THE LOBSTER COMMUNITY ON THE AMAZON CONTINENTAL SHELF

Crustaceana 86 (9) 1084-1102

STRUCTURE AND DIVERSITY OF THE LOBSTER COMMUNITY ONTHE AMAZON CONTINENTAL SHELF

BY

K. C. A. SILVA1), R. CRUZ2,4), I. H. A. CINTRA1) and F. A. ABRUNHOSA3)1) Instituto Socioambiental e dos Recursos Hídricos, Universidade Federal Rural da Amazônia,

Av. Tancredo Neves 2501, Montese, Belém, Pará, Brazil2) Instituto de Ciências do Mar (Labomar), Av. da Abolição 3207,

Meireles Fortaleza, Universidade Federal do Ceará, Brazil3) Instituto de Estudos Costeiros, Faculdade de Engenharia de Pesca, Universidade Federal do Pará,

Alameda Leandro Ribeiro, s/n, Aldeia, Bragança, Pará, Brazil

ABSTRACT

The present study analyses the diversity and community structure of the lobsters captured usingbottom trawl shrimp nets at depths of 41 to 626 m during fishery prospection cruises conductedbetween 1996 and 1998. The study area encompasses the coasts of the Brazilian states of Amapá andPará, which were divided into two sectors, to the north and to south of Cape North. The 44 lobsterspecimens identified belonged to nine species: Acanthacaris caeca (A. Milne-Edwards, 1881),Nephropsis aculeata Smith, 1881, Nephropsis rosea Bate, 1888, Palinustus truncatus A. Milne-Edwards, 1880, Panulirus argus (Latreille, 1804), Parribacus antarcticus (Lund, 1793), Polychelestyphlops Heller, 1862, Scyllarides delfosi Holthuis, 1960 and Stereomastis sculpta (Smith, 1880).The specimens were captured preferentially on muddy sand substrates in the northern sector andgravelly and muddy sand in the southern sector. All species were more common in the northern sectorduring the dry season (June-November), and in the southern sector in the rainy season (December-May).

RESUMO

O presente estudo aborda a diversidade e a estrutura da comunidade de lagostas capturadasdurante os cruzeiros de prospecção, entre 1996 e 1998, com rede de arrasto para camarão emprofundidade de 41 a 626 m. A área estudada compreende a costa dos estados do Amapá e Pará,a qual foi dividida em dois setores, norte e sul do Cabo Norte. Os 44 espécimes identificadospertencem a nove espécies: Acanthacaris caeca (A. Milne-Edwards, 1881), Nephropsis aculeataSmith, 1881, Nephropsis rosea Bate, 1888, Palinustus truncatus A. Milne-Edwards, 1880, Panulirusargus (Latreille, 1804), Parribacus antarcticus (Lund, 1793), Polycheles typhlops Heller, 1862,Scyllarides delfosi Holthuis, 1960, e Stereomastis sculpta (Smith, 1880). O grupo foi encontradopreferencialmente em fundo de areia lamosa ao norte, e em areia cascalhosa e areia lamosa ao sul.Na área ao norte as espécies foram mais comuns no período seco (junho a novembro) e ao sul nochuvoso (dezembro a maio).

4) Corresponding author; e-mail: [email protected]

© Koninklijke Brill NV, Leiden, 2013 DOI:10.1163/15685403-00003227

DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF 1085

INTRODUCTION

The lobsters are a diverse group of crustaceans, which vary not only in bodysize, but also in their geographic distribution and habitat preferences, being foundthroughout the World’s oceans at practically all depths.

The marine lobsters form the suborder Macrura Reptantia Bouvier, 1917, whichincludes four infraorders — Astacidea Latreille, 1802, Glypheidea Winkler, 1882,Achelata Scholtz & Richter, 1995 and Polychelida Scholtz & Richter, 1995(Chan, 2010). In Brazil, the harvesting of achelatan lobsters is an extremelyimportant activity from both a social and economic viewpoint. The species withthe highest commercial value belong to the families Palinuridae Latreille, 1802and Scyllaridae Latreille, 1825, and include the spiny lobsters, Panulirus argus(Latreille, 1804), Panulirus laevicauda (Latreille, 1817), Panulirus echinatusSmith, 1869 and the slipper lobsters, Scyllarides brasiliensis Rathbun, 1906 andScyllarides delfosi Holthuis, 1960.

