The First Zoeal Stage of Two Species of Grapsidae (Decapoda, Brachyura) and a Key to SuchLarvae from the Brazilian CoastAuthor(s): A. Fransozo, J. A. Cuesta, M. L. Negreiros-FransozoSource: Crustaceana, Vol. 71, No. 3 (Apr., 1998), pp. 331-343Published by: BRILLStable URL: http://www.jstor.org/stable/20105993Accessed: 22/02/2010 14:46

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THE FIRST ZOEAL STAGE OF TWO SPECIES OF GRAPSIDAE

(DECAPODA, BRACHYURA) AND A KEY TO SUCH LARVAE FROM THE BRAZILIAN COAST

BY

A. FRANSOZO1), J. A. CUESTA2) and M. L. NEGREIROS-FRANSOZO1)

l) NEBECC (Group of Studies on Crustacean Biology, Ecology and Culture), Departamento de Zoologia, Instituto de Bioci?ncias, Centro de Aq?icultura, Universidade Estadual Paulista,

C. P. 510, Cep 18618-000 Botucatu, S?o Paulo, Brazil

2) Departamento de Ecolog?a, Facultad de Biolog?a, Universidad de Sevilla, Apdo. 1095, E-41080 Sevilla, Spain

ABSTRACT

First zoeal stages of the grapsinid Goniopsis cruentata (Latreille, 1803) and the sesarminid

Aratus pisonii (H. Milne Edwards, 1837), are described and illustrated. Grapsinae zoeae can be

distinguished from the other grapsid larvae by the absence of lateral spines on the carapace and

the reduction of the antennal exopod to a small seta. A key to the first zoeal stage of the Brazilian

coast Grapsidae is provided.

R?SUM?

Les premiers stades zo? du grapsinide Goniopsis cruentata (Latreille, 1803) et du sesarminide

Aratus pisonii (H. Milne Edwards, 1837) sont d?crits et figur?s. Les zo?s de Grapsinae peuvent

?tre distingu?es de celles des autres larves de Grapsidae par l'absence d'?pines lat?rales sur la

carapace et la r?duction de l'?caill? exopodiale de l'antenne ? une petite soie. Une clef du premier

stade zo? des Grapsidae de la c?te br?silienne est fournie.

INTRODUCTION

Decapod classification is based on adult characters which are influenced by

adaptations to different life styles. There are many taxonomic uncertainties re

sulting from the difficulty of recognizing apomorphic features in adult crus

taceans. However, according to Rice (1980), the adaptations of adult brachyurans are not reflected in larval development.

Most marine decapods have comparable planktonic larvae, and giving more

consideration to larval characters would hopefully solve many of the problems of existing classifications.

? Koninklijke Brill NV, Leiden, 1998 Crustaceana 71 (3)

332 A. FRANSOZO ET AL.

The known Brazilian fauna is composed of 302 brachyuran species (Melo,

1996), of which only 25% have their complete larval development described.

Among the Brachyura, there are 22 species of littoral Grapsidae, distributed

in four subfamilies. The species with reference to larval descriptions, are the

following:

Grapsinae: Pachygrapsus gracilis (Saussure, 1858) studied by Ingle (1987)

and Brossi-Garcia & Rodrigues (1993), Pachygrapsus transversus (Gibbes, 1850)

by Ingle (1987) and Cuesta & Rodr?guez (1994), Planes cyaneus Dana, 1852 by Konishi & Minigawa (1990), and Geograpsus lividus (H. Milne Edwards, 1837)

by Cuesta (unpubl. data).

Sesarminae: Chasmagnathus granulatus Dana, 1851 studied by Boschi et al.

