Prognathodes basabei Pyle & Kosaki

Pyle, Richard L. & Kosaki, Randall K., 2016, Prognathodesbasabei, a new species of butterflyfish (Perciformes, Chaetodontidae) from the Hawaiian Archipelago, ZooKeys 614, pp. 137-152 : 139-146

Taxon classification Animalia Perciformes Chaetodontidae

Prognathodes basabei Pyle & Kosaki sp. n. Figs 1, 2, 3, 4, 5

Prognathodes sp. 1; Allen et al. 1998: 250.

Prognathodes " basabei " ; Randall 2007: 291.

Type locality.

Northwestern Hawaiian Islands, Pearl and Hermes Atoll, southwest side, " Prognathodes Point", 27.7641°N, 175.9859°W.

Holotype.

BPBM 41290 , female, GenBank KX783257 , Barcode of Life PROBA001-16 , 93.4 mm SL, Northwestern Hawaiian Islands , Pearl and Hermes Atoll, southwest side, " Prognathodes Point ", 27.7641°N, 175.9859°W, 61 m, 13 September 2015, R. L. Pyle, aboard NOAA ship Hi‘ialakai ( Cruise : HA-15-05), hand nets, under limestone ledge (ancient seashore). Collected as part of a group of three associated individuals (along with CAS 242132 and USNM 440272 ).

Paratypes.

BPBM 41285 , 3 specimens: 97.7-106.3 mm SL, same location, habitat, collector, vessel and collecting method as holotype, 55 m, 17 August 2009, Cruise: HI-09-06;

CAS 242132 , GenBank KX783255 , Barcode of Life PROBA003-16 , 102.5 mm SL, same location, depth, habitat, collector, vessel, cruise and collecting method as holotype, 14 September 2015;

USNM 440272 , GenBank KX783256 , Barcode of Life PROBA002-16 , same data as holotype.

Non-type specimen.

BPBM 38441, 82 mm SL, Hawaiian Islands, O‘ahu, south shore, 116 m, 31 May 1998, R. L. Pyle, hand nets, along limestone ledge (specimen died in captivity and partially deteriorated).

Diagnosis.

A species of Prognathodes (sensu Smith et al. 2003) distinguished by the following combination of characters: dorsal-fin soft rays 21 or 22; anal-fin soft rays 16 or 17; head 2.63-2.80 in SL; body depth 1.58-1.69 in SL; pelvic-fin spine length 3.63-4.07 in SL; color in life pale yellow dorsally fading to white ventrally (sometimes entirely white) with three black bands with narrow white margins on each side of the body, the first band originating at and including the first dorsal-fin spine, extending diagonally to the eye and continuing horizontally as an orangish brown stripe from the eye to the tip of the snout, the second band originating at and including the fourth to sixth dorsal-fin spines, extending vertically at a slightly posterior angle to the ventral surface of the abdomen just anterior to the anus, tapering slightly and curving slightly posteriorly below the pectoral fin, and the third band originating at and including the last four to five dorsal-fin spines and first four to five dorsal-fin soft rays, extending vertically at a slightly posterior angle to and including the first several anal-fin soft rays, a narrow orange band on the dorso-posterior margin of the operculum, extending ventrally the posterior angle of the operculum, an oblong orange spot with some dark pigmentation on the upper one-third of the pectoral-fin axis, pelvic fins white on the spine and anterior one-third of fin, and bright orange on the posterior two-thirds of fin, a bright orange submarginal band with narrow white posterior margin extending along the posterior margins of the soft portions of the dorsal and anal fins, and continuing across the caudal peduncle.

Description.

Dorsal fin XIII (XII in two paratypes),21 (22 in one paratype), last soft ray branched to base; anal fin III,16 (17 in one paratype), last soft ray branched to base; pectoral-fin rays 16 (15 in one paratype); pelvic-fin rays I,5; principal branched caudal rays 15, upper procurrent unbranched caudal rays 4, lower procurrent unbranched caudal rays 3; pored lateral-line scales 26 (24-28); scale rows above lateral line to origin of dorsal fin 10 (11 in all but one paratype); scale rows below lateral line to origin of anal fin 24 (21-27); gill rakers on upper limb 6, on lower limb 9 (10 in one paratype); vertebrae 24.

Body deep, the depth 1.58 (1.61-1.69) in SL, and compressed, the width 4.05 (3.80-4.33) in depth; head length 2.63 (2.65-2.80) in SL; snout produced, its length 2.35 (2.19-2.62) in head; orbit diameter 3.59 (3.50-3.83) in head; interorbital slightly convex, the least bony width 4.18 (3.85-4.28) in head; least depth of caudal peduncle 4.33 (4.00-4.33) in head.

