Description of a New Hirsutiid (N.G., N.Sp.) and Reassignement of This Family from Order Mictacea to the New Order, Bochusacea (Crustacea, Peracarida)

Trav. Mus. natl. Hist. nal. «Grigore Antipa>.> Vol. XL

DESCRIPTION OF A NEW HIRSUTIID (N.G., N.SP.) AND REASSIGNEMENT OF THIS FAMILY FROM ORDER MICTACEA TO THE NEW ORDER, BOCHUSACEA (CRUSTACEA, PERACARIDA)

MODEST GUrU, THOMAS M. ILIFFE

Confonnement aux plusieurs exemplaires provenant dans les anchialine et submarine grottes de Bahamas, Thetispelecaris remex n.g., n.sp. appartenant a la famille Hirsutiidae Sanders, Hessler et Gamer, 1985, est decrit. Ensuite une analyse detaille des caracteres morphologiques nous avons constate que 1'inclusion de la famille mentionee dans I'ordre Mictacea Bowman, Gamer, Hessler, Tliffe et Sanders, 1985 est artificiel. Parce-que les caracteres des especes de Hirsutiidae ne se trouvent pas dans aucun ordre de crustaces peracarides nous decrivons Ie nouveau ordre Bochusacea, avcc trois especes: Hirsutia bathyalis Sanders, Hessler et Gamer, 1985, H. sandersetalia Just et Poore, 1988, et Thetispelecaris remex, n.g., n.sp. Keywords: Bochusacea, new order (Crustacea, Peracarida), Hirsutiidae, Thetis pelecaris remex n.g., n.sp., cave, Bahamas.

In 1985 Sanders, Hessler and Garner described Hirsutia bathyalis, and Bowman and Iliffe, Mictocaris halope, each species representing a new genus and a new family of peracarid crustaceans. The analysis of the morphological features made by the above-mentioned authors led them to the conclusion that the two species have some common features which did not occur in other peracaridan orders, and, therefore, they described order Mictacea being formed of the mentioned species. Whereas the description of the species Mictocaris halope was made studying numerous specimens (males, females, juveniles and manca), that of Hirsutia bathyalis was based on very limited material. The authors had at their disposal a single, partially damaged female, which, in addition "was embedded in a medium that later proved to be so caustic that serious deterioration resulted" (Sanders et al., 1985, p. 34) and a "molt of one 94 MODEST GUrU, THOMAS M. ILIFFE juvenile". Thus, minute and comparative observations on multiple specimens was impossible. Three years later, Just and Poore (1988) described a new species of Hirsutia (H.sandersetalia), also based on a single specimen, which "was damaged during dredging" (Just and Poor,1988, pA83). In 1993 and 1996 one of the authors of this papcr (T. I1iffe) collected 21 specimens (16 females, 2 of them very damaged, and 5 manca) from anchialine and submarine caves in Bahamas that belong to a species very much like those from the genus Hirsutia. The good condition of most of the collected specimens permitted us to make reexamine the so-called "oostegites", which confirm the opinion of Sanders et al. (1985, p. 55), about necessity "to erect a new order" for the species of Hirsutiidae. This new order is described herein.

