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The larval development of Hymenosoma orbiculareDesmarest, 1825 (Crustacea: Decapoda: Brachyura:Hymenosomatidae)Maria Dornelas a , José Paula a & Adriano Macia ba IMAR, Laboratório Marítimo da Guia , Faculdade de Ciências da Universidade deLisboa , Estrada do Guincho, 2750 Cascais, Portugal E-mail:b Departamento Ciências Biológicas , Universidade Eduardo Mondlane , CP 257, Maputo,MozambiquePublished online: 26 May 2010.

To cite this article: Maria Dornelas , José Paula & Adriano Macia (2003) The larval development of Hymenosomaorbiculare Desmarest, 1825 (Crustacea: Decapoda: Brachyura: Hymenosomatidae), Journal of Natural History, 37:21,2579-2597, DOI: 10.1080/00222930210155684

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J N H10 N 2003, 37, 2579–2597

The larval development of Hymenosoma orbiculare Desmarest, 1825(Crustacea: Decapoda: Brachyura: Hymenosomatidae)

MARIA DORNELAS†, JOSE PAULA† and ADRIANO MACIA‡

†IMAR, Laboratorio Marıtimo da Guia, Faculdade de Ciencias daUniversidade de Lisboa, Estrada do Guincho, 2750 Cascais, Portugal;e-mail: maad@netcabo.pt‡Departamento Ciencias Biologicas, Universidade Eduardo Mondlane,CP 257, Maputo, Mozambique

(Accepted 23 April 2002)

The zoeal stages of Hymenosoma orbiculare Desmarest, 1825 are described andillustrated from materials hatched in the laboratory and plankton collectedin the field. A comparison with other hymenosomatid larvae is tabulated anddiscussed. The present description agrees with typical larval characters of theHymenosomatidae, and shows a number of differences in relation to previousdescriptions.

K: Brachyura, Hymenosomatidae, Hymenosoma orbiculare, larvaldescription.

IntroductionHymenosoma orbiculare Desmarest, 1825 (Crustacea: Brachyura: Hymeno-

somatidae) is a crab common in brackish and fresh water in south-east Africa. Thisspecies inhabits shallow soft sediment zones, where it spends the day buried andemerges during the night to forage for food (Hill and Forbes, 1979). The genusHymenosoma was established for H. orbiculare by Desmarest (1825) and currentlyincludes two more species, H. depressum Jaquinot in Hombron and Jacquinot, 1846and H. hodgkini Lucas, 1980, which occur in New Zealand and eastern Australia,respectively (Lucas, 1980).

The family Hymenosomatidae is composed of more than 90 species distributedin 16 genera. Larvae of only 29 species of the family have been described so far,and from these only 15 descriptions include detail on appendages.

The typical larval development of hymenosomatid crabs includes three zoealstages which are not followed by a typical megalopa (reviewed by Lucas, 1980;Guinot and Richer de Forges, 1997). However some freshwater species such asAmarinus lacustris (Chilton, 1882), A. angelicus (Holthius, 1968) and Elamenopsisbovis (Barnard 1950), have suppressed larval development (Barnard, 1950; Lucas,

Journal of Natural HistoryISSN 0022-2933 print/ISSN 1464-5262 online © 2003 Taylor & Francis Ltd

http://www.tandf.co.uk/journalsDOI: 10.1080/00222930210155684

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1971, 1980). Terada (1977) describes the larval developnment of several hymenosom-atids as comprising three zoeae followed by a megalopa. However, only the zoeaeare described in Terada (1977); the megalopae are said to be described in a sub-sequent report, but to our knowledge it has never been published. Boschi et al.(1969) included in their larval development of Halicarcinus planatus (Fabricius,1775) a description of a megalopa that more closely resembles a first crab stagethan a typical megalopa.

