Lucifuga lucayana, Møller, Peter R., Schwarzhans, Werner, Iliffe, Thomas M. & Nielsen, Jørgen G., 2006

Møller, Peter R., Schwarzhans, Werner, Iliffe, Thomas M. & Nielsen, Jørgen G., 2006, Revision of the Bahamian cave­fishes of the genus Lucifuga (Ophidiiformes, Bythitidae), with description of a new species from islands on the Little Bahama Bank, Zootaxa 1223, pp. 23-46 : 27-34

Lucifuga lucayana , new species

( 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 & Böhlke 1991: 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).

Holotype: ANSP 146475, 99 mm SL, female, Grand Bahama Island, Lucayan Caverns, depth 12 m, collected by Dennis Williams, 1 November 1980.

Paratypes: ANSP 148497, 44 mm SL, male, Abaco Island, inland sink hole, precise location and depth unknown, collected by Dennis Williams, date unknown; BMNH 1984.12.1.20, 80 mm SL, female, Grand Bahama Island, Pisces Cave, Zodiac Caverns, depth 15 m, collected by Robert Palmer, 4 July 1984 (see Palmer 1985a); ZMH 9517, ca. 85 mm SL, head missing, female, Abaco Island, 26°14'30''N, 77°11'25''W, collected by Dennis Williams, 7 June 1984; ZMH 9522, ca. 110 mm SL, head missing, male, Grand Bahama Island, Lucayan Caverns, other data missing; ZMH 9566, ca. 125 mm SL, head missing, male, Grand Bahama Island, Lucayan Caverns, other data missing.

Diagnosis. Lucifuga lucayana is 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 gill­rakers 3, pale.

Similarity. Lucifuga lucayana resembles L. spelaeotes in 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 gill­rakers (pale vs. dark) and the broadly scaled occiput (vs. occiput scaled only in central part) (fig. 4). Furthermore, L. lucayana in 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 lucayana resembles two Cuban species L. subterranea and L. teresinarum in 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. dentata and L. simile by 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 mid­body, 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) medio­lateral neuromasts posteriorly. Many small sensory papillae on head.

L. lucayana L. spelaeotes

HT HT and 5 PTs1 HT HT, PT and 42 nontypes

ANSP Mean and range USNM Mean and range 146475 204603

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 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 New Providence Island 1 1 1 2 7 5 3 3 2 2 2 2 1 1 1 34 Number of pectoral finrays Number of total vertebrae

17 18 19 20 21 n 50 51 52 53 54 55 n, examined Vertebrae / Anal

finrays

Pectoral finrays Caudal finrays Palatine teeth line neuromasts Total Gill­rakers gill­raker pigmentation n Dorsal Lateral Occiput squamation Eye (% SL size) Long

Bahamas

L. lucayana sp. n. 6 12–13 [phone] / 17–18 10 Absent 12–13/ 13–17 Pale Dense 0.9–1.0

=50–52 63–69 34–35

L. spelaeotes 44 13–14+38–42 86–109/ 17–20 10 Present 12–19/ 15–23 Dark Weak 0.7–1.8

=51–55 66–82 3–56 30 –47

Cuba

L. dentata 1, 2 6 11 [phone] / 15–17 8 Present 13/35 15–21 Pale Absent 0.0

=46–48 69–78 23–24

L. simile 2, 3 1 11 [phone] / 11–16 8 Present 12/ 18 Pale Absent 0.0

=45–46 57–64 25 20–24

L. subterranea 1, 2 3 11 [phone] / 12–14 8 Absent? 14–15 Pale Weak 0.0–0.3

=46–48 61–66

L. teresinarum 2,4 0? 78– 80 / 10–11 8 Absent? 13–14 Pale Weak 0.0

61–64

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.

Dentition: Premaxilla with 6 (5–7) rows of granular teeth, slightly larger in inner row. Vomer horseshoe­shaped, 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 from vertebra 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 wing­shaped 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.

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 gill­rakers.

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 type­locality, Lucayan Caverns, which they used as a graveyard ( Palmer, 1985a)

FIGURE 4. Head squamation of Bahamian Lucifuga spp . (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 type­locality of the crustacean class Remipedia ( Yager 1981) and several other cave­dwelling 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. lucayana was collected in the 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.

2

1.8

1.6

1.4

)

SL

1.2

%

(

diameter 1 Eye 0.8

0.6

0.4

0.2

0

0 20 40 60 80 100 120 140 160

Standard Length (SL)

Figures

FIGURE 2. Lucifuga lucayana. Holotype, ANSP 146475, female, 99 mm SL, Grand Bahama Island. Drawn by Birgitte Rubaek, ZMUC.