The lobsters that inhabit the Brazilian coast have been studied by a number ofresearchers. The most important publications include the guideline to samplingmethods produced by Cruz et al. (2011), the Melo (1999) identification handbook,and the genetic studies of Diniz et al. (2004, 2005). Other studies include thoseof Cutrim et al. (2001), Porto et al. (2005) and Silva et al. (2002, 2003, 2008) innorthern Brazil, Fausto-Filho (1966) in the northeast, Serejo et al. (2006, 2007) inthe southeast and D’Incao (1999) in the state of Rio Grande do Sul, in the extremesouth. Santana et al. (2007) described the larvae, and Abrunhosa et al. (2007, 2008)raised larvae in the laboratory in the state of Ceará. Recently, Cavalcante et al.(2011) and Silva & Fonteles-Filho (2011) analysed the strategies adopted in thecountry for the harvesting of spiny lobsters.

The Brazilian government created the Revizee Program [Evaluation of thePotential Sustainability of the Natural Resources of the Exclusive Economic Zone]with the aim of managing and conserving natural resources for their sustainableexploitation. The present study examines the lobster specimens collected duringfishery prospection cruises of the northern coast of Brazil, with the principalobjective of analysing the spatial distribution of these crustaceans.

MATERIAL AND METHODS

The study area encompasses the continental shelf of the Brazilian states ofAmapá and Pará, part of the Exclusive Economic Zone of northern Brazil. Basedon the physiographic characteristics of this area, a line starting from Cabo Norte(00°59′09′′N 49°57′02′′W) was traced at an angle of 45°, in order to divide study

1086 K. C. A. SILVA ET AL.

area into two sectors: northern sector (NS) and souther sector (SS), using ArcGIS9.3 program (Esri, 2008).

The sampling was done during fishery prospection cruises carried out between1996 and 1998, using bottom trawl shrimp nets similar to those used by the localindustrial shrimping fleet, with rectangular wooden doors and iron shoes. The21 trawls lasted between 0.5 and 5.5 h, and were conducted on different typesof sediment bottoms, at depths varying from 41 to 626 m. Diurnal trawls wereconducted between 06:00 h and 17:59 h and nocturnal catches between 18:00 hand 05:59 h.

The lobsters captured were stored on ice in individually-identified baskets,labelled with date, trawl number and geographic position. The initial and finaldepth of each trawl, and the type of sediment were also recorded. As most of thesediments were classified as “indeterminate”, however, the classification proposedby Figueiredo-Júnior et al. (2008) was used for the analysis of the data. Oncelanded, the specimens were kept in a cold storage chamber until analysis.

The specimens were identified in the CEPNOR Crustaceans Laboratory and inthe Carcinology Laboratory of the Federal Rural University of Pernambuco using anumber of publications, including Takeda (1983) and Holthuis (1991). The specieswere classified according to the taxonomy of Chan (2010). All specimens weredeposited in the Carcinology Collection of the CEPNOR Crustaceans Laboratory.

The relative abundance of taxa is given by their percentual contribution to thesample:

A = n × 100

N,

where n is the number of individuals by superfamily, family or species, and N thetotal number of lobsters captured.

The frequency of occurrence of the different species, expressed as a percentage,was provided by the following formula:

F = p × 100

P,

where p is the number of trawls containing the species, and P the total number oftrawls containing lobsters.

The relative abundance and occurrence frequency were classified accordingto the scheme of Santos (2000). For abundance, the categories were dominant(above 50%), abundant (30-50%), uncommon (10-29%) and rare (below 10%). Forfrequency, they were very frequent (above 70%), frequent (30-70%), infrequent(10-30%) and sporadic (below 10%).

Differences in the total number of specimens captured between the two studysectors (NS and SS) were tested using χ2, with α = 0.05. Species richness


was determined on the basis of the simple count of the number of species, assuggested by Krebs (1985) and Ricklefs (2003). The Shannon diversity index wascalculated considering the study period as a whole. This method assumes that allthe specimens were collected randomly from an infinite population, and thus, thatall the species are represented in the sample (Tischer & Santos, 2003).