The larvae of a number of species of Halicarcinus have been described including:H. messor (Stimpson, 1858) by Aikawa, 1929 (first zoea); H. coralicola (Rahtbun,1909) by Aikawa, 1929 (first zoea) and Terada, 1977 (complete development),H. varius (Dana, 1851) by Wear, 1965 (first zoea); H. cookii (Filhol, 1885) byMelrose, 1975 (first zoea) and Wear and Fielder, 1985 (complete development);H. planatus (Fabricius, 1775) by Boschi et al., 1969 (complete development);H. ovatus Stimpson, 1958 and H. rostratus (Haswell, 1882) by Lucas, 1971 (completedevelopment); H. orientalis Sakai, 1932 by Terada, 1977 (complete development);H. afecundus Lucas, 1980 by Lucas, 1980 (complete development); H. innominatusRichardson, 1949 and H. withei (Meirs, 1876) by Wear and Fielder, 1985 (first zoea).

Aikawa (1933) described a ‘Hymenozoea abdominalis’, identified by Lucas (1980)as zoeae of Elamena truncata (Stimpson, 1858), and Terada (1977) later describedthe complete larval development of this species. Other larvae of this genus weredescribed: Elamena mathaei (Desmarest, 1825) by Gurney, 1938 (complete develop-ment) and Al-Kholy, 1959 (complete development); Elamena sindensis Alcock, 1900,by Prasad and Tampi, 1957 (first zoea), Tirmizi and Kazmi, 1987 (complete develop-ment) and Siddique et al., 1987 (complete development); Elamena cristatipesGravely, 1927, by Hashmi, 1969 (complete development); E. longirostris Filhol,1885, E. momona Melrose, 1975 and E. producta Kirk, 1879, by Wear and Fielder,1985 (complete development briefly).

The larvae of other Hymenosomatidae genera are also known: Trigonoplaxunguiformis (de Haan, 1839) by Aikawa, 1929 (first zoea) and Fukuda, 1981 (com-plete development); Trigonoplax cimex Kemp, 1915, by Krishnan and Kannupandi,1988 (complete development); Elamenopsis kempi (Chopra Das, 1930) by Salmanand Ali, 1996 (complete development); Amarinus laevis (Targioni Tozzetti, 1877)and A. paralacustris (Lucas, 1970) by Lucas, 1971 (complete development);Neohymenicus pubescens (Dana, 1851) by Wear and Fielder, 1985 (completedevelopment briefly).

Wear and Fielder (1985) describe the larval external morphology of Hymenosomadepressum. Broekhuysen (1955) and Hill and Forbes (1979) figure lateral views ofzoeas I and III of Hymenosoma orbiculare. However no detail of the appendages isgiven, nor are the larvae described. No complete larval description is thus knownfor a species of Hymenosoma. Three zoeal stages of H. orbiculare have beenrecognized from plankton samples (Hill and Forbes, 1979; Paula et al., 2000),and correspond to the larval development typical of this family. The purpose ofthe present paper is to describe the complete larval development of Hymenosomaorbiculare, and to compare it with that of other species of this family.

Materials and methodsOne ovigerous female of Hymenosoma orbiculare was collected at Inhaca Island,

Mozambique, with a bottom trawl at 5 m depth in the Saco Mangrove creek on9 December 2000. The female was kept unfed in an individual, gently aerated 500 ml

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beaker with sea water changed daily until 12 December 2000, when the larvaehatched. After larvae were seen swimming, females and zoeas I were fixed in 4%formalin and later transferred to 70% ethanol. Zoeas I, II and III were earliercollected from plankton trawls at the same mangrove creek on 15 November 1997,preserved in 4% formalin and later transferred to 70% ethanol.

Ten zoeas of each stage were used for measurements with a micrometer under aWild M5 dissecting microscope. Carapace length was measured from the base ofthe rostral spine to the most posterior carapace margin; carapace width above theeyes and distance from the tip of the dorsal spine to the tip of the rostral spine werealso measured. Five larvae of each stage were dissected and mounted in polyvinyllactophenol, with the cover slips sealed by clear nail varnish. Observations anddrawings were made with an Olympus BH-2 compound microscope equipped witha camera lucida. The larval stages were described and illustrated as recommendedby Clark et al. (1998). The zoeal series and the spent females were deposited at TheNatural History Museum of London registration number 2002:91.

