This article is part of the special series Nauplius offered by the Brazilian Society the Journal of the in honor to Nilton José Hebling in recognition BraZIlIan crustacean socIety of his dedication and contributions to the development of carcinology in Brazil.

orIgInal artIcle e-ISSN 2358-2936 www.scielo.br/nau www.crustacea.org.br First zoeal stage of the Domecia acanthophora (Desbonne, in Desbonne & Schramm, 1867) (, Brachyura) and revision of the larval morphology of superfamily

Douglas Fernandes Rodrigues Alves1,4, João Alberto Farinelli Pantaleão2,4, Samara de P. Barros-Alves1,4, Rogério Caetano da Costa2,4 and Valter José Cobo3,4

1 Laboratório de Carcinologia, Universidade Federal de Sergipe, UFS. 49100-000 São Cristóvão, Sergipe, Brazil. DFRA E-mail: [email protected] sPBa E-mail: [email protected]

2 Laboratório de Biologia de Camarões Marinhos e de Água Doce – LABCAM, Universidade Estadual Paulista, UNESP. 17033-360 Bauru, São Paulo, Brazil. JafP E-mail: [email protected] rcc E-mail: [email protected]

3 Laboratório de Biologia Marinha – LABBMAR, Universidade de Taubaté, UNITAU. 12030-180 Taubaté, São Paulo, Brazil. VJc E-mail: [email protected]

4 Group of Studies on Crustacean Biology, Ecology and Culture – NEBECC, Departamento de Zoologia, Universidade Estadual Paulista, UNESP. 18618-970 Botucatu, São Paulo, Brazil.

ZooBanK htt p://zoobank.org/urn:lsid:zoobank.org:pub:16FC6416-35D0-4147- CORRESPONDING AUTHOR 9219-3246E57F55BF Samara de P. Barros-Alves [email protected]

SUBMITTED 24 February 2016 ACCEPTED 13 July 2016 aBstract PUBLISHED 21 November 2016 Th e morphology of the fi rst zoeal stage ofDomecia acanthophora (Desbonne, Guest Editors in Desbonne & Schramm, 1867) was described from laboratory-hatched Maria Lúcia Negreiros-Fransozo and Adilson Fransozo material obtained from ovigerous females collected at Vitória Island on the southeastern Brazilian coast. We compared the larval morphology

DOI 10.1590/2358-2936e2016021 Nauplius, 24 e2016021 1

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(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 .

Key words Domeciidae, larval description, southeastern Brazil, , .

