Crustaceana 86 (9) 1084-1102
STRUCTURE AND DIVERSITY OF THE LOBSTER COMMUNITY ON
THE 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, Brazil
2 ) Instituto de Ciências do Mar (Labomar), Av. da Abolição 3207,
Meireles Fortaleza, Universidade Federal do Ceará, Brazil
3 ) 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 using
bottom trawl shrimp nets at depths of 41 to 626 m during fishery prospection cruises conducted
between 1996 and 1998. The study area encompasses the coasts of the Brazilian states of Amapá and
Pará, which were divided into two sectors, to the north and to south of Cape North. The 44 lobster
specimens identified belonged to nine species: Acanthacaris caeca (A. Milne-Edwards, 1881),
Nephropsis aculeata Smith, 1881, Nephropsis rosea Bate, 1888, Palinustus truncatus A. MilneEdwards, 1880, Panulirus argus (Latreille, 1804), Parribacus antarcticus (Lund, 1793), Polycheles
typhlops Heller, 1862, Scyllarides delfosi Holthuis, 1960 and Stereomastis sculpta (Smith, 1880).
The specimens were captured preferentially on muddy sand substrates in the northern sector and
gravelly and muddy sand in the southern sector. All species were more common in the northern sector
during the dry season (June-November), and in the southern sector in the rainy season (DecemberMay).
RESUMO
O presente estudo aborda a diversidade e a estrutura da comunidade de lagostas capturadas
durante os cruzeiros de prospecção, entre 1996 e 1998, com rede de arrasto para camarão em
profundidade 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 identificados
pertencem a nove espécies: 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), Polycheles typhlops Heller, 1862,
Scyllarides delfosi Holthuis, 1960, e Stereomastis sculpta (Smith, 1880). O grupo foi encontrado
preferencialmente 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 no
chuvoso (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 body
size, but also in their geographic distribution and habitat preferences, being found
throughout the World’s oceans at practically all depths.
The marine lobsters form the suborder Macrura Reptantia Bouvier, 1917, which
includes 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 extremely
important activity from both a social and economic viewpoint. The species with
the highest commercial value belong to the families Palinuridae Latreille, 1802
and Scyllaridae Latreille, 1825, and include the spiny lobsters, Panulirus argus
(Latreille, 1804), Panulirus laevicauda (Latreille, 1817), Panulirus echinatus
Smith, 1869 and the slipper lobsters, Scyllarides brasiliensis Rathbun, 1906 and
Scyllarides delfosi Holthuis, 1960.
The lobsters that inhabit the Brazilian coast have been studied by a number of
researchers. The most important publications include the guideline to sampling
methods produced by Cruz et al. (2011), the Melo (1999) identification handbook,
and the genetic studies of Diniz et al. (2004, 2005). Other studies include those
of Cutrim et al. (2001), Porto et al. (2005) and Silva et al. (2002, 2003, 2008) in
northern Brazil, Fausto-Filho (1966) in the northeast, Serejo et al. (2006, 2007) in
the southeast and D’Incao (1999) in the state of Rio Grande do Sul, in the extreme
south. 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 the
country for the harvesting of spiny lobsters.
The Brazilian government created the Revizee Program [Evaluation of the
Potential Sustainability of the Natural Resources of the Exclusive Economic Zone]
with the aim of managing and conserving natural resources for their sustainable
exploitation. The present study examines the lobster specimens collected during
fishery prospection cruises of the northern coast of Brazil, with the principal
objective of analysing the spatial distribution of these crustaceans.
MATERIAL AND METHODS
The study area encompasses the continental shelf of the Brazilian states of
Amapá and Pará, part of the Exclusive Economic Zone of northern Brazil. Based
on 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
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K. C. A. SILVA ET AL.
area into two sectors: northern sector (NS) and souther sector (SS), using ArcGIS
9.3 program (Esri, 2008).