(1967), Aratus pisonii (H. Milne Edwards, 1837) by Warner (1968), Cyclograp sus integer (H. Milne Edwards, 1837) by Gore & Scotto (1982), Metasesarma

rubripes (Rathbun, 1897) by Diaz & Ewald (1968) and Mont? et al. (1990), Sesarma ricordi H. Milne Edwards, 1853 by Diaz & Ewald (1968), Sesarma

rectum Randall, 1840 by Fransozo & Hebling (1986), and Armases angustipes

(Dana, 1852) by Kowalczuk (1994); the larvae described by Aikawa (1937) as

Grapsus grapsus (Linnaeus, 1758) should probably be ascribed to a species of

this subfamily (Cuesta et al., in press). Varuninae: Cyrtograpsus altimanus Rathbun, 1914 studied by Scelzo & Bas

tida (1979), Euchirograpsus americanus A. Milne Edwards, 1880 by Wilson

(1980), and Cyrtograpsus angulatus Dana, 1851 by Boschi (1981).

Plagusiinae: Plagusia depressa (Fabricius, 1775) studied by Wilson & Gore

(1980), and Percnon gibbesi (H. Milne Edwards, 1853) by Paula & Hartnoll

(1989). At present there are no larval data of Pachygrapsus corrugatus (Von Martens,

1872), Armases benedicti (Rathbun, 1897), Sesarma crassipes Cano, 1889, Cyr

tograpsus affinis (Dana, 1851), and Euchiro grapsus kingsleyi (Miers, 1885).

Abele (1992) reviewed all the known American species of Sesarma and some

species were moved to the new genus Armases. For instance, Sesarma angustipes

changed to Armases angustipes, and Sesarma ricordi to Armases ricordi. Later,

Niem (1993) also referred Metasesarma rubripes to the genus Armases.

Only more descriptions of reared larvae will lead to identifications of plank

tonic zoeae and a more valid classification of decapod crustaceans. This paper

aims to facilitate the identification of planktonic grapsid larvae from the Brazilian

coast by providing the first description of zoea I of Goniopsis cruentata and a

redescription of the same stage of Aratus pisonii, both hatched in the laboratory.

BRAZILIAN GRAPSID ZOEAE 333

MATERIAL AND METHODS

According to Melo (1996) the grapsid crab Goniopsis cruentata is distributed

in the western Atlantic from Florida, the Bermudas, and the Gulf of Mexico, to Central America, the north of South America, the Guyanas, and Brazil (i.e.,

Fernando de Noronha and from Para to Santa Catarina). Aratus pisonii is dis

tributed in the western Atlantic from Florida and the Gulf of Mexico, to Central

America, the Antilles, the north of South America, the Guyanas, and Brazil (i.e.,

from Paraiba to Santa Catarina), and even to the eastern Pacific (from Nicaragua to Peru).

Ovigerous females of G. cruentata and A. pisonii were collected during No

vember 1993, in an estuarine mangrove area in Ubatuba (23?S 45?W), along the

northern coast of Sao Paulo State, Brazil.

The specimens obtained were transported to the laboratory (Botucatu, S?o

Paulo) in thermic boxes of 20 1 capacity, with 1 1 of water from the collecting site. In the laboratory, the ovigerous females were maintained at a constant

temperature (24 ? 1?C). Each specimen was isolated in aquaria with aerated

natural sea water (27%o), until hatching. Adult females were fed on fish muscles and mangrove leaves, offered twice

a week. They were daily checked for the occurrence of hatching larvae or pre zoeae.

Larvae (zoea I) were preserved in a mixture of alcohol (96%) and glycerin in

a 1:1 proportion. The illustrations were made from live and fixed material, based on 20 spec

imens or more. Drawings and measurements were made under a Zeiss micro

scope, provided with a camera lucida. In the morphological descriptions, the

setation is listed from proximal to distal segments. The terminology adopted in the descriptions is based on Ingle (1992). The antennal type mentioned in the table was based on the peduncle/exopod

length ratio (Aikawa, 1929):

Type A, peduncle as long as exopod, similarly modified or smooth, and longer than rostrum or antennule.

Type B, medium size, in many cases longer than rostrum; exopod about half

as long as peduncle or even a little longer.