BOCHUSACEA, new order Diagnosis. Cylindrical body. Cephalon fused with one thoracic somite. Pereon formed of seven free pereonites. Pleon with six pleonites. Telson free. Eye lobes absent. Antennule (antenna 1) with three-segmented peduncle and two flagella. Antenna (antenna 2) with squama. Mouthparts, completely uncovered (laterally unprotected by the carapace), large sized (in comparison with other appendages), with numerous long setae together being brush-like antero-ventrally directed. Mandibles with three-segmented palpus; lacinia mobilis present in left mandible; setal row present; pars molaris well developed. Labium (paragnaths) having on each part, very long and thin spiniform or palpiform lobe. Maxillule (maxilla 1) with two endites and without palpus; the outer (distal) endite rostrally widened with numerous setiform-spiniform elements of different fornls and lengths; inner (proximal) endite smaller than the outer one, with five distal spiniform elements, outer one being brush-like. Maxilla (maxilla 2) with strong and long external seta; outer lobe bilobed, with long setae on rostral margin; inner lobe with setae spread up to base. Maxilliped with long and wide basis, rostrally elongated with enditc with numerous setae on mesiodistal margin; palpus formed of five thin segments. Pereopod 1 palpifonn (resembling a maxilliped), seven-segmented, very long, thin and equally thick, with or without exopodite; pereopods 2-7 ambulatory with different dimensions and morphology, only last one lacking exopodite; pereopodal 2 exopodite three-segmented; pereopodal 3-6 exopodite four-segmented; at least, females have inner epipodites on pereopods 2-6, assisting in sustaning eggs. Pleopods very small, uniramous, in five pairs. Uropods A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 95 biramed, with cylindrical rami; exopodite bisegmented and the endopodite multi-segmented. During development, manca stage characterized by the absence of the last pair of pereopods and epipodites; pleopods present. Etymology. From the Latin bochus, "which has a prominent mouth" (referring to the mouthparts completely uncovered and prominent) and acea, the ending occurred in many crustacean orders. Composition. New order consists of three marine species (of two genera), all of them belonging to family Hirsutiidae Sanders, Hessler and Gamer, 1985, as follows: Hirsutia bathyalis Sanders,Hessler and Gamer, 1985, H.sandersetalia Just and Poor, 1988 and Thetispelecaris remex n.g., n.sp. The first two are deep water species (1,000 m and, respectively, 1,500 m), and the third one of shallow water, originating in anchialine and submarine caves. Distribution. The Tasmanian Sea for H.sandersetalia and the West Central Atlantic for the other two species. Remarks. In Peracarida, with few exceptions, oostegites consist of fine oval membranes, devoid of setae. They develop continuously as the moment of egg laying approaches. Therefore, various females in a collection tend to have oostegites unequal in size. Rarely will two females have equally large oostegites. Finally, oostegites "fuse" forming a closed pouch (marsupium) where the eggs and manca states reside and subsequently, the marsupium detaches. In contrast with this process which occurs in most of peracarids, the two known species of Hirsutia, as well as in the 16 adult specimens of our species, "oostegites" (corresponding to a certain pereopod) have almost the same dimentions. This would be as if all the females of the three species (collected in various places and at different time periods) reached sexual maturity simultaneously, this being highly unlikely. (Compare the "oostegites" of pereopods 5, for example, from the figures presented by Sanders et al., 1985, Fig. 14 with those of Just and Poor, 1988, Fig. 3, and with those from the present paper, Fig. 11 A, to which only the length and number of the setae are different). It is true that we have not remarked the presence of "oostegites" in the five manca specimens that we had at our disposal. Four of them had not developed the last pair of pereopods, while the other one had thcse pereopods in the final state of their development, with propodus and dactylus still undeveloped. However, we found normal "oostegites" in a small sized specimen (slightly bigger than a manca), but with the last pereopod already formed. So, "oostegites" are formed immediately after the development of the last pair of pereopods, that is in the juvenile state, being probably permanent structures (epipodites), which have different functions. Possible functions 96 MODEST GUrU, THOMAS M. ILIFFE

for these epipodites could include respiration or supporting egg laying, without these "oostegites" fusing and forming a marsupium, as in other peracarids. At most, the plumose setae could develop and form a kind of "net" for retaining the eggs. If this hypothesis is correct, the method of egg laying would resemble that of most decapod crustaceans, excepting that, in decapods, eggs are retained on the abdomen and are supported by pleopods. Therefore, we believe that what in Hirsutiidae seems to be "classical" oostegites (temporary developed for sustaining the eggs), are permanent epipodites. Besides the aforementioned functions (respiratory and of support of egg laying), epipodites could assist in drawing the water currents created by the pereopods. Another argument supporting the existence of the epipodite could be that in peracarids, the last oostegite pair (excepting mysidaceans) develops on the corresponding pereopod to thoracomere six, no matter the respective species have the bead fused with a thoracic somite (as in amphipods, isopods or the species Mictocaris halope) or with two-three somites (as in tanaidaceans and cumaceans). However in Hirsutiidae, the last "oostegites" pair is at the level of thoracic somite seven. The exception from mysidaceans is probably due to the presence of a highly developed carapace that "pushed" the oostegites on the last thoracic somite and thus, it . cannot be compared with the Hirsutiidae. Finally, the inclusion of family Hirsutiidae in the same order with Mictocaris halope does not seem reasonable. In support of this conclusion we call attention to the completely different form of pereopods and their exopodites, of uropods, etc. Thus in Hirsutiidae, pereopod 1 is different from the following pereopods (but also from pereopod 1 of Mictocaris) both from morphological and functional point of view and it has the following particularities: - it is long and thin, with uniform thickness; - basis is curved antero-ventrally, following the convex form of the cephalothorax, thus permitting free movements (back-forth and left-right) without disturbing or being disturbed by the other closed appendages (maxilliped and pereopod 2); - from the level of the joint ischium-merus, it changes its direction, carpus is taking an antero-Iateral direction, parallel with the ventral side of cephalothorax and almost perpendicular to the vertical plan of the basis; - carpus and propodus, very long and slightly curved towards the longitudinal axis of the body, give it a form like a sickle directed latero rostrally, which makes lateral movements in an horizontal plan; A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 97