Equitability was also calculated for the study period as a whole. This indexvaries from 0 to 1, with values above 0.5 indicating that the specimens arewell distributed (proportionally equal across the species) and values below 0.5indicating a poor distribution of the specimens across species (Tischer & Santos,2003).

In order to evaluate whether the species found in both study sectors presentedthe same distribution patterns, the following variables were analysed: (i) depth,based on the following categories (Nittrouer et al., 1986): inner shelf (up to40 m), mid-shelf (40-60 m), outer shelf (60-100 m), break slope (100-200 m)and mesopelagic slope (200-1000 m) (Fischer & Haimovici, 2007); (ii) type ofsediment — gravel, gravelly sand, sand, muddy sand and mud (Figueiredo-Júnioret al., 2008); and (iii) season — rainy season, from December to May, and dryseason, from June to November (Oliveira et al., 2007).

The specimens were also sexed and the total length (TL) was measured from theanterior edge of the rostral plate to the posterior extremity of the telson. The mean(standard deviation) and range of values were determined separately for males andfemales. When only one specimen was available, its length was considered to bethe maximum value.

The specimens were sexed based on the diagnostic morphological characteris-tics for each family. In some species of the family Palinuridae, sexual dimorphismis found in the morphology of the pleopods, which are uniramous in the malesand biramous in the females, which also present a subchela on the fifth pair ofpereopods. In the Scyllaridae, the males have copulatory organs on the first pairof pleopods, while the females have oviduct openings in the coxae of the thirdpair of pereopods (D’Incao, 1999). In the family Nephropidae Dana 1852, the firstpair of pleopods in the males has been transformed into a rigid copulatory sty-lus, while the females also present oviduct openings in the coxae of the third pairof pereopods (D’Incao, 1999). In Polycheles, there is a chela on the fifth pair ofpereopods in the females, while in the males, this dactyl may be either simple ora subchela. The second to sixth pairs of pleopods are biramous, with an appendixmasculina in the males (Galil, 2000).

The sex ratio was not evaluated in the species found in both study sectors dueto the small numbers of males and females collected, given that, according toLegendre & Legendre (1998), a reliable use of the χ2 requires a total sample sizeat least 10-times larger than the number of cells in the contingency table.


RESULTS

Catches of species of lobsters

A total of 21 trawls were conducted during the nine cruises carried out overthe sector NS (n = 8) and SS (n = 13). These trawls were made over differenttypes of sediments, at depths varying from 41 to 626 m. A total of 44 lobsterspecimens were captured, with between one and seven lobsters being captured pertrawl (table I).

Representatives of three infraorders were collected, Achelata, Astacidea, andPolychelida, three superfamilies — Nephropoidea Dana, 1852, Palinuroidea La-treille, 1802 and Eryonoidea De Haan, 1841, and four families — Nephropidae,Palinuridae, Polychelidae Wood-Mason, 1875 and Scyllaridae. Eight genera wererecorded — Acanthacaris Bate, 1888, Nephropsis Wood-Mason, 1872, Palinustus

TABLE IFishery survey (FS), haul (H), sector (S), latitude and longitude, sediment type, mean depth, trawlingduration (TD), period of the day (P), i.e., night (N) or day (D) and number of lobsters (N ) captured inthe northern and southern study sectors with bottom shrimp trawl nets during the Northern Revizee

Score program between 1996 and 1998

FS H Date S Latitude Longitude Sediment Depth TD P N

(m)

