Lucifuga spelaeotes Revision of the Bahamian cave-fishes of the genus Lucifuga (Ophidiiformes, Bythitidae), with description of a new species from islands on the Little Bahama Bank. Peter R. Møller Werner Schwarzhans Thomas M. Iliffe Jørgen G. Nielsen Zootaxa 2006 1223 23 46 3WBPN Bythitidae Lucifuga CoL Animalia Ophidiiformes 27 Chordata species lucayana  (Figs. 2-6, Tables 1-3)   Lucifuga spelaeotes(not Cohen & Robins, 1970): Yager 1981: 328; Cunliffe 1985: 108, photo; Palmer 1985a: 114, photo 51; Yager & Williams 1988: 102; Wilkens et al. 1989: 127 (eyes); Smith -Vaniz & Boehlke1991: 202 (part); Cohen & McCosker 1998: 184 (part); Proudlove 2001: 207 (part).  Material examined (6 specimens, of which 3 lack the head, 44-99 mm SL, due to earlier studies of the eye development (H. Wilkens pers. comm.). Identification of headless specimens based on finray and vertebrae counts).  Diagnosis. Lucifuga lucayanais distinguished from other members of the genus by the following combination of characters: Vertebrae 12-13+37-39=50-52, dorsal finrays 84-91, anal finrays 63-69, pectoral finrays 17-18, caudal finrays 10; head profile above eye strongly depressed, scales on occiput, eyes present (0.9-1.0 % SL); palatine teeth absent; color uniformly light brown, long gill-rakers 3, pale. Similarity. Lucifuga lucayanaresembles L. spelaeotesin all but a few characters listed in Tables 1-3. It differs in the lack of palatine teeth (vs. 3-56) (fig. 6), color of elongated gill-rakers (pale vs. dark) and the broadly scaled occiput (vs. occiput scaled only in central part) (fig. 4). Furthermore, L. lucayanain average have fewer dorsal finrays (84- 91, x=89.2 vs. 86-109, x=97.5), anal finrays (63-69, x=66.2 vs. 66-82, x=73.8) (Tables 1- 3) and perhaps smaller eyes (0.9-1.0, x=1.0 vs. 0.7-1.8, x = 1.3 % SL, fig. 5).  Lucifuga lucayanaresembles two Cuban species L. subterraneaand L. teresinarumin the lack of palatine teeth, but it differs from these and the two other Cuban species in several characters e.g. larger eye (0.9-1.0 vs. 0.0-0.3 % SL), and in the higher number of vertebrae(12-13 +37-39 = 50-52 vs. 11+34-37 = 45-48 (unknown for L. teresinarum) and caudal finrays (10 vs. 8) (Table 3). It further differs from the Cuban species L. dentataand L. simileby the lack of palatine teeth and number of anal finrays (63-69 vs. 69-78 and 57- 60, respectively).  Description. Meristic and morphometric characters are given in Tables 1-3. Body moderately elongate, compressed. Head profile strongly depressed (figs. 2-4). Eyes relatively small, 0.9-1.0 % SL (fig. 5). Upper jaw slightly protruding. Anterior nostril tubeshaped, placed low on snout near upper lip; posterior nostril a mere hole, larger, about midway between snout and eye. Maxilla expanded posteriorly, not sheathed by skin flap. Opercular spines absent. Anterior gill arch with 3 elongate rakers and 13 (10-14) broad plates arranged in the following configuration: Upper branch with 3 (3-5) broad plates; the bend between upper and lower gill arch with one long raker, and lower gill arch with 1 (0- 1) small plate, 2 elongate rakers interspersed with 1 small plate and followed by 8 (6-8) small plates. Pseudobranchial filaments 2 (1-2). Branchiostegal rays 7. Scales on body large, oval (in HT about 1.6 mm horizontally at mid-body, and about 20 horizontal rows above anal fin origin); vertical