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Microphrys Bicornutus

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Microphrys Bicornutus 7/ JOURNAL OF CRUSTACEAN BIOLOGY, 2(4): 514-534, 1982 MICROPHRYS BICORNUTUS (LATREILLE, 1825): THE COMPLETE LARVAL DEVELOPMENT UNDER LABORATORY CONDITIONS WITH NOTES ON OTHER MITHRACINE LARVAE (DECAPODA: BRACHYURA: MAJIDAE) Robert H. Gore, Liberta E. Scotto, and Won Tack Yang ABSTRACT The complete larval development, consisting of two zoeal stages and a megalopa, and the first crab stage is described for the shallow-water western Atlantic spider crab Microphrys bicornulus. Data from laboratory cultures indicate that the species can complete its plank- tonic development in less than a week, and is able to attain first crab stage in as few as 10 days. The zoeal stages of M. bicornulus show a great many similarities to known zoeae in other genera within the subfamily Mithracinae, including species of the American genus Milhrax, and to Macrocoeloma, and to a lesser extent the Indo-West Pacific genera Tiarinia and Micippa. Morphological features shared among both the zoeal and megalopal stages of the various mithracine genera are compared, and phylogenetic relationships within Micro- phrys, Milhrax, and Macrocoeloma are proposed. Microphrys bicornulus is a small (ca. 3CM0 mm carapace width, cw) intertidal and shallow subtidal spider crab in the subfamily Mithracinae, which occurs from Bermuda and North Carolina, southward to Brazil (Williams, 1965; Markham and McDermott, 1981). The species is ovigerous throughout the year over a large part of its range, but in spite of its abundance and the ease of its collection there has been no complete study published on the larval development of this crab. Lebour (1944) gave an abbreviated description and illustration of 2 zoeal stages from Bermuda, and Hartnoll (1964) provided an expanded description and illustration of the prezoea and 2 zoeal stages from Jamaica. Neither author's study is suffi- ciently detailed to allow meaningful comparison with larval stages of other mith- racine crabs. Ironically, the complete larval development had been worked out for Microphrys bicornulus by Yang (1967), but the study forms part of a volu- minous dissertation which is not readily available, and has remained unpublished. Inasmuch as several discrepancies in description and illustrations occur between the publications of Lebour and Hartnoll and that of Yang, we recultured the larvae of M. bicornulus and compared our material with the detailed study provided by Yang. In this paper we redescribe and illustrate the zoeal, megalopal, and first crab stage of M. bicornulus. We compare the larvae of this species with those known from other genera and species in the Mithracinae, pointing out possible relationships among them. METHODOLOGY AND REARING EXPERIMENT RESULTS An ovigerous female crab was collected from near Bodden Town, Grand Cay- man Island, on 15 July 1980. The specimen was returned to the laboratory at Fort Pierce, and maintained in a 19 cm diameter bowl with nonflowing seawater (34%, salinity) until hatching occurred on 17 July. A total of 96 larvae were cultured at 20° and 25°C in controlled temperature units using methodology described by Gore (1968). Measurements follow those proposed by Yang (1967) and are the arithmetic means for the number of specimens examined. Specimens and/or molts of the two zoeal stages are deposited in the National Museum of Natural History, 514 GORE ETAL.\ LARVAL DEVELOPMENT OF MICROPHRYS B1CORNUWS 515 5 10 15 20 25 30 35 40 45 DAYS IN STAGE Fig. I. Number of animals and stage duration of Microphrys bicornutus reared at 25°C. (After Yang, I967.) Washington, D.C., (USNM 190705), the British Museum (Natural History), Lon- don (1982-136), and the Indian River Coastal Zone Museum, Fort Pierce, Florida (IRCZM 89-5223). As noted by Hartnoll (1964), Microphrys bicornutus possesses one prezoeal and two zoeal stages. Data from this study, and previous studies by Yang (1967), show that a single megalopal stage occurs approximately 1 week after hatching. Zoeae usually lasted 3, rarely 4, days in stage I at 25°C, and from 5-6 days at 20°C before molting to stage II. The sole megalopal stage in this study was ob- tained after 3 days in stage II at 25°C; other larvae that survived for longer than 3 days died without molting. In this study we used Artemia nauplii as food, but the zoeae may not have fed very well on this, a fact also noted by Hartnoll (1964) in his attempt to rear M. bicornutus. However, Yang (1967) also used newly hatched Artemia nauplii and was considerably more successful in his culture experiment, obtaining sixth crab stage before terminating his study (Fig. 1). In Yang's program at 25°C, both first and second stages required a minimum of 3 days, and the megalopal stage lasted at least 4 days before attaining first crab stage. Results from these 2 culture programs suggest that Microphrys bicornutus is able to complete its planktonic development in as few as 10 days. The survival curve illustrated in Fig. 1 is similar to others obtained for majid crabs in laboratory culture (see e.g., Yang, 1967; Wilson el al., 1979; Scotto and Gore, 1980). In Yang's study approximately 60% of first stage larvae reached stage II, and nearly 516 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 2, NO. 4. 