First zoeal stage of the crab Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867) (Decapoda, Brachyura) and revision of the larval morphology of superfamily Trapezioidea

ABSTRACT

The morphology of the first zoeal stage of Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867) was described from laboratory-hatched material obtained from ovigerous females collected at Vitória Island on the southeastern Brazilian coast. We compared the larval morphology (zoea I) of fourteen species of the superfamily Trapezioidea, which Domecia glabra Alcock, 1899 is the only congeneric representative of the species described in this study. The morphological characteristics of the first zoea that distinguish D. acanthophora from D. glabra are: three aesthetascs on the exopod antennule; three pairs of lateral spines on carapace; bilobed basial endite of maxilla, with four plumodenticulate setae on each lobe; and telson furcae distally spinulated. It also provides information that may enhance some phylogenetic hypotheses within Trapezioidea crabs.

Key words
Domeciidae; larval description; southeastern Brazil; Tetraliidae; Trapeziidae

Introduction

The most recent Brachyura classification (Ng et al., 2008; De Grave et al., 2009; Ahyong et al., 2011) recognizes the superfamily Trapezioidea Miers, 1886, which consists of three families: Domeciidae Ortmann, 1893, Tetraliidae Castro, Ng & Ahyong, 2004 and Trapeziidae Miers, 1886. Crabs of the family Domeciidae are grouped into four genera: DomeciaEydoux & Souleyet, 1842, JonesiusSankarankutty, 1962, MaldiviaBorradaile, 1902 and PalmyriaGalil & Takeda, 1986. The species Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867) has a wide geographical distribution in the Western Atlantic, extending from North Carolina, Bermuda, Florida, the Gulf of Mexico, the Antilles, northeastern South America and Brazil (St. Peter and St. Paul Archipelago, Rocas Atoll, Fernando de Noronha, and from states of Ceará and Pernambuco to São Paulo) (Alves et al., 2006; Coelho-Filho, 2006; Melo, 1998).

In recent decades, several studies about the morphology of larval stages have been used to elucidate the taxonomic status of these crabs, of which there is still no consensus with respect to the superfamily Trapezioidea (e.g., Clark and Guerao, 2008; Clark and Ng, 2010). About 60 species of the superfamily Trapezioidea are known (see Ng et al., 2008), but the morphology of the first zoeal stage is described only for 14 species (see Tab. 1).

Table 1
Comparison of some larval characters of the first zoeal stage of Domecia acanthophor a (Desbonne, in Desbonne & Schramm, 1867) and other species of the superfamily Trapezioidea.

Species/original larval description

Domeciidae Ortmann, 1893

Tetrallidae Castro, Ng & Ahyong, 2004

Trapeziidae Miers, 1886

Calocarcininae Števčić, 2005

Quadrellinae Števčić, 2005

Trapeziinae Miers, 1886

Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867)1

Domecia glabra Alcock, 18992

Tetralia cavimana Heller, 18613

Tetralia glaberrima (Herbst, 1790)4

Tetralia rubridactyla Garth, 19715

Calocarcinus africanus Calman, 19096

Quadrella maculosa Alcock, 18985

Quadrella serenei Galil, 19865

Trapezia bidentata (Forskål, 1775)4,7

Trapezia cymodoce (Herbst, 1801)3,4,7

Trapezia digitalis Latreille, 1828)4,7

Trapezia richtersi Galil & Lewinsohn, 19835

Trapezia rufopunctata (Herbst, 1799)4

Trapezia septata Dana, 18524

Trapezia tigrina Eydoux & Souleyet, 18427

Carapace

Rostral

1sp (smooth)

1sp (smooth)

1sp (spinulate)

1 sp (spinulate)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (smooth)

Dorsal

1sp (smooth)

1sp (smooth)

1sp (spinulate)

1 sp (spinulate)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (smooth)

1sp (smooth)

1sp (spinulate)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

1sp (smooth)

1sp (spinulate)

Lateral

6sp (3 pairs) (with 2 spinulates)