The sampling was done during fishery prospection cruises carried out between
1996 and 1998, using bottom trawl shrimp nets similar to those used by the local
industrial shrimping fleet, with rectangular wooden doors and iron shoes. The
21 trawls lasted between 0.5 and 5.5 h, and were conducted on different types
of sediment bottoms, at depths varying from 41 to 626 m. Diurnal trawls were
conducted between 06:00 h and 17:59 h and nocturnal catches between 18:00 h
and 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 final
depth of each trawl, and the type of sediment were also recorded. As most of the
sediments were classified as “indeterminate”, however, the classification proposed
by Figueiredo-Júnior et al. (2008) was used for the analysis of the data. Once
landed, the specimens were kept in a cold storage chamber until analysis.
The specimens were identified in the CEPNOR Crustaceans Laboratory and in
the Carcinology Laboratory of the Federal Rural University of Pernambuco using a
number of publications, including Takeda (1983) and Holthuis (1991). The species
were classified according to the taxonomy of Chan (2010). All specimens were
deposited in the Carcinology Collection of the CEPNOR Crustaceans Laboratory.
The relative abundance of taxa is given by their percentual contribution to the
sample:
A=
n × 100
,
N
where n is the number of individuals by superfamily, family or species, and N the
total 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 of
trawls containing lobsters.
The relative abundance and occurrence frequency were classified according
to the scheme of Santos (2000). For abundance, the categories were dominant
(above 50%), abundant (30-50%), uncommon (10-29%) and rare (below 10%). For
frequency, 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 study
sectors (NS and SS) were tested using χ 2 , with α = 0.05. Species richness
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1087
was determined on the basis of the simple count of the number of species, as
suggested by Krebs (1985) and Ricklefs (2003). The Shannon diversity index was
calculated considering the study period as a whole. This method assumes that all
the specimens were collected randomly from an infinite population, and thus, that
all the species are represented in the sample (Tischer & Santos, 2003).
Equitability was also calculated for the study period as a whole. This index
varies from 0 to 1, with values above 0.5 indicating that the specimens are
well distributed (proportionally equal across the species) and values below 0.5
indicating a poor distribution of the specimens across species (Tischer & Santos,
2003).
In order to evaluate whether the species found in both study sectors presented
the same distribution patterns, the following variables were analysed: (i) depth,
based on the following categories (Nittrouer et al., 1986): inner shelf (up to
40 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 of
sediment — gravel, gravelly sand, sand, muddy sand and mud (Figueiredo-Júnior
et al., 2008); and (iii) season — rainy season, from December to May, and dry
season, from June to November (Oliveira et al., 2007).
The specimens were also sexed and the total length (TL) was measured from the
anterior 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 and
females. When only one specimen was available, its length was considered to be
the maximum value.
The specimens were sexed based on the diagnostic morphological characteristics for each family. In some species of the family Palinuridae, sexual dimorphism
is found in the morphology of the pleopods, which are uniramous in the males
and biramous in the females, which also present a subchela on the fifth pair of
pereopods. In the Scyllaridae, the males have copulatory organs on the first pair
of pleopods, while the females have oviduct openings in the coxae of the third
pair of pereopods (D’Incao, 1999). In the family Nephropidae Dana 1852, the first
pair of pleopods in the males has been transformed into a rigid copulatory stylus, while the females also present oviduct openings in the coxae of the third pair
of pereopods (D’Incao, 1999). In Polycheles, there is a chela on the fifth pair of
pereopods in the females, while in the males, this dactyl may be either simple or
a subchela. The second to sixth pairs of pleopods are biramous, with an appendix
masculina in the males (Galil, 2000).
The sex ratio was not evaluated in the species found in both study sectors due
to the small numbers of males and females collected, given that, according to
Legendre & Legendre (1998), a reliable use of the χ 2 requires a total sample size
at least 10-times larger than the number of cells in the contingency table.
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K. C. A. SILVA ET AL.
RESULTS
Catches of species of lobsters
A total of 21 trawls were conducted during the nine cruises carried out over
the sector NS (n = 8) and SS (n = 13). These trawls were made over different
types of sediments, at depths varying from 41 to 626 m. A total of 44 lobster
specimens were captured, with between one and seven lobsters being captured per
trawl (table I).
