Bermudamysis G. N., Platyops G. N. and Other Mysids from Bermudian Caves

Spgologia 2 (112) 1986, E. J. Brill, Leiden

BERMUDAMYSIS G. N., PLATYOPS G. N. AND OTHER MYSIDS FROM BERMUDIAN CAVES

MIHAI BACESCU Muste dYHistoireNaturelle "Grigore Antipa", 1 ChaussCe Kisselef, ~ucuresti111, Romania

THOMAS M. ILIFFE Bermuda Biological Station for Research, Ferry Reach 1- 15, Bermuda

SUMMARY One of the authors (T. Iliffe) collected mysids from 9 caves, within the framework of a faunistic survey of Bermudian caves. A brief characterization of the caves and the collecting technique is presented. Two new genera, each with a new species - i.e. Bewnudamysis speluncola n.g., n.sp. and Platyops stnrm' n.g., n.sp. - are described. The first species was collected in dozens of specimens from numerous caves; the second shows dorsoventrally flattened eyes, a convergency with the genus Heteromysoides, although the new genus seems to belong to the Mysini tribe. Morphological and zoogeographical remarks are made on another mysid common in those caves, Heteromysis bmnudmsis Sars, 1885, as well as on a fourth species, H. guitarti Bacescu, 1978.

L'un des auteurs (T. Iliffe) a collectt des MysidacCs dans 9 grottes dans le cadre d'une etude faunistique des grottes des Bermudes. On donne une brkve description des grottes en question et de la technique de capture des MysidacCs. On fournit des descriptions de deux genres nouveaux, chacun avec une nouvelle espkce, i savoir Bmnudamysis speluncola n.g., n.sp. et Platyops stmeri n.g., n.sp. La premikre espsce a CtC capturCe en des dizaines d'exemplaires dans un grand nombre de grottes; la seconde a des yeux aplatis dorsoventralement, convergence avec le genre Heteromysoi&s, bien que Platyops semble ap- partenir B la tribu des Mysini. On fait des remarques morphologiques et zooghographiques concernant deux autres MysidacCs, Heteromysis bewnudmsis Sars, 1885 et H. guitarti Bacescu, 1978.

INTRODUCTION The extensive inland marine caves of Bermuda constitute a diverse habitat containing over 200 species of macro-invertebrates (Sket 81 Iliffe, 1980; Iliffe et al., 1983). The caves are situated in the Pleistocene eolian limestones which cap a mid ocean volcanic sea mount. The most unusual and specialized animals have been found in those tidal cave pools and underwater chambers 94 BERMUDA MARINE CAVE SYMPOSIUM most isolated from the sea. Utilizing advanced cave diving techniques, many of the animals have been collected from the deeper waters of the pools and often at considerable penetrations into the underwater caves - areas previously unknown or thought to be inaccessible. The three areas of zonation applicable to terrestrial caves - a twilight zone near the entrance, a middle zone of complete darkness and variable temperature, and the deep interior zone of darkness and constant temperature (Poulson & White, 1969) - are found in these marine caves. While the diversi- ty and abundance of organisms decrease with distance inland, away from the sea, it is in these far reaches of the caves that the most significant troglobitic species have been discovered. New species described from the Bermuda caves include Atlantusellus caver- nicolus, an isopod representing a new family (Sket, 1979); Miostephos leamingtonensis, a new calanoid copepod (Yeatman, 1980); Apseudes bermudeus, a new hermaphroditic tanaidacean (Bacescu, 1980); Cocoharpinia ilzffei, a gammaridean amphipod from a new genus (Karaman, 1980a); Idunella sketi, a new species of gammaridean amphipod (Kararnan, 1980b); Somersiella sterr& and Typhlatya iltffei, two new species of caridean shrimp (Hart & Manning, 1981); Mesonerilla prospers, a new archiannelid polychaete (Sterrer & Iliffe, 1982); and Bermudalana aruboides, representing a new genus of cirolanid isopods (Bowman & Iliffe, 1983), and Mictocaris halope described by Bowman & Iliffe (1985), a representative of a new order (Mictacea) of Peracarida.