Type C, exopod less than 1/4 the length of the protopodite. The telson type adopted was that proposed by Aikawa (1929):

Type A, fork well developed, one to two-thirds as long as telson, the pointed ends extending straight outwards; telson triangular, bearing on either side 1-3

short outer denticulations on lateral and dorsal surfaces, varying in number ac

cording to species.

334 A. FRANSOZO ET AL.

Type B, differing from above by the absence of outer teeth.

RESULTS

Goniopsis cruentata (Latreille, 1803) Zoea I (figs. 1A-D, 2A-D)

Total length 0.87 ? 0.04 mm; carapace length 0.41 ? 0.02 mm.

Carapace (fig. 1 A). Globose, smooth, and without tubercles. Rostral and dorsal

spine stout. Lateral spines minute, rounded, knob-like projections. Pair of dorso

lateral simple setae. Postero-lateral margin without setae. Eyes sessile.

Antennule (fig. 1C). Uniramous. Endopod absent. Unsegmented exopod with

2 long aesthetascs and 2 simple setae.

Antenna (fig. ID). Well developed protopod slightly shorter than rostral spine,

spinous process with minute denticles. Exopod present as small setae.

Mandible. Endopod palp absent.

jc.o

Fig. 1. Goniopsis cruentata (Latreille, 1803), zoea I. A, lateral view; B, abdomen; C, antennule;

D, antenna.

BRAZILIAN GRAPSID ZOEAE 335

A,B_t

Fig. 2. Goniopsis cruentata (Latreille, 1803), zoea I. A, maxillule; B, maxilla; C, first maxilliped;

D, second maxilliped. In C and D, natatory setae are shown truncated.

Maxillule (fig. 2A). Coxal and basal endites with 5 plumodenticulate setae

each; seta on external margin of basis absent. Endopod 2-segmented with 1

simple seta on proximal segment and 1 subterminal and 4 terminal sparsely setose setae on distal one.

Maxilla (fig. 2B). Coxal and basal endites bilobed with 4 + 3 and 5 + 4

plumodenticulate setae on inner and outer lobes, respectively. Unsegmented

endopod distally bifurcate with 2 + 2 long setose setae. Scaphognathite with 4

marginal setose setae and a very stout posterior process.

First maxilliped (fig. 2C). Basis with 8 medial setae arranged 2 + 2 + 2 + 2.

Endopod 5-segmented with 1, 2, 1, 2, 5 setae. Exopod 2-segmented with 4

terminal plumose setae.

Second maxilliped (fig. 2D). Basis with 4 setae arranged 1 +1 +1 +1. Endopod

3-segmented with 0, 1, 5 setae. Exopod 2-segmented with 4 terminal plumose setae.

Third maxilliped. Absent.

Pereiopods. Absent.

336 A. FRANSOZO ET AL

Abdomen (fig. 1A, B). Five abdominal somites; segments 2 and 3 with dorso

lateral processes; postero-lateral process on segment 3 and 4 as subacute lobes.

Pleopods absent. Telson (fig. IB) with lateral margin straight with a broad

U-shaped cleft; each half with 3 plumose setae of equal length. Outer spines absent.

Aratus pisonii (H. Milne Edwards, 1837) Zoea I (figs. 3A-D, 4A-D)

Aratus pisonii: Warner, 1968: 251 (fig. 1).

Total length 0.69 ? 0.03 mm; carapace length 0.41 ? 0.02 mm.

Fig. 3. Aratus pisonii (H. Milne Edwards, 1837), zoea I. A, lateral view; B, abdomen; C, antennule;

D, antenna.

BRAZILIAN GRAPSID ZOEAE 337

Carapace (fig. 3A). Globose, smooth, and without tubercles. Rostrum stout;

dorsal spine recurved.

Lateral spines absent. A pair of dorso-lateral simple setae and two pairs of

mid-dorsal setae.

Postero-lateral margin without setae. Eyes sessile.

Antennule (fig. 3C). Uniramous. Endopod absent. Unsegmented exopod with

3 long aesthetascs and 2 simple setae.

Antenna (fig. 3D). Well developed protopod slightly longer than rostral spine,

spinous process with several denticules. Exopod almost reaching tip of protopod

and bearing 3 simple setae.