4sp (2 pairs) (with 1 spinulate)

4sp (2 pairs) (with 1 spinulate)

4sp (2 pairs) (spinulates)

4sp (2 pairs) (with 1 spinulate)

6sp (3 pairs) (smooth)

4sp (2 pairs) (with 1 spinulate)

4sp (2 pairs) (smooth)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

2sp (1 pair) (spinulate)

Antennule

Exopod

3a+1s

4a+1-2s

4a+1s, 1sp

2a+2s*

4a+1s

5a+1s*

4a+1s

4a+1s

3a+1s

4a+1s, 1sp

2a+3s

4a+1s

2a+3s

2a+2s

2a+3s?

Antenna

Exopod

3s

3s

3s

3s

3s

3s

3s+1spp

3s

3s

3s

3s

3s

3s

3s

3s

Protopod

spinulate bilaterally

spinulate bilaterally

spinulate bilaterally

spinulate bilaterally

spinulate bilaterally (variable size)

spinulate bilateral and distaly

multispinulate distally

multispinulate distally

spinulate distally

spinulate distally

spinulate distally

spinulate bilaterally

spinulate distally

spinulate distally

spinulate distally

Maxillule

Coxal endite

(1s+5pd)

(2+5)p*

(2+5)?

(2+5)p

(2+5)p*

(1+6)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p*

(2+5)p

Basial endite

(2c+3pd), m

(3c+2p)*

(4c*+1?)

(5p+2c)

(4c+1s)*

(3pd+2c)*

(4c+1p)*

(4c+1p)*

(3p+2c)*

(3c+2p)*

(3p+2c)*

(4c+1p)*

(3p+2c)*

(3p+2c)*

(3p+2c)

Endopod

0, (3pd+1s)

0, 4p*

1s*, 4?

1p, (1+4)p

1s, (1+4)p*

1pd*, (2+4)pd*

1p, (2+4)p*

1p, (2+4)p*

1s*, (1+4)s*

1s*, (3p+2s)

1s*, (1+4)p*

1p, (1+4)p*

1s, (1s+4p)*

1s*, (2p+3s)*

1, (1+4)p

Maxilla

Coxal endite

(4+3)pd, 1sp

(4+3)p*, 1sp

(4+3)?, 1sp

(4+3)p*

(4+3)p*, 1sp

(6+4)p*

(5+3)p*, 1sp

(5+3)p*, 1sp

(5+3)p*

(5-6+3)p*, 1sp

(5+3)p*

(5+3)p*, 1sp

(5+3)p*

(5+3)p*

(5+3)p

Basial endite

(4+4)pd, m

(5+3)p*

(4+4)?

(4+4)p*

(4p, 1sp+4p)*

(5+4)p*

(5+4)p*

(5+4)p*

(4+4)p*

(4+4)p*, 1sp

(4+4)p*

(4+4)p*, 1sp

(5+4)p*

(4+4)p*

(4+4)p

Endopod

(1pd,1s+2pd, 1s)

(3+2)p*

(2+3)?

(2+3)p*

(2+3)p*

(3+5)p*

(3+5)p*

(3+5)p*

(3+2)p*

(3-4+2)p*

(3+2)p*

(3+2)p*

(3+2)p*

(3+2)p*

(3+2)p

First maxilliped

Coxa

no seta

1p*

?

?

1p*

1p*

no seta

no seta

?

?

?

1p*

?

?

no seta

Basis

(2+2+3+2)s

(2+2+3+2)p*

(2+2+3+3)?

(2+2+3+3)p*

(2+2+3+3)p*

(2+2+3+3)p*

(2+2+3+3)p*

(2+2+3+3)p*

(2s+2s+3p+2p)*

(2+2+3+3)p

(2+2+3+3)p*

(2+2+3+3)p*

(2+2+3+2)p

(2+2+3+3)p

(2+2+3+3)?

Endopod

1s, (1s+1pd), 0, 2pd, (2+2)pd

1p, 2p, 0, 2p, (4+1)p*

2?, 2?, 1?, 2?, (4+1)?