COLLECTION SITES In order to avoid repetition when indicating the materials and describing the species, we shall henceforth use only the numbers designating each of the 9 stations.

St. 1: Green Bay Cave, 13 June 1982, specimens taken from a white sponge collected using scuba in the Connection Passage near the Rat Trap at 16 m water depth. St. 2: Green Bay Cave, 31 January 1984, specimens collected using scuba with a fine mesh hand net from bottom silt in the "Desert" in 15 m water depth. St. 3: Castle Grotto, 20 July 1982, specimens collected with a suction bottle from bottom silt of the rear portion of the main pool in 0.5 to 1 m water depths. St. 4: Cheny Pit Cave, 12 January 1984, specimens collected using scuba with a suction bottle from the sloping silt floor in 8 to 12 m water depths. St. 5: Palm Cave, 13 & 16 March 1982, specimens collected using scuba with a suction bottle and fine mesh hand net from bottom silt in the underwater Palm Cave Room at 16 m water depth. St. 6: Walsingharn Cave, 17 February 1984, specimens collected with a fine mesh hand net from bottom rocks in the main entrance pool at 2 to 3 m water depths. St. 7: Leamington Cave, 14 August 1984, specimens taken in a plastic bottle trap baited with squid and left for 4 days at 5 m water depth in the main pool. St. 8: Green Bay Cave, 15 August 1984, specimens collected using scuba with a suction bottle and fine mesh hand net from bottom silt in the "Desert" at 15 m water depth. St. 9: Grenadier Pool, 15 August 1984, specimens collected using scuba with fine a mesh hand net and bottles from under a rock ledge in 5 m water depth. STYGOLOGZA 2 (112) 1986 95

HABITAT Green Bay Cave is Bermuda's longest cave at over 2 km in length, despite being almost totally underwater. It consists of 2 principal passages, the Green Bay Passage and the North Shore Passage, which extend across a peninsula of land separating the nearly enclosed Harrington Sound from the open North Lagoon. Linking these 2 segments of the cave are the Trunk Passage and its extension, the Connection Passage. The Rat Trap is a low area with stronger currents just inside the Green Bay entrance where the Green Bay and Connec- tion Passages branch. The "Desert" is a segment of the Connection Passage with a broad silty floor. Castle Grotto is a former commercial cave with a large shallow lake and was shown to tourists by boat during the 1930's. It is located on the edge of Castle Harbour and has direct tidal connections with it. Cherry Pit Cave is in the Walsingham area and consists of a steeply sloping fissure extending from the surface to an underwater room. Palm Cave is part of a large, multi-entrance underwater cave system extending inland from Harrington Sound towards Castle Harbour. Walsingham Cave is similarly part of another extensive cave system located near Castle Harbour. A lake in the dimly illuminated entrance section of the cave has tidal currents passing through it. Leamington Cave is presently shown to tourists. An artificially illuminated lake within the cave connects to Harrington Sound. Grenadier Pool is a nearly circular water-filled collapse sink located about 75 m from the North Shore of Bermuda. Underwater, the sink extends horizontal- ly beneath a rock ledge for about 10 m to a dimly illuminated section.

DESCRIPTIONS In the material from these caves, we found four species of Mysidacea: one described from Bermuda by G. 0. Sars in 1885 (i.e. Heteromysis bemudensis) whose male we now describe and whose diagnose we complete; one known from Cuba (i.e. Heteromysisguitartt); and two representing new genera as well as new species.

Bermudamysis new genus Diagnosis. - Small translucent red mysids (2.8 - 4.0 mm) of the Mysini tribe, distinguished within the tribe through the lack of pleopodal dimorphism; long and fine appendages; very elongate pereiopods IV-V (0,Q) with thin articles and $.-segmented tarsus, the first pereiopods shorter and a bit thicker, with 3-segmented tarsus. Globular eyes: the corneal part smaller than the cylindrical peduncle. 96 BERMUDA MARINE CAVE SYMPOSIUM

Two marsupial laminae with 2 or 4 eggs; lanceolated, antenna with feathered setae all around. Telson armed with 5-6 spines only in its distal half, with a deep sinus bordered by long apical spines and 6-7 lamina. Uropodal endopodite without spines.