I 7 02 September 1996 NS 03°23′N 49°06′W Gravelly sand 90 1 h D 2I 10 05 September 1996 SS 02°09′N 48°19′W Muddy sand 46 5 h N 1I 11 05 September 1996 SS 01°20′N 48°00′W Mud 48 5 h 30 min N 2I 18 09 September 1996 SS 01°28′N 46°43′W Gravelly sand 75 1 h 30 min D 2II 6 06 October 1996 NS 04°08′N 49°20′W Muddy sand 394 1 h D 2III 1 15 November 1996 NS 03°24′N 49°07′W Gravelly sand 92 1 h N 1III 2 15 November 1996 NS 03°43′N 48°53′W Muddy sand 186 1 h D 1III 5 16 November 1996 NS 03°44′N 48°33′W Gravel 459 1 h D 2III 6 16 November 1996 NS 03°11′N 48°54′W Gravelly sand 102 30 min N 1III 7 16 November 1996 NS 03°04′N 48°59′W Muddy sand 76 1 h N 1IV 1 07 December 1996 SS 02°44′N 47°39′W Gravel 621 1 h D 1IV 3 08 December 1996 SS 02°29′N 47°34′W Muddy sand 429 1 h D 2V 7 11 May 1997 SS 01°03′N 46°21′W Muddy sand 69 30 min D 2VII 3 20 March 1998 SS 02°43′N 47°39′W Gravel 626 1 h D 4VII 4 20 March 1998 SS 02°39′N 47°43′W Muddy sand 456 1 h D 3VIII 1 31 March 1998 NS 04°09′N 49°22′W Muddy sand 422 1 h D 1IX 5 01 May 1998 SS 02°08′N 48°19′W Muddy sand 51 1 h D 4IX 6 01 May 1998 SS 01°41′N 48°20′W Mud 41 1 h D 2IX 8 02 May 1998 SS 01°27′N 46°43′W Gravelly sand 75 1 h N 2IX 11 03 May 1998 SS 01°58′N 47°33′W Gravelly sand 76 1 h D 7X 5 01 June 1998 SS 01°28′N 46°46′W Gravelly sand 75 1 h N 1Total 44

NS, northern sector; SS, southern sector.


A. Milne-Edwards, 1880, Panulirus White, 1847, Parribacus Dana, 1852, Poly-cheles Heller, 1862, Scyllarides Gill, 1898 and Stereomastis Bate, 1888.

The infraorder Achelata was the most abundant in both study sectors, repre-senting 63.64% of specimens in the northern sector and 68.75% in the southernsector. The Astacidea contributed the remaining 36.36% of specimens collected inthe northern sector and 18.75% in the southern sector, while Polychelida were onlycollected in the southern sector, where they constituted 12.50% of specimens.

The most abundant families in the northern sector were Nephropidae and Pal-inuridae, both providing 36.36% of the specimens, with Scyllaridae contributing27.28%. In the southern sector, more than half of the specimens (54.55%) werepalinurids, followed by nephropids (18.18%), scyllarids (15.15%) and polychelids(12.12%).

Acanthacaris and Palinustus were recorded only in the northern sector, withboth genera contributing 18.18% of the specimens. Nephropsis and Panulirus bothalso contributed 18.18% of the specimens in this sector, while Scyllarides provided27.28%. In the southern sector, Parribacus (6.06% of the specimens), Polycheles(6.06%) and Stereomastis (6.06%) were exclusively found (i.e., only in that sector),while Nephropsis (18.18%), Panulirus (54.55%) and Scyllarides (9.09%) were alsopresent.

A total of nine lobster species were identified — Acanthacaris caeca (A.Milne-Edwards, 1881) (n = 2), Nephropsis aculeata Smith, 1881 (n = 4),Nephropsis rosea Bate, 1888 (n = 4), Palinustus truncatus A. Milne-Edwards,1880 (n = 2), Panulirus argus (n = 20), Parribacus antarcticus (Lund, 1793)(n = 2), Polycheles typhlops Heller, 1862 (n = 2), Scyllarides delfosi (n = 6) andStereomastis sculpta (Smith, 1880) (n = 2) (fig. 1).

The relative contribution of the different species to the specimens collected inthe present study was 45.48% for P. argus, 13.64% for S. delfosi, 9.09% for N.aculeata, 9.09% for N. rosea, 4.54% for A. caeca, 4.54% for P. antarcticus, 4.54%for P. truncatus, 4.54% for P. typhlops and 4.54% for S. sculpta.

Diversity indices

In the present study, species richness was lower (n = 5) in northern sector incomparison with southern sector (n = 7). The species N. aculeata, P. argus and S.delfosi were the only ones captured in both sectors (fig. 2).