1982 50% of the initial population developed to megalopa, a stage with a mean duration of 5 days. Greatest mortality occurred either in stage I or megalopa, a situation similar to that observed in our study. Crab stages showed enhanced survival, and lasted varying lengths of time from 4-11 days within each instar (Yang, 1967). Yang stated that lowering of water temperatures immediately prior to hatching would at least prolong the prezoeal stage, suggesting that Hartnoll's (1964) ob- servation of lengthy prezoeal development might have been a consequence of lower water temperatures. On the other hand, we tend to agree with Hartnoll's statement that most Anemia nauplii are simply too large a food item for the rather small first zoeae of M. bicomulus (carapace length, cl 0.66-0.75 mm). Yang's successful culturing of this species is apparently a result of obtaining newly hatched Anemia filtered through a stainless steel sieve (105 x 105 mesh, 0.003 wire) thus producing a food item more easily ingested by the majid zoeae. DESCRIPTION OF ZOEAL, MEGALOPA, AND FIRST CRAB STAGES The following description is based primarily on larvae cultured from the ovi- gerous female collected at Grand Cayman Island. Descriptive text and illustra- tions of these zoeae and megalopae were compared with those provided by Yang (1967) from specimens cultured from females collected in Biscayne Bay, Florida. Important variation is noted under the appropriate appendage description. The description and illustration of the first crab stage is rewritten from that of Yang (1967). All setal formulas in the following descriptions progress proximally to distally unless otherwise stated. First Zoea Carapace length, 0.66 mm; 5 specimens examined. Carapace (Fig. 2A).—Cephalothorax smooth, globose, with short ventrally di- rected rostrum, posteriorly curved dorsal spine, median dorsal tubercle; pair of minute setae posterior to tubercle, second pair posterolateral to dorsal spine; posteroventral border with 6 setae, first stoutest (=anterior seta); pterygostomial region produced as bluntly rounded tooth; frontal organ on eyes talk extremely minute (often indiscernible in preserved specimens); eyes sessile. Abdomen (Fig. 2A, B).—Five somites; first with 2 dorsomedial setae; second with 2 posterodorsal setae, 2 lateral anteriorly curved knobs; third to fifth with 2 posterodorsal setae plus pair of short posteroventral spines. Telson (Fig. 2A, B, b).—Trapezoidal; elongate furcae covered with fine hairs, pair of movable lateral spines; posterior margin bearing 6 spines armed with spinules. Antennule (Fig. 2C).—Conical rod, 2 long, stout, 2 short, thinner aesthetascs plus I hair (3 aesthetascs, Yang, 1967). Antenna (Fig. 2D).—Protopodite an elongate tapered process armed with 2 rows of spinules; exopodite similar in form, equal to or slightly longer than protopodite with 2 rows of spinules on distal one-third, 1 naked seta and I serrate spine at base of spinules; endopodite bud one-third length of protopodite. Mandibles (Fig. 2E).—Asymmetrically scoop-shaped, dentate processes; left with 3 bluntly rounded teeth on posterior margin, right with 1 tooth; incisor margins evenly and bluntly dentate; molar processes irregularly dentate. GORE ETAL.: LARVAL DEVELOPMENT OF MICROPHRYS BICORNUTUS 517 A,B 1.0mm b 0.5mm C-l 0.25mm Fig. 2. Microphrys bicornutus, first zoea: A, Lateral view; B, Abdomen; b, Telson; C, Antennule; D, Antenna; E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped I; I, Maxilliped 2. 518 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 2, NO. 4, 1982 Majcf/Mf (Fig. 2F).—Endopodite 2-segmented, short proximal article with i seta, longer distal article with 2 subterminal, 4 terminal setae; basal endite bearing 5 spines, 2 setae; coxal endite with 7 processes. Maa(//a (Fig. 2G).—Unilobate endopodite with 5 setae; bilobed basal and coxal endites each with 5, 4 processes progressing distally; scaphognathite bearing 12 or 13 thin, plus I stout, apical plumose setae; pubescence as illustrated. Ma^rY/iped / (Fig. 2H).—Coxopodite with 1 seta; basipodite ventral setae 2,2,3,3; endopodite 5-segmented, 3,2,1,2,4 + I (Roman numeral denoting dorsal seta); exopodite with 4 natatory setae. Ma;c///;/W2 (Fig. 21).—Coxopodite naked; basipodite with 3 naked ventral setae; endopodite 3-segmented 0,1,5 setae (2 long, 1 medium, 2 very short); exopodite with 4 natatory setae. Co/or.—Carapace, basipodites of first and second maxillipeds, and abdomen, lime green. Single melanophore on mandibles, basipodites of first and second maxil- lipeds, paired melanophores distoventrally on abdominal somites 2-5. Gastric region and intestine to abdominal somite 3 shaded with black. Orange rose chro- matophore above eyes, on surface of carapace at posteroventral angle, at base of antenna, on proximal portion of basipodite of maxilliped 2, and on dis- toventral margins of somites 3-5.
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