THE LARVAL STAGES of HOMOLA BARBATA (FABRICIUS) (CRUSTACEA, DECAPODA, HOMOLIDAE) REARED in the Laboratoryl
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BIOLOGICAL RESULTS OF THE UNIVERSITY OF MIAMI DEEP-SEA EXPEDITIONS. 55. THE LARVAL STAGES OF HOMOLA BARBATA (FABRICIUS) (CRUSTACEA, DECAPODA, HOMOLIDAE) REARED IN THE LABORATORyl A. L. RICE British Museum (Natural History), Cromwell Road, London, S.W. 7, England AND ANTHONY J. PROVENZANO, JR. University of Miami, Rosenstiel School of Marine and Atmospheric Sciences ABSTRACT Larvae yielded by a female collected off Yucatan were reared to the seventh, and final, zoeal stage at 20°C and 36.2%0 salinity. The larvae, the first complete series available of any species of the family, are described and illustrated. Comparison of the reared larvae with others attributed to the same species but captured in Mediterranean and South African waters confirmed the morphological differences which previously had led to sug- gestions of specific or subspecific separation of the parent populations. Examination of adult material failed to reveal positive evidence of specific status of the separated populations. The setation of the uropodal endopods in Homola appears in the third zoeal stage, earlier than for any other decapod known. Recent studies of larval forms tend to confirm the view that the Homolidae are related to the Raninidae and are close to the line of descent of the true crabs, and that the family is well separated from both the typical Anomura and the much more primitive Dromiidea. INTRODUCTION Homola barbata has been recorded in the eastern and western North Atlantic, in the Mediterranean, and in the South Atlantic near South Africa. Throughout its range (Fig. 1), H. barbata is the only recognized repre- sentative of the genus, except off the east coast of North America where a closely related species, H. vigil Milne Edwards, also occurs. Zoeal larvae attributed to Homola barbata have been described from the Bay of Naples (Cano, 1893; Pike & Williamson, 1960) and from Mes- sina (Thiele, 1905). Although none of these larvae was hatched from a known adult or molted to a certainly identifiable stage, the fact that H. barbata is the only species of the genus recorded from this fairly well studied area makes an erroneous identification rather unlikely. 1 Contribution No. 1194 from the University of Miami, Rosenstiel School of Marine and Atmospheric Sciences. This work was supported by research grant GB-7075X and ship support grant GB-7082 from the National Science Foundation; by research grant GM-I1244 from the National Institute of General Medical Sciences, U.S. Department of Health, Education and Welfare; and by the National Geographic Society-University of Miami Deep-Sea Biology Program. This paper is one of a series resulting from the National Geographic Society-University of Miami Deep-Sea Biology Program. 1970] Rice & Provenzano: Larval Stages of Homola barbata 447 FIGURE 1. Distribution of Homola barbata (circles) and H. vigil (triangles). Data from the literature and from unpublished records in the British Museum (N atural History) and the Rosenstiel School of Marine and Atmospheric Sciences, University of Miami. In most cases each symbol corresponds to one record, but in some areas, such as the east coast of the United States and in the Caribbean, there are more records than are indicated here. Five zoeal stages taken in the plankton off South Africa and described by Rice & von Levetzow (1967) are very similar to the Mediterranean larvae and probably belong to the same species. Megalopal stages and terminal zoeae taken in the plankton of the Straits of Florida off Miami in 1963 were linked, through molts in the laboratory, with a first crab stage which was also identified as H. barbata (Rice, 1964). However, those Florida larvae differed from the Mediterranean and South African zoeae in a number of details, which suggests that what is con- sidered to be H. barbata may include more than one subspecies, or even species, with distinct larvae. 448 Bulletin of Marine Science [20(2) ..0 •...Q) •...'"Q) •...Q) Q) ~ •... Q) p. '"0.. .~ -5 .5 '"0 Q) .D .;::; u VJ Q) '"0 Q) •...'"> ~ Q) -5 ..r:: .S! ..r:: ~ ~ E .•..••...0 Q) E'" .•..