Representatives of three infraorders were collected, Achelata, Astacidea, and
Polychelida, three superfamilies — Nephropoidea Dana, 1852, Palinuroidea Latreille, 1802 and Eryonoidea De Haan, 1841, and four families — Nephropidae,
Palinuridae, Polychelidae Wood-Mason, 1875 and Scyllaridae. Eight genera were
recorded — Acanthacaris Bate, 1888, Nephropsis Wood-Mason, 1872, Palinustus
TABLE I
Fishery survey (FS), haul (H), sector (S), latitude and longitude, sediment type, mean depth, trawling
duration (TD), period of the day (P), i.e., night (N) or day (D) and number of lobsters (N ) captured in
the 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
I
I
I
I
II
III
III
III
III
III
IV
IV
V
VII
VII
VIII
IX
IX
IX
IX
X
Total
7
10
11
18
6
1
2
5
6
7
1
3
7
3
4
1
5
6
8
11
5
NS
SS
SS
SS
NS
NS
NS
NS
NS
NS
SS
SS
SS
SS
SS
NS
SS
SS
SS
SS
SS
03°23′ N
02°09′ N
01°20′ N
01°28′ N
04°08′ N
03°24′ N
03°43′ N
03°44′ N
03°11′ N
03°04′ N
02°44′ N
02°29′ N
01°03′ N
02°43′ N
02°39′ N
04°09′ N
02°08′ N
01°41′ N
01°27′ N
01°58′ N
01°28′ N
02 September 1996
05 September 1996
05 September 1996
09 September 1996
06 October 1996
15 November 1996
15 November 1996
16 November 1996
16 November 1996
16 November 1996
07 December 1996
08 December 1996
11 May 1997
20 March 1998
20 March 1998
31 March 1998
01 May 1998
01 May 1998
02 May 1998
03 May 1998
01 June 1998
NS, northern sector; SS, southern sector.
49°06′ W
48°19′ W
48°00′ W
46°43′ W
49°20′ W
49°07′ W
48°53′ W
48°33′ W
48°54′ W
48°59′ W
47°39′ W
47°34′ W
46°21′ W
47°39′ W
47°43′ W
49°22′ W
48°19′ W
48°20′ W
46°43′ W
47°33′ W
46°46′ W
Gravelly sand
Muddy sand
Mud
Gravelly sand
Muddy sand
Gravelly sand
Muddy sand
Gravel
Gravelly sand
Muddy sand
Gravel
Muddy sand
Muddy sand
Gravel
Muddy sand
Muddy sand
Muddy sand
Mud
Gravelly sand
Gravelly sand
Gravelly sand
Depth TD
(m)
90
46
48
75
394
92
186
459
102
76
621
429
69
626
456
422
51
41
75
76
75
1h
5h
5 h 30 min
1 h 30 min
1h
1h
1h
1h
30 min
1h
1h
1h
30 min
1h
1h
1h
1h
1h
1h
1h
1h
P N
D
N
N
D
D
N
D
D
N
N
D
D
D
D
D
D
D
D
N
D
N
2
1
2
2
2
1
1
2
1
1
1
2
2
4
3
1
4
2
2
7
1
44
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1089
A. Milne-Edwards, 1880, Panulirus White, 1847, Parribacus Dana, 1852, Polycheles Heller, 1862, Scyllarides Gill, 1898 and Stereomastis Bate, 1888.
The infraorder Achelata was the most abundant in both study sectors, representing 63.64% of specimens in the northern sector and 68.75% in the southern
sector. The Astacidea contributed the remaining 36.36% of specimens collected in
the northern sector and 18.75% in the southern sector, while Polychelida were only
collected in the southern sector, where they constituted 12.50% of specimens.
The most abundant families in the northern sector were Nephropidae and Palinuridae, both providing 36.36% of the specimens, with Scyllaridae contributing
27.28%. In the southern sector, more than half of the specimens (54.55%) were
palinurids, followed by nephropids (18.18%), scyllarids (15.15%) and polychelids
(12.12%).
Acanthacaris and Palinustus were recorded only in the northern sector, with
both genera contributing 18.18% of the specimens. Nephropsis and Panulirus both
also contributed 18.18% of the specimens in this sector, while Scyllarides provided
27.28%. 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 also
present.