- dactylus (as thick as the other segments) resembles a terminal segment of a palpus, having mesiodistally, an agglomeration of fine setae (curved inwards), which underlines the sikle form. From a ventral view, the pair of pereopods I have the form of two sickles opening face-to-face, inside the maxillipeds (Fig. 6 C). Thus, the lateral movements from the horizontal plan double the movements of the maxilliped palpus, helping the latter from the functional point of view. The strong musculature of the body (as of the telson and uropods) visible through the transparency as well as the morphology of the other pereopods lead us to believe that the Hirsutiidae are swimmers. Within this context, pereopod 1 has the role of generating a water currents flowing towards the mouthparts. Very small food "particles" are drawn with the water and retained by the numerous long, fine setae of the mouthparts (placed as a thick brush, directed antero-ventrally) which acts as a filter. In other words, pereopod 1 have a similar role to that of the maxilliped palpus but with the difference that they can move with a larger amplitude, thus assisting in obtaining food. When the animal does not swim, but floats at rest or sits on the substratum, pereopod 1 draws the food particles by a permanent movement, either due to the water current or detaching them from the substratum and sending them to the mouth. In consequence, we have no doubt that in Hirsutiidae pereopod 1 is specialized in feeding, not for locomotion; it could be considered maxilliped 2. In Mictocaris halope pereopod 1 resembles the other pereopods and is used in locomotion. In this way, the species of Hirsutiidae have six pairs of locomotor pereopods, and Mictocarididae, seven. Also, pereopod 2 is equally sized and similar in structure to the following ones in Mictocarididae, while in Hirsutiidae, it is the largest and different. Regarding pereopods, we note the resemblance between Mictocaris and Hirsutiidae, due to the presence of exopodites. But, making a careful analysis, we can remark two essential differences, at least, as concern the segment number which forms these exopodites and their presence or absence on the last two pairs of locomotor pereopods. While in Mictocaris the exopodites are bisegmented (with long second segment) and are absent on the last two pairs of pereopods, in Hirsutiidae, they are absent only on the last pereopod (and accidentaly on the first one) and the segment number is variable: two (on pereopod I as in the new genus described in this paper), three (on pereopods 2) or four (pereopods 3-6). Other particular features concern the uropods. In Hirsutiidae, the endopod is cylindrical and multisegmented (as in tanaidaceans, although 98 MODEST GUTU, THOMAS M. ILIFFE they are filamentous), while in Mictocaris it is widened and formed of only two segments, as in other many peracaridans. In addition to the above-mentioned assertions, we add the different form ofthe labium. In Hirsutiidae, each side of the lower lip has a very long rostral prolongation, which seems to be jointed at its base, as in the labium lobe from tanaidaceans, with the difference that in the latter, it is much smaller and of a different form. In the new genus described by us, this labial "lobe" is like a unisegmented palpus, very long with numerous hairs, which suggests us that it also directs the food "particles" to the masticatory pieces ofthe mandibles. Also, at the level ofmouthparts in Hirsutiidae, the maxilla has a very long strong seta (with long hairs), jointed at the base of the outer lobe that, in our opinion, has the same role, of "pushing" food particles to the mouth. As we have mentioned before, as a consequence of what we have presented, we cannot include Hirsutiidae in the same order with the species Mictocaris halope. As a matter of fact, the senior author (Gutu,1998) considers that Mictocaris halope is closely allied with Spelaeogriphacea. More than that, the way in which the main morphological features of the three species of Hirsutiidae are "gathered" proves that they belong to another order and, in the same time, to a new order. At present, it is very difficult to make definitive assertions on the relationship with other orders. According to Bowman and Abele's classification (1982), which we agree, we includ this new order among peracarids. Only the future examination ofmales and ovigerous females can fully clarify the problem of their relationship. In any event, the presence of the epipodites in females which, maybe, have also play the role of oostegites (correlated with the other morphological features) makes us to belive that we are dealing with the most primitive order of Peracarida, placed at the "border" with Syncarida. Therefore Sanders et al. (1985, p.55) could be right when they ask themselves ifHirsutiidae are really peracarids.

Family HIRSUTIIDAE Sanders, Hessler and Gamer, 1985 Remarks. The new diagnosis of the family coincides with that of the new order, Bochusacea.

Thetispelecaris n.g. Type species: Thetispelecaris remex n.sp. Diagnosis. Labium (paragnaths) with palpiform rostral prolongation distally covered with long hairs. Pereopod I with exopodite. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 99

Etymology. From the Greek words Thetis, "mythological marine god", spelaeon, "cave" and karis "shrimp", making reference to the fact that the type species is a "shrimp" originating in the anchialine caves, in comparison with the other genus known only from the open sea. Gender, feminine. Remarks. The most obvious difference between the new genus, Thetispelecaris and Hirsutia is the absence, in the latter, of the pereopod I exopodite. Another differentiating feature is present at the labium level, which in Hirsutia has a spiniform prolongation, devoid of hairs in the distal half, while in the new genus, labium has a rostral prolongation resembling a very long palpus, with numerous hairs at the distal end. Pereopod 2 has only four distal spines on propodus in Thetispelecaris n.g., and in Hirsutia it has nine. Also, other differences can be remarked at the level of the other pereopods.