•Q) <::I' ~ ~... ~<::I <::I C ~ c ::t:: N I.lJ c.: ~ 0 ~ 1970) Rice & Provenzano: Larval Stages of Homola barbata 449 TABLE 1 NUMBERS AND PERCENTAGES OF LARVAE OF Homola barbata IN SUCCESSIVE STAGES Stage No. of specimens Percentage I 140 100.0 11 115 82.5 III 108 76.5 IV 100 70.0 y 76 55.0 VI 24 17.5 V11 1 0.7 Obviously, it would be desirable to rear the larvae from adults taken in various parts of the known range. This paper presents the results of one such rearing made at the Rosenstiel School of Marine and Atmospheric Sciences of the University of Miami. We are indebted to Barbara Stolen for the illustration of the female crab and to C. Edith Marks and R. A. Feigenbaum for assistance in the laboratory. MATERIAL AND METHODS On 23 May 1967, an ovigerous female was taken at station P-582 by the R/V JOHN ELLIOTT PILLSBURY at Arrowsmith Bank in the Yucatan Channel (21°10'N, 86°18'W). The depths sampled by the otter trawl during this tow varied from 250 to 20 fathoms (457 to 36 meters), so there is no way of knowing accurately the depth at which the female was collected. The specimen was maintained alive on board the ship and was returned to the laboratory. One hundred and forty of the larvae hatched on 6 June 1967 were reared individually in compartmented plastic trays at 20°C and 36.27<'0 salinity. Newly hatched nauplii of Artemia were provided as food for the larvae. Details of the rearing method and treatment of the resulting ma- terial were similar to those described earlier (Rice & Provenzano, 1964), except that exuviae and dead animals were preserved in ethylene glycol. The female from which larvae were obtained (Fig. 2) is deposited in the museum of the Rosenstiel School of Marine and Atmospheric Sciences, and its catalogue No. is UMML 32: 3824. Carapace length of larvae was measured from the tip of the rostrum to the middle of the posterior margin of the carapace. RESULTS None of the animals reached the megalopa stage, and only one individual reached the terminal zoeal stage, after molting six times. The numbers and percentages surviving into each successive stage are given in Table 1, 450 Bulletin of Marine Science [20(2) FIGURE 3. Homola barbata, first zoea. which shows that the main mortalities occurred after the third molt and particularly between the fifth and sixth stages. The duration of the larval stages was rather more variable than is usual in laboratory-reared larvae, but molts occurred fairly regularly with peaks at about 9.5, 17.5, 26.5, 37, and 48 days after hatching. MORPHOLOGY In the following descriptions of the larval stages, the illustrations are largely self-explanatory, but are supplemented by a summary of the arma- ture of selected features in Table 2. Otherwise, the descriptions are gen- erally restricted to the major changes in the anatomy from stage to stage and variations within a single stage. TABLE 2 ARMATURE OF SELECTED FEATURES OF SUCCESSIVE ZOEAL STAGES OF Homola barbata Marginal Artic- Setation of Setation setae of Natatory setae ulated Uropods of Zoeal Ant. 2 processes -~- scaphog- stage scale Mxp. I Mxp.2 Mxp.3 of tel son Exopod Endopod nathite I 9 4 4 10 5 II 12 6 6 4-5 11-12 11-12 1II 18-19 7-9 8-9 7-8 16-18 11-13 4-6 20-21 IV 24 10-11 11-13 11-12 23-25 20-24 13-18 38-41 V 27-30 13-15 16-18 14-17 29-34 25-35 22-30 55-62 VI 31-33 16-18 20-22 18-21 36-41 40-42 34-37 80-86 VII 34 21-22 25-26 25-26 45 45-47 40-42 100-103 1970] Rice & Provenzano: Larval Stages of Homola barbata 451 , '':.:: FIGURE 4. Homola barbata, first zoea: a, b, dorsal and lateral views of ab- domen, respectively; c, antennule; d, antenna; e, basal endite of first maxilla; f, endopod and scaphognathite of second maxilla; g, h, first and second maxil- lipeds, respectively. Bar scale represents 0.3 mm for a and b; 0.2 mm for c, d, g, and h; and 0.1 mm for e and f. Stage [.-SIZE: Carapace length (CL) 0.67-0.76 mm. (4 spec.) DESCRIPTION (Figs. 3 and 4.): The most striking features of the larvae are the rows of teeth on the winglike extensions of the carapace which seem to be typical of early-stage homolids. The length of the rostmm 452 Bulletin of Marine Science [20(2) 1.0 mm FIGURE 5. Homola barbata: a, b, second zoea; c, d, third zoea. varied considerably in the specimens examined and could be much shorter than that shown in the illustration; this variation was largely responsible for the range in carapace length. The only other major variation at this 1970] Rice & Provenzano: Larval Stages of Homola barbata 453 --l- "/ .~l/\\~ J(("l\ \ ~ \J ~ 'I ., .;, a , "~ 1.,1! \J 11l' t- - .-----------I I-------i 0.1 mm 0.5mm (b,c,d,el (0) FIGURE 6. Homola barbata, second zoea: a, abdomen; b, antennule; c, an- tenna; d, third maxilliped; e, portion of a telson inside which that of the third zoea is visible. stage was that the third maxillipeds either were represented by unarmed, unsegmented buds or were entirely lacking. Stage II.-SIZE: CL 1.19-1.25 mm. (2 spec.) DESCRIPTION (Figs. 5, a, b, and 6.): The eyes are now free, and the carapace carries a pair of supraorbital spines.