J N H 10 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, Laborato´rio Marı´timo da Guia, Faculdade de Cieˆncias da Universidade de Lisboa, Estrada do Guincho, 2750 Cascais, Portugal; e-mail:
[email protected] ‡Departamento Cieˆncias Biolo´gicas, Universidade Eduardo Mondlane, CP 257, Maputo, Mozambique (Accepted 23 April 2002) The zoeal stages of Hymenosoma orbiculare Desmarest, 1825 are described and illustrated from materials hatched in the laboratory and plankton collected in the field. A comparison with other hymenosomatid larvae is tabulated and discussed. The present description agrees with typical larval characters of the Hymenosomatidae, and shows a number of differences in relation to previous descriptions. K: Brachyura, Hymenosomatidae, Hymenosoma orbiculare, larval description.
Introduction Hymenosoma orbiculare Desmarest, 1825 (Crustacea: Brachyura: Hymenosomatidae) is a crab common in brackish and fresh water in south-east Africa. This species inhabits shallow soft sediment zones, where it spends the day buried and emerges during the night to forage for food (Hill and Forbes, 1979). The genus Hymenosoma was established for H. orbiculare by Desmarest (1825) and currently includes two more species, H. depressum Jaquinot in Hombron and Jacquinot, 1846 and 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 distributed in 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 zoeal stages which are not followed by a typical megalopa (reviewed by Lucas, 1980; Guinot and Richer de Forges, 1997). However some freshwater species such as Amarinus lacustris (Chilton, 1882), A. angelicus (Holthius, 1968) and Elamenopsis bovis (Barnard 1950), have suppressed larval development (Barnard, 1950; Lucas, Journal of Natural History ISSN 0022-2933 print/ISSN 1464-5262 online © 2003 Taylor & Francis Ltd http://www.tandf.co.uk/journals DOI: 10.1080/00222930210155684
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1971, 1980). Terada (1977) describes the larval developnment of several hymenosomatids as comprising three zoeae followed by a megalopa. However, only the zoeae are described in Terada (1977); the megalopae are said to be described in a subsequent 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 stage than 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) by Melrose, 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 (complete development); H. orientalis Sakai, 1932 by Terada, 1977 (complete development); H. afecundus Lucas, 1980 by Lucas, 1980 (complete development); H. innominatus Richardson, 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 described the complete larval development of this species. Other larvae of this genus were described: Elamena mathaei (Desmarest, 1825) by Gurney, 1938 (complete development) and Al-Kholy, 1959 (complete development); Elamena sindensis Alcock, 1900, by Prasad and Tampi, 1957 (first zoea), Tirmizi and Kazmi, 1987 (complete development) and Siddique et al., 1987 (complete development); Elamena cristatipes Gravely, 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: Trigonoplax unguiformis (de Haan, 1839) by Aikawa, 1929 (first zoea) and Fukuda, 1981 (complete development); Trigonoplax cimex Kemp, 1915, by Krishnan and Kannupandi, 1988 (complete development); Elamenopsis kempi (Chopra Das, 1930) by Salman and 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 (complete development briefly). Wear and Fielder (1985) describe the larval external morphology of Hymenosoma depressum. Broekhuysen (1955) and Hill and Forbes (1979) figure lateral views of zoeas I and III of Hymenosoma orbiculare. However no detail of the appendages is given, nor are the larvae described. No complete larval description is thus known for a species of Hymenosoma. Three zoeal stages of H. orbiculare have been recognized from plankton samples (Hill and Forbes, 1979; Paula et al., 2000), and correspond to the larval development typical of this family. The purpose of the present paper is to describe the complete larval development of Hymenosoma orbiculare, and to compare it with that of other species of this family. Materials and methods One ovigerous female of Hymenosoma orbiculare was collected at Inhaca Island, Mozambique, with a bottom trawl at 5 m depth in the Saco Mangrove creek on 9 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 larvae hatched. 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 earlier collected 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 a Wild M5 dissecting microscope. Carapace length was measured from the base of the rostral spine to the most posterior carapace margin; carapace width above the eyes and distance from the tip of the dorsal spine to the tip of the rostral spine were also measured. Five larvae of each stage were dissected and mounted in polyvinyl lactophenol, with the cover slips sealed by clear nail varnish. Observations and drawings were made with an Olympus BH-2 compound microscope equipped with a camera lucida. The larval stages were described and illustrated as recommended by Clark et al. (1998). The zoeal series and the spent females were deposited at The Natural History Museum of London registration number 2002:91. Description Zoea I Size. 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 as long as dorsal spine; lateral spines absent; anterodorsal setae absent; one pair of posterodorsal setae; ventral margin with two pairs of lateral plumose setae and one pair of posterior plumose setae; carapace with posterolateral ventral fold, and dorsal zone with a hollow section inside; eyes sessile. Antennule (figure 3A). Uniramous; endopod absent; exopod unsegmented with two terminal aesthetascs and one seta. Antenna (figure 3A). Protopod smaller than antennule; endopod and exopod absent; no setation. Maxillule (figure 4A). Epipod absent, coxal endite with four terminal and one subterminal setae; basial endite with five setal processes; endopod two-segmented, proximal segment with one seta, distal segment with one subterminal and four terminal setae; exopod setae absent. Maxilla (figure 5A). Coxal endite simple, lobed with two setae; basial endite bilobed with 4+4 setae; endopod simple with five (2 subterminal+3 terminal ) setae; exopod (scaphognathite) margin with four plumose setae plus distal stout plumose process. First maxilliped (figure 6A). Coxal endite with one seta; basial endite with 10 setae (2, 2, 3, 3); endopod five-segmented with 3, 2, 1, 2, 5 (1 subterminal+4 terminal ) setae; exopod two-segmented with four terminal natatory setae. Second maxilliped (figure 7A). Coxal endite naked; basial endite with four setae; endopod three segmented with 1, 1, 6 (2 subterminal+1 almost terminal+3 terminal ) 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 of stout spinulate setae; the most lateral pair curved laterally, and the medial pair shorter than the other two pairs. Zoea II Size. 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) second zoea; (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 III Size. 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) first zoea; (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) first zoea; (B) second zoea; (C ) third zoea. Scale: 0.3 mm.