Bermudamysis speluncola new species. (fig. 1 A. - J.)

Material - Dozens of 0 and Q specimens from the following stations: St. 1 (2 Q , 3 W, 1juv.); St. 2 (40 Q , 50 W, 5 juv.); St. 3 (1 Q); ST. 4 (4 Q ov., 1 Q P, 3 a); ST. 5 (14 Q , 12 0,1 Q p.); St. 6 (27 specimens among which 5 were ovigerous 9); St. 7 (1 Q , 2 a j.); St. 8 (3 expl.); St. 9 (117 a & Q : a few juv.).

Description of the male and the female - Size: 2.3 - 3.0 mm. Integument slightly transparent, having a red tint; several specimens fixed some time ago show only the brown-golden visual part of the eyes. But during the life span the mysid has a bright and beautiful colour in spite of keeping its transparency, a colour dominated by intense red; this colour covers the whole cephalothorax (excepting the golden eggs), the basis of the thoracic appendices, joints of the abdominal segments and especially the last joint binding the pleotelson to the last abdominal segment. Reddish-orange starlets are also present on the scales, antennae, base of the long flagellum Al , telson and uropods. Yellowish-orange occular peduncles, corneal part externally translucent with a bright brown nucleus. After preservation in alcohol for as little as 4 months, this crustacean becomes milky-white in colour and half-transparent all over; cephalothorax dull-white; only occular peduncles keep their yellow colour, while the visual part shines due to its brownish-red colour. Carapace with frontal side slightly prominent reaching the base of the eyes (fig. 1A). Eyes short and cylindrical (Fig. 1A). Corneal part almost as big as peduncle with small omatidia in deeply brown-reddish cups whose color remains un- changed in alcohol for 2 years. Antenna 1 with a tiny male lobe from which starts a narrow fascicle of more than 15 fine, closely gathered, ventrally directed setae (fig. 1B) and a very long flagellum slightly exceeding the body length. Antenna 2 with a shorter base than of antenna 1, its exopodite (scale) exceeding them in length and somewhat lanceolate, being slightly narrowed toward the distal article (fig. 1A). The buccal parts do not show any remarkable differences as compared to the representatives of the Mysini tribe. Labrum slightly triangular; mandible with strong pars incisiva (5 teeth), with 3-segmented long palp. Maxillule with 12 spines on the distal article of palp, as in fig. 1C. STYGOLOCIA 2 (112) 1986 97

Maxilliped I1 with 6 typical strong double-serrated phanera (fig. IF) biser- rated, slightly pointed at the apex in Q , with a prolongation in 0' (fig. 1 f) in- serted like the fingers of a hand on the antero-inner border of dactylum. Peraeopod I (thoracopod 111) strong, short (fig. lH), provided with 4 tarsal articles (dactylus included), while the rest of the pereiopods are exquisitely fine, much thinner and longer, similar to those of the genus Erythrops. The others peraeopods (fig. 1G) are alike as far as phanerotaxis is concerned, but are different in length in comparison with pereiopod I, firstly due to the almost twice longer meros, then due to the proximal article of the carpo-propodus, equal in length with pereiopod 11, but growing gradually up to pereiopod I, where, in the female, it is almost as long as the other two. Pleopods do not show any kind of dimorphism: more or less triangular plates on pairs I-IV (fig. 1 I, J 0,Q ), short, hardly exceeding the posterior margin of the pleonite, about two times longer on the V-th pair (fig. 1J). Uropods with slightly shorter endopodite, small statolith and no spine (fig. 1E). Telson (fig. ID) provided only in its distal half with 5-6 lateral spines (2 long apical ones as long as the sinus), all fine and very pointed; on the inner side of the apical spines, some tiny subapically attached spines hardly longer than 8-10 laminae ornamenting the shallow bottom of the sinus (fig. Id). The female is slightly larger than the male: 3-4 mm. It is provided with only 1 pair of oostegites and bears 2 or 4 genital produces. Its flagellum on antenna 2 is shorter, hardly reaching the telson.