Mouth small, the upper jaw 2.77 in head, slightly diagonal, the gape forming an angle of about 20° to the horizontal, the upper jaw slightly protruding; teeth in jaws densely setiform, the longest 7.8 in orbit diameter; nostrils anterior to the eye horizontally in line with the top of the iris, the anterior in a short membranous tube with a well-developed posterior flap, the posterior slightly larger, ovate, with a low fleshy rim. Lower edge of lacrimal smooth; margin of preopercle finely serrate; margins of other opercular bones smooth.

Lateral line forming a broad arc, ending below the base of the third to fifth soft dorsal ray and within the second black band on the body. Scales ctenoid, moderately large on body except for chest and near origins of dorsal and anal fins, where small; head fully scaled except anterior portions of both jaws and around nostrils, the scales on the head small; scales on fleshy sheath surrounding base of dorsal and anal fins moderately large anteriorly and proximally, reducing in size posteriorly and distally; scales on caudal peduncle and covering base of caudal fin small.

Origin of dorsal fin slightly posterior to upper end of gill opening, its base 1.45 (1.43-1.52) in SL; first dorsal-fin spine the shortest, its length 3.09 (2.49-3.68) in head; second dorsal-fin spine 1.27 (1.42-1.98, broken in one paratype) in head; third dorsal-fin spine the longest, its length 0.93 (0.94-1.11, broken in one paratype) in head; fourth dorsal-fin spine nearly as long as the third, its length 1.04 (1.03-1.13, broken in one paratype, deformed in one paratype) in head; fifth dorsal-fin spine shorter, its length 1.20 (1.14-1.23, broken in one paratype) in head; dorsal-fin spines progressively shorter posteriorly, the last 1.80 (1.80-2.06) in head; membranes between anterior dorsal-fin spines deeply incised, progressively less so posteriorly; first dorsal-fin soft ray the longest, approximately the same length as the last dorsal-fin spine, 1.80 (1.70-1.88) in head, dorsal-fin soft rays progressively shorter posteriorly; first anal-fin spine the shortest, its length 2.54 (2.44-3.10) in head; second anal-fin spine the longest, its length 1.31 (1.16-1.31) in head; third anal-fin spine 1.52 (1.39-1.62) in head; first anal-fin soft ray the longest, its length 1.35 (1.39-1.62) in head, anal-fin soft rays progressively shorter posteriorly; caudal fin slightly convex with a slight concavity at the mid-line, its length 1.79 (1.83-2.00) in head; pectoral fins 1.37 (1.26-1.43) in head; pelvic spine 1.38 (1.36-1.51) in head; first soft ray of pelvic fin with a filamentous extension, its length 1.08 (1.19, broken in all but one paratype) in head.

Color in life as in Figures 1-5. Body pale yellow dorsally fading to white ventrally and on the thorax and lower head (color of body sometimes lacking pale yellow coloration); three prominent black bands on each side of the body with narrow white margins, the first band originating at and including the first dorsal-fin spine, extending diagonally to the eye and continuing horizontally as an orangish brown stripe from the eye to the tip of the snout, the second band originating at and including the fourth to sixth dorsal-fin spines, extending vertically at a slightly posterior angle to the ventral surface of the abdomen just anterior to the anus, tapering slightly and curving slightly posteriorly below the pectoral fin, and the third band originating at and including last four to five dorsal-fin spines and first four to five soft dorsal rays, extending vertically at a slightly posterior angle to and including the first several anal soft rays, the bands becoming dark orangish brown distally on the dorsal fin; a narrow orangish brown stripe extending from the dorsal side of the snout broadening dorsally on the nape to a point just above the interorbital space, becoming darker dorsally; a narrow orange band on the dorso-posterior margin of the operculum, extending ventrally to the posterior angle of the operculum, an oblong orange spot with some dark pigmentation on the upper one-third of the pectoral-fin axis; pelvic fins white on the spine and anterior one-third of fin, and bright orange on the posterior two-thirds of fin; a bright orange submarginal band with narrow white posterior margin extending along the posterior margins of the soft portions of the dorsal and anal fins and continuing across the caudal peduncle; caudal fin and pectoral fins translucent.

Color in alcohol similar to life color, except body a uniform dull yellow, bands dark brown, and orange areas pale brown.

A single juvenile, about 25 mm SL, was observed by RLP at a depth of 120 m during a dive off Pearl Harbor, O‘ahu on 16 August 1998. The general body shape and color pattern were the same as for adults.

Morphometric data for selected characters of type specimens are provided in Table 1.