Description

Zoea ISize. Carapace width 0.57±0.05 mm; carapace length 0.66±0.02 mm; distance

from tip of rostral spine to tip of dorsal spine 2.83±0.09 mm.Carapace (figures 1A, 2A). Rostral and dorsal spines very long; dorsal spine

slightly curved, twice the carapace length; rostral spine spinulate and twice aslong as dorsal spine; lateral spines absent; anterodorsal setae absent; one pair ofposterodorsal setae; ventral margin with two pairs of lateral plumose setae and onepair of posterior plumose setae; carapace with posterolateral ventral fold, and dorsalzone with a hollow section inside; eyes sessile.

Antennule (figure 3A). Uniramous; endopod absent; exopod unsegmented withtwo terminal aesthetascs and one seta.

Antenna (figure 3A). Protopod smaller than antennule; endopod and exopodabsent; no setation.

Maxillule (figure 4A). Epipod absent, coxal endite with four terminal and onesubterminal setae; basial endite with five setal processes; endopod two-segmented,proximal segment with one seta, distal segment with one subterminal and fourterminal setae; exopod setae absent.

Maxilla (figure 5A). Coxal endite simple, lobed with two setae; basial enditebilobed with 4+4 setae; endopod simple with five (2 subterminal+3 terminal ) setae;exopod (scaphognathite) margin with four plumose setae plus distal stout plumoseprocess.

First maxilliped (figure 6A). Coxal endite with one seta; basial endite with 10setae (2, 2, 3, 3); endopod five-segmented with 3, 2, 1, 2, 5 (1 subterminal+4terminal ) setae; exopod two-segmented with four terminal natatory setae.

Second maxilliped (figure 7A). Coxal endite naked; basial endite with foursetae; endopod three segmented with 1, 1, 6 (2 subterminal+1 almost terminal+3terminal ) setae.

Third maxilliped. Absent.Pereiopods. Absent.Abdomen (figures 8A, 9A). Five somites; somite 2 with a pair of short rounded

dorsolateral processes, directed laterally; somite 5 with a pair of ventrolateral spines;somites 2–5 with a pair of dorsomedial setae; pleopods absent.

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F. 1. Hymenosoma orbiculare Desmarest, 1825. Anterior view of carapace: (A) first zoea;(B) second zoea; (C) third zoea. Scale: 0.3 mm.

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F. 2. Hymenosoma orbiculare Desmarest, 1825. Setation of ventral carapace margin:(A) first zoea; (B) second zoea; (C) third zoea. Scale: 0.1 mm.

Telson (figure 10A). Rectangular, small furca distally spinulate; three pairs ofstout spinulate setae; the most lateral pair curved laterally, and the medial pairshorter than the other two pairs.

Zoea IISize. Carapace with 0.69±0.06 mm; carapace length 0.84±0.04 mm; distance

from tip of rostral spine to tip of dorsal spine 4.28±0.23 mm.Carapace (figures 1B, 2B). Ventral margin with three pairs of lateral plumose

setae and one pair of posterior plumose setae; eyes stalked; otherwise unchanged.Antennule (figure 3B). Exopod with four terminal aesthetascs and one seta;

otherwise unchanged.Antenna (figure 3B). One lateral seta near base (exopod seta); otherwise

unchanged.Maxillule (figure 4B). Basial endite with seven setal processes; exopod seta

present; otherwise unchanged.Maxilla (figure 5B). Scaphognathite with five plumose setae and two plumose

setae on distal process; otherwise unchanged.First maxilliped (figure 6B). Exopod with seven terminal natatory setae;

otherwise unchanged.

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F. 3. Hymenosoma orbiculare Desmarest, 1825. Antennule and antenna: (A) first zoea;(B) second zoea; (C) third zoea. Scale: 0.1 mm.