Key of genera and species of family Hirsutiidae and in the same time of Bochusacea, new order

1 - Pereopod 1 with exopodite Thetispelecaris (T.remex n.sp.) - Pereopod 1 without exopodite Hirsutia 2 2 - The third segment of maxillipedal palpus do not exceed the length of the rostral margin of endite H.sandersetalia - The third segment of maxillipedal palpus exceed much the length of the rostral margin of endite H. bathyalis

Thetispelecaris remex n. sp. (Figs 1-12) Material: 21 specimens (16 females, 2 of them very damaged, and 5 manca); Holotype female, No. BOC 001 ("Grigore Antipa" National Museum of Natural History in Bucharest, Romania), from Station 93-004, Norman's Pond Cave, Norman's Pond Cay, 6 May 1993, collected with a 93 ).lm plankton from fine silt on ledges in the dark, rear section of the first room at 6 m depth of this inland blue hole cave, leg. T. Iliffe. Paratypes: 3 females, No. BOC 002, from Station 93-004; 4 females and 2 manca, No. BOC 003, from Station 96-031, Master Harbour Cave, Great Exuma Island, 9 September 1996, collected with a 93 ).lm plankton from the ceiling in 12-15 m depth of this ocean blue hole cave; 4 females and 2 manca from Station 93-004 were deposited in the collections of the National Museum of Natural History, Washington; 2 females and I manca from the Station 100 MODEST GUTU, THOMAS M. ILIFFE

06-030, Mystery Cave, Stocking Island, 8 September 1996, collected with a 93 !-tm plankton from the sandy bottom and walls in 50 m depth of this ocean blue hole cave are in the senior author's collection; leg. T. Ilitle.

Description ofthe female Body (Fig. 1) semi-transparent, cylindrical, seven times longer than maximum width. Length (with telson), 1.2-1.6 mm. Head (Figs 1 and 2), slightly longer than wide; anterior part, rostrally narrowed, like a dorsal shield without laterally covering mouthparts; posterior part (fused with first thoracomeres) wider and partially covers lateral sides; anterior and posterior parts having between them narrow slightly visible depression. Rostrum pointed, slightly curved ventrally. Eye lobes absent. Pereon (Fig. 1) formed of seven free pereonites, wider than long and rounded on sides; first pereonite shortest, practically invisible under optical microscope; pereonites four-six equal, slightly longer than others. Pleon (Figs 1 and 8 B) formed of six free pleonites, cylindrical, almost equal, slightly thinner, but longer than pereonites; last pleonite slightly swollen. Telson (Figs land 3), almost as long as a pleonite, is flattened dorso ventrally, narrowed terminally, and with four spines on each side; four terminal setae, two dorsal, smaller, and two ventral, stronger and longer. Antennule or antenna J (Figs 2 and 4 A, B), biramous, with the peduncle formed of three short segments, decreasing in length towards last segment; distally, each segment has long setae. Exopodite short, formed of four small segments, first two being longer than last; terminally, next to last segment with two aesthetascs, last one with two setae. Endopodite, longer than exopodite, formed of three-four thin segments, first one being almost as long as last segment of peduncle; following two segments slightly shorter than first, last one smallest. Antenna or antenna 2 (Figs 2 and 4 C, D), biramous, and longer than antennule; peduncle formed of five segments, each of first three being shorter than any of last two; squama of small sizes, with four terminal setae, two of them very long; last segment, peduncular, with mediodistal prominence and five-six long setae. Flagellum, formed of five-six cylindrical segments, is almost as the last two segments of peduncle; last segment of flagellum has three setae, one of them being very long. Labrum (Fig. 5 A ), trapezoidal, exceeding anterior margin of carapace (Fig. 2). A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 101

Fig. 1 -Thetispelecaris remex, n.g.,n.sp., female, paratype: body, dorsal view. 102 MODEST GUrU, THOMAS M. ILIFFE

Fig. 2 - Thetispelecaris remex, n.g.,n.sp., female, paratype: head and first two free pereonite, with the appendages, dorsal view. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 103

Fig. 3 - Thetispelecaris remex, n.g.,n.sp., female, paratype: last pleonite, with tclson and uropods, dorsal view. 104 MODEST GUrU, THOMAS M. ILIFFE

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Fig. 4 - Thetispelecaris remex, n.g.,n.sp., female, paratypes: A and B, antennule of two different specimens; C and D, antenna of two different specimens. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 105