Introduction Ng et al., 2008) in a comparative table, in order to The most recent Brachyura classification (Nget al., facilitate the identification of these larvae. 2008; De Grave et al., 2009; Ahyong et al., 2011) recognizes the superfamily Trapezioidea Miers, Material and methods 1886, which consists of three families: Domeciidae Two ovigerous females of D. acanthophora were Ortmann, 1893, Tetraliidae Castro, Ng & Ahyong, collected in September 2006 by scuba divers at Vitória 2004 and Trapeziidae Miers, 1886. Crabs of the family Island (23°44’04”S 45°01’35”W) in the southeastern of Domeciidae are grouped into four genera: Domecia the São Paulo State, Brazil. The crabs were kept isolated Eydoux & Souleyet, 1842, Jonesius Sankarankutty, in aquaria filled with seawater from the collecting site, 1962, Maldivia Borradaile, 1902 and Palmyria Galil at constant temperature (24 ± 1°C), salinity (35 ± 1) & Takeda, 1986. The speciesDomecia acanthophora and moderate aeration, until the larvae hatched. Newly (Desbonne, in Desbonne & Schramm, 1867) has a hatched zoeae were preserved in a 1:1 mixture of 70% wide geographical distribution in the Western Atlantic, ethyl alcohol and glycerin. For detailed examination, extending from North Carolina, Bermuda, Florida, the larvae and appendages were dissected under a Zeiss the Gulf of Mexico, the Antilles, northeastern South Stemi 200C trinocular stereomicroscope and prepared America and Brazil (St. Peter and St. Paul Archipelago, on semi-permanent slides with glycerin. Rocas Atoll, Fernando de Noronha, and from states Drawings and measurements were done using of Ceará and Pernambuco to São Paulo) (Alves et al., a Leica DM750 microscope equipped with camera 2006; Coelho-Filho, 2006; Melo, 1998). lucida. The illustrations and measurements were based In recent decades, several studies about the on at least 10 specimens. The carapace length (CL) morphology of larval stages have been used to elucidate the taxonomic status of these crabs, of which there was considered from the eyes (base of the rostrum) is still no consensus with respect to the superfamily to the posterolateral cephalothorax margin, and the Trapezioidea (e.g., Clark and Guerao, 2008; Clark rostrodorsal length (RDL) was considered from the and Ng, 2010). About 60 species of the superfamily tip of the rostral spine to the tip of the dorsal spine. Trapezioidea are known (see Ng et al., 2008), but the The long terminal plumose natatory setae on distal morphology of the first zoeal stage is described only exopod segments of the first and second maxillipeds for 14 species (see Tab. 1). were drawn truncated. Considering the morphology of the first larval Larval description and terminology of setae are stage is unknown for most species of the superfamily based on Clark et al. (1998) and Garm (2004). The Trapezioidea, new descriptions could contribute with parental females of D. acanthophora were deposited in knowledge of the relationship within this taxon. Thus, the crustacean collection of the Museum of Zoology we provided here a detailed description of the first zoea of the University of São Paulo (MZUSP-16705). of the crab D. acanthophora, hatched in the laboratory. The larvae ofD. acanthophora were deposited in the Additionally, we reviewed and compared the larval scientific collection of the Marine Biology Laboratory morphology (zoea I) of Trapezioidea species (sensu of the University of Taubaté (UNITAU-201237).

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Diagramação e XML SciELO Publishing Schema: www.editoraletra1.com Alves et al. Description of the first zoea of D. acanthophora 7 ? ? 3s no seta 2a+3s? (4+1)? (3+2)p (4+4)p (5+3)p (2+5)p (3p+2c) 1, (1+4)p Eydoux & Eydoux Clark and and Clark (spinulate) 2sp (1 pair) 2sp (1 pair) (2+2+3+3)? 2?, 1?, (1+1+1+0)p 3 1sp (smooth) 0, 1s, (1p+3s) 1sp (spinulate) Trapezia tigrina tigrina Trapezia Souleyet, 1842 Souleyet, spinulate distally spinulate spinulate distally spinulate 4 ? 3s 2a+2s absent (3+2)p* (4+4)p* (5+3)p* (2+5)p* spinulate (4p+1s)* (3p+2c)* developed (spinulate) 2sp (1 pair) 2sp (1 pair) Dana, 1852 Dana, 2s, 2s, 1s, 2p, 2s, 1s, 2p, 6sp (3 pairs) (1+1+1+1)p (2+2+3+3)p 1s*, (2p+3s)* 1sp (smooth) 0, 1s, (1p+3s)* 1sp (spinulate) septata Trapezia spinulate distally spinulate 4

