J. Mar. Biol. Ass. U.K. (2006), 86, 1123^1132 Printed in the United Kingdom
Larval development of the eastern Paci¢c anomuran crab Porcellana cancrisocialis (Crustacea: Decapoda: Anomura: Porcellanidae) described from laboratory reared material Marcelo U. Garcia-Guerrero*O, Antonio Rodr|¤ guezP and Michel E. Hendrickx*$ *Unidad Acade¤mica Mazatla¤n, Instituto de Ciencias del Mar y Limnolog|¤ a, Universidad Nacional Auto¤noma de Me¤xico, PO Box 811, Mazatla¤n, Sinaloa, 82000 Me¤xico. OCentro Interdisciplinario de Investigacio¤n para el Dessarrollo Integral Regional ^ Instituto Polite¤cnico Nacional, PO Box 280, Guasave, Sinaloa, 81000, Mexico. PInstituto de Ciencias Marinas de Andaluc|¤ a (CSIC), Avenida Repu¤blica Saharaui, 2, E-11510 Puerto Real, Ca¤diz, Spain. $ Corresponding author, e-mail:
[email protected]
The two zoeal and megalopal stages of Porcellana cancrisocialis are described and illustrated from laboratory-reared larvae. At an average temperature of 258C, Zoea I of P. cancrisocialis lasted seven days before moulting to Zoea II which, in turn, lasted eight days to reach the megalopa. Larval features of P. cancrisocialis are compared with those of other species of Porcellana. Porcellana cancrisocialis exhibits similar but distinguishable features from those observed in Porcellana sigsbeiana and Porcellana platycheles. This species resembles P. sigsbeiana more than P. platycheles in the setation and aesthetasc pattern of Zoea I and II, but the relationships of these three species are not clear.
INTRODUCTION Porcellanid zoeas are easily distinguished from other decapod zoeas because of their elongated carapace, rostral spine and paired posterior spines. Among porcelain crabs, Petrolisthes larvae have been extensively studied, with at least 29 species developments partially or totally described (Garc|¤ a-Guerrero et al., 2005). In contrast, only a few larval stages of Porcellana species have so far been described. The complete larval development of Porcellana sigsbeiana A. Milne-Edwards, 1880, was described by Gore (1971), that of Porcellana gravelei Sankolli, 1963, by Shenoy & Sankolli (1977), and that of Porcellana platycheles (Pennant, 1777), the type species of the genus, was redescribed by Gonza¤les-Gordillo et al. (1996) who reviewed previous descriptions of the larvae of this species. In addition, ¢rst zoeal stages of two other species of Porcellana from the Indo-West Paci¢c (Porcellana ornata Stimpson, 1858) and the western Atlantic [Porcellana sayana (Leach, 1820)] was described by Sankolli (1967) and Brooks & Wilson (1881), respectively. Porcellana ornata was later separated from the genus Porcellana and is now included as the type species in the genus Enosteoides, endemic to the Indo-West Paci¢c (Haig, 1978). Haig (1978) reviewed the heterogeneous grouping of species assigned to Porcellana and redistributed nine species into four new genera. Consequently, the genus Porcellana was restricted at that time to 11 species (and two subspecies). The addition of Porcellana lillyae Lemaitre & Campos, 2000, from the Caribbean coast of Colombia and the synonymization of Porcellana paivacarvalhoi Rodrigues da Costa, 1968 (from the west Atlantic) with P. platycheles (see Veloso & Melo, 1993; Lemaitre & Campos, 2000) has maintained the number of the American known species of the genus to seven, of which three are presently known from the eastern Paci¢c: Porcellana cancrisocialis Journal of the Marine Biological Association of the United Kingdom (2006)
Glassell, 1936; Porcellana paguriconviva Glassell, 1936; and Porcellana hancocki Glassell, 1938. The former two species are by far the most abundantly reported in their distribution area (Haig, 1960). Porcellana cancrisocialis is known from San Juanico Bay, western coast of Baja California, and throughout the Gulf of California, Mexico, to Tumbes, Peru (Hendrickx & Harvey, 1999). As indicated by the speci¢c name, it is often found to be associated with ‘crabs’, usually with the hermit crab Petrochirus californiensis Bouvier, 1895, and less frequently with another species of hermit crab, Dardanus sinistripes (Stimpson, 1859). However, large series of specimens collected at stations where no commensalism was noted indicates that juveniles and adults of Porcellana cancrisocialis are probably freeliving most of the time (Haig, 1960; Brusca, 1980). This paper describes in detail on the morphology of the larval stages of P. cancrisocialis and compares the features with those of the congeneric species for which larval developments are known.