2p,2p,1s, 2s, (4p+1s)*

3p, 2p, 1p, 2p, (4p+1s)*

(1s+2p), 2p, 1p, 2p, (4+1)p*

3p, 2p, 1p, 2p, (4p+1s)*

3p, 2p, 1p, 2p, (4p+1s)*

2s, 2s, 1s, 2s, (4p+1s)*

2s, 2s, 1s, 2s, (4+1)s*

2s, 2s, 1p, 2p, (4p+1s)*

2p, 2p, 1p, 2p, (4p+1s)*

2s, 2s, 1s, 2s, (4p+1s)*

2s, 2s, 1s, 2p, (4p+1s)*

2?, 2?, 1?, 2?, (4+1)?

Second maxilliped

Basis

(1+1+1+1)s

(1+1+1+1)p*

(1+1+1+1)?

(1+1+1+1)s*

(1+1+1+1)p*

(1+1+1+1)p*

(1+1+1+1)p*

(1+1+1+1)p*

(1+1+1+0)p*

(1+1+1+1)p

(1+1+1+0)p

(1+1+1+0)p*

(1+1+1+0)p

(1+1+1+1)p

(1+1+1+0)p

Endopod

0, 0, (2+2)p

0, 0, (2+2)p*

1?, 1?, (1+3)?

1s,1s, (2p+2s)*

1p, 1p, (2p+2s)*

1p, 1p, (2s,1p+3s)*

1p, 1p, (2s+2p)*

1p, 1p, (3p+1s)*

0, 1s, (1c+2s)*

0, 1s, (3p+1s)*

0, 1s, (1p+3s)*

0, 1p, (3p+1s)*

0, 1s, (2p+2s)*

0, 1s, (1p+3s)*

0, 1s, (1p+3s)

Telson

Furcae

spinulate 4sp (2 pairs)

smooth 6sp (3 pairs)

smooth 6sp (3 pairs)

spinulate 6sp (3 pair)

spinulate 6sp (3 pairs)

smooth 6sp (3 pairs)

spinulate 6sp (3 pairs)

spinulate 6sp (3 pairs)

smooth 6sp (3 pairs)

spinulate 6sp (3 pairs)

spinulate 6sp (3 pairs)

smooth 6sp (3 pairs)

smooth 4sp (2 pairs)

spinulate 6sp (3 pairs)

spinulate distally

Spine 2

absent

vestigial

vestigial

vestigial

vestigial

developed

developed

developed

developed

developed

vestigial

developed

vestigial

developed

?

Abdomen

Dorsolateral process on somites 4 and 5

absent

absent

absent

absent

absent

present

4 present 5 absent

absent

absent

absent

absente

present

absent

absent

?

• a = aesthetasc; s = simple seta; p = plumose seta; c = cuspidate seta; pd = plumodenticulate seta; sp = spine; spp = spinule; m = microtrichia. References: ¹Present study; ²Clark and Ng (2010); ³Clark and Galil (1988); ⁴Shikatani and Shokita (1990); ⁵Clark and Ng (2006); ⁶Clark and Guerao (2008); ⁷Al-Aidaroos (1992). ? The author did not mention the setae type. *Data from figures observation.

Considering the morphology of the first larval stage is unknown for most species of the superfamily Trapezioidea, new descriptions could contribute with knowledge of the relationship within this taxon. Thus, we provided here a detailed description of the first zoea of the crab D. acanthophora, hatched in the laboratory. Additionally, we reviewed and compared the larval morphology (zoea I) of Trapezioidea species (sensuNg et al., 2008) in a comparative table, in order to facilitate the identification of these larvae.