Mandibles (Fig. 5 C, D) having well developed body, exceeding lateral margin of carapace (Fig. 2). Palpus present, three-segmented, ending with three strong and long setae. Pars incisiva thin, ending with three denticles in both mandibles. Lacinia mobilis, also thin and three-dented, present only at left mandible (Fig. 5 D). Setiferous row different in the two mandibles: left mandible with four fine setae, followed by an area with hairs and then group of 12 long setae (Fig. 5 D); right mandible with three spiniform thick setae and 14-15 long and fine setae, without space between two kinds of setae (Fig. 5 C). Pars molaris well developed (Fig. 5 C). Labium or paragnaths (Fig. 5 B) with two sides oval, about 2.4-2.5 times longer than wide. Narrow distally, with long, thin process (covered with long hairs), palpus-like, seemingly jointed with labium body; subterminally, on inner margin, having two setiform-spiniform processes and prominence resembling (by its face-to-face position with its symetrical) processus molaris from two mandibles; rostral-inner area has dense, very long hairs; outer margin with short hairs. Palpi (corresponding to lobes in tanaidaceans), considerably exceeds length of anterior margin of carapace (Fig. 2). Maxillule or maxilla 1 (Fig. 6 A) without palpus; endites well developed. Outer endite (distal), bigger than inner one (proximal), with six setae on laterodistal margin, thick and widened terminally, continuing on rostral margin with other four setae like piano hammers and six spiniform setae, bifid terminally; subterminally, two plumose setae. Inner endite (proximal), thinner and shorter than distal endite; five spiniform setae on rostral margin, outer one brush-like (Fig. 6 A); following three setae with a distal gap each, fifth one simple. Maxilla or maxilla 2 (Fig. 6 B), longer than wide, has the outer (mobile) endite formed of two lobes with very many setae destributed in three rows on rostral edge; near the base of outer lobe, very long, thick, plumose seta joints (considerably exceeding length of rostral setae oflobe); inner (fixed) endite shorter than outer one, rostrally cut, with numerous setae, different in shape, continuing on inner margin of maxilla, up to base. Inner endite also seems formed of two lobes, partially superposed, difficult to be seen with optical microscope. Maxilliped (Figs 6 C and 7 A, B) without endopodite. Basis long and narrow, elongated rostrally with endite with different kinds of setae distally (Fig. 7 B); inner margin of basis also with long setae, one distaly placed seems wider and membranous. Palpus, thin and thick uniformly, fanned of five unequal segments, each with one-three simple setae; first segment (ischium) shortest, while second (merus) equal with fifth one (dactylus) 106 MODEST GUTU, THOMAS M. lLIFFE

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Fig. 5 - Thetispelecaris remex, n.g.,n.sp., female, paratype: A, labrum; B, labium; C, left mandible; D, right mandible, rostral half. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 107

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Fig. 6 - Thetispelecaris remex, n.g.,n.sp., female, paratype: A, maxillule; B, maxilla, C, head and first two pereonite with maxilliped and pereopod 1, ventral view (the mouthpart was removed). 108 MODEST GUru, THOMAS M. ILIFFE

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Fig. 7 - Thetispelecaris remex, n.g.,n.sp., female, paratype: A, maxilliped; B, ditto, rostral end of endite; C, pereopod l; D, ditto, dactylus. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 109 and, slightly shorter than the fourth segment (propodus); third segment (carpus) twice longer than first two, measured together. Pereopod 1 (Figs 2, 6 C and 7 C, D) long and thin, thick uniformly, formed of seven segments. Coxa short. Basis, slightly curved, medianly directed; proximally having one seta, and, distally, two; laterally, having short exopodite, formed of two segments, second one being much smaller, ending with two setae. Ischium continues curved shape of basis, about three times shorter than latter. Merus short; by type of joint with ischium, it changes direction of carpus from middle of body to lateral side so that carpus forms right angle with vertical plan of basis. Carpus long (almost as basis) with three unequal and long setae on antero-inner margin. Propodus, almost as long as carpus or basis (but slightly thinner), very small spines on inner margin, three-four setae, distally. Dactylus short (almost as ischium) having same thickness at base as propodus, thinner at terminal end; mesiodistaly having five thin, unequal satae, as well as two setiform wide and membranous (Fig. 7 D). Carpus, propodus and dactylus together sickle like which moves horisontally (Figs 6 C and 7 C). Pereopod 2 (Figs 1 and 8 A) thicker, longer and different from others. Coxa small, having on inner side short and narrow epipodite-oostegite with four plumose setae. Basis, about 3.5 times longer than thick; in first half of posterior margin on three-segmented exopodite with long plumose setae. Ischium short, with two distal setae. Merus (twice longer than ischium, but shorter than the other segments) having five long setae distally, two anterior, and three posterior. Carpus (shorter than basis, but longer than other segments) having distally three strong spines and long seta. Propodus, slightly shorter and thinner than carpus, having distally four strong spines, short ciliated seta and two long, simple setae. Dactylus, as long as propodus, but thinner, having two strong spines (one slightly curved) placed terminally, as well as two very short setae. Pereopod 3 (Fig. 9) smaller than previous one, and with several very long fine setae. Coxa, small, having a well-developed epipodite-oostegite on the inner side, only slightly shorter than basis; epipodite-oostegite on inner side, only slightly shorter than basis; epipodite with four long plumose setae on outer side, other two terminal and one subtem1inal on inner side. Basis (3.5 times longer than thick) having two long plumose setae in first half of anterior margin, and antero-distally four simple setae; proximally, large exopodite, four-segmented (slightly shorter than basis), with very long plumose setae. Ischium short, with three antero-distal setae. Merus, longer but thinner than ischium, having six spines and seven long setae spread as illustrated in Fig. 9. Propodus, as long as carpus but thinner, 110 MODEST GUTU, THOMAS M. ILIFFE