? 3s Clark and Ng (2010); Ng and Clark 2a+3s absent 2 smooth vestigial (3+2)p* (5+4)p* (5+3)p* (2+5)p* Trapezia Trapezia (4p+1s)* (4p+1s)* (3p+2c)* (spinulate) 2sp (1 pair) 2sp (1 pair) 2s, 1s, rufopunctata 1s, (1s+4p)* 4sp (2 pairs) (1+1+1+0)p (2+2+3+2)p 1sp (smooth) 0, 1s, (2p+2s)* 1sp (spinulate) (Herbst, 1799) (Herbst, spinulate distally spinulate 5 and other species of the superfamily species other and 3s 1p* 4a+1s Galil & present smooth (3+2)p* (2+5)p* spinulate spinulate (4p+1s)* (4c+1p)* bilaterally developed (spinulate) 2sp (1 pair) 2sp (1 pair) 1p, (1+4)p* 1p, Present study; Present 6sp (3 pairs) (4+4)p*, 1sp (5+3)p*, 1sp 2p, 2p, 1p, 2p, 2p, 1p, 2p, 2p, 1sp (smooth) (1+1+1+0)p* (2+2+3+3)p* 1 0, 1p, (3p+1s)* 0, 1p, 1sp (spinulate) Trapezia richtersi Trapezia Lewinsohn, 1983 4,7 Trapeziinae Miers, 1886 Miers, Trapeziinae ? 3s 2a+3s absente vestigial (3+2)p* (4+4)p* (5+3)p* (2+5)p* spinulate (4p+1s)* (3p+2c)* (spinulate) 2sp (1 pair) 2sp (1 pair) 1s*, (1+4)p* 6sp (3 pairs) (1+1+1+0)p 2s, 2s, 1p, 2p, 2p, 2s, 1p, 1sp (smooth) (2+2+3+3)p* 0, 1s, (1p+3s)* 1sp (spinulate) Trapezia digitalis digitalis Trapezia spinulate distally spinulate Latreille, 1828) Latreille, 3,4,7 ? 3s Trapeziidae Miers, 1886 Miers, Trapeziidae absent (4+1)s* (2+5)p* spinulate (3c+2p)* developed (3-4+2)p* 4a+1s, 1sp (spinulate) 2sp (1 pair) 2sp (1 pair) 2s, 1s, 1s*, (3p+2s) 6sp (3 pairs) (1+1+1+1)p (2+2+3+3)p (4+4)p*, 1sp 0, 1s, (3p+1s)* (5-6+3)p*, 1sp 1sp (spinulate) 1sp (spinulate) spinulate distally spinulate (Herbst, 1801) (Herbst, Trapezia cymodoce cymodoce Trapezia 4,7 ? 3s 3a+1s absent smooth (3+2)p* (4+4)p* (5+3)p* (2+5)p* (4p+1s)* (3p+2c)* developed (spinulate) 2sp (1 pair) 2sp (1 pair) 1s*, (1+4)s* 2s, 1s, 6sp (3 pairs) 1sp (smooth) (1+1+1+0)p* 0, 1s, (1c+2s)* 1sp (spinulate) spinulate distally spinulate (2s+2s+3p+2p)* (Forskål, 1775) (Forskål, bidentata Trapezia 5 (Desbonne, in Desbonne & Schramm, 1867) & Schramm, in Desbonne (Desbonne, 3s 4a+1s absent no seta distally (3+5)p* (5+4)p* (2+5)p* spinulate (4p+1s)* (4c+1p)* (smooth) developed Galil, 1986 1p, (2+4)p* 1p, 4sp (2 pairs) 4sp (2 pairs) 6sp (3 pairs) (5+3)p*, 1sp 3p, 2p, 1p, 2p, 2p, 1p, 2p, 3p, 1sp (smooth) 1sp (smooth) (1+1+1+1)p* (2+2+3+3)p* multispinulate multispinulate 1p, 1p, (3p+1s)* 1p, 1p, serenei Quadrella 5 Quadrellinae Števčić, 2005 Števčić, Al-Aidaroos (1992). ? The author did not mention the setae type. * Data from figures observation.figures from Data type.setae the