fins and pectoral fin naked except for scales on pectoral fin peduncle. Predorsal area, operculum and top of head, including the occiput scaled. Snout, interorbital, area below eyes and narrow supraorbital and postorbital bands naked (figs. 3A, 4A -B). Origin of dorsal fin above tip of pectoral fins. Pelvic fin with a single ray reaching about one third from its base to origin of anal fin. Pectoral fin on the middle of body, peduncle short and narrow. Caudal fin free, not fused with dorsal and anal fins. Head sensory pores: Supraorbital pores 4, anteriormost 3 on snout and posteriormost above opercular flap; infraorbital pores 6 (3 large anteriorly and smaller 3 posteriorly); mandibular pores 6 (3 anterior and 3 posterior), the 2nd anterior very long, the posterior ones increasing in size; preopercular pores 2 (2 lower and 0 upper) (fig. 3A). Lateral line with 13 (12-13) dorsal neuromasts anteriorly and 35 (34-35) medio-lateral neuromasts posteriorly. Many small sensory papillae on head.   TABLE1. Meristic and morphometric characters of Lucifuga lucayana sp. n.and L. spelaeotes.     L. lucayana  L. spelaeotes  HT HT and 5 PTs1 HT HT, PT and 42 nontypes  ANSP 146475 Mean and range USNM 204603 Mean and range  SL (mm) 99 74.3(44-99) 110 106.3(42-166)  Meristic characters  Dorsal finrays 91 89.2(84-91) 92 97.5(86-109)  Caudal finrays 10 10 10 10  Anal finrays 67 66.2(63-69) 71 73.8(66-82)  Pectoral finrays 17 17.2(17-18) 18 18.5(17-20)  Precaudal vertebrae 13 12.8(12-13) 13 13.2(13-14)  Caudal vertebrae 39 38.6(37-39) 39 39.5(38-42)  Total vertebrae 52 51.3(50-52) 52 52.7(51-55)  Rakers on anterior gill arch 16 15.3(13-17) 21 18.8(15-23)  Pseudobranchial filaments 2 1.5(1-2) 0 1.3(0-4)  Anterior dorsal fin ray above vertebra no. (D/V) 8 8.5(8-9) 9 9.0 (8-10)  Anterior anal fin ray below dorsal fin ray no. (D/A) 27 26.3(23-28) 25 27.2(24-32)  Anterior anal fin ray below vertebra no. (V/A) 19 18.8(18-20) 19 18.9(18-20)  Palatine teeth rows 0 0 5 3.7(l-7)2  Palatine teeth number 0 0 ? 22.6(3-56)  Premaxillary teeth rows 7 5-7 8 6.8(4-10)  Morphometric characters in % of SL  Head length 28.8 28.4(27.1-29.3) 29.1 28.8(26.2-31.3)  Head width 14.2 12.3(10.5-14.2) 14.2 14.5(11.4-17.0)  Head height 16.8 15.1(13.9-16.8) 16.3 16.4(13.7-19.4)  Snout length 7.3 7.3(7.1-7.6) 7.5 7.4(6.2-8.7)  Upper jaw length 14.9 14.2(13.2-14.9) 16.2 14.7(12.4-16.8)  Maxillary height 4.3 4.1(3.9-4.3) 5.4 5.0(3.5-6.3)  Diameter of pigmented eye 1.0 1.0(0.9-1.0) 1.3 1.3(0.7-1.8)  Interorbital width 6.9 6.0(4.3-6.9) 5.5 5.6(4.1-7.6)  Postorbital length 20.9 20.5(19.3-21.3) 21.0 20.4(18.4-22.6)  Preanal length 55.6 55.0(51.7-57.6) 54.1 54.2(48.2-60.5)  Predorsal length 37.0 36.4(35.5-37.0) 39.3 38.1(34.7-41.1)  Body depth at origin of anal fin 14.6 13.4(9.9-15.8) 14.3 13.5(10.9-15.4)  Pectoral fin length 13.3 12.6(11.3-13.3) 12.5 12.7(11.1-14.4)  Pelvic fin length 11.8 10.6(9.4-11.8) 14.7 14.0(8.0-20.0)  Base of pelvic fin to anal fin origin 34.9 33.7(29.4-36.9) 31.3 31.2(27.0-36.9)  Caudal fin length 9.6 10.9(9.6-12.2) ? 9.5(6.4-12.3)  1 Head missing in three paratypes.  2 Bony projection without teeth on left palatine and damaged palatine teeth on right palatine in one specimen (USNM 274737).   TABLE2. Meristc frequency.    Number of dorsal finrays  84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 n   L. lucayana n. sp.  