Mandible. Endopod palp absent.

Fig. 4. Aratus pisonii (H. Milne Edwards, 1837), zoea I. A, maxillule; B, maxilla; C, first

maxilliped; D, second maxilliped. In C and D, natatory setae are shown truncated.

338 A. FRANSOZO ET AL.

Maxillule (fig. 4A). Coxal and basal endites with 5 plumodenticulate setae

each. Endopod 2-segmented with 1 simple seta on proximal segment, 1 subter

minal and 4 terminal sparsely setose setae on distal one.

Maxilla (fig. 4B). Coxal and basal endites bilobed with 4 + 4 and 4 + 4

plumodenticulate setae in inner and outer lobes. Unsegmented endopod distally bifurcate with 2 + 3 long setose setae. Scaphognathite with 4 marginal setose

setae and very stout posterior process.

First maxilliped (fig. 4C). Basis with 10 medial setae arranged 2 + 2 + 3 + 3.

Endopod 5-segmented with 2, 2, 1, 2, 5 setae. Exopod 2-segmented with 4

terminal plumose setae.

Second maxilliped (fig. 4D). Basis with 4 medial setae arranged 1 + 1 + 1 + 1.

Endopod 3-segmented with 0, 1, 6. Exopod 2-segmented with 4 terminal plumose setae.

Third maxilliped. Absent.

Pereiopods. Absent.

Abdomen (fig. 3A, B). Five abdominal somites; segments 2 and 3 with dorso

lateral processes; postero-lateral processes on segments 2 to 5 as subacute lobes.

Pleopods absent. Telson (fig. 3B) with posterior margin with a broad U-shaped

cleft; each half with 3 plumose setae of equal length. Outer spines absent.

DISCUSSION

The meroplankton constitutes an important part of the marine community as

a whole, but it is still poorly studied for the Brazilian coast. The crustacean

larvae, mainly decapods, are very common in the marine meroplankton, but

comparatively few are described. This fact probably is one of the causes of the

lack of ecological studies on meroplanktonic organisms in Brazilian waters.

The Grapsidae, in particular, are very difficult to distinguish among the brachy uran crabs but, according to Rice (1980), they are characterized by: telson furcae

almost always unarmed but occasionally with 1 or 2 very small setae or spines;

distal segment of the endopod of the maxillule always armed with 4 terminal

and 1 subterminal setae, and basal segment almost always with a simple seta;

maxillar endopod with 2 + 2 or 2 + 3 setae; first maxilliped basis with 8, 10,

or 12 medial setae, endopod segments usually with 2, 2, 1, 2, 5 setae, second

maxilliped basis with 4 medial setae. The rest of the appendages present more

variable features.

In table I, the first zoeal features of our material and previously studied larvae

are compared.

BRAZILIAN GRAPSID ZOEAE 339

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Table II Differences between the first zoeal stage of Aratus pisonii (H. Milne Edwards,

1837) described by Warner (1968), and the present study

Appendages Warner (1968) Present study

Antennule 3 a 3 a, 2 s

Antenna

Exopod Is 3 s Maxilla

Coxal endite 7 sp 4 + 4 sp First maxilliped

Basis no data 2 + 2 + 3 + 3

Endopod 2,2,1,1,5 2,2,1,2,5

Second maxilliped Basis no data 1 + 1 + 1 + 1

Endopod_OU_0,1,6

a, aesthetacs; s, simple setae; sp, plumodenticulate setae.

Larvae of the Grapsinae are distinguishable by: antenna with vestigial endopod

seta; abdominal somites 3-5 with remarkable postero-lateral projections, and

telson usually with 2 short outer spines on outer margin. This is the first description of a zoea of the genus Goniopsis. Its morphological

characters are similar to those of the rest of the grapsinine larvae (Cuesta et al.,

in press), reinforcing the genus status in this subfamily. The first zoeal stage of

Goniopsis cruentata differs from other Grapsinae zoeae mainly by the absence

of outer spines at the base of each fiircal ramus.