Material and Methods

Two ovigerous females of D. acanthophora were collected in September 2006 by scuba divers at Vitória Island (23°44'04"S 45°01'35"W) in the southeastern of the São Paulo State, Brazil. The crabs were kept isolated in aquaria filled with seawater from the collecting site, at constant temperature (24 ± 1°C), salinity (35 ± 1) and moderate aeration, until the larvae hatched. Newly hatched zoeae were preserved in a 1:1 mixture of 70% ethyl alcohol and glycerin. For detailed examination, the larvae and appendages were dissected under a Zeiss Stemi 200C trinocular stereomicroscope and prepared on semi-permanent slides with glycerin.

Drawings and measurements were done using a Leica DM750 microscope equipped with camera lucida. The illustrations and measurements were based on at least 10 specimens. The carapace length (CL) was considered from the eyes (base of the rostrum) to the posterolateral cephalothorax margin, and the rostrodorsal length (RDL) was considered from the tip of the rostral spine to the tip of the dorsal spine. The long terminal plumose natatory setae on distal exopod segments of the first and second maxillipeds were drawn truncated.

Larval description and terminology of setae are based on Clark et al. (1998) and Garm (2004). The parental females of D. acanthophora were deposited in the crustacean collection of the Museum of Zoology of the University of São Paulo (MZUSP-16705). The larvae of D. acanthophora were deposited in the scientific collection of the Marine Biology Laboratory of the University of Taubaté (UNITAU-201237).

Results

Description of first zoea of D. acanthophora (Figs. 1-3)

Dimensions: CL: 0.38 ± 0.01 mm; RDL: 0.76 ± 0.02 mm (n = 10).

Cephalothorax (Fig. 1a, b): dorsal spine smooth, gently curved backward distally, almost equal in length to rostral spine. Pair of posterodorsal simple seta near the basis of the dorsal spine, posterodorsal protuberance present. Three pairs of lateral spines: dorsal pair curved ventrally, with one ventral and one dorsal spinules; second pair smaller, smooth, slightly curved ventrally; third pair smallest and smooth near carapace margin. Sessile eyes.

Figure 1
Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867), Zoea I. a. Cephalothorax, frontal view; b. Cephalothorax, lateral view; c. Antennule; d. Antenna (scale bars: a, b = 0.1 mm; c, d = 0.05 mm).

Figure 2
Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867), Zoea I. a. Mandible; b. Maxillule; c. Maxilla; d. First maxilliped; e. Second maxilliped (scale bars: a-e = 0.05 mm).

Figure 3
Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867), Zoea I. a. Pleon, dorsal view; b. Pleon, lateral view; c. Detail of spinulation of the telson furcae (scale bars: a, b = 0.1 mm; c = 0.02 mm).

Antennule (Fig. 1c): uniramous, smooth. Endopod absent; exopod unsegmented, with three long terminal aesthetascs and one simple seta.

Antenna (Fig. 1d): biramous; well-developed and long protopod, with one row of short, equal spines in each lateral margin of similar sizes. Endopod absent. Exopod unsegmented, with three unequal terminal simple setae, the longest almost equal in length to exopod.

Mandibles (Fig. 2a): incisor and molar processes as illustrated; palp absent.

Maxillule (Fig. 2b): coxal endite with 1 (subterminal) simple and 5 (1 subterminal and 4 terminal) plumodenticulate setae. Basial endite with 5 plumodenticulate setae and 2 small protuberances, microtrichia on proximal margin. Endopod 2-segmented, proximal segment without seta; distal segment with 3 (1 subterminal, 2 terminal) sparsely plumodenticulate setae, 1 shorter subterminal simple seta. Exopod seta absent.

Maxilla (Fig. 2c): coxal endite bilobed, with 4 plumose setae on proximal lobe, 3 plumose setae and 1 terminal small spine on distal lobe. Basial endite bilobed, with 4 plumodenticulate setae on each lobe. Unsegmented endopod bilobed, with 2 (1 plumodenticulate, 1 simple) setae on proximal lobe, 3 (1 subterminal, 2 terminal) setae on distal lobe. Exopod (scaphognathite) margin with 4 plumose setae and a long distal process. Microtrichia on both proximal and distal margins of coxal and basial endites, endopod and distal process of the scaphognathite.