In the present study, all the species were classified as uncommon in the northernsector, whereas in the southern sector most species were considered to be rare,except for P. argus, which was classified as dominant (table II). Similarly, mostspecies recorded in the northern sector were considered to be infrequent, with theexception of S. delfosi, which was classified as frequent. In the southern sector,


Fig. 1. The species of lobster identified during the present study: A, Acanthacaris caeca (A. Milne-Edwards, 1881), 30.0 cm; B, Nephropsis aculeata Smith, 1881, 11.0 cm; C, Nephropsis rosea Bate,1888, 11.5 cm; D, Polycheles typhlops Heller, 1862, 7.6 cm; E, Stereomastis sculpta (Smith, 1880),6.3 cm; F, Palinustus truncatus A. Milne-Edwards, 1880, 4.4 cm; G, Parribacus antarcticus (Lund,1793), 6.3 cm; H, Panulirus argus (Latreille, 1804), 29.0 cm and I, Scyllarides delfosi Holthuis,

1960, 23.3 cm.

most species were classified as sporadic, except for P. argus, which was frequent(table III).


Fig. 2. Georeferenced collected points for Acanthacaris caeca (A. Milne-Edwards, 1881), Nephrop-sis aculeata Smith, 1881, Nephropsis rosea Bate, 1888, Palinustus truncatus A. Milne-Edwards,1880, Panulirus argus (Latreille, 1804), Parribacus antarcticus (Lund, 1793), Polycheles typhlopsHeller, 1862, Scyllarides delfosi Holthuis, 1960, and Stereomastis sculpta (Smith, 1880), capturedin the northern (NS) and southern (SS) sectors of the study area collected using bottom shrimp trawlnets during the Northern Revizee Score program between 1996 and 1998. This figure is publishedin colour in the online edition of this journal, which can be accessed via http://booksandjournals.

brillonline.com/content/15685403.

A significant difference (χ2 = 11.0, df = 1, p<0.001) was found in thenumber of specimens captured between the two study sectors. Shannon’s index


TABLE IIRelative abundance and classification of the species captured in the northern (NS) and southern (SS)study sectors with bottom shrimp trawl nets during the Northern Revizee Score program between

1996 and 1998

Species Relative Classificationabundance (%)

NS SS NS SS

Acanthacaris caeca 18.18 UncommonNephropsis aculeata 18.18 6.06 Uncommon RareNephropsis rosea 12.12 UncommonParribacus antarcticus 6.06 RarePanulirus argus 18.18 54.55 Uncommon DominantPalinustus truncatus 18.18 UncommonPolycheles typhlops 6.06 RareScyllarides delfosi 27.27 9.09 Uncommon RareStereomastis sculpta 6.06 Rare

was 2.2999 bits ind−1 for the northern sector and 2.1410 bits ind−1 for the southernsector, while equitability was 0.9905 for the north and 0.7626 for the south.Both equitability values were over 0.5, indicating a relatively good distribution ofindividuals among the species in the two sectors. There was, however, a dominantspecies in the southern sector, where P. argus constituted 54.55% of the specimens.

Sex and length composition

In the present study, two female N. aculeata were captured in the northernsector of the study area, and a male and a female in the southern sector. Only

TABLE IIIFrequency of occurrence and classification of the species captured in the northern (NS) and southern(SS) study sectors with bottom shrimp trawl nets during the Northern Revizee Score program

between 1996 and 1998

Species Frequency of Classificationoccurrence (%)

NS SS NS SS

Acanthacaris caeca 25.00 InfrequentNephropsis aculeata 12.50 7.69 Infrequent SporadicNephropsis rosea 7.69 SporadicParribacus antarcticus 7.69 SporadicPanulirus argus 25.00 61.54 Infrequent FrequentPalinustus truncatus 12.50 InfrequentPolycheles typhlops 7.69 SporadicScyllarides delfosi 37.50 15.38 Frequent InfrequentStereomastis sculpta 7.69 Sporadic


TABLE IVSex distribution of lobsters (Nephropsis aculeata, Panulirus argus and Scyllarides delfosi) capturedin the northern (NS) and southern (SS) study sectors with bottom shrimp trawl nets during the

Northern Revizee Score program between 1996 and 1998

Species S Number of individuals

Male Female Total

Nephropsis aculeata NS 0 2 2Nephropsis aculeata SS 1 1 2Panulirus argus NS 1 1 2Panulirus argus SS 10 8 18Scyllarides delfosi NS 2 1 3Scyllarides delfosi SS 0 3 3

one male and one female P. argus were collected in the northern sector, whereas10 males and eight females were captured in the southern sector. Two male andone female S. delfosi were collected in the northern sector, but only three femalesin the southern sector (table IV). An ovigerous female N. aculeata was collectedin the northern sector at a depth of 459 m over a gravel bottom during the dryseason. Two ovigerous female P. argus were collected in the southern sector, onein shallower water (46 m) over muddy sand during the dry season, and the other inslightly deeper water (76 m) on gravelly sand during the rainy season.