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F. 4. Hymenosoma orbiculare Desmarest, 1825. Maxillule: (A) first zoea; (B) second zoea;(C) third zoea. Scale: 0.1 mm.

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F. 5. Hymenosoma orbiculare Desmarest, 1825. Maxilla: (A) first zoea; (B) second zoea;(C) third zoea. Scale: 0.1 mm.

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F. 6. Hymenosoma orbiculare Desmarest, 1825. First maxilliped: (A) first zoea; (B) secondzoea; (C) third zoea. Scale: 0.1 mm.

Second maxilliped (figure 7B). Exopod with seven terminal natatory setae;otherwise unchanged.

Third maxilliped (figure 11A). Present as uniramous bud.Pereiopods (figure 11A). Present as small buds, chela bilobed.Abdomen (figures 8B, 9B). Somite one with one plumose seta.

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F. 7. Hymenosoma orbiculare Desmarest, 1825. Second maxilliped: (A) first zoea;(B) second zoea; (C) third zoea. Scale: 0.1 mm.

Telson (figure 10B). Unchanged.

Zoea IIISize. Carapace width 0.95±0.05 mm; carapace length 1.07±0.03 mm; distance

from tip of rostral spine to tip of dorsal spine 5.97±0.24 mm.

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F. 8. Hymenosoma orbiculare Desmarest, 1825. Dorsal view of the abdomen: (A) firstzoea; (B) second zoea; (C) third zoea. Scale: 0.3 mm.

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F. 9. Hymenosoma orbiculare Desmarest, 1825. Lateral view of the abdomen: (A) firstzoea; (B) second zoea; (C) third zoea. Scale: 0.3 mm.

Carapace (figures 1C, 2C). Ventral margin with four pairs of lateral plumosesetae and one pair of posterior plumose setae. Otherwise unchanged.

Antennule (figure 3C). Unchanged.Antenna (figure 3C). Unchanged.Maxillule (figure 4C). Unchanged.Maxilla (figure 5C). Scaphognathite now with five plumose setae and four

plumose setae on distal process; otherwise unchanged.First maxilliped (figure 6C). Exopod with eight natatory setae; otherwise

unchanged.Second maxilliped (figure 7C). Exopod with eight natatory setae; otherwise

unchanged.

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F. 10. Hymenosoma orbiculare Desmarest, 1825. Dorsal view of the telson: (A) first zoea;(B) second zoea; (C) third zoea. Scale: 0.1 mm.

Third maxilliped (figure 11B). Biramous.Pereiopods (figure 11B). Present as large buds.Abdomen (figures 8C, 9C). Unchanged.Telson (figure 10C). With one pair of dorsomedial setae.

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F. 11. Hymenosoma orbiculare Desmarest, 1825. Third maxilliped and pereiopods:(A) second zoea; (B) third zoea. Scale: 0.1 mm.

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DiscussionIn spite of great controversy regarding the phylogenetic position of the family

Hymenosomatidae among the Brachyura, the unity of the family has never beenquestioned (Guinot and Richer de Forges, 1997). Several larval characters certainlycontribute to this fact. Aikawa (1929) was the first to describe three species ofhymenosomatid zoeas and to select several characters as typical of this family: allcarapace spines absent, except rostral spine; antenna as an inconspicuous spine,often distally spinulate; telson short, trapezoidal, wider anteriorly, with three pairsof stout setae on posterior margin; and the presence of primary chromatophores inthe maxillipeds. After the descriptions of several other species, Lucas (1980) selectedthe typical form of the telson, the small antennae, naked, with one eccentric terminalor one basial seta, and added the absence of pleopods on the abdomen, and themaxilla with single coxal and basial endites, coxal endite vestigial, with one or morerarely two setae, as diagnostic characters of the family. The larvae describedsince then, and the present study confirm all these characters as typical of theHymenosomatidae. Two characters are exclusive to Hymenosoma orbiculare, namelythe setation of the maxillule and the ventrolateral spines on somite 5 of the abdomen.