author * did not mention (1992). ? The Al-Aidaroos 7 4a+1s no seta distally 5 absent (3+5)p* (5+4)p* (2+5)p* 3s+1spp maculosa spinulate (4p+1s)* 4 present (4c+1p)* Quadrella Quadrella developed 1p, (2+4)p* 1p, 4sp (2 pairs) 4sp (2 pairs) 6sp (3 pairs) (5+3)p*, 1sp 3p, 2p, 1p, 2p, 2p, 1p, 2p, 3p, (1+1+1+1)p* (2+2+3+3)p* Alcock, 1898 multispinulate multispinulate 1sp (spinulate) 1sp (spinulate) 1p, 1p, (2s+2p)* 1p, 1p, (with 1 spinulate) 6 Domecia acanthophora 3s 1p* 1p, 1p, 1p, 1p, 5a+1s* present smooth (3+5)p* (5+4)p* (6+4)p* (1+6)p* africanus (smooth) (smooth) developed (3pd+2c)* and distaly and (2s,1p+3s)* 2p, (4+1)p* 2p, Calocarcinus Calocarcinus 6sp (3 pairs) 6sp (3 pairs) 6sp (3 pairs) Števčić, 2005 Števčić, 1sp (smooth) 1sp (smooth) (1+1+1+1)p* (2+2+3+3)p* Calman, 1909 Calman, Calocarcininae Calocarcininae 1pd*, (2+4)pd* (1s+2p), 2p, 1p, 1p, 2p, (1s+2p), spinulate bilateral bilateral spinulate 5 3s Clark and Guerao (2008); Guerao and Clark 1p* 6 4a+1s absent Tetralia Tetralia vestigial (2+3)p* (2+5)p* (4c+1s)* spinulate spinulate spinulate (4p+1s)* bilaterally bilaterally 1s, (1+4)p* rubridactyla 1 spinulate) 6sp (3 pairs) Garth, 1971 (4+3)p*, 1sp 3p, 2p, 1p, 2p, 2p, 1p, 2p, 3p, (1+1+1+1)p* (2+2+3+3)p* (4p, 1sp+4p)* (4p, (variable size) (variable 1sp (spinulate) 1sp (spinulate) 1p, 1p, (2p+2s)* 1p, 1p, 4sp (2 pairs) (with 4sp (2 pairs) 4 ? 3s absent 2a+2s* (2+5)p vestigial (2+3)p* (4+3)p* (4+4)p* (5p+2c) spinulate spinulate spinulate (4p+1s)* bilaterally 1p, (1+4)p 1p, 6sp (3 pair) 2p,2p,1s, 2s, 2p,2p,1s, (spinulates) 4sp (2 pairs) 4sp (2 pairs) (1+1+1+1)s* (2+2+3+3)p* 1s,1s, (2p+2s)* 1 sp (spinulate) 1 sp (spinulate) (Herbst, 1790) (Herbst, Tetralia glaberrima glaberrima Tetralia 3 Clark and Ng (2006); Ng and Clark 5 ? Tetrallidae Castro, Ng & Ahyong, 2004 Ng Castro, Tetrallidae 3s 1s*, 4? absent (4+1)? (2+3)? (4+4)? (2+5)? smooth vestigial spinulate spinulate (4c*+1?) bilaterally bilaterally 4a+1s, 1sp (4+3)?, 1sp 1 spinulate) (1+1+1+1)? 6sp (3 pairs) (2+2+3+3)? 2?, 1?, Heller, 1861 Heller, 1?, (1+3)? 1sp (spinulate) 1sp (spinulate) Tetralia cavimana cavimana Tetralia 4sp (2 pairs) (with 4sp (2 pairs) 2 3s 1p* 0, 4p* absent smooth 4a+1-2s vestigial (4+1)p* (3+2)p* (5+3)p* (2+5)p* spinulate spinulate (3c+2p)* bilaterally 1p, 2p, 0, 2p, 0, 2p, 2p, 1p, 6sp (3 pairs) 4sp (2 pairs) 4sp (2 pairs) (4+3)p*, 1sp 0, (2+2)p* 1sp (smooth) 1sp (smooth) (1+1+1+1)p* (2+2+3+2)p* Alcock, 1899 glabra Domecia (with 1 spinulate) 1