The only male N. aculeata specimen was collected in the southern sector ofthe study area. The largest female was collected in the northern sector, and thesmallest in the southern sector. Both the largest and the smallest male P. arguswere collected in the southern sector, as were the females. All the male S. delfosiwere captured in the northern sector; the smallest female was also collected in thissector, whereas the largest female was captured in the southern sector (table V).

Depth and benthic substrate

The lobsters were captured at depths of between 41 m and 626 m within thestudy area, but were more common on the mesobenthic slope (200-1000 m) in thenorthern sector and on the outer shelf (60-100 m) in the southern sector (fig. 3).Nephropsis aculeata was captured only on the mesobenthic slope (200-1000 m) inboth sectors. In the northern sector, P. argus occurred only on the outer shelf (60-100 m), whereas in the southern sector it was more abundant, but not exclusive tothis zone. Scyllarides delfosi was more abundant on the outer shelf (60-100 m) inthe northern sector and on the mid-shelf (40-60 m) in the southern sector (fig. 4).

Nephropsis aculeata was captured from gravelly substrates in the northernsector of the study area, but on muddy sand in the south. Panulirus argus wascollected from gravelly and muddy sand in the northern sector, but presented a


TABLE VDescriptive parameters for the total length (mean and range of values, in cm) of lobsters (Nephropsisaculeata, Panulirus argus and Scyllarides delfosi) captured in the northern (NS) and southern (SS)study sectors with bottom shrimp trawl nets during the Northern Revizee Score program between

1996 and 1998

Species S Male Female

Mi M Ma SD N Mi M Ma SD N

Nephropsis aculeata NS 1 10.20 10.60 11.00 0.71 2Nephropsis aculeata SS 8.50 1 7.60 1Panulirus argus NS 26.40 1 22.00 1Panulirus argus SS 10.10 25.53 35.00 7.64 8 9.20 25.57 39.00 15.11 3Scyllarides delfosi NS 22.80 23.10 23.40 0.42 2 22.80 1Scyllarides delfosi SS 0 23.30 1

Abbreviations: S, sector; Mi, minimum; M, mean; Ma, maximum; SD, standard deviation; N ,number of lobsters.

preference for gravelly sand in the southern sector. Scyllarides delfosi occurred ongravelly sand in the northern sector, but occurred preferentially on muddy sandin the southern sector (fig. 5). No systematic pattern was observed in relation topossible substrate preferences in the species recorded in both sectors.

Lobster behaviour by season

The lobsters were more abundant in the northern sector during the dry season,but during the rainy season in the southern sector (fig. 6). Nephropsis aculeata

Fig. 3. Relative abundance of all the lobster species captured in the study area (NS and SS) duringthe prospection cruises conducted between 1996 and 1998 by depth.


Fig. 4. Number of individuals of the lobster species common to both sectors (NS and SS) of the studyarea captured during the prospection cruises conducted between 1996 and 1998 by depth.

occurred only in the rainy season in the northern sector, but only during the dryseason in the southern sector. By contrast, P. argus was collected in the northernsector only during the dry season, and was more abundant during the rainy seasonin the southern sector. Similarly, S. delfosi occurred only during the dry season inthe northern sector, and only in the rainy season in the southern sector (fig. 7).

Fig. 5. Number of individuals of the lobster species common to both sectors (NS and SS) of the studyarea captured during the prospection cruises conducted between 1996 and 1998 by sediment type.


Fig. 6. Relative abundance of all the lobster species captured in the study area (NS and SS) duringthe prospection cruises conducted between 1996 and 1998 by pluviometric season.

DISCUSSION

Lobsters species catches

In Brazil, Serejo et al. (2006) recorded a relative abundance of 0.10% and afrequency of occurrence of 1.30% for the palinurids.

Fig. 7. Number of specimens of the lobster species common to both sectors (NS and SS) of the studyarea captured during the prospection cruises conducted between 1996 and 1998 by pluviometric

season.