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) and
Stereomastis sculpta (Smith, 1880) (n = 2) (fig. 1).
The relative contribution of the different species to the specimens collected in
the 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 in
comparison 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 northern
sector, whereas in the southern sector most species were considered to be rare,
except for P. argus, which was classified as dominant (table II). Similarly, most
species recorded in the northern sector were considered to be infrequent, with the
exception of S. delfosi, which was classified as frequent. In the southern sector,
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K. C. A. SILVA ET AL.
Fig. 1. The species of lobster identified during the present study: A, Acanthacaris caeca (A. MilneEdwards, 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).
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1091
Fig. 2. Georeferenced collected points for 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), Polycheles typhlops
Heller, 1862, Scyllarides delfosi Holthuis, 1960, and Stereomastis sculpta (Smith, 1880), captured
in the northern (NS) and southern (SS) sectors of the study area collected using bottom shrimp trawl
nets during the Northern Revizee Score program between 1996 and 1998. This figure is published
in colour in the online edition of this journal, which can be accessed via http://booksandjournals.
brillonline.com/content/15685403.
A significant difference (χ 2 = 11.0, df = 1, p<0.001) was found in the
number of specimens captured between the two study sectors. Shannon’s index
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K. C. A. SILVA ET AL.
TABLE II
Relative 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
abundance (%)
NS
Acanthacaris caeca
Nephropsis aculeata
Nephropsis rosea
Parribacus antarcticus
Panulirus argus
Palinustus truncatus
Polycheles typhlops
Scyllarides delfosi
Stereomastis sculpta
18.18
18.18
18.18
18.18
27.27
Classification
SS
6.06
12.12
6.06
54.55
6.06
9.09
6.06
NS
SS
Uncommon
Uncommon
Uncommon
Uncommon
Uncommon
Rare
Uncommon
Rare
Dominant
Rare
Rare
Rare
was 2.2999 bits ind−1 for the northern sector and 2.1410 bits ind−1 for the southern
sector, 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 of
individuals among the species in the two sectors. There was, however, a dominant
species 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 northern
sector of the study area, and a male and a female in the southern sector. Only
TABLE III
Frequency 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
occurrence (%)
NS
Acanthacaris caeca
Nephropsis aculeata
Nephropsis rosea
Parribacus antarcticus
Panulirus argus
Palinustus truncatus
Polycheles typhlops
Scyllarides delfosi
Stereomastis sculpta
25.00
12.50
25.00
12.50
37.50
SS
7.69
7.69
7.69
61.54
7.69
15.38
7.69
Classification
NS
Infrequent
Infrequent
Infrequent
Infrequent
Frequent
SS
Sporadic
Sporadic
Sporadic
Frequent
Sporadic
Infrequent
Sporadic
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1093
TABLE IV
Sex distribution of lobsters (Nephropsis aculeata, 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
Nephropsis aculeata
Nephropsis aculeata
Panulirus argus
Panulirus argus
Scyllarides delfosi
Scyllarides delfosi
S
NS
SS
NS
SS
NS
SS
Number of individuals
Male
Female
Total
0
1
1
10
2
0
2
1
1
8
1
3
2
2
2
18
3
3
one male and one female P. argus were collected in the northern sector, whereas
10 males and eight females were captured in the southern sector. Two male and
one female S. delfosi were collected in the northern sector, but only three females
in the southern sector (table IV). An ovigerous female N. aculeata was collected
in the northern sector at a depth of 459 m over a gravel bottom during the dry
season. Two ovigerous female P. argus were collected in the southern sector, one
in shallower water (46 m) over muddy sand during the dry season, and the other in
slightly deeper water (76 m) on gravelly sand during the rainy season.
The only male N. aculeata specimen was collected in the southern sector of
the study area. The largest female was collected in the northern sector, and the
smallest in the southern sector. Both the largest and the smallest male P. argus
were collected in the southern sector, as were the females. All the male S. delfosi
were captured in the northern sector; the smallest female was also collected in this
sector, 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 the
study area, but were more common on the mesobenthic slope (200-1000 m) in the
northern 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) in
both sectors. In the northern sector, P. argus occurred only on the outer shelf (60100 m), whereas in the southern sector it was more abundant, but not exclusive to
this zone. Scyllarides delfosi was more abundant on the outer shelf (60-100 m) in
the 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 northern
sector of the study area, but on muddy sand in the south. Panulirus argus was
collected from gravelly and muddy sand in the northern sector, but presented a
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K. C. A. SILVA ET AL.