3s 3a+1s absent absent no seta Domecia spinulate spinulate spinulate Domeciidae Ortmann, 1893 Domeciidae (1s+5pd) bilaterally (4+4)pd, m 0, (3pd+1s) 0, (2+2)p (1+1+1+1)s (2+2+3+2)s 4sp (2 pairs) 2 spinulates) Desbonne & Desbonne (2c+3pd), m (2c+3pd), acanthophora (4+3)pd, 1sp 1sp (smooth) 1sp (smooth) 2pd, (2+2)pd (Desbonne, in (Desbonne, Shikatani and Shokita (1990); Shokita and Shikatani 1s, (1s+1pd), 0, 1s, (1s+1pd), 4 (1pd,1s+2pd, 1s) Schramm, 1867) Schramm, 6sp (3 pairs) (with 6sp (3 pairs) Comparison of some larval characters of the first zoeal stage of stage zoeal of the first larval of some Comparison characters

Furcae Endopod Basis Endopod Basis Coxa Endopod Basial endite Basial Coxal endite Coxal Endopod Basial endite Basial Coxal endite Coxal Protopod Dorsolateral on process 5 4 and somites Exopod Exopod Spine 2 Spine Lateral Dorsal Rostral Telson Second Second maxilliped First maxilliped First Maxilla Maxillule Abdomen Antenna Antennule Galil (1988); Table 1. Table Species/original larval description Carapace microtrichia.spine;References: = = m sp spinule; = spp seta; plumodenticulate = pd seta; cuspidate = c seta; plumose = p seta; simple = s aesthetasc; = a Trapezioidea.

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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).

Results gently curved backward distally, almost equal in Description of first zoea ofD. acanthophora (Figs. 1–3). length to rostral spine. Pair of posterodorsal simple Dimensions: CL: 0.38 ± 0.01 mm; RDL: 0.76 ± seta near the basis of the dorsal spine, posterodorsal 0.02 mm (n = 10). protuberance present. Three pairs of lateral spines: Cephalothorax (Fig. 1a, b): dorsal spine smooth, dorsal pair curved ventrally, with one ventral and one

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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).

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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). dorsal spinules; second pair smaller, smooth, slightly segment with 3 (1 subterminal, 2 terminal) sparsely curved ventrally; third pair smallest and smooth near plumodenticulate setae, 1 shorter subterminal simple carapace margin. Sessile eyes. seta. Exopod seta absent. Antennule (Fig. 1c): uniramous, smooth. Endopod Maxilla (Fig. 2c): coxal endite bilobed, with 4 absent; exopod unsegmented, with three long terminal plumose setae on proximal lobe, 3 plumose setae aesthetascs and one simple seta. and 1 terminal small spine on distal lobe. Basial Antenna (Fig. 1d): biramous; well-developed and endite bilobed, with 4 plumodenticulate setae on long protopod, with one row of short, equal spines in each lobe. Unsegmented endopod bilobed, with 2 (1 each lateral margin of similar sizes. Endopod absent. plumodenticulate, 1 simple) setae on proximal lobe, 3 Exopod unsegmented, with three unequal terminal (1 subterminal, 2 terminal) setae on distal lobe. Exopod simple setae, the longest almost equal in length to (scaphognathite) margin with 4 plumose setae and a exopod. long distal process. Microtrichia on both proximal and Mandibles (Fig. 2a): incisor and molar processes distal margins of coxal and basial endites, endopod and as illustrated; palp absent. distal process of the scaphognathite. Maxillule (Fig. 2b): coxal endite with 1 First maxilliped (Fig. 2d): coxa without setae. Basis (subterminal) simple and 5 (1 subterminal and 4 with 9 simple setae arranged 2+2+3+2. Endopod terminal) plumodenticulate setae. Basial endite with 5-segmented with 1 simple seta on first segment; 2 (1 5 plumodenticulate setae and 2 small protuberances, simple, 1 plumodenticulate) setae on second segment; microtrichia on proximal margin. Endopod no seta on third segment; 2 plumodenticulate setae 2-segmented, proximal segment without seta; distal on fourth segment; 4 (2 subterminal, 2 terminal)

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Diagramação e XML SciELO Publishing Schema: www.editoraletra1.com Alves et al. Description of the first zoea of D. acanthophora plumodenticulate setae and one subterminal simple rubridactyla Garth, 1971 present two rows of variable seta on distal segment. Exopod slightly 2-segmented, size distal spines (Clark and Galil, 1988; Clark and Ng, distal segment with 4 long terminal plumose natatory 2006); Trapezia cymodoce (Herbst, 1799) and Trapezia setae. richtersi Galil & Lewinsohn, 1983 present two rows of Second maxilliped (Fig. 2e): coxa without setae. distal spines arranged sparsely (Clark and Galil, 1988; Basis with 4 simple setae arranged 1+1+1+1. Endopod Shikatani and Shokita, 1990; Al-Aidaroos, 1992; Clark 3-segmented, with 0, 0 and 4 (2 simple, 2 long sparsely and Ng, 2006). plumodenticulate) setae, respectively. Exopod slightly The morphology of the larval stages of decapod 2-segmented, distal segment with 4 long terminal is used in studies to evaluate phylogenetic plumose natatory setae. questions (e.g. Marques et al., 2003; Barros-Alves et al., Third maxilliped: absent. 2013; Guerao et al., 2014). According to phylogenetic Pereiopods: absent. analysis, Clark and Ng (2010) suggested that the genera Pleon (Fig. 3a, b): five somites. First somite smooth. Domecia, Quadrella Dana, 1851, Tetralia Dana, 1851 Second to fifth somites with one pair of posterodorsal and Trapezia Latreille, 1828 are nested in one clade simple seta. Second and third somites with one pair that is defined by the absence of seta 3.5 and seta of dorsolateral processes. Fourth and fifth somites 3.4 on the third endopodal segment of the second with long and acute posterolateral processes; pleopods maxilliped (for details, see Clark and Guerao, 2008), absent. and should be according to a synapomorphy defines Telson (Fig. 3a–c): telson furcae distally spinulated the Domecia + Tetralia + Trapezia clade: the presence and slightly curved outward, with 1 pair of well- of one subterminal seta on the distal endopod segment developed lateral spines (spinulated only on the of the maxillule (vs. two subterminal setae). Other posterior margin), 1 pair of plumose setae posteriorly synapomorphies define the sister groupDomecia + to the well-developed spines, and one pair of dorsal Tetralia, including: the absence of a subterminal seta on spines (spinulated only on the proximal inner margin); the distal endopod lobe of the maxilla; and the absence inner margin with 3 pairs of plumose setae. of dorsolateral process on abdominal somites fourth and fifth (Clark and Ng, 2010). However,D. acanthophora Discussion presents two subterminal seta on the distal endopod According to Clark and Ng (2010), the characters segment of the maxillule and one subterminal seta on that separate the early stages of zoeae of the species the distal endopod lobe of the maxilla. Thus, these of superfamily Trapezioidea are: the spinulation of features contradict the synapomorphies mentioned the antennal protopod, the terminal setation of the above, since they differ from observed by Clark and antennule, the setation of the maxillule, maxilla, first Ng (2010). and second maxillipeds, and medial and dorsolateral As previously discussed by Clark and Guerao (2008) processes of the abdominal somites (see Tab. 1). The and Clark and Ng (2010), Calocarcinus Calman, 1909 antennal morphology of D. acanthophora follows appears not to be related to the trapezioids. However, a that observed for the other genera of the superfamily similar morphological characteristic was observed for Trapezioidea (Tab. 1). However, the distal spinulation C. africanus and D. acanthophora: the presence of three of the antennal protopod in these genera is variable. pairs of lateral spines on the carapace. Therefore, the The antennal protopod and its distal spinulation of presence of this structure seems to be evidence that D. acanthophora is similar to that of Domecia glabra supports Calocarcinus relationship with the trapezioids Alcock, 1899 and Calocarcinus africanus Calman, 1909, (see Tab. 1), but further analysis is needed about this which is formed by two rows of similar size spines. relationship. Other trapezioids have a distinct spinulation of the Clark and Ng (2010) suggested, as autapomorphic antennal protopod: Quadrella maculosa Alcock, 1898 features of D. glabra, the absence of setae on the and Quadrella serenei Galil, 1986 show an antennal proximal endopod segment of the maxillule, four protopod distally multispinulated (Clark and Ng, terminal setae and one subterminal seta on the distal 2006); Tetralia cavimana Heller, 1861 and Tetralia endopod segment of the maxillule, the basial setation

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Diagramação e XML SciELO Publishing Schema: www.editoraletra1.com Alves et al. Description of the first zoea of D. acanthophora of the first maxilliped arranged as 2, 2, 3, 2, the first Acknowledgements endopod segment of the first maxilliped bearing only The authors are indebted to Dr. Maria Lucia Negreiros- one seta, and the endopod of the second maxilliped Fransozo, who gave us valuable suggestions for with only four (two subterminal and two terminal) improving the manuscript. DFRA is grateful to CNPq seta in the distal segment. These characters were also - Conselho Nacional de Desenvolvimento Científico e observed for the first zoea ofD. acanthophora in this Tecnológico (134950/2007-0) for his Master of Science study, except the number of subterminal seta on the fellowship. JAFP thanks Capes for a PhD scholarship. distal endopod segment of the maxillule (two for D. RCC is grateful to CNPq (PQ 305919/2014-8). We acanthophora vs. one for D. glabra). Thus, we suggest thank the “Omni Mare” Dive Center for the facilities that other characters could be synapomorphies of the provided during fieldwork. All sampling was conducted genus Domecia. in accordance with applicable Brazilian state and federal Some characters of the first zoeal stage that could laws with respect wild . be used to distinguish D. acanthophora and D. glabra are: the number of aesthetascs in the antennula exopod References (three and four, respectively); the number of spines Ahyong, S.H.; Lowry, J.S.; Alonso, M.; Bamber, R.N.; Boxshall, in the lateral carapace (three pairs and two pairs, G.A.; Castro, P.; Gerhen, S.; Karaman, G.S.; Goy, J.W.; Jones, D.S.; Meland, K.; Rogers, C. and Svavarsson, J. 2011. 2011. respectively), wherein the lower pair is not ventrally Subphylum Crustacea Brünnich, 1772. In: Z.Q. Zhang (ed), deflected as inD. glabra; the number of setae in the basial biodiversity: An outline of higher-level classification endite of maxilla (4+4 and 5+3, respectively); the pairs and survey of taxonomic richness. Zootaxa, 3148: 165–191. of spines on the telson (two and three, respectively); Al-Aidaroos, A. 1992. The first zoeal stage of four species of and furcae distally spinulate in D. acanthophora and Trapezia (Crustacea, Brachyura, Xanthidae). Journal of King Abdulaziz University Marine Science, 3: 57–66. smooth in D. glabra (see Tab. 1). Alcock, A. 1898. Materials for a carcinological fauna of India. No. The first zoeal stage of D. acanthophora exhibits 3. The Brachyura Cyclometopa. Part I. The family Xanthidae. some characters that differ from the other Trapezioidea Journal of the Asiatic Society of Bengal, 67(2): 67–233. species, such as the presence of simple setae at the Alcock, A. 1899. Natural History notes from the Royal Indian basis of the first maxiliped vs( . plumose setae); the Marine Survey Ship ‘Investigator,’ Commander T. H. Heming, presence of only two pairs of spines on the telson furcae R. N., commanding. Series III. No. 3. On some notable new and rare species of Crustacea. Journal of the Asiatic Society of (vs. three pairs); and telson furcae with one pair of Bengal, 68(2): 111–119. plumose setae posteriorly to the well-developed spines Alves, D.F.R.; Cobo, V.J. and Melo, G.A.S. 2006. Extension of the (vs. absent) (see Tab. 1). Thus, these characters can geographical distribution of some brachyuran and porcellanid be considered as autapomorphy of D. acanthophora. decapods (Crustacea) to the coast of the State of São Paulo, Therefore, to prove this hypothesis, it is necessary Brazil. Revista Brasileira de Zoologia, 23(4): 280–283. further zoea I descriptions for the remaining species Barros-Alves, S.P.; Alves, D.F.R.; Bolla, E.A.; Cobo, V.J. and Negreiros-Fransozo, M.L. 2013. First zoeal stage of of the superfamily Trapezioidea. Cataleptodius parvulus (Fabricius, 1793) and Xanthodius The current study expands the number of species denticulatus (White, 1848) (Decapoda: Brachyura): larval with first zoeal morphology known in Trapezioidea. evidences and systematic position. Zootaxa, 3731(2): 234– Thus, we assert in favor of future studies to increase 242. the number of larval descriptions, and to improve Borradaile, L.A. 1902. Marine Crustaceans III. The Xanthidae and some other crabs. In: J. Stanley Gardiner (ed) The Fauna and larval culture techniques in the laboratory, enabling geography of the Maldive and Laccadive Archipelagoes being the culture of larval stages for the species that the the account of the work carried on and the collections made cultivation techniques currently used not guarantee by an expedition during the years 1899–1900. Cambridge survival and complete larval development, thereby University Press, Cambridge 1: 237–271. enabling knowledge of a greater number of stages Calman, W.T. 1909. On a new crab taken from a deep-sea telegraph-cable in the Indian Ocean. Annals and Magazine for larval species. This would certainly help to solve of Natural History, 3(13): 30–33. taxonomic and phylogenetic problems in the infraorder Castro, P.; Ng, P.K.L. and Ahyong, S.T. 2004. Phylogeny and Brachyura. systematics of the Trapeziidae Miers, 1886 (Crustacea:

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Brachyura), with the description of a new family. Zootaxa, Carsten Niebuhr. Adjuncta est materia Medica Kahirina. 643: 1–70. Hauniae, Heineck et Faber, 164p. Clark, P.F.; Calazans, D.D. and Pohle, G.W. 1998. Accuracy and Galil, B.S. 1986. Quadrella (Brachyura: : Trapeziidae) standardization of brachyuran larval descriptions. Invertebrate - review and revision. Journal of Crustacean Biology, 6(2): Reproduction and Development, 33(2–3): 127–144. 275–293. Clark, P.F. and Galil, B.S. 1988. Redescription of Tetralia cavimana Galil, B.S. and Lewinsohn, C. 1983. Researches on the coast Heller, 1861 and Trapezia cymodoce (Herbst, 1799) first of Somalia. Trapezia richtersi n. sp., a new trapezid crab stage zoeas with implications for classification within the (Decapoda Brachyura). Monitore Zoologico Italiano, 18(1): superfamily Xanthoidea (Crustacea: Brachyura). Proceedings 159–166. of the Biological Society of Washington, 101(4): 853–860. Galil, B.S. and Takeda, M. 1986. Resurrection of the genus Jonesius Clark, P.F. and Guerao, G. 2008. A description of Calocarcinus and establishment of a new genus; commensal crabs associated africanus Calman, 1909 (Brachyura, Xanthoidea) first with from the Indo-Pacific.Bulletin of the National zoeal stage morphology with implications for Trapeziidae Science Museum Tokyo ser. A, 12: 163–171. systematics. Proceedings of the Biological Society of Washington, Garm A. 2004. Revising the definition of the crustacean seta 121(4): 475–500. and setal classification systems based on examinations of Clark, P.F. and Ng, P.K.L. 2006. First stage zoeas of Quadrella the mouthpart setae of seven species of decapods. Zoological Dana, 1851 [Crustacea: Decapoda: Brachyura: Xanthoidea: Journal of the Linnean Society, 142(2): 233–252. Trapeziidae] and their affinities with those ofTetralia Dana, Garth, J.S. 1971 [1969]. Borradaile’s Maldivian collections 1851, and Trapezia Latreille, 1828. Hydrobiologia, 560(1): revisited. 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