The results of the Revizee Program included the first records of six lobsterspecies in Brazilian waters: Ramos-Porto et al. (2000) reported Stereomastissculpta (Smith, 1880) from Pará, while Silva et al. (2002) recorded Acanthacariscaeca (A. Milne-Edwards, 1881); Nephropsis aculeata Smith, 1881; Nephropsisrosea Bate, 1888 and Polycheles typhlops Heller, 1862 on the northern coast. WhileSerejo et al. (2007) claimed the first Brazilian records of N. neglecta and N. rosea,the latter had been reported by Silva et al. (2002). Serejo et al. (2006) reported onlya single species of palinurid lobster, Justitia longimana (H. Milne Edwards, 1837)in the central coast of Brazil. In the same region, Serejo et al. (2007) reported fivenephropid lobster species, A. caeca, N. aculeata, Nephropsis agassizii A. Milne-Edwards, 1880, N. neglecta and N. rosea, four polychelids, Pentacheles laevisBate, 1878, Pentacheles validus A. Milne-Edwards, 1880, Polycheles typhlops andS. sculpta, and three scyllarids, S. brasiliensis, Scyllarus depressus (Smith, 1881)and Bathyarctus ramosae (Tavares, 1997), but only one palinurid, P. argus.

Takeda (1983) identified a total of 12 lobster species in a survey of FrenchGuiana and Suriname, including A. caeca, N. aculeata, Nephropsis neglectaHolthuis, 1974, N. rosea, P. argus, P. laevicauda, P. truncatus, P. typhlops, S.delfosi, Scyllarus chacei Holthuis, 1960, Scyllarus depressus (Smith, 1881) (asScyllarus nearctus Holthuis, 1960) and S. sculpta. Using a number of differentfishing devices, Silva et al. (2002) collected the same species, whereas Cutrimet al. (2001) collected only P. antarticus in the area known as Lixeira, in Pará(0°20′-1°10′N 47°00′-47°55′W). Fausto-Filho (1966) reported the occurrence offour species — P. argus, P. echinatus, P. laevicauda and S. brasiliensis — innortheastern Brazil.

Panulirus argus was also the most abundant species collected in northern Brazilby Silva et al. (2003) and in Amapá, the northern most coastal state of Brazil, bySilva et al. (2007). It was also the most common species collected by Porto et al.(2005) in the area off northern Brazil where the species is harvested commercially(between latitudes 3°50′N in Amapá and 1°30′N in Pará).

Diversity indices

The species richness (n = 9) recorded in the present study was lower than thevalues recorded by Takeda (1983) in French Guiana (n = 12) and Serejo et al.(2007) in the central coast of Brazil (n = 15); equal to that found by Silva et al.(2002) (n = 9) in North of Brazil, and greater than the values reported by Fausto-Filho (1966) (n = 2), Cutrim et al. (2001) (n = 1) and Serejo et al. (2006) (n = 1)in central coast of Brazil.

Understanding which species occur within a given area and how speciesrichness varies among sites is an important baseline for the management of natural


resources, which requires reliable information on the areas that are capable ofsupporting the largest number of species or which species are most abundant in agiven area, in order to develop an appropriate management strategy. The interest ofthe general public in the problems facing the planet’s natural diversity has grownconsiderably in recent decades, stimulated by the imminent extinction of manyspecies resulting from pollution and the destruction of habitats (Ricklefs, 1987).

Severino-Rodrigues et al. (2007) classified Nephropsis aculeata as occasional(captured in less than 19.9% of hauls) based on commercial bottom trawls thattargeted the langoustine Metanephrops rubellus (Moreira, 1903).

Sex and length composition

The maximum total lengths recorded in the present study were lower than thosereported by Holthuis (1991) for the species N. aculeata (14.5 cm), P. argus (45 cm)and S. delfosi (25 cm).

Silva et al. (2008) report that the stock of P. argus off the coast of Amapá andPará is made up of relatively large individuals, with a carapace length of up to22.8 cm, based on specimens collected between 2001 and 2003. These animalscontribute the largest numbers of eggs to the population (Kanciruk, 1980; Cruz& Bertelson, 2009). Cruz et al. (2011) found that the spawning biomass of thepopulation can be found at depths between 50 m and 100 m, and consider thesedepths to be an essential reference point for the production of larvae and theconservation of lobster populations. Based on this finding, these depths shouldnot be fished anywhere in Brazil.