Grand Bahama Island 1 1 2 4  Abaco Island 1 1   L. spelaeotes  Berry Islands 1 1  New Providence Island 1 1 1 2 1 6 2 3 2 4 6 1 1 1 1 0 1 34  Eleuthera Island 1 1  Great Exuma Island 1 1  Long Island 1 1 2 1 5    Number of anal finrays  63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 n   L. lucayana n. sp.  Grand Bahama Island 1 1 1 1  Abaco Island 1 1   L. spelaeotes  Berry Islands 1 1  New Providence Island 1 1 1 2 7 5 3 3 2 2 2 2 1 1 1 34  Eleuthera Island 1 1  Great Exuma Island 1 1  Long Island 1 1 2 1 5    Number of pectoral finrays Number of total vertebrae  17 18 19 20 21 n 50 51 52 53 54 55 n   L. lucayana n. sp.  Grand Bahama Island 3 1 4 1 2 3  Abaco Island 2 2 1 1   L. spelaeotes  Berry Islands 1 1 1 1  New Providence Island 1 16 15 2 34 12 17 4 1 34  Eleuthera Island 1 1 1 1  Great Exuma Island 1 1 1 1  Long Island 1 2 2 5 4 1 5   TABLE3. Selected characters of all species of Lucifuga.    n, examined Vertebrae Dorsal/ Anal finrays Pectoral finrays Caudal finrays Palatine teeth Lateral line neuromasts Total Gill-rakers Long gill-raker pigmentation Occiput squamation Eye size (% SL)  Bahamas   L. lucayana sp. n. 6 12-13+37-39 =50-52 84-91/ 63-69 17-18 10 Absent 12-13/ 34-35 13-17 Pale Dense 0.9-1.0   L. spelaeotes 44 13-14+38-42 =51-55 86-109/ 66-82 17-20 10 Present 3-56 12-19/ 30-47 15-23 Dark Weak 0.7-1.8  Cuba   L. dentata1,2 6 11+36-37 =46-48 83-95/ 69-78 15-17 8 Present 23-24 13/35 15-21 Pale Absent 0.0   L. simile2, 3 1 11+34-35 =45-46 70-78/ 57-64 11-16 8 Present 25 12/ 20-24 18 Pale Absent 0.0   L. subterranea1,2 3 11+36-37 =46-48 80-86/ 61-66 12-14 8 Absent ? 14-15 Pale Weak 0.0-0.3   L. teresinarum2, 4 0 ? 78-80/ 61-64 10-11 8 Absent ? 13-14 Pale Weak 0.0  1 Data from Cohen and Robins (1970) included.  2 Data from Cohen and McCosker (1998) included.  3 Data from Nalbant (1981) included.  4 Data from Diaz Perez (1988) included.  FIGURE 2. Lucifuga lucayana. Holotype, ANSP 146475, female, 99 mm SL, Grand Bahama Island. Drawn by Birgitte Rubaek, ZMUC. Dentition: Premaxilla with 6 (5-7) rows of granular teeth, slightly larger in inner row. Vomer horseshoe-shaped, with 3 (2-3) teeth rows and a total of 31 (24-43) teeth. Palatines without teeth (fig. 6). Dentary with 6 (3-6) outer rows of granular teeth and an inner row of long, pointed teeth. First neural spine less than half the length of second spine; spine 2-4 slightly longer and more slender than spines 5-10; spines 4-9 slightly depressed. Parapophyses present fromvertebra 7-13, increasing in length. Pleural and epipleural ribs on vertebrae 2-13. Last precaudal vertebra without ribs. Male copulatory organ completely integrated in the fleshy genital hood, similar to the configuration in L. spelaeotes(fig. 8C -D; Cohen & Robins 1970). Penis not easily visible underneath oval accessory organ. Soft lateral lips covering over accessory organ, distally with fleshy wing-shaped expansion not unlike the outer pseudoclasper in Dinematiychtyini, but clearly attached to the hood and thus not homologous. Otolith thin, elongate, with pointed anterior and posterior tips in smaller specimens (ANSP 146475) and strongly expanded broad anterior region and broad postdorsal concavity in large specimens (ZMH 9566, fig. 3B -C). Otolith length to height ratio 2.2 to 2.4. Sulcus very short, with single fused colliculum, between 28% and 35% of otolith length and located nearly at centre of inner face. Ventral furrow on inner face indistinct, dorsal depression very feeble or absent.  