The larvae of Hymenosoma orbiculare figured by Broekhuysen (1955) are some-what different from the ones here described, and resemble the estuarine type figuredby Hill and Forbes (1979). The present larvae instead resemble the freshwater laketype figured in that study. However, the present larvae were captured in a highlysaline mangrove area, with no freshwater input apart from groundwater flow ( Kalk,1995), thus contradicting the hypothesis of freshwater and seawater types of larvae.Further, adult morphological differences were reported by Hill and Forbes (1979),and, in spite of the lack of consistency, should be further investigated in order toclarify specific classification.

A ‘hymenosomide’ zoea was figured by Frontier (1963), showing pleopod budson the abdomen. Although the telson is remarkably similar to hymenosomatidlarvae, and the abdomen shows lateral projections on somite 5, also present inElamena spp., the larvae described by Frontier (1963) were collected in the plankton,and no detail of the appendages was given. Consequently, it is not clear whether ornot this larva actually belongs to the Hymenosomatidae.

Those hymenosomatid zoeal descriptions with detail of appendages (table 1)suggest that setation of the maxillule and the distal segment of the endopod on bothfirst and second maxillipeds is fairly constant within this family. Some divergenceoccurs in the case of Hymenosoma orbiculare and among Elamena spp., as well asTrigonoplax spp. (the latter genus formerly considered a subgenus of Elamena).These two genera also possess distinctive large lateral projections on the fifth somiteof the abdomen. However, these species should be re-examined to confirm thedifferences in such easily overlooked characters prior to evaluating their significancein divergence within the Hymenosomatidae.

For the 16 species in which detailed appendages have been described, 10 havea single lateral seta on the antenna. Further, the figures of Halicarcinus cookii,H. varius, H. whithei, H. planatus, Neohymenicus pubescens and Hymenosomadepressum by Wear and Fielder (1985) all show the presence of a seta in this position.The location of this seta is remarkably close to the place where the exopod apparentlywould be, suggesting that it might be a vestigial exopod, and it therefore is referredto in this paper as the exopod seta.

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Table 1. A comparison of first-stage zoeal carapace, antenna, maxillule, maxilla, first and second maxilliped, and abdomen characters from knownhymenosomatid descriptions (characters taken from references under species names).

Antennaexopod First Second

Carapace spines seta Maxillule Maxilla maxilliped maxilliped Abdomen

Halicarcinus messor Short rostral spine Present 4, 5, 1+4 1, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral(Stimpson, 1858) processes

Aikawa (1929)(Terada, 1977)H. coralicola (Rahtbun, Very short rostral Absent 4, 5, 1+4 1, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral

1909) spine processes(Aikawa, 1929)(Terada, 1977)H. planatus (Fabricius, Short rostral spine Present 4, 5, 1+4 1, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral

1775) processes(Boschi et al., 1969)H. ovatus Stimpson, 1858 Short rostral spine Present 4, 5, 1+4 – – – No processes(Lucas, 1971)H. rostratus (Haswell, 1882) Very short rostral Absent 4, 5, 1+4 – – – No processes(Lucas, 1971) spineH. orientalis Sakai, 1932 Short rostral spine Present 4, 5, 1+4 1, 8, 6 1+4 3+3 Somite 2 with paired dorsolateral(Terada, 1977) processesElamena truncata Very short rostral Present 4, 5, 1+4 1, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral

(Stimpson, 1858) spine processes; somite 5 with a pair(Aikawa, 1933) of lateral projections(Terada, 1977)Elamena mathaei Very short rostral Absent 4, 4, 1+4 Unclear 0+2 2+3 Somite 2 with paired dorsolateral

(Desmarest, 1825) spine (1, 7, 6) processes; somite 5 with a pair(Gurney, 1938) of lateral projections(Al-Kholy, 1959)

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Table 1. (Continued ).