Distribution.

Prognathodes basabei has been observed or collected at depths of 45-187 m at several locations throughout the Hawaiian Archipelago, including both the main Hawaiian Islands ( Hawai‘I, O‘ahu, Penguin Banks) and the Northwestern Hawaiian Islands (NWHI; French Frigate Shoals, Lisianski, Pearl and Hermes Atoll, Midway Atoll, Kure Atoll). No observations of this species were made during 61 submersible dives or eight mixed-gas rebreather dives to appropriate depths at Johnston Atoll ( Randall et al. 1985; Wagner et al. 2014), nor has any similar fish been observed or collected anywhere in the central or eastern Pacific. Thus, it appears that Prognathodes basabei is endemic to the Hawaiian Archipelago (although further exploration of MCEs in nearby regions may yet reveal its presence elsewhere). This is consistent with the observation that fish assemblages on deep coral reefs have proportionally more endemic species than on shallow reefs ( Pyle 1996, Kane et al. 2014, Kosaki et al. 2016).

Habitat.

Pyle and Chave (1994: 92) described the habitat for this species based on videotaped observations from submersibles as follows:

Eighteen (56%) of the observed [fish] were in areas of basalt substrata (e.g., basalt talus, blocky lava, lava tubes and pillows, basalt boulders), 13 (41 %) were in limestone habitats (primarily limestone holes and ledges), and one fish was sighted on a large (2-m diam.) water pipe. Four of the fish were in the vicinity of an unidentified antipatharian coral, three near Cirrhipathes spiralis (Linnaeus), and one near Antipathes dichotoma Pallas.

Subsequent observations of this species by divers and submersible dives, totaling several dozen individuals mostly off O‘ahu and various sites within the NWHI, were all found in association with limestone ledges and discontinuities representing ancient shorelines (Figures 3-5). In almost all cases, the fish were found underneath, inside of, or in close proximity to small undercut overhangs or caves, often swimming upside-down in association with the roof of the overhangs and caves. There are no obvious associations with other species, such as antipathinarian corals, other corals and sessile invertebrates, or particular fish species; although certain other fish species, such as than anthias Odontanthias fuscipinnis (Jenkins, 1901) and the wrasse Bodianus sanguineus (Jordan & Evermann, 1903), tend to occupy the same depth and habitat.

Etymology.

We take great pleasure in naming this species basabei , in honor of Peter K. Basabe, long-time diver, aquarium fish collector and resident of Kona, Hawai‘i, both for his role in the collection of the first specimen of this new species in 1998, and more generally for his extensive contributions and assistance to many researchers (especially the authors) in the ichthyological community.

Morphological comparisons.

Prognathodes basabei appears to be most similar in color and morphology to an undescribed Prognathodes species collected at a similar depth in Palau. These two species differ from each other in number of dorsal-fin soft rays (21-22 for basabei , compared to 17-19 for the Palau species) and anal-fin soft rays (16-17, compared to 15). Prognathodes basabei also has a smaller head (2.63-2.80 in SL, compared to 2.48-2.49 in SL), deeper body (1.58-1.69 in SL, compared to 1.71-1.76 in SL), and shorter pelvic-fin spine (3.63-4.07 in SL, compared to 4.18-4.46 in SL) than the Palau species. The two species also differ in certain aspects of life color. The anterior edge of the second black band of the Palau species originates at the third dorsal-fin spine, whereas this band originates on the fourth dorsal-fin spine in Prognathodes basabei . Moreover, both of the dark bands on the Palau species are proportionally broader dorsally, tapering more substantially ventrally than in Prognathodes basabei . Also, the orangish coloration on the pelvic fins and posterior margin of the soft dorsal and anal fins of the Palau species are much darker and brownish than in Prognathodes basabei .

Prognathodes basabei is also similar in color and morphology to Prognathodes guezei from the western Indian Ocean. It differs from that species morphologically in number of dorsal-fin soft rays (21-22 for basabei , compared to 20 for guezei ), head size (2.63-2.80 in SL, compared to 2.47-2.48 in SL), body depth (1.58-1.69 in SL, compared to 1.87-1.95 in SL), and shorter pelvic-fin spine (3.63-4.07 in SL, compared to 4.21-4.33 in SL). There are also several differences in life color between the two species. In particular, Prognathodes guezei (Figure 6) has more pronounced and discrete yellow bars on the body between the black bands, compared with more diffuse and paler yellow in Prognathodes basabei . As with the Palau species, the anterior edge of the second black band of the Prognathodes guezei originates at the third dorsal-fin spine, whereas this band originates on the fourth dorsal-fin spine in Prognathodes basabei , and the two black bands on the body of Prognathodes guezei taper even more substantially than they do in the Palau species, with the dorsal end of the posterior band in Prognathodes guezei covering the last five dorsal-fin spines, compared with the last four dorsal-fin spines on Prognathodes basabei . Also, the orangish coloration on the pelvic fins and posterior margin of the soft dorsal and anal fins of Prognathodes guezei are much paler and yellowish than in Prognathodes basabei .