Ecological and ethological remarks - This is the most widely distributed and most abundant of the mysids inhabiting Bermuda's caves being found in all the 7 explored caves: 1 specimen (St. 3) to 117 specimens (St. 9). B. speluncola, the most common mysid from Green Bay Cave, and Grenadier Pool, is an epibenthic species observed in large numbers and always remaining in close proximity to the surface of silty sediments. It is primarily found in those parts of the cave which have more dirkct water exchange with the open sea. On one occasion, many individuals were noted around a dead fish in the Connection Passage. The internal organs are pale yellow to white with a few red-orange pigment granules in the cephalothorax, at the joints of the abdominal segments in the telson and uropods. The visual part of the eyes is golden-brown and consists of a reduced brown section possibly the screening pigment, with a distinct red central portion. Reduced or absent screening pigment is a characteristic of both cave and bathypelagic crustaceans. In the darker parts of Grenadier Pool, B. speluncola is free swimming and oc- curs in swarms, together with the bright red calanoid copepod Ridgewayia marki. In January, almost all the females were ovigerous. The sex ratio is ap- proximately equal in Green Bay Cave on 31/1/84. The males can be distinguished by the white shining cylindric penis with the fascicles (fig. 1 H) 98' BERMUDA MARINE CA VE SYMPOSIUM

Fig. 1. A-J, Bennudamysis spelun~olan. g., n. sp. 0 A, eye, antennae 1 and 2; B, detail of antenna 1; C, palp of maxilla; D, telson 0 ; d, tip of telson of another specimen ( Q ); E, uropod; F, tip of maxilliped 11; f, f, a dactylar dimorphic phanerae (9) magnified of maxiliped 11, impressive claws resembling those of the strong mandibular palp, to latch the prey; G, peraeopod I Q ; H, peraeopod V; I, pleopod IV; J. pleopod V Q , (but the same in 0). K-P, Heteromysis aff. guitarti (Bacescu, 1968). K, phanerae complex ofantennula; L, peraeopod IV; M, endopod of uropod; N. telson; 0, pleopod IV 0;P, penis. STYGOLOGZA 2 (1/2) 1986 99

visible of spermatozoa. All dead specimens have exopodites closely gathered in the posterior and dorsal part of the body of these mysids, while the endopodites are crouched, pressed under the thorax as if holding something between them. The suppleness of the body of Bermudamysis, as well as the large numbers in which it was collected in some caves. indicate a free life and an agglomeration of its population in swarms. This way of life is comparable to Mysidium rubroculatum just discovered in the Cuban caves (Bacescu & Ortiz, 1984). These mysids have a deep reddish color of the eyes. Nevertheless, the small size of the thoracopodal exopodites and its occurrence in the company of Heteromysis guitarti (and in a white sponge) (St. 1) or of H. bermudensis (St. 2 and 3), are in- teresting; the capture of Bermudamysis speluncola in a plastic bottle trap baited with squid, left for 4 days at 5 m water depth in the main pool of Leamington cave is also remarkable. The shape of the telson is reminiscent of an Anisomysis (e.g. A. pelewensis Panampunail). The similarity of pleopods in both sexes - differentiated only by the short branch of the endopodite - is similar to some Heteromysini, e.g. Plalymysis Brattegard.

Heteromysis (Olivaemysis) bermudensis Sars, 1885 Syn .?H. antilensis Ver- rill, 1923, H. b. cesari Bacescu, 1968 (?) (fig. 2, M-R) The most common mysid found at St. 3 (Castle Grotto) is Heteromysis bermudensis, the second species of this genus to be described from the tropical western Atlantic. The difficulty in identifying this species arises from the fact that in- itially it had been described after a single Q specimen. Verrill (1923) and Clarke (1955) had only the Q , with the homologation of H. antilensis with H. bermudensis based only on a drawing of the telson (Verrill, 1923 - fig. 5). Brat- tegard (1973) also had Q , but he illustrated the group of phanerae of antenna 1 that characterize the subgenus Olivaemysis. Bacescu (1968) was the first to describe the CY of this species, creating the subspecies cesari for the Cuban form.