Antennaexopod First Second

Carapace spines seta Maxillule Maxilla maxilliped maxilliped Abdomen

E. sindensis Alcock, 1900 No spines Absent 3, 5, 1+4 1, 8, 5 0+4 3+3; 2+3; Somite 2 with paired dorsolateral(Prasad and Tampi, 1957) (1, 7, 5) 1+3 processes; somite 5 with a pair(Tirmizzi and Kazmi, 1987) of lateral projections(Siddique et al., 1987)E. cristatipes Gravely, 1927 Short rostral spine Absent 4, 5, 1+4 1, 6, 5 0+4 2+3 Somite 2 with paired dorsolateral(Hashmi, 1970) (1, 7, 5) processes; somite 5 with a pair

of lateral projectionsTrigonoplax unguiformis Short rostral spine Present 4, 5, 1+4 1, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral

(de Haan, 1839) processes; somite 5 with a pair(Aikawa, 1929) of lateral projections(Fukuda, 1981)T. cimex Kemp, 1915 No spines Absent 4, 5, 1+4 1, 6–8, 5 1+3 2+3 Somite 2 with paired dorsolateral( Krishnan and processes; somite 5 with a pair

Kannupandi, 1988) of lateral projectionsAmarinus laevis (Targioni Rostral and dorsal Present 4, 5, 1+4 1, 8, 5 1+4 2+3 No processes

Tozzetti, 1877) spinesA. paralacustris (Lucas, Short rostral spine Absent 4, 5, 1+4 – – – No processes

1970)(Lucas, 1971)Elamenopsis kempi (Chopra Long rostral and Terminal 4, 5, 1+4 2, 8, 5 2+4 3+3 Somite 2 with paired dorsolateral

Das, 1930) dorsal spines seta processes(Salman and Ali, 1996)Hymenosoma orbiculare Long rostral and Present 5, 5, 1+5 2, 8, 5 1+4 3+3 Somite 2 with paired dorsolateral

Desmarest, 1825 dorsal spines processes; somite 5 with a pair(this study) of ventro lateral spinesD

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AcknowledgementsThe authors wish to thank Catia Bartilotti for identifying the plankton samples.

The authors are grateful to the Estacao de Biologia Marinha da Inhaca for supportduring field work. This research was supported by Fundacao para a Ciencia eTecnologia grant to M.D.

ReferencesA, H., 1929, On larval forms of some Brachyura, Records of Oceanographic Work in

Japan, II, 17–55.A, H., 1933, On larval forms of some Brachyura, Paper II; a note on indeterminable

zoeas, Records of Oceanographic Work in Japan, V, 124–254.A-K, A. A., 1959, Larval stages of four brachyuran Crustacea (from the Red Sea),

Publications Marine Biology Station of Al-Ghardaqa, 10, 239–246.B, K. H., 1950, Descriptive catalogue of South African Decapod Crustacea (crabs

and shrimps), Annals of the South African Museum, XXXVIII, 1–864.B, E. E., S, M. A. and G, B., 1969, Desarrollo larval del cangrejo,

Halicarcinus planatus (Fabricius), en el laboratorio, con observaciones sobre la distribui-cion de la especie, Bulletin of Marine Science, 19, 224–242.

B, G. J., 1955, The breeding and growth of Hymenosoma orbiculare Desm.(Crustacea, Brachyura), Annals of the South African Museum, 41, 313–343.

C, P. F., C, D. K. and P, G. W., 1998, Accuracy and standardization ofbrachyuran larval descriptions, Invertebrate Reproduction and Development, 33, 127–144.

D, A. J., 1825, Considerations Generales sur la Classes des Crustaces, et Descriptionsdes Especes de ces Animaux, qui vivent dans la Mer, sur les Cotes, ou dans les eauxdouces de la France (Paris and Strasbourg), xix+446 pp., 5 tables, pls 1–56.