Based on data from the Cuban archipelago (Cruz & Phillips, 2000), the coralbarrier reef of Santa Catalina in Colombia (Cruz et al., 2007) and the northern coastof Brazil (Silva et al., 2008), Cruz et al. (2011) found that the mean length of thecarapace increases with depth, but differentially between the two sexes. In addition,the females in reproductive condition tended to be larger than the immature femalesand, as the depth increased (27 m), their size became equal to or larger than that ofthe males.

Depth and benthic substrate

In the study of Serejo et al. (2006), palinurid lobsters were collected only atdepths between 50 m and 100 m.

A similar distribution pattern was recorded in the two sectors for two ofthe species common to both areas, that is, N. aculeata (captured only on themesopelagic slope at 200-1000 m) and P. argus (preferably in the externalcontinental shelf, at 60-100 m). Panulirus argus undertakes two types of migration(Fonteles-Filho & Ivo, 1980). One involves random movements parallel to the


coast, primarily during the second half of the year, when the animals search for newfeeding grounds. The second type of migration occurs during the first half of theyear, when the lobsters range in search of breeding grounds, moving systematicallyto deeper waters, perpendicular to the coast.

The lobsters were captured preferentially on muddy sand bottoms in thenorthern sector, and on gravelly and muddy sand in the southern sector. The naturalhabitat of the lobsters is a substrate of benthic calcareous algae (Fonteles-Filho,1992), formed by red algae of the family Rhodopliceae and green algae of theChlorophyceae.

Kowsmann & Costa (1979) described carbonate facies consisting of molluscs,benthic foraminifers and holocenic algae typical of shallow waters, as well asbiodetritic sands. The muddy substrates adjacent to the river are mainly composedof silt. The sediments overlaying the continental shelf to the northeast of themouth of the Amazon are rich in clays. This distribution of sediments probablyreflects the effects of the reworking of the substrates by tidal currents throughoutthe Amazon estuary, as well as the transportation of clay to the northwest by thecoastal currents, reaching a distance of some 1600 km, well beyond the mouth ofthe Orinoco River, forming the longest continuous mud deposit anywhere in theWorld (Fass, 1986).

Lobster behaviour by season

No clear pattern was observed in relation to the pluviometric season, consider-ing the species common to both sectors, due in particular to the fact that only P.argus was captured during both seasons, but only in the southern sector. The othertwo species, N. aculeata and S. delfosi, were captured in only one of the sectorsduring each season.

The pluviometric patterns observed by Curtrin (1986) in the study area can beunderstood in terms of the variation in the turbidity of the water which generatesthe Amazon plume, which runs to the northwest from the mouth of the river,running 500 km along the coast, with a width of 200 km. During the rainyseason, which occurs between December and May, the output of freshwater andthe discharge of sediments onto the continental shelf increase considerably. Bycontrast, during the dry season, which spans the other half of the year (from Juneto November), the salinity in the inner coastal waters increases and the NorthernBrazilian Current becomes more intense (Oliveira et al., 2007).

However, even during this period of reduced discharge there is an intense mixingof the oceanic waters with the freshwater discharge of the Amazon River provokedby the waves generated by the winds blowing over the waters of the continentalshelf (Silva et al., 2001). Minimum precipitation levels are reached in October


and November, when there is a marked reduction in the discharge of the Amazon,leading to an increase in the salinity of the waters of the internal continental shelf,but even so, a wedge of brackish water forms over the oceanic waters (Silva et al.,2001).

ACKNOWLEDGEMENTS

The authors express their thanks to the Centro de Pesquisa e Gestão dosRecursos Pesqueiros do Litoral Norte (CEPNOR, acronym in Portuguese) for themotivation and logistic support at the Research Group. We would also like toexpress our thanks to Italo José Araruna Vieira (“in memória”) and Élcio da PaulaRocha for their support during the implementation of the Revizee Program. Weexpress our gratefulness to anonymous referees and J. C. von Vaupel Klein forvery useful comments, which encouraged the authors to improve an earlier versionof this manuscript.

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First received 27 June 2012.Final version accepted 3 June 2013.

STRUCTURE AND DIVERSITY OF THE LOBSTER COMMUNITY ON THE AMAZON CONTINENTAL SHELF

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