~emarks.- The males collected from Castle Grotto correspond to those of the Cuban population as indicated by the armature of the telson (fig. IN) and the uropod (fig. lM), the existence of sexual dimorphism only at pleopod IV CY (fig. 1, 0)but not also at pleopod 111, and the presence of the largest number of laminae (26-30 Bacescu, 35 Bowman) known within the genus lato sensu, i.e. , non-flagellate short laminae (fig. 1P). We therefore conclude that the Cuban population may be a separate subspecies. The fact that Heteromysis species were also found in the Bermuda caves, with only a few specimens (even just one pair, 0 and 9 of H. bermudensis) shows that they are not free-living like Ber- 100 BERMUDA MARINE CAVE SYMPOSIUM mudamysis, but occur in association with other living beings i.e. with sponges (St. 1). H. bermudensis is a member of the H. guitarti-type of the subgenus Olivaemysis, which, as with other members - e.g. H. actiniae and H. dispar - shows dimorphism only at pereiopod IV. The pair of setae bordering the flat flagellate lamellar phanera are proximal- ly thick, but thinner and strongly feathered in the other parts (fig. 1K).

Geographic distribution. - H. bermudensis was collected by the Challenger ex- pedition from shallow waters around Bermuda and described by G. 0. Sars in 1885. It was summarily studied by Verril1(1923), also from Bermuda, and by Tattersall (l951), Clarke (1955), and Bowman (1981). Bacescu (1968) found it in Cuban water, and Brattegard (1973) and Bowman (1981) in Columbian waters.

Platyops n.g. Diagnosis. - Small mysids (2.5 mm) with rectangular eyes, seen from above, strongly dorso-ventrally flattened with corneal side tiny in a lateral external position. Pereiopod I bulky, with 3-articulated tarsus, pereiopod I1 almost 3 times longer and thin, with 6 articulated tarsus. Telson vaguely linguiform provided, only in the distal 114, with 3 lateral spines. On the narrow truncated apical side are 2 long lateral spines with 2 median ones, half as long. Pleopods small, 1-articulated, all similar. I Platyops sterreri n.sp. (fig. 2 A-L) j Description of the female - Dumpy, reminiscent of Heteromysis. i Frontal side of carapace with 2 laterally pointed apophyses and a weak 1 triangular prolongation between the eyes. Antenna 1 (fig. 2C) with the peduncle a bit longer than that of antenna 2, but slightly shorter than the scale of the latter. Peduncle provided with 2 setae at the end of the middle joint on the inner side and 2 others on the terminal joint. Between the flagelli, an oval, slightly crenulated apophysis with 2 sensitive hairs. Antenna 2 with base shorter than its scale (fig. 2D). The latter is oval, three times longer than its maximum width, with well delimited, rounded apical article. Its flagellum hardly exceeds the cephalothorax. The eyes, seen from above, immediately draw one's atten- tion to the shape of their white, vaguely rectangular plates that circumscribe a tiny visual, antero-lateral, shiny brown part (fig. 2B). In lateral view, cornea is oval and peduncle anteriorly strongly flattened chisel-like (fig. 2A). Labrum obtuse triangular. Mandible with common 3-segmented palp. Ex- opod and palp of maxilla like in fig. 2E, and endite of maxillule with 7 terminal spines and 2 fine fidate setae (fig. 2e). STYGOLOGZA 2 (112) 1986 101

Fig. 2 A-L, PlatyoPs stmeri n. g., n. sp., Q : A, eye in lateral view; B, eye seen from above and I scale of antena 2; C,antennula Q ; D, antenna 2; E, endite of maxillule; e, palp of maxilla; F, ' peraeopod I; G, peraeopod 11; H, peraeopod 111; J, pleopod IV; L, uropodal endopodite; K. telson. M-R Hetcromysri bmnudmris Sars, 1885, adult a = 6 mm: M, complex of phanerae of antenna 1; N, uropod; 0, telson; P, pleopod IV; R, pleopod 111. 102 BERMUDA MARINE CA VE SYMPOSIUM