Genetic comparisons.

Genetic comparisons provide another compelling justification for regarding Prognathodes basabei as distinct from the Palau species. The vertebrate mtDNA barcode (cytochrome oxidase I) sequences obtained from the holotype and two paratypes of Prognathodes basabei , compared to specimens of Prognathodes sp. collected in Palau, reveal 8% uncorrected sequence divergence. This is consistent with species-level divergences in other fish taxa ( Johns and Avise 1998, Bellwood et al. 2004, Fessler and Westneat 2007, Randall and Rocha 2009, Rocha 2004, Rocha et al. 2008). The accepted mtDNA clock rate of approximately 2% per million years in fishes ( Bowen et al. 2001, Reece et al. 2010) indicates divergence between these species on the order of 4 million years.

No tissue samples or DNA sequences have been reported for Prognathodes guezei , but given the geographic distributions of Prognathodes guezei in the western Indian Ocean, the Palau species, and Prognathodes basabei , we anticipate that the genetic divergence between Prognathodes basabei and Prognathodes guezei will prove to be even deeper than that between Prognathodes basabei and the Palau species.

Discussion.

Prognathodes basabei is an example of the conspicuous new fish species that have been discovered on deep coral reefs over the past two decades, mostly involving the use of modern mixed-gas closed-circuit rebreather diving technology ( Pyle 1996, 2000). There has been increased attention focused on mesophotic coral ecosystems (MCEs), coral-reef habitat at depths of approximately 30-150 m in tropical regions worldwide ( Hinderstein et al. 2010, Baker et al. 2016).

One particularly unusual characteristic of this species is the tendency for it to be found in groups of three individuals. Although Pyle and Chave (1994) reported that most videotaped observations from submersibles involved apparent pairs or solitary individuals, in most cases these observations were incidental to the research focus on the submersible dives, so no concerted effort was made to determine the total number of individuals at each sighting. Every observation of adults of this species by the authors during mixed-gas dives in both the main Hawaiian Islands and NWHI (nearly two dozen instances), as well as observations by RLP during several submersible dives off south O‘ahu in 2011, involved groups of three individuals; the only exception was the solitary juvenile observed by RLP in 1998. Only two individuals of the Palau species were observed together, and none of the approximately ten individuals of Prognathodes guezei observed by RLP at depths of 115-120 m off Sodwana Bay in 2011 were found in a group of three. Butterflyfishes ( Chaetodontidae ) in general are known to display a variety of social and mating systems, including monogamous pairs, harems, and schools ( Reese 1975, Yabuta and Berumen 2013). Territoriality and the distribution of food resources are important determinants of these social systems ( Hourigan 1988, Pratchett et al. 2013). Groups of three individuals as a primary social grouping have not been noted in other chaetodontid species. The six type specimens of Prognathodes basabei were found as two groups of three individuals. In both cases, the groupings included a single female and two males. More samples are necessary to determine whether such associations represent loose social groupings, territorial behavior, a mating system, or coincidence.

Another interesting aspect of this new species is the strikingly similar color pattern it shares with both the Palau species, and with Prognathodes guezei , in contrast to the deep genetic divergence that exists between the Hawaiian and Palauan specimens. It will be interesting to compare the genetics of Prognathodes guezei once tissue samples can be obtained, and as previously noted, we expect the genetic divergence to be similarly deep. A more thorough analysis and discussion of genetic comparisons between Prognathodes basabei and the Palauan species will be included in the forthcoming description of the latter species.

Prognathodes basabei is the twelfth recognized member of the genus, a group generally inhabiting deeper habitats than most other chaetodontid species. Nalbant (1995) suggested that the group may have an antitropical distribution, which would apply to Prognathodes basabei in Hawai‘i and Prognathodes guezei in the southwest Indian Ocean, but less so in light of the undescribed species in Palau. One potential explanation for such disjunct distributions is that these are relics of a once more widely distributed genus (paleoendemics; Bellwood and Meyer 2009). Another, perhaps more likely explanation is that the dearth of mesophotic exploration across the tropical central and western Pacific and Indian Oceans has left significant gaps in our understanding of Prognathodes distribution, and that additional populations and species await discovery.