MATERIALS AND METHODS A berried female of Porcellana cancrisocialis was found in the shell of the hermit crab Petrochirus californiensis Bouvier, 1895 in February 2005. The hermit crab was collected by a gill-net in the entrance of Mazatla¤n Harbour at about 18 m depth. The porcellanid was taken to laboratory facilities and maintained in a seawater-¢lled 3-l jar until hatching of larvae. After two days, approximately 120 larvae hatched and were separated in two equal groups. Every group was placed in a 12-l £at-tip conical tank maintained at ambient temperature 24^268C 0.58C. A third of the water volume was changed daily and larvae were fed through development with newly hatched Artemia nauplii. Moulting of the larvae in tanks was
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recorded for determining the duration of each stage. Prezoea were not considered since this pre-stage does not have distinctive taxonomic features. Samples of ten larvae of each stage per tank were ¢xed in 4% formaldehyde and then preserved in 70% ethanol. The carapace of both zoeal stages and megalopa were measured following the standard method proposed for porcelanid larvae by Herna¤ndez et al. (2000). All measurements are expressed as the arithmetic average (and corresponding standard variation) taken during this study. The following abbreviations were used: CL, carapace length; RSL, rostral spine length; PSL, length of the posterior spines of the carapace; CW, carapace width. Descriptions and measurements were based on ten specimens for Zoea I and II, and eight specimens for megalopa. Measurements were made with an ocular micrometer. Drawings were made using a Leica Ortholux compound microscope equipped with a camera lucida. The long natatory setae on the exopod distal segments of the ¢rst and second maxillipeds were drawn truncated. Body somites are described from anterior to posterior, appendages from endopod to exopod, and segments and setae from proximal to distal. The spent female and complete larval series are deposited in the crustacean reference collection (Coleccio¤n de Referencia de Crusta¤ceos) at Universidad Nacional Auto¤noma de Me¤xico, Mazatla¤n, Sinaloa, Mexico, under the voucher no. EMU-2677.
RESULTS The larvae of Porcellana cancrisocialis passed through two zoeal and one megalopal stages. At the experimental temperature, larvae needed an average of seven days from hatching to reach the second zoea, and eight days to complete the second zoea and to reach the megalopa stage.