Carapace (figures 1C, 2C ). Ventral margin with four pairs of lateral plumose setae 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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Discussion In spite of great controversy regarding the phylogenetic position of the family Hymenosomatidae among the Brachyura, the unity of the family has never been questioned (Guinot and Richer de Forges, 1997). Several larval characters certainly contribute to this fact. Aikawa (1929) was the first to describe three species of hymenosomatid zoeas and to select several characters as typical of this family: all carapace spines absent, except rostral spine; antenna as an inconspicuous spine, often distally spinulate; telson short, trapezoidal, wider anteriorly, with three pairs of stout setae on posterior margin; and the presence of primary chromatophores in the maxillipeds. After the descriptions of several other species, Lucas (1980) selected the typical form of the telson, the small antennae, naked, with one eccentric terminal or one basial seta, and added the absence of pleopods on the abdomen, and the maxilla with single coxal and basial endites, coxal endite vestigial, with one or more rarely two setae, as diagnostic characters of the family. The larvae described since then, and the present study confirm all these characters as typical of the Hymenosomatidae. Two characters are exclusive to Hymenosoma orbiculare, namely the setation of the maxillule and the ventrolateral spines on somite 5 of the abdomen. The larvae of Hymenosoma orbiculare figured by Broekhuysen (1955) are somewhat different from the ones here described, and resemble the estuarine type figured by Hill and Forbes (1979). The present larvae instead resemble the freshwater lake type figured in that study. However, the present larvae were captured in a highly saline 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 to clarify specific classification. A ‘hymenosomide’ zoea was figured by Frontier (1963), showing pleopod buds on the abdomen. Although the telson is remarkably similar to hymenosomatid larvae, and the abdomen shows lateral projections on somite 5, also present in Elamena 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 or not 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 both first and second maxillipeds is fairly constant within this family. Some divergence occurs in the case of Hymenosoma orbiculare and among Elamena spp., as well as Trigonoplax spp. (the latter genus formerly considered a subgenus of Elamena). These two genera also possess distinctive large lateral projections on the fifth somite of the abdomen. However, these species should be re-examined to confirm the differences in such easily overlooked characters prior to evaluating their significance in divergence within the Hymenosomatidae. For the 16 species in which detailed appendages have been described, 10 have a single lateral seta on the antenna. Further, the figures of Halicarcinus cookii, H. varius, H. whithei, H. planatus, Neohymenicus pubescens and Hymenosoma depressum 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 apparently would be, suggesting that it might be a vestigial exopod, and it therefore is referred to 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 known hymenosomatid descriptions (characters taken from references under species names).
Carapace spines
Maxillule
Maxilla
First maxilliped
Second maxilliped
Abdomen
Short rostral spine
Present
4, 5, 1+4
1, 8, 5
1+4
3+3
Somite 2 with paired dorsolateral processes
Very short rostral spine
Absent
4, 5, 1+4
1, 8, 5
1+4
3+3
Somite 2 with paired dorsolateral processes
Short rostral spine
Present
4, 5, 1+4
1, 8, 5
1+4
3+3
Somite 2 with paired dorsolateral processes
Short rostral spine
Present
4, 5, 1+4
–
–
–
No processes
Very short rostral spine Short rostral spine
Absent
4, 5, 1+4
–
–
–
No processes
Present
4, 5, 1+4
1, 8, 6
1+4
3+3
Very short rostral spine
Present
4, 5, 1+4
1, 8, 5
1+4
3+3
Very short rostral spine
Absent
4, 4, 1+4
Unclear (1, 7, 6)
0+2
2+3
Somite 2 with paired dorsolateral processes Somite 2 with paired dorsolateral processes; somite 5 with a pair of lateral projections Somite 2 with paired dorsolateral processes; somite 5 with a pair of lateral projections
M. Dornelas et al.
Halicarcinus messor (Stimpson, 1858) Aikawa (1929) (Terada, 1977) H. coralicola (Rahtbun, 1909) (Aikawa, 1929) (Terada, 1977) H. planatus (Fabricius, 1775) (Boschi et al., 1969) H. ovatus Stimpson, 1858 (Lucas, 1971) H. rostratus (Haswell, 1882) (Lucas, 1971) H. orientalis Sakai, 1932 (Terada, 1977) Elamena truncata (Stimpson, 1858) (Aikawa, 1933) (Terada, 1977) Elamena mathaei (Desmarest, 1825) (Gurney, 1938) (Al-Kholy, 1959)
Antenna exopod seta
Table 1.
Carapace spines
First maxilliped
Second maxilliped
Maxilla
Abdomen
Absent
3, 5, 1+4
1, 8, 5 (1, 7, 5)
0+4
Absent
4, 5, 1+4
1, 6, 5 (1, 7, 5)
0+4
2+3
Short rostral spine
Present
4, 5, 1+4
1, 8, 5
1+4
3+3
No spines
Absent
4, 5, 1+4
1, 6–8, 5
1+3
2+3
Rostral and dorsal spines Short rostral spine
Present
4, 5, 1+4
1, 8, 5
1+4
2+3
Absent
4, 5, 1+4
–
–
–
Long rostral and dorsal spines
Terminal seta
4, 5, 1+4
2, 8, 5
2+4
3+3
Somite 2 with paired dorsolateral processes
Long rostral and dorsal spines
Present
5, 5, 1+5
2, 8, 5
1+4
3+3
Somite 2 with paired dorsolateral processes; somite 5 with a pair of ventro lateral spines
3+3; 2+3; Somite 2 with paired dorsolateral 1+3 processes; somite 5 with a pair of lateral projections Somite 2 with paired dorsolateral processes; somite 5 with a pair of lateral projections Somite 2 with paired dorsolateral processes; somite 5 with a pair of lateral projections Somite 2 with paired dorsolateral processes; somite 5 with a pair of lateral projections No processes No processes
Larval development of Hymenosoma orbiculare
Maxillule
E. sindensis Alcock, 1900 No spines (Prasad and Tampi, 1957) (Tirmizzi and Kazmi, 1987) (Siddique et al., 1987) E. cristatipes Gravely, 1927 Short rostral spine (Hashmi, 1970) Trigonoplax unguiformis (de Haan, 1839) (Aikawa, 1929) (Fukuda, 1981) T. cimex Kemp, 1915 ( Krishnan and Kannupandi, 1988) Amarinus laevis ( Targioni Tozzetti, 1877) A. paralacustris (Lucas, 1970) (Lucas, 1971) Elamenopsis kempi (Chopra Das, 1930) (Salman and Ali, 1996) Hymenosoma orbiculare Desmarest, 1825 (this study)
Antenna exopod seta
(Continued ).
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Acknowledgements The authors wish to thank Ca´tia Bartilotti for identifying the plankton samples. The authors are grateful to the Estac¸a˜o de Biologia Marinha da Inhaca for support during field work. This research was supported by Fundac¸a˜o para a Cieˆncia e Tecnologia grant to M.D. References A, 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 distribuicion 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 of brachyuran larval descriptions, Invertebrate Reproduction and Development, 33, 127–144. D, A. J., 1825, Conside´rations Ge´ne´rales sur la Classes des Crustace´s, et Descriptions des Espe`ces de ces Animaux, qui vivent dans la Mer, sur les Coˆtes, ou dans les eaux douces de la France (Paris and Strasbourg), xix+446 pp., 5 tables, pls 1–56. F, S., 1963, Notes sur quelques larves atypiques de de´capodes brachyoures re´colte´es dans 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, Affinite´s 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, Some brachyuran 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 of Scientific and Industrial Research, 12, 279–285. H, B. J. and F, A. T., 1979, Biology of Hymenosoma orbiculare Desm. in Lake Sibaya, 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) with particular reference to Australian species: systematics and biology, Records of the Australian 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), in Macrobenthos of Eastern African Mangroves: Life Cycles and Reproductive Biology of Exploited Species, Final Report. ERBIC 18-CT96-0127, Part B (Florence), pp. 97–111.
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P, R. R. and T, P. R. S., 1957, Notes on some decapod larvae, Journal of the Zoological Society of India, 9, 22–39. S, S. D. and A, M. H., 1996, Larval development of Elamenopsis kempi (Chopra and Das) (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 Elamena sindensis 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, Zoological Magazine (Japan), 86, 174–184. T, N. M. and K, Q. B., 1987, Larval development of two spider crabs reared in the laboratory, families Hymenosomatidae and Majidae, Crustaceana, 53, 281–291. W, R. G., 1965, Zooplankton of Wellington Harbour, New Zealand, Zoology Publications from Victoria University of Wellington, 38, 1–31. W, R. G. and F, D. R., 1985, The marine fauna of New Zealand: larvae of the Brachyura (Crustacea, Decapoda), New Zealand Oceanographic Institute Memoirs, 92, 1–90.