FIGURE 3. Lucifuga lucayana. (A) Lateral view of head, Holotype, ANSP 146475, female, 99 mm SL, Grand Bahama Island; (B) Median view of right otolith, ZMH 9566, male, ca. 125 mm SL, Grand Bahama Island; (C) Ventral view of same otolith, ZMH 9566. Coloration. Live color uniformly pale brown (Palmer 1985a, fig. 51). Preserved color light brown, with lighter fins (figs. 2, 4A-B). No pigment on vertical fin bases or elongated gill-rakers.  Etymology. The specific epithet lucayana is in honor of the Lucayan Indians, who inhabited the Bahamas for more than 2000 years, before they were eliminated by European invaders. They gave name to the type-locality, Lucayan Caverns, which they used as a graveyard (Palmer, 1985a)    FIGURE4. Head squamation of Bahamian Lucifugaspp. (A-B) Lucifuga lucayana, Paratype, BMNH 1984.12.1.20, 80 mm SL female, Grand Bahama Island, Pisces Cave, Zodiac Caverns; (C-D) Lucifuga spelaeotes, ZMUC P771363, 90 mm SL, female, Great Exuma Island; (E-F) L. spelaeotes, AMNH 57448, 96 mm SL, male, Bahamas, southern Long Island, Hard Bargain, Alphonso Dean’s Blue Hole. Note the difference in squamation on the lateral occipital parts (indicated by arrows).  Distribution and habitat. Known from at least three inland cave systems on Grand Bahama and Abaco Islands, located on the Little Bahama Bank, northern Bahamas (fig. 1): 1. Lucayan caverns, an anchialine cave located on the central southern part of Grand Bahama Island, about 1 km from the ocean. The cave is connected to the sea via a submerged passage which opens into a tidal saltwater creek. The upper about 14 m in the cave consists of freshwater (22 °C), separated from a lower, slightly warmer (23-25 °C), salty layer by a distinct halocline (Yager 1981). According to the ANSP museum label, the holotype was caught in a depth of 12 m, which must then most likely have been in fresh water. The Lucayan Caverns is the type-locality of the crustacean class Remipedia (Yager 1981) and several other cave-dwelling crustaceans (see e.g. Carpenter 1994; Pesce & Iliffe 2002). 2. Zodiac Caverns, north of Sweetings Cay Settlement, eastern Grand Bahama. The cave system was described by Palmer (1985a, b) and seems to have connections to the Sea via Zodiac Creek (see maps in Palmer (1985a, b)). One specimen of L. lucayanawas collected inthe cave Pisces at 15 m and more specimens have been observed in the caves Aquarius and Sagittarius in depths down to about 20 m in salty water (Cunliffe 1985; Palmer 1985b). 3. Abaco Island. Collected from one or two inland sink holes, but the physical conditions are unknown. 923905663 1980-11-01 ANSP Dennis Williams Bahamas Lucayan Caverns, depth 12 m ANSP 146475 Grand Bahama Island Holotype 923905665 ANSP Dennis Williams Bahamas inland sink hole, precise location and depth unknown ANSP 148497 Abaco Island Paratype 923905668 1984-07-04 BMNH Robert Palmer Bahamas Pisces Cave, Zodiac Caverns, depth 15 m BMNH 1984.12.1.20 Grand Bahama Island Paratype 923905666 1984-06-07 ZMH Dennis Williams Bahamas 26.241667 Abaco Island -77.19028 ZMH 9517 Abaco Island Paratype 923905671 ZMH Bahamas Lucayan Caverns ZMH 9522 Grand Bahama Island Paratype 923905669 ZMH Bahamas Lucayan Caverns ZMH 9566 Grand Bahama Island Paratype