First maxilliped (Fig. 2d): coxa without setae. Basis with 9 simple setae arranged 2+2+3+2. Endopod 5-segmented with 1 simple seta on first segment; 2 (1 simple, 1 plumodenticulate) setae on second segment; no seta on third segment; 2 plumodenticulate setae on fourth segment; 4 (2 subterminal, 2 terminal) plumodenticulate setae and one subterminal simple seta on distal segment. Exopod slightly 2-segmented, distal segment with 4 long terminal plumose natatory setae.

Second maxilliped (Fig. 2e): coxa without setae. Basis with 4 simple setae arranged 1+1+1+1. Endopod 3-segmented, with 0, 0 and 4 (2 simple, 2 long sparsely plumodenticulate) setae, respectively. Exopod slightly 2-segmented, distal segment with 4 long terminal plumose natatory setae.

Third maxilliped: absent.

Pereiopods: absent.

Pleon (Fig. 3a, b): five somites. First somite smooth. Second to fifth somites with one pair of posterodorsal simple seta. Second and third somites with one pair of dorsolateral processes. Fourth and fifth somites with long and acute posterolateral processes; pleopods absent.

Telson (Fig. 3a-c): telson furcae distally spinulated and slightly curved outward, with 1 pair of well-developed lateral spines (spinulated only on the posterior margin), 1 pair of plumose setae posteriorly to the well-developed spines, and one pair of dorsal spines (spinulated only on the proximal inner margin); inner margin with 3 pairs of plumose setae.

Discussion

According to Clark and Ng (2010), the characters that separate the early stages of zoeae of the species of superfamily Trapezioidea are: the spinulation of the antennal protopod, the terminal setation of the antennule, the setation of the maxillule, maxilla, first and second maxillipeds, and medial and dorsolateral processes of the abdominal somites (see Tab. 1). The antennal morphology of D. acanthophora follows that observed for the other genera of the superfamily Trapezioidea (Tab. 1). However, the distal spinulation of the antennal protopod in these genera is variable. The antennal protopod and its distal spinulation of D. acanthophora is similar to that of Domecia glabraAlcock, 1899 and Calocarcinus africanusCalman, 1909, which is formed by two rows of similar size spines. Other trapezioids have a distinct spinulation of the antennal protopod: Quadrella maculosaAlcock, 1898 and Quadrella sereneiGalil, 1986 show an antennal protopod distally multispinulated (Clark and Ng, 2006); Tetralia cavimanaHeller, 1861 and Tetralia rubridactylaGarth, 1971 present two rows of variable size distal spines (Clark and Galil, 1988; Clark and Ng, 2006); Trapezia cymodoce (Herbst, 1799) and Trapezia richtersiGalil & Lewinsohn, 1983 present two rows of distal spines arranged sparsely (Clark and Galil, 1988; Shikatani and Shokita, 1990; Al-Aidaroos, 1992; Clark and Ng, 2006).

The morphology of the larval stages of decapod crustaceans is used in studies to evaluate phylogenetic questions (e.g. Marques et al., 2003; Barros-Alves et al., 2013; Guerao et al., 2014). According to phylogenetic analysis, Clark and Ng (2010) suggested that the genera Domecia, QuadrellaDana, 1851, Tetralia Dana, 1851 and TrapeziaLatreille, 1828 are nested in one clade that is defined by the absence of seta 3.5 and seta 3.4 on the third endopodal segment of the second maxilliped (for details, see Clark and Guerao, 2008), and should be according to a synapomorphy defines the Domecia + Tetralia + Trapezia clade: the presence of one subterminal seta on the distal endopod segment of the maxillule (vs. two subterminal setae). Other synapomorphies define the sister group Domecia + Tetralia, including: the absence of a subterminal seta on the distal endopod lobe of the maxilla; and the absence of dorsolateral process on abdominal somites fourth and fifth (Clark and Ng, 2010). However, D. acanthophora presents two subterminal seta on the distal endopod segment of the maxillule and one subterminal seta on the distal endopod lobe of the maxilla. Thus, these features contradict the synapomorphies mentioned above, since they differ from observed by Clark and Ng (2010).