TABLE V
Descriptive parameters for the total length (mean and range of values, in cm) of lobsters (Nephropsis
aculeata, 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
Mi
Nephropsis aculeata
Nephropsis aculeata
Panulirus argus
Panulirus argus
Scyllarides delfosi
Scyllarides delfosi
M
NS
SS
NS
SS 10.10 25.53
NS 22.80 23.10
SS
Ma
Female
SD
N
8.50
26.40
35.00 7.64
23.40 0.42
1
1
1
8
2
0
Mi
M
Ma
10.20 10.60 11.00
7.60
22.00
9.20 25.57 39.00
22.80
23.30
SD
N
0.71
2
1
1
3
1
1
15.11
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 on
gravelly sand in the northern sector, but occurred preferentially on muddy sand
in the southern sector (fig. 5). No systematic pattern was observed in relation to
possible 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) during
the prospection cruises conducted between 1996 and 1998 by depth.
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1095
Fig. 4. Number of individuals of the lobster species common to both sectors (NS and SS) of the study
area 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 dry
season in the southern sector. By contrast, P. argus was collected in the northern
sector only during the dry season, and was more abundant during the rainy season
in the southern sector. Similarly, S. delfosi occurred only during the dry season in
the 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 study
area captured during the prospection cruises conducted between 1996 and 1998 by sediment type.
1096
K. C. A. SILVA ET AL.
Fig. 6. Relative abundance of all the lobster species captured in the study area (NS and SS) during
the 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 a
frequency 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 study
area captured during the prospection cruises conducted between 1996 and 1998 by pluviometric
season.
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1097
The results of the Revizee Program included the first records of six lobster
species in Brazilian waters: Ramos-Porto et al. (2000) reported Stereomastis
sculpta (Smith, 1880) from Pará, while Silva et al. (2002) recorded Acanthacaris
caeca (A. Milne-Edwards, 1881); Nephropsis aculeata Smith, 1881; Nephropsis
rosea Bate, 1888 and Polycheles typhlops Heller, 1862 on the northern coast. While
Serejo 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 only
a 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 five
nephropid lobster species, A. caeca, N. aculeata, Nephropsis agassizii A. MilneEdwards, 1880, N. neglecta and N. rosea, four polychelids, Pentacheles laevis
Bate, 1878, Pentacheles validus A. Milne-Edwards, 1880, Polycheles typhlops and
S. 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 French
Guiana and Suriname, including A. caeca, N. aculeata, Nephropsis neglecta
Holthuis, 1974, N. rosea, P. argus, P. laevicauda, P. truncatus, P. typhlops, S.
delfosi, Scyllarus chacei Holthuis, 1960, Scyllarus depressus (Smith, 1881) (as
Scyllarus nearctus Holthuis, 1960) and S. sculpta. Using a number of different
fishing devices, Silva et al. (2002) collected the same species, whereas Cutrim
et 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 of
four species — P. argus, P. echinatus, P. laevicauda and S. brasiliensis — in
northeastern Brazil.
Panulirus argus was also the most abundant species collected in northern Brazil
by Silva et al. (2003) and in Amapá, the northern most coastal state of Brazil, by
Silva 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 the
values 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 FaustoFilho (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 species
richness varies among sites is an important baseline for the management of natural
1098
K. C. A. SILVA ET AL.
resources, which requires reliable information on the areas that are capable of
supporting the largest number of species or which species are most abundant in a
given area, in order to develop an appropriate management strategy. The interest of
the general public in the problems facing the planet’s natural diversity has grown
considerably in recent decades, stimulated by the imminent extinction of many
species 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 that
targeted the langoustine Metanephrops rubellus (Moreira, 1903).