Fig. 8 - Thetispelecaris remex, n.g.,n.sp., female, paratype: A, pereopod 2; B, pleon with pleopods and telson, ventral view. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA III

Fig. 9 - Thetispelecaris remex, n.g.,n.sp., female, paratype: pereopod 3. 112 MODEST GUrU, THOMAS M. ILIFFE

Fig. 10 - Thetispelecaris remex, n.g.,n.sp., female, paratype: pereopod 4. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 113

Fig. II - Thetispelecaris remex, n.g.,n.sp., female, paratype: A, pereopod 5; B, pleon with pleopods, lateral view. 114 MODEST GUrU, THOMAS M. ILIFFE

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Fig. 12 - Thetispelecaris remex, n.g.,n.sp., female, paratype: A and B, pereopods 6 and 7; C-G, pleopods 1-5, lateral view. A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 115 having on anterior margin four spiniform long elements and six postero distal setae, very long. Dactylus thin, as long as merus, having a setiform spine, also very long, terminally. Pereopods 4 and 5 (Figs 10 and 11 A) resemble pereopod 3. Small differences in size of the epipodite-oostegite and exopodite (which are bigger), in the number of spines on carpus and propodus, as well as in that of setae. Pereopod 6 (Fig. 12 A) is smaller than previous one, with propodus much longer (according to its thickness, or in comparison with other segments) and with less setae. The epipodite-oostegite and exopodite smaller than the previous ones. Pereopod 7 (Fig. 12 B) almost as long as the previous one, but thinner, with less setae and without epipodite-oostegite on coxa or exopodite on basis. Exopodites ofthe thoracopods (Figs 7 C, 8 A, 9, 10, 11 A and 12 A), counting six (on pereopods 1-6), are fonned, as we have already mentioned, of two segments (pereopod 1), of three (pereopod 2) or four (pereopods 3 6). Exopodites of pereopods 3-5 are largest, while exopodite of pereopod 1 is smallest. Epipodites-oostegites (Figs 8 A, 9,10,11 A and 12 A) present on pereopods 2-6, least developed being those of pereopod 2, largest belong to pereopod 3-6. Pleopods (Figs 8 B, 11 and 12 C-G), in five pairs, are very small, uniramous and unisegmented, with one-three small setae. First three, tubercle-like and not visibly jointed. Uropods (Fig. 3) biramous, with cylindrical and long rami. Protopodite thick (slightly shorter than length of last pleonite, which it is articulated) having flagellated spine mesiodistally, and latero-outerly, two short setae. Exopodite formed of two thin cylindrical segments, almost equal and as long as the protopodite; first segment with two setae (one inner and one outer), and two laterodistal spines, while second segment has seven fine and long setae, three of them apical. Endopodite, slightly longer than exopudite, is formed of five cylindrical segments having flagellated robust spine each, mesiodistally and seta each, distalo-outwardly; last segment with four long setae terminally. Manca stage. Smallest specimens (0.80-0.87 mm long) lack last pair of locomotor pereopods and of epipodites-oostegites. Both exopodites of the thoracopods and the pleopods are present. In four specimens, pair of 116 MODEST GUTU, THOMAS M. ILIFFE small tubercules on the last pereonite corresponding to last pair of pereopod. In slightly larger specimen (measuring 0.95 mm) last pair of pereopods formed almost entirely: left pereopod 7 with dactylus devoid of apical spiniform seta and its pair, right one, with respective seta short, claw-like. Data on inner morphology. Although our study was not aimed at examining the inner morphology, the semi-transparency of the specimens allowed us to make some observations which we think are worth mentioning. Thus, we observed the presence of a very well developed muscular system, represented by a muscular band on each side of the body (with thin fascicles which are jointed on the level of every segment of the body), visible from the head to the inside of the telson and ofuropods (Fig. 3), with ramifications at the level of each pereopod. Alimentary canal simple (possible devoid of diverticles) goes through the entire body to the anal opening, which is placed at the distal end of the telson. Breathing organs are represented, in all probability as we have shown - by epipodites-oostegites. Female organs of reproduction consists of two bag like ovaries which stretch from the midpoint of thoracomere seven to the middle of the third pleonite. Observations made on the 19 specimens demonstrated the following aspects: the absence of the ovarian bags in the manca specimens with dimension of 0.80 and 0.95 mm; reduced dimension of the ovarian bags (corresponding to the last free pereonite and to the first two pleonites) in females of a length between 1.20 and 1.32 mm, which may not have reached maturity; and ovarian bags, completely developed, which can be observed from the level ofthe last free pereonite to the middle of the third pleonite (Fig. 1) in larger specimens, with a length of 1.47 1.62 mm. Variability occurs in the number of the segments of antennule flagella; which can be three instead of four in some specimens, or in antenna flagellum, which can be five instead of six (Figs 1, 2 and 4 A-D). At the level ofpereopod 3, we observed in a specimen the presence of three spiniform elements on propodus and not four as in its symmetrical. Etymology. From the Latin remex, "oarsman", refering to the fact the species is an active swimmer, to all appearances. Type locality. Norman's Pont Cay, Exuma Islands, Bahamas. Remarks. Because of the very small size of the species, we could not establish accurately the shape of the carapace and of some microstructures, especially those of the second thoracomere (the first free pereonite). In three specimens (with molt aspect), we observed seven free pereonite, but A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 117 in all other ones, the first free pereonite is, practically, invisible. This disagreement is present, too, in the papers of Sanders et al. (1985) and Just and Poor (1988). While Sanders et al. (1985, Fig.l B) illustrates the pereon with seven pereonites (the first well sketched, but very short, as it is mentioned in their description, p.31), in Just and Poor's drawing (1988, Fig. 1) only six free pereonites appear, the smallest one (that is the first, according to Sanders et al.) being completely absent. We hope that future collections will succeed in obtaining males and ovigerous females that will permit us to continue to study this species, using SEM photos to clarify some aspects that remain uncertain. Habitat. The Exuma Island are situated along the eastern rim of the Great Bahama Bank, a shallow water carbonate platform also including the islands of Andros, New Providence, Eleuthera, Cat, and Long. On the east side of the Exuma Islands is the Exuma Sound, a steep sided submarine valley reaching oceanic depths to more than 1800 m. Extensive anchialine and submarine cave systems are present along the margins of the platform. The developement of these caves may be related to joint patterns in the Pleistocene limestone. Viewed from the air, the deep blue color of the circular sinkhole cave entrances, has resulted their being named "blue holes". These blue holes can occur either on islands (referred to as inland blue holes) or in shallow waters of the bank or shelf (referred to as ocean blue holes). Inland blue holes are circular, often deep, water-filled shafts that bell out beneath the surface into a wide underwater cavern. A few have solutional cave passages associated with them. Due the terrestrial influences of such sites, inland blue holes are properly classified as anchialine caves (Stock et al., 1986). Ocean blue holes are opening to extensive, strongly tidal, submerged cave systems. Most of the caves associated with ocean blue holes are fault caves with parallel wells and collapse rock on the floor. Since they lack terrestrial influences, ocean blue holes are classified as submarine caves. Due to the extremely strong, reversing tidal currents that flush ocean blue holes on a diurnal cycle, few troglobitic species are typically found in them. Norman's Pont Cave is an inland blue hole cave located near the northern tip of Norman's Pont Cay. The entrance is a 2 m wide by 8 m long sinkhole situated just above the high tide line. The cave has been explored horizontally for 210m, reaching a depth of 86 m (Brian Kakuk, pers. commun.). The cave consists of a collapse-floored fissure up to 8 m wide that extends under the island and trends toward the open waters of Exuma 118 MODEST GUrU, THOMAS M. ILIFFE