Maxilla (Figure 3D). Coxal endite bilobed, with 6+6 plumodenticulate setae. Basial endite bilobed, with 5+9 plumodenticulate setae. Endopod unsegmented, with 3+2+4 sparsely plumose setae; scaphognathite (exopod) with seven or eight plumose marginal setae, long setose posterior process, series of setules present each on mesial and lateral margins of posterior lobe. First maxilliped (Figure 4A). Biramous. Coxa with two ventral setae; basis with eight ventral, sparsely plumose setae arranged 1+2+2+3, ventroproximal angle sharply pointed; endopod 4-segmented, segments 1^3 with 2, 3, 3 sparsely setose setae ventrally, segment 4 with nine plumodenticulate distal setae and a long plumose seta on proximodorsal margin; exopod 2-segmented, distal segment with four terminal, plumose natatory setae. Second maxilliped (Figure 4D). Biramous. Coxa naked; basis with three plumodenticulate setae arranged 1+2; endopod 4-segmented, segments 1^3 with 2, 2, 3 sparsely plumose setae, respectively, segment 4 with 2+2+4 distal plumodenticulate setae and one long plumose seta at dorsoproximal angle. Third maxilliped (Figure 5K). Biramous. Small unsegmented buds. Pereiopods. Present as small buds. Abdomen (Figure 5A). Five somites, ¢fth somite longest; distinct posterolateral spines on somites 2^5, pleopods absent. Telson (Figure 5A). Slightly longer than broad; posterior margin produced into broad trapezoid with ¢ve pairs of long, stout, plumose setae, bearing hooklets distally; posteromedial region narrow, concave, bearing a pair of short, naked setae; lateral angles each with stout spine and a short plumose setae at mesial base of spine; one pair of short, simple setae dorsally; anal spine present on ventral surface. Zoea II
Porcellana cancrisocialis Glassell, 1936 (Figures 1^5) Zoea I
Size. Carapace length 1.44 to 1.72 mm (mean 1.45 mm, N¼10); RSL 4.36 to 5.04 mm (mean 4.7 mm, N¼18); PSL 0.96 to 1.24 mm (mean 1.1mm, N¼10). Carapace (Figure 1A). Typical porcellanid zoea. Elongated rostral spine about 4^5 times carapace length, armed with rows of tiny spinules along entire length. Two posterior spines, each about as long as carapace, with a single row of ventral spines on proximal half. Antennule (Figure 2A). Exopod unsegmented, with ¢ve aesthetascs and one short seta distally. Endopod absent. Antenna (Figure 2D). Biramous. Basis with a subdistal seta; endopod with apex sharply pointed and a subterminal long plumose seta. Exopod tapering, slightly longer than endopod, with a subterminal simple seta. Mandible (Figure 2G). Incisor and molar process well developed, palp absent. Maxillule (Figure 3A). Coxal endite with ¢ve terminal + ¢ve subterminal simple and plumodenticulate setae. Basial endite with six plumodenticulate cuspidate setae and four plumodenticulate submarginal setae. Endopod 2-segmented, distal segment with 1+2 terminal plumodenticulate setae. Journal of the Marine Biological Association of the United Kingdom (2006)
Size. carapace length 1.94 to 2.17 mm (mean 2.05 mm, N¼10); RSL 4.22 to 5.19 mm (mean 4.70 mm, N¼10); PSL 1.11 to 1.24 mm (mean 1.16 mm, N¼10). Carapace (Figure 1B). Similar to ¢rst zoea; rostral spine with fewer ventral spinule; eyes stalked. Antennule (Figure 2B). Biramous. Endopod naked. Exopod with 12 aesthetascs arranged 2, 3, 2, 4 terminally. Antenna (Figure 2E). Endopod greatly lengthened, about 2.3 times as long as exopod, otherwise unchanged. Mandible (Figure 2H). Incisive process more developed, no palp. Maxillule (Figure 3B). Coxal endite with seven terminal + ¢ve subterminal plumodenticulate setae. Basial endite with seven plumodenticulate cuspidate setae and ¢ve subterminal plumodenticulate setae. Endopod unchanged. Maxilla (Figure 3E). Coxal endite now with 8+6 plumodenticulate setae. Basial endite with 11+8 plumose or plumodenticulate setae. Endopod unchanged. Scaphognathite with 23 marginal plumose setae plus posterior, stout setose process, without microtrichia. First maxilliped (Figure 4B). Basis unchanged. Setal formula (ventral+dorso basal plumose setae) of endopodal segments 1^3 as 3+1, 3+1, 4+1, segment 4 with one dorsal seta and ten distal, plumose setae. Exopod distal segment with 12 plumose natatory setae terminally.
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Figure 1. Porcellana cancrisocialis Glassell, 1945. Larval stages. (A) Zoea I, lateral view; (B) Zoea II, lateral view; (C) Megalopa, dorsal view; (D) Zoea I, rostrum, detail. Scale bar in mm.