The larval development of Aratus pisonii was described by Warner (1968). His

description of the first zoea and our present description differ markedly (table II).

Grapsus tenuicrustatus (Herbst, 1783) rather than G. grapsus (L., 1758) is

the Japanese representative of this genus. However, Aikawa (1937) apparently misidentified his material, since his descriptions probably refer to a sesarminid

(Cuesta et al., in press). The larvae of Chasmagnathus show affinities with Varuninae larvae mainly

by the presence of lateral spines on the carapace and the endopod setation of the

maxilla.

Plagusiinae larvae are characterized by dorsal, rostral, and lateral carapace

spines carrying tubercles, spinules, and/or setae, and by the presence of a single seta on the basal segment of the second maxilliped's endopod. The degree of

difficulty in recognizing species is higher among larvae of the same genus, par

ticularly, on the Brazilian coast, for Cyrtograpsus, Armases, and Pachygrapsus. As can be seen in table I, Euchiro grapsus americanus larvae are more similar

to Plagusiinae than to Varuninae. Besides, the first zoea of E. americanus has

BRAZILIAN GRAPSID ZOEAE 341

lateral, rostral, and dorsal spines on the carapace with several sparse teeth or

setae, has a type A telson in the first zoea, but type B in subsequent stages

(Wilson, 1980). New genetic evidence (mt-DNA sequence data from 16s rRNA)

suggests that the genus Euchiro grapsus is very closely related to Plagusia and

does not show affinity to any of the sequenced Varuninae (cf. Schubart et al.,

unpubl. data).

The present analysis allows the presentation of the key as below.

KEY TO THE FIRST ZOEAL STAGE OF GRAPSIDAE FROM THE BRAZILIAN COAST

1. Carapace lateral spines present.Varuninae and Plagusiinae (plus Chasmagnathus), 2 -

Carapace lateral spines absent.Grapsinae and Sesarminae, 6

2. Antenna type B.Varuninae (plus Chasmagnathus), 3 - Antenna type C.Plagusiinae (plus Euchirograpsus), 4

3. Basipodite of the first maxilliped with 10 medial setae .Cyrtograpsus spp. -

Basipodite of the first maxilliped with 8 medial setae.Chasmagnathus granulata

4. Telson type B.Plagusiinae, 5 -

Telson type A.Euchirograpsus americanus

5. Endopod of the second maxilliped with 1,1, and 5 setae.Percnon gibhesi -

Endopod of the second maxilliped with 1,1, and 6 setae.Plagusia depressa

6. Antennal exopod longer than 1/2 of spinous process.Sesarminae, 7 - Antennal exopod reduced to a small seta.Grapsinae, 10

7. Antenna type A.Cyclograpsus integer -

Antenna type B.8

8. Endopod of the second maxilliped with 0, 1, 5 setae.Sesarma rectum -

Endopod of the second maxilliped with 0, 1, 6 setae.9

9. Antennal exopod with 3 terminal setae.Aratus pisonii - Antennal exopod with 2 terminal setae.Armases spp.

10. Telson furca unarmed (type B).Goniopsis cruentata - Telson furca with minute outer seta-like spines (type A).11

11. Fourth abdominal segment with medio-lateral and postero-lateral process well developed

.Planes cyaneus - Fourth abdominal segment with minute medio-lateral process.12

12. Fourth abdominal segment laterally expanded.Pachygrapsus spp. - Fourth abdominal segment not laterally expanded.Geograpsus lividus

ACKNOWLEDGEMENTS

The second author's travel costs to Brazil and his stay at Botucatu (SP) at the

NEBECC Laboratory, were supported by a grant from the Junta de Andaluc?a

(Ayudas a la Investigaci?n) and the facilities of the NEBECC. The FAPESP

provided transportation during the work (#94/4878-8). The authors wish to thank

especially Augusto Alberto Valero Flores, for his English suggestions.

342 A. FRANSOZO ET AL.

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Received for publication 30 January 1997.