Sound. Because of its proximity to the coast, waters in the cave are fully marine. Remipedes, amphipods (Bahadzia sp.), cyclopoid, harpacticoid, and calanoid copepods, ostracods (Spelaeoecia styx and Danielopolina exuma), cumaceans and archiannelids have been collected from the cave (Kornicker and Iliffe, 1998). Mystery Cave is located on the southwest side of Stocking Island in an almost totally enclosed bay that faces Elizabeth Harbour. The entrance to this ocean blue hole is situated at 6 m depth in a vertical rock wall forming one edge of the island. The cave trends gradually downward, extending under the island, as a 5 m diameter fissure passage, somewhat similar in configuration to Norman's Pond Cave. Very strong, revising tidal currents sweep through this section of the cave and all exposed rock surfaces are covered by sponges and other encmsting species. At depth exceeding 50 m, the passage becomes much narrower and partially filled with breakdown boulders. Recently, an upper level passage has been discovered which leads to more hydrologically isolated section of the cave where anchialine fauna including remipedes have been observed. Salinity in the cave is fully marine. Ostracodes, cumaceans, mysids, amphipods and nebaliaceans none apparently troglomorphic - were also collected. Master Harbour Cave is located in Master Harbour, about 4 km southeast of George Town on the northeast coast of Great Exuma Island. The cave entrance consists of an elongated fissure at 6-8 m depths in a submerged sinkhole. The fissure becomes roofed over at one end and continues as a narrow, 1-3 m wide by up to 10 m high rift. Strong tidal currents move through the cave such that divers can only enter it at slack tide when the water flow decreases and changes direction. The cave has been explored horizontally for more than 300 m, reaching water depths of 30 m. Like Mystery Cave, all exposed rock surfaces are covered with encmsting organisms. Salinity in the cave is fully marine. Ostracodes, copepods, mysids, brachiopods, polychaetes, amphipods, shrimps, bryozoans and cumaceans - none apparently troglomorphic - were also collected.