Sex and length composition
The maximum total lengths recorded in the present study were lower than those
reported 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 and
Pará is made up of relatively large individuals, with a carapace length of up to
22.8 cm, based on specimens collected between 2001 and 2003. These animals
contribute the largest numbers of eggs to the population (Kanciruk, 1980; Cruz
& Bertelson, 2009). Cruz et al. (2011) found that the spawning biomass of the
population can be found at depths between 50 m and 100 m, and consider these
depths to be an essential reference point for the production of larvae and the
conservation of lobster populations. Based on this finding, these depths should
not be fished anywhere in Brazil.
Based on data from the Cuban archipelago (Cruz & Phillips, 2000), the coral
barrier reef of Santa Catalina in Colombia (Cruz et al., 2007) and the northern coast
of Brazil (Silva et al., 2008), Cruz et al. (2011) found that the mean length of the
carapace increases with depth, but differentially between the two sexes. In addition,
the females in reproductive condition tended to be larger than the immature females
and, as the depth increased (27 m), their size became equal to or larger than that of
the males.
Depth and benthic substrate
In the study of Serejo et al. (2006), palinurid lobsters were collected only at
depths between 50 m and 100 m.
A similar distribution pattern was recorded in the two sectors for two of
the species common to both areas, that is, N. aculeata (captured only on the
mesopelagic slope at 200-1000 m) and P. argus (preferably in the external
continental shelf, at 60-100 m). Panulirus argus undertakes two types of migration
(Fonteles-Filho & Ivo, 1980). One involves random movements parallel to the
DIVERSITY OF LOBSTER COMMUNITY ON AMAZON SHELF
1099
coast, primarily during the second half of the year, when the animals search for new
feeding grounds. The second type of migration occurs during the first half of the
year, when the lobsters range in search of breeding grounds, moving systematically
to deeper waters, perpendicular to the coast.
The lobsters were captured preferentially on muddy sand bottoms in the
northern sector, and on gravelly and muddy sand in the southern sector. The natural
habitat 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 the
Chlorophyceae.
Kowsmann & Costa (1979) described carbonate facies consisting of molluscs,
benthic foraminifers and holocenic algae typical of shallow waters, as well as
biodetritic sands. The muddy substrates adjacent to the river are mainly composed
of silt. The sediments overlaying the continental shelf to the northeast of the
mouth of the Amazon are rich in clays. This distribution of sediments probably
reflects the effects of the reworking of the substrates by tidal currents throughout
the Amazon estuary, as well as the transportation of clay to the northwest by the
coastal currents, reaching a distance of some 1600 km, well beyond the mouth of
the Orinoco River, forming the longest continuous mud deposit anywhere in the
World (Fass, 1986).
Lobster behaviour by season
No clear pattern was observed in relation to the pluviometric season, considering 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 other
two species, N. aculeata and S. delfosi, were captured in only one of the sectors
during each season.
The pluviometric patterns observed by Curtrin (1986) in the study area can be
understood in terms of the variation in the turbidity of the water which generates
the 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 rainy
season, which occurs between December and May, the output of freshwater and
the discharge of sediments onto the continental shelf increase considerably. By
contrast, during the dry season, which spans the other half of the year (from June
to November), the salinity in the inner coastal waters increases and the Northern
Brazilian Current becomes more intense (Oliveira et al., 2007).
However, even during this period of reduced discharge there is an intense mixing
of the oceanic waters with the freshwater discharge of the Amazon River provoked
by the waves generated by the winds blowing over the waters of the continental
shelf (Silva et al., 2001). Minimum precipitation levels are reached in October
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K. C. A. SILVA ET AL.
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 dos
Recursos Pesqueiros do Litoral Norte (CEPNOR, acronym in Portuguese) for the
motivation and logistic support at the Research Group. We would also like to
express our thanks to Italo José Araruna Vieira (“in memória”) and Élcio da Paula
Rocha for their support during the implementation of the Revizee Program. We
express our gratefulness to anonymous referees and J. C. von Vaupel Klein for
very useful comments, which encouraged the authors to improve an earlier version
of this manuscript.
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First received 27 June 2012.
Final version accepted 3 June 2013.