As previously discussed by Clark and Guerao (2008)) and Clark and Ng (2010), CalocarcinusCalman, 1909 appears not to be related to the trapezioids. However, a similar morphological characteristic was observed for C. africanus and D. acanthophora: the presence of three pairs of lateral spines on the carapace. Therefore, the presence of this structure seems to be evidence that supports Calocarcinus relationship with the trapezioids (see Tab. 1), but further analysis is needed about this relationship.

Clark and Ng (2010) suggested, as autapomorphic features of D. glabra, the absence of setae on the proximal endopod segment of the maxillule, four terminal setae and one subterminal seta on the distal endopod segment of the maxillule, the basial setation of the first maxilliped arranged as 2, 2, 3, 2, the first endopod segment of the first maxilliped bearing only one seta, and the endopod of the second maxilliped with only four (two subterminal and two terminal) seta in the distal segment. These characters were also observed for the first zoea of D. acanthophora in this study, except the number of subterminal seta on the distal endopod segment of the maxillule (two for D. acanthophora vs. one for D. glabra). Thus, we suggest that other characters could be synapomorphies of the genus Domecia.

Some characters of the first zoeal stage that could be used to distinguish D. acanthophora and D. glabra are: the number of aesthetascs in the antennula exopod (three and four, respectively); the number of spines in the lateral carapace (three pairs and two pairs, respectively), wherein the lower pair is not ventrally deflected as in D. glabra; the number of setae in the basial endite of maxilla (4+4 and 5+3, respectively); the pairs of spines on the telson (two and three, respectively); and furcae distally spinulate in D. acanthophora and smooth in D. glabra (see Tab. 1).

The first zoeal stage of D. acanthophora exhibits some characters that differ from the other Trapezioidea species, such as the presence of simple setae at the basis of the first maxiliped (vs. plumose setae); the presence of only two pairs of spines on the telson furcae (vs. three pairs); and telson furcae with one pair of plumose setae posteriorly to the well-developed spines (vs. absent) (see Tab. 1). Thus, these characters can be considered as autapomorphy of D. acanthophora. Therefore, to prove this hypothesis, it is necessary further zoea I descriptions for the remaining species of the superfamily Trapezioidea.

The current study expands the number of species with first zoeal morphology known in Trapezioidea. Thus, we assert in favor of future studies to increase the number of larval descriptions, and to improve larval culture techniques in the laboratory, enabling the culture of larval stages for the species that the cultivation techniques currently used not guarantee survival and complete larval development, thereby enabling knowledge of a greater number of stages for larval species. This would certainly help to solve taxonomic and phylogenetic problems in the infraorder Brachyura.

Acknowledgments

The authors are indebted to Dr. Maria Lucia Negreiros-Fransozo, who gave us valuable suggestions for improving the manuscript. DFRA is grateful to CNPq - Conselho Nacional de Desenvolvimento Científico e Tecnológico (134950/2007-0) for his Master of Science fellowship. JAFP thanks Capes for a PhD scholarship. RCC is grateful to CNPq (PQ 305919/2014-8). We thank the "Omni Mare" Dive Center for the facilities provided during fieldwork. All sampling was conducted in accordance with applicable Brazilian state and federal laws with respect wild animals.

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• 1
  This article is part of the special series offered by the Brazilian Crustacean Society in honor to Nilton José Hebling in recognition of his dedication and contributions to the development of carcinology in Brazil.

• 2
  Guest Editors: Maria Lúcia Negreiros-Fransozo and Adilson Fransozo

• Zoobank:

  http://zoobank.org/urn:lsid:zoobank.org:pub:16FC6416-35D0-4147-9219-3246E57F55BF

Publication Dates

• Publication in this collection
  2016

History

• Received
  24 Feb 2016
• Accepted
  13 July 2016