ACKNOWLEDGEMENTS

This research was supported by grants from the Caribbean Marine Research Center (CMRC) of the National Oceanic and Atmospheric Administration (NOAA) and from the National Science Foundation (NSF #9870219). We thank John Pohlman and Brett A NEW HIRSUTIID (N. G., N. SP.) AND A NEW ORDER, BOCHUSACEA 119

Dodson (Texas A&M University) and Brian Kakuk (CMRC) for assistance with cave diving collections. We also thank Mrs. Adriana Stoica and Mrs. Petruta Dumitridi, both from "Grigore Antipa" National Museum ofNatural History, Bucharest; the first for inking the illustrations and the second, for word-processing. The authors wish to sincerely thank Dr. Mary Wicksten of Texas A & D University and Dr. Frederick Schram of the University of Amsterdam for reviewing the manuscript and providing helpful remarks. However, we must note that after careful consideration, we did not follow all their suggestions.

DESCRIEREA UNUI NOU HIRSUTlID (N.G.,N.SP.) SI TRECEREA ACESTEI FAMILII DIN ORDINUL MICTACEA IN NOUL ORDIN, BOCHUSACEA (CRUSTACEA, PERACARIDA)

REZUMAT

Descrierea familiei Hirsutiidae de ditre Sanders, Hessler ~i Gamer (1985) in baza unui singur exemplar (partial deteriorat) dintr-o noua specie nu a permis observatii complete. Dispunand de mai multe exemplare dintr un nou gen ~i 0 noua specie (Thetispelecaris remex, descris in aceasta lucrare) provenind dintr-o groai cu apa marina din Bahamas ~i apartinand aceleia~i familii Hirsutiidae, autorii au ajuns la concluzia di incadrarea ei in ordinul Mictacea Bowman, Gamer, Hessler, Iliffe ~i Sanders, 1985 (alaturi de familia Mictocarididae Bowman ~i Iliffe, 1985) este artificiala. Totodata autorii au constatat ca familia Hirsutiidae reune~te specii ale caror caractere nu se regasesc la alte ordine de crustacee, ceea ce a condus la descrierea unui ordin nou, Bochusacea, care cuprinde in prezent trei specii.

REFERENCES

BOWMAN, T.E., ABELE, L.G., 1982 - Classification of the Recent Crustacea. In: D.E. Bliss (Ed.), The Biology of Crustacea, 1:1-27. BOWMAN, T.E., GARNER, S.P., HESSLER, R.R., ILIFFE, T.M., SANDERS, H.L., 1985 - Mictacea, a new order of Crustacea Peracarida. J Crus t. Bio!., 5, 1:74-78. BOWMAN, T.E., ILIFFE, T.M., 1985 Mictocaris halope, a new unusual peracaridan crustacean from marine caves on Bennuda. JCrust.Biol., 5, 1:58-73. 120 MODEST GUTU, THOMAS M. ILIFFE

GUTU, M., 1998 Spelaeogriphacea and Mictacea (partim), suborders of a new order, Cosinzeneacea (Cmstacea,Peracarida). Trav. Mus. natl.Hist. nat. "Grigore Antipa ", 40: 121-129. JUST, J., POORE, G.c.B., 1988 Second record of Hirsutiidae (Peracarida:Mictacea): Hirsutia sandersetalia, new species, from southeastern Australia. 1. Crust.Biol.,S, 3:483-488. KORNICKER, L.S., ILIFFE, T.M., 1998 - Myocopcopid Ostracoda (Halocypridina, Cladocopina) from anchialine caves in the Bahamas, Canary Islands, and Mexico. Smithson. Contrib. Zool., 599: 1-93. SANDERS, H.L., HESSLER, R.R., GARNER, S.P., 1985 - Hirsutia bathyalis, a new unusual deep-sea benthic peracaridan crustacean from the tropical Atlantic. 1. Crust.BioI., 5, 1:30-57. STOCK, J.H., ILIFFE, T.M., WILLIAMS, D., 1986 - The concept "anchialine" reconsidered. Stygologia, 2, 1/2:90-92.

Received: July 20, 1998 Modest Gutu Accepted: August 13, 1998 Muzeul National de Istorie Naturalii "Grigore Antipa" $os. KiselefJ, 1; 79744 Bucure~·ti, 2 Romania

Thomas M.Iliffe Texas A & M University Department ofMarine Biology Galveston, TX 77553-1675; US.A.