Second maxilliped (Figure 4E). Coxa with one setae. Basis unchanged. Setal formula of endopodal segments 1^3 as 3+1, 3+1, 4+1, segment 4 with 4+6 terminal setae plus a single, dorsal plumose seta. Exopod distal segment with 12 large plumose natatory terminal setae. Third maxilliped (Figure 5L). Buds elongated and swollen; endopod naked; exopod 2-segmented, with four terminal setae. Pereiopods. Unsegmented, ¢rst pair distally bilobed, showing chelate appearance. Abdomen (Figure 5B). Pair of biramous pleopod buds present each on somites 2^4. Otherwise unchanged. Telson (Figure 5B). Posteromedian margin with additional pair of setae, thus posterior margin with six pairs of long, stout, plumose setae; dorsal setae absent. Megalopa
Size. Carapace length 1.73 to 2.11mm (mean 1.86 mm, N¼10); CW 1.37 to 1.55 mm (mean 1.42 mm, N¼10). Carapace (Figure 1C). About 1.25 times as long as broad; front broad, trilobate, nearly smooth on dorsal surface, median lobe denticulated on frontal margin; lateral lobe with two small teeth; dorsal surface of carapace with scattered setae, more abundant near lateral and posterior margins; branchial margin with acute spine at anterior angle. Antennule (Figure 2C). Peduncle 3-segmented; ¢rst segment broad, rounded, bearing two distolateral teeth and 12^14 simple setae; second segment without setae; third segment with two subterminal setae; exopod 6-segmented, ¢rst segment naked, row of aesthetascs present on segments 2^5, numbering proximal to distal 4, Journal of the Marine Biological Association of the United Kingdom (2006)
6, 4, 3 and 0, 1, 3, 1 setae respectively; distal segment with three large and three short, sparsely setulated setae. Endopod 3-segmented, with 1, 1, 5+2 simple setae. Antenna (Figure 2F). Peduncle 3-segmented, with 2, 4, 3 short setae respectively, £agellum composed of 20^22 articles, setation as ¢gured. Mandible (Figure 2I). Subsymmetrically scoop-like process, palp 3-segmented, proximal segment with two simple setae, median segment naked, distal segment with seven stout serrate setae terminally and eight subterminal setae. Maxillule (Figure 3C). Coxal endite with 22^24 stout, cuspidate or plumodenticulate setae, some subterminal. Basial endite with 12 stout cuspidate and 11 plumodenticulate setae, lateral margin with three plumodenticulate setae. Endopod unsegmented, slender, sharply pointed terminally, with one short, distal simple seta. Maxilla (Figure 3F). Coxal endite with 16 terminal and 28 subterminal + eight terminal and four subterminal + nine terminal plumodenticulate setae. Basial endite with 15 terminal and ¢ve subterminal + 19 terminal and ten subterminal plumodenticulate setae. Endopod unsegmented, with two proximal plumodenticulate setae; scaphognathite with 57 ^62 marginal plumose setae and ¢ve minute ventral setae. First maxilliped (Figure 4C). Coxal endite with 16 terminal and 15 subterminal setae. Basial endite with 47 terminal and subterminal setae. Endopod unsegmented, with one proximal and three subterminal simple setae. Exopod unsegmented, slightly constricted at mid-length, terminating in slender acute process, with three lateral and two subterminal setae.
