AnimaliaPerciformesGrammatidaeBaldwinCarole C.RobertsonD. RossNonakaAiTornabeneLukeTwo new deep-reef basslets (Teleostei, Grammatidae, Lipogramma), with comments on the eco-evolutionary relationships of the genusZookeys71220162016638458210.3897/zookeys.638.10455 Lipogramma levinsoni http://zoobank.org/C12172C1-B3BF-48B8-B267-61D845EDCC63 Baldwin, Nonaka & Robertsonsp. n.Figure 2 Lipogramma evidesRobins & Colin, 1979: 43, fig. 2, table 1, ANSP 134332, paratype from Jamaica (photograph, counts, measurements). Lipogramma evidesRobins & Colin, 1979, fig. 3 in Gilmore and Jones (1988: 441), IRCZM 107:07660 from San Salvador, Bahama Islands (illustration, habitat information).Type locality.

Curaçao, southern Caribbean.

Holotype.

USNM 406139, 28.3 mm SL, tissue no. CUR11139, Curasub submersible, sta. CURASUB11-02, Curaçao, off Substation Curaçao, 12.083197 N, 68.899058 W, 137–146 m depth, 23 May 2011, C. Baldwin, D. Robertson & B. Van Bebber.

Paratypes.

BONAIRE: USNM 426784, 24.2 mm SL, tissue no. CUR13183, Curasub submersible, Bonaire, Bonaire City Dock, Kralendijk, Dive 2, 12.15 N, 68.2829 W, 121-137 m depth, 30 May 2013, B. Van Bebber, A. Schrier, C. Baldwin, T. Christiaan; CURAÇAO: ANSP 201863, 24.0 mm SL, Curasub submersible, Curaçao, off Substation Curaçao, 12.083197 N, 68.899058 W, no depth data available; UF 238589, 25.0 mm SL, tissue no. CUR11018, Curasub submersible, sta. CURASUB11-22, Curaçao, off of Substation Curaçao downline, 12.083197 N, 68.899058 W, no depth data available, 27 February 2011, C. Baldwin & L. Weigt; USNM 406393, 25.7 mm SL, tissue no. CUR11393, Curasub submersible, sta. CURASUB11-06, Curaçao, 132 m depth, 31 May 2011, C. Baldwin, A. Driskell, A. Schrier & B. Van Bebber; USNM 414877, 25