F, S., 1963, Notes sur quelques larves atypiques de decapodes brachyoures recolteesdans le plankton de Nosy Be, Cahiers O.R.S.T.O.M., oceanographie, 3, 31–59.

F, Y., 1981, Larval development of Trigonoplax unguiformis (De Haan) (Crustacea;Brachyura) reared in the laboratory, Zoological Magazine Japan, 90, 164–173.

G, D. and R F, N., 1997, Affinites entre les Hymenosomatidae Macleay,1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura), Zoosystema,19, 453–502.

G, R., 1938, Notes on some decapod Crustacea from the Red Sea, VI–VIII. VI, Somebrachyuran larvae, Proceedings of the Zoological Society of London, 108B(1), 73–84,pls 1–6.

H, S. S., 1969, The larvae of Elamena (Hymenosomidae) and Pinnotheres(Pinnotheridae) hatched in the laboratory (Decapod: Crustacea), Pakistan Journal ofScientific and Industrial Research, 12, 279–285.

H, B. J. and F, A. T., 1979, Biology of Hymenosoma orbiculare Desm. in LakeSibaya, South African Journal of Zoology, 14, 75–79.

K, M., 1995, A Natural History of Inhaca Island, Mozambique (Johannesburg:Witwatersrand University Press).

K, T. and K, T., 1988, Larval development of Elamena (Trigonoplax)cimex Kemp, 1915 in the laboratory: the most unusual larvae known in the Brachyura(Crustacea: Decapoda), Bulletin of Marine Science, 43, 215–228.

L, J. S., 1971, The larval stages of some Australian species of Halicarcinus (Crustacea,Brachyura, Hymenosomatidae). I. Morphology, Bulletin of Marine Science, 21, 471–490.

L, J. S., 1980, Spider crabs of the family Hymenosomatidae (Crustacea; Brachyura) withparticular reference to Australian species: systematics and biology, Records of theAustralian Museum, 33, 148–247.

M,M. J., 1975, The marine fauna of New Zealand (Crustacea, Decapoda, Brachyura),New Zealand Oceanographic Institute Memoir, 34, 1–116.

P, J., D, T., N M, R., B, C., M, A. and Q, H.,2000, Larval fluxes at Saco Mangrove Creek, Inhaca Island (South Mozambique), inMacrobenthos of Eastern African Mangroves: Life Cycles and Reproductive Biology ofExploited Species, Final Report. ERBIC 18-CT96-0127, Part B (Florence), pp. 97–111.

Dow

nloa

ded

by [

Yor

k U

nive

rsity

Lib

rari

es]

at 1

8:27

15

Nov

embe

r 20

14

Larval development of Hymenosoma orbiculare 2597

P, R. R. and T, P. R. S., 1957, Notes on some decapod larvae, Journal of theZoological Society of India, 9, 22–39.

S, S. D. and A,M. H., 1996, Larval development of Elamenopsis kempi (Chopra andDas) (Brachyura, Hymenosomatidae) reared in the laboratory, Scientia Marina, 60,407–416.

S, F. A., K, Q. B. and T, N. M., 1987, Complete development of Elamenasindensis Alcock, 1900 (Brachyura, Hymenosomatidae) under laboratory conditions,Proceedings of the 7th Pakistan Journal of Zoology, 181–191.

T,Y., 1977, On the zoea larvae of four crabs of the family Hymenosomatidae, ZoologicalMagazine (Japan), 86, 174–184.

T, N. M. and K, Q. B., 1987, Larval development of two spider crabs reared inthe laboratory, families Hymenosomatidae and Majidae, Crustaceana, 53, 281–291.

W, R. G., 1965, Zooplankton of Wellington Harbour, New Zealand, Zoology Publicationsfrom Victoria University of Wellington, 38, 1–31.

W, R. G. and F, D. R., 1985, The marine fauna of New Zealand: larvae of theBrachyura (Crustacea, Decapoda), New Zealand Oceanographic Institute Memoirs, 92,1–90.

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Yor

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