Pereiopod I (third thoracic limb) short (fig. ZF), with 3-jointed tarsus, 3 serrated external spines at the carpus. Pereiopod I1 is exceptionally long (fig. ZG), almost three times longer than the others, with 6-segmented tarsus - the proximal segment twice as long as the others. Pereiopods 111-V of pereiopod I type, but with 5-segmented tarsus; exopods short, 8- to 9-segmented. Pleopods tiny, one-segmented (fig. 25). Uropods with branches equal in length and with endopodite lacking spine. Statolith small, perfectly spherical (fig. 2L). Telson vaguely triangular (fig. ZK), armed on its distal quarter with fine pointed spines: 3 lateral, 2 long (almost 114 of telson length) latero-apical, and 2 median on the narrow truncated apex of telson. An alveolar tissue on the sides of the distal half. Length of specimens: 2.5 mm.

Material. - Two Q (2.5 and 3 mm long), each of them with 2 embryos, from Castle Grotto (St. 3) and 3 specimens (1 Q with 2 embryos and 2 Q - 2 and 2.5 mm) from Green Bay Cave (St. 2). Specimens of Heteromysis bermudensis and Bermudamysis speluncola were also found at St. 2.

Remarks. - As we do not have males at our disposal, we assign this genus, with a certain hesitation, to the Mysini tribe. At least the only equivalent to the structure of the eye is in the genus Heteromysoides Bacescu, 1978. i.e. H. speluncola (Bacescu, 1968). The main feature of Platyops - whence the name - is certainly the widened eye which is strongly dorso-ventrally flattened, with latero-external visual part. Accordingly, the genus achieves an impressive convergence with the genus Heteromysoides. The element which brought about this lateral vision is still unknown.

Heteromysis (Olivaemysis) aff. guitarti Bacescu, 1978 (fig. 1, K-0).

This mysid, which was found in Green Bay Cave (St. 1, 1 Q and St. 3, 1 0 and 2 9 ), is closely related to H. (0livaemysis)guitarti from Cuban waters. They have in common the phanerae of antenna 1 (fig. lK), the armature of telson all along its sides, and the morphology of pleopod IV 0 (fig. 1 0) as well as the deep sinus of the telson, but this is provided in the Bermuda specimens with much longer and more numerous laminae - more than 25 (fig. 1P). The complex of phanerae of antenna 1 is of the Olivmysis type (fig. 1K). Eyes short, cylindrical, with peduncle twice as high as the cornea, ending in an antero- lateral apophysis. Conical penis, with a split apex (fig. 1P). Q St. 1 with 6 embryos. The Bermuda specimens also differ from the Cuban population by having more tarsal articles (4-8) at peraeopods, more lateral spines at telson (12 instead of 9-10), more laminae (27 instead of 16-21), more flagellate setae at pleopod IV a (fig. 1 0)(8 instead of 2-3), and a smaller size (6 instead of 4.5 mm) - perhaps the small number of phanerae comes from this small size (maybe a local subspecies). Tarsus of pereiopod IV (fig. 1L) 9-articulated.

Geographic distribution. - H. guitarti typica was described by M. Bacescu (1968) from sponges in Cuban waters. Subsequently, it was found by B. Brat- tegard (1970) in the Bahamas (Andros Island), by Modlin (1984) at the Florida Middle Ground, and now by us from a sponge in Green Bay Cave, Bermuda.

ACKNOWLEDGEMENTS This research was supported by the National Science Foundation (Grant BSR-8215672 to T. Iliffe). Long term scientific exchange visits by T. Iliffe to Romania in 1981 and 1983 for joint consultations and studies were made possible by the U.S. National Academy of Science and the Academy of the Socialist Republic of Romania. We thank Robert Power and Mary van Soeren for assistance with cave diving collections.

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Received 23 April 1985