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Figure 2. Porcellana cancrisocialis Glassell, 1945. (A^C) Antennule; (D^F) antenna and mandible. (A,D,G) Zoea I; (B,E,H) Zoea II; (C,F,I) Megalopa. Scale bar in mm. Journal of the Marine Biological Association of the United Kingdom (2006)
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Figure 3. Porcellana cancrisocialis Glassell, 1945. (A^C) Maxillule; and (D^E) maxilla. (A,D) Zoea I; (B,E) Zoea II; (C,F) Megalopa. Scale bar in mm. Journal of the Marine Biological Association of the United Kingdom (2006)
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Figure 4. Porcellana cancrisocialis Glassell, 1945. (A^C) Maxilliped 1; and (D^F), maxilliped 2. (A,D) Zoea I; (B,E) Zoea II; (C,F) Megalopa. Scale bar in mm. Journal of the Marine Biological Association of the United Kingdom (2006)
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Figure 5. Porcellana cancrisocialis Glassell, 1945. (A^C) Abdomen; (K^M) third maxilliped; and (D^J) pleopods and pereiopods of megalopa. (A,K) Zoea I; (B,L) Zoea II; (C,M) Megalopa; (D,E,F) pleopods of somites 2, 3, 4 respectively; (G) cheliped; (H) third pereiopod; (I) ¢fth pereiopod; (J) propodus and dactylus of ¢fth pereiopod. Scale bar in mm. Journal of the Marine Biological Association of the United Kingdom (2006)
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Table 1. Morphological characters of the larval stages of Porcellana sigsbeiana (Gibbes, 1850) (after Gore, 1971), P. platycheles Haig, 1968 (after Gonza¤lez-Gordillo et al., 1996) and P. cancrisocialis Glassell, 1945 (present study).
Species Zoea I Antennule exopod a, s. Antenna exopod s. Maxillule coxa s. basis s. endopod s. Maxilla coxa s. basis s. endopod s. scaphognathite Second maxilliped basis s endopod s. Telson minute s. pairs
P. sigsbeiana (Gibbes, 1850)
P. platycheles P. cancrisocialis (Glassell, (Haig, 1945) 1968)
3,3 3^5
3,3 3
5,1 1
9
10
10
10 2+1 13
11 2+1 15
10 2+1 12
16 10 7^8
19 11 6^7
14 9 7^8
3 2,2,2,7
3 2,3,3,9+1
3 2,2,3,8+1
2
1
2
Zoea II Antennule exopod a, s. 3,3,2,3 3,3,2,3 2,3,2,4 Antenna exopod s. 1 2 1 Maxillule coxa s. 11 12 12 basis s. 12 13 12 Maxilla coxa s. 19 19 14 basis s. 20 24 19 scaphognathite 22^24 22 23 First maxilliped endopod s. 3+1, 3+1, 3+1, 3+1, 5+1, 7+1, 3+1, 3+1, 4+1, 7+2 11+1 10+1 Second maxilliped endopod s. 2+1, 2+1, 2+1, 2+1,3+1, 3+1,3+1, 7+1 3+1,9+1 4+1, 10+1 exopod 9 12 12 terminal s.
Species Zoea II (cont.) Megalopa Antennule peduncle s. exopod a, s.
P. sigsbeiana (Gibbes, 1850)
0,0,8+0, 8+2, 5+1, 3+0
P. platycheles P. cancrisocialis (Glassell, (Haig, 1945) 1968)
17,0,2
12,14,0,2 0, 0+4, 0+6, 0+4, 1+3, 2+0, 1
Antenna peduncle s. n.d. 5+4+5 2+4+3 £agellum 24^29 17 20^22 segments Mandible stout setae 18 n.d. 7 Maxillule coxa s. 35 27 22^24 basis s. 36 26 26 endopod s. 1 Maxilla coxa s. 25,21,12+8, 4,12,10+14 16,28+8,4,9 3,12 (proximal+ coxal) 21,3+50 15+28 15,5+19,10 basis s. (proximal+ coxal) endopod s. 3+2 2 scaphognathite s. 70 50 57^62 First maxilliped coxa s. 19 19 31 basis s. 21 28 47 endopod s. 8 2 4 exopod s. 19 6 5 Second maxilliped endopod s. 9,10,18,20 4,8,5,30,14 4,9,11,21,19 exopod s. 19,12 7,7 9,17 Third maxilliped coxal s. 12 19 25 basial s. 7 6 5 endopod s. 22,6,20,12,10 17,15,7,25,14 20,5+7,14+5, 9+10+2+1, 5+9 exopod s. n.d. 1,2,1,2 1,2,1,4 Pereiopods spines on 4 2 2 dactylus
s, setae; a, aesthetascs; n.d., no data.
Second maxilliped (Figure 4F). Coxa and basis undi¡erentiated, with 11 plumose setae on mesial margin; endopod 5-segmented, with 4, 9, 11, 21 and 19 setae, respectively; exopod 2-segmented, with six mesial and three lateral setae on proximal segment and 17 (two subterminal, simple+ten terminal, plumose) setae on distal segment. Third maxilliped (Figura 5M). Coxa with 25 short simple setae; basis with ¢ve short simple setae; endopod 5-segmented, ischium with 20 plumose setae, merus with ¢ve plumose setae on ventro£exor margin and seven simple setae, four on dorso£exor margin and Journal of the Marine Biological Association of the United Kingdom (2006)
three on ventral surface, carpus with 14 or 15 long plumose and serrate setae on disto£exor margin diminishing in size terminally, and ¢ve short simple setae, one on ventral surface and four in the extensor margin, propodus with nine long plumose setae on £exor margin, ten simple and serrate setae on lateral margin, two simple setae laterodistal, 1 laterodorsal and 1 mediodorsal, dactylus with ¢ve long plumose and nine simple setae; exopod 4-segmented, with 1, 2, 1 lateral and four terminal setae respectively. Pereiopods (Figure 5G ^ J). All legs fully functional, with numerous short and long simple setae as ¢gured. Cheliped
Porcellana cancrisocialis larval development £attened dorsoventrally, dactyli of ambulatory legs (2^4) with three subequal spines on £exor margin. Fifth pereiopod short, slender, chelate, propodus with three or four long hook-like serrate setae, and many scattered, simple setae. Abdomen (Figure 5C). Six somites with numerous short, simple and plumose setae. Pleopods (Figure 5D ^ F). Biramous, present on somites 2^5, endopods increasing in length posteriorly, each with three cincinuli, somites 3^5 with one subterminal plumose setae, exopods with 10, 13, 12 and 14 long plumose setae, respectively. Telson (Figure 5C). Semi-oval, with a deep median notch on posterior margin but without separation of plates, lateral margins each with nine plumose setae on distal third, increasing in length posteriorly, dorsal surface with seven pairs of simple setae; uropod biramous, endopod with row of 15 plumose marginal setae plus four simple submarginal setae; exopod with row of 21 long plumose marginal setae.
DISCUSSION At an average temperature of 258C, Zoea I of Porcellana cancrisocialis lasted seven days before moulting to Zoea II which, in turn, lasted eight days to reach the megalopa. Gore (1971) reported a very variable development period for Zoea I and II of P. sigsbeiana, depending primarily upon water rearing temperature. At 258C, this species lasted seven and 12 days to moults to Zoea I and II, respectively, and its developmental time is very similar to what was observed during our study. Gonza¤lez-Gordillo et al. (1996) reported that P. platycheles had one prezoea (its duration is about 1.5 hours), and zoeal stages of 6^9 (Zoea I) and 11^20 (Zoea II) days, at a water temperature of 208C. The setation of the basis of each the ¢rst and second maxillipeds for Zoea I and II in the Anomura is normally constant between congeneric species and this is also the case with Porcellana. When the three species of Porcellana for which the complete larval developments are known are compared (Table 1), however, most appendages show di¡erences in the setation pattern. Although there are similarities in setation and aesthetascs pattern between Zoea I and II of P. cancrisolicalis and P. sigsbeiana (7 and 2 similarities in Zoea I and II, respectively) more than between P. cancrisolicalis and P. platycheles (5 and 1 in the two zoeal stages), there is no clear a⁄nity among these species. According to Haig (1960), P. cancrisocialis is very closely related to P. sayana (Leach, 1820) from the western Atlantic in the adult morphology but di¡er only by the number of epibranchial spinules (2 vs 1 in the two species) and the shape of spinules on the median frontal lobe in larger specimens. However, the larval development of P. sayana is not known at present. The larval study of this species is needed for assessing the relationships of Porcellana species. When comparing Zoea I of P. cancrisocialis with those of both P. sigsbeiana and P. platycheles, the main di¡erences are observed in mouthparts setation and minor distinctions in antenna and antennule setation. A similar trend is also observed in Zoea II, with additional di¡erences in setation of exopods of the ¢rst and second maxillipeds. In the case Journal of the Marine Biological Association of the United Kingdom (2006)
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of the megalopa, setation pattern in maxilla, maxillula and maxillipeds is inconsistent among the three species. A particular variation is noticed in the number of segments of the antenna; 17 segments in P. platycheles, while 24^29 segments in P. sigsbeiana and 20^22 segments in P. cancrisocialis. Gonza¤lez-Gordillo et al. (1996) emphasized that the lateral spines on abdominal somites of Zoea I and II are important as the interspeci¢c characters. In this respect, Porcellana cancrisocialis resembles P. sigsbeiana in having the spines on the somites 2^5, but di¡ers in having the spines only on somites 4 and 5 in P. platycheles (spines on somites 4 and 5). All the three Porcellana species have a distinct ventroproximal spine on the basis of the ¢rst maxilliped, a character not known for the other porcellanid zoeas. The megalopa of P. cancrisocialis features a 4-segmented exopod of the third maxilliped, thus strikingly di¡erent from what has been observed in other species of Porcellanidae where this appendage is at most 2-segmented (usually indistinctly segmented). Observations made by Gonza¤lezGordillo et al. (1996) in P. platycheles, however, indicate that this appendage has up to ¢ve segments; its structure in megalopa of other species of Porcellana should be carefully reviewed in order to con¢rm this character which seems to be unique for Porcellanid megalopas as well as for other decapod megalopas. We are in debt to Manuel Ayo¤n P. for collecting the specimen of Porcellana cancrisolicalis and bringing to our attention the presence of the berried female. We are also in debt to the anonymous referees for their valuable observations. This study was partly supported by Consejo Nacional de Ciencia y Tecnolog|¤ a, Mexico, project 44675.
REFERENCES Brooks, W.K. & Wilson, E.B., 1881. The ¢rst zoea of Porcellana. Studies from the Biological Laboraotry. Johns Hopkins University. Baltimore, 3, 58^64. Brusca, R.C., 1980. Common intertidal invertebrates of the Gulf of California, 2nd edn. Arizona: University of Arizona Press. Garc|¤ a-Guerrero, M., Cuesta, J., Hendrickx, M.E. & Rodr|¤ guez, A., 2005. Larval development of the eastern Paci¢c anomuran crab Petrolisthes robsonae (Crustacea: Decapoda: Anomura: Porcellanidae) described from laboratory material. Journal of the Marine Biological Association of the United Kingdom, 85, 1^11. Gonza¤les-Gordillo, J.I., Cuesta, J.A. & Rodr|¤ guez, A., 1996. Studies on the larval development of northeastern Atlantic and Mediterranean Porcellanidae (Decapoda, Anomura). I Redescription of the larval stages of Porcellana platycheles (Pennant, 1777) reared under laboratory conditions. Helgola«nder Meeresuntersuhungen, 50, 517^531. Gore, R.H., 1971. The complete larval development of Porcellana sigsbeiana (Crustacea: Decapoda) under laboratory conditions. Marine Biology, 11, 344^355. Haig, J., 1960. The Porcellanidae (Crustacea: Anomura) of the Eastern Paci¢c. Allan Hancock Paci¢c Expedition, 24, 1^440. Haig, J., 1978. Contribution towards a revision of the Porcellanid genus Porcellana (Crustacea: Decapoda: Anomura). Proceedings of the Biological Society of Washington, 91, 706^714. Hendrickx, M.E., 1996. Habitats and biodiversity of decapod crustaceans in the SE Gulf of California, Mexico. Revista de Biolog|¤ aTropical, 44, 603^617. Hendrickx, M.E. & Harvey, A., 1999. Checklist of anomuran crabs (Crustacea: Decapoda) from the eastern tropical Paci¢c. BelgianJournal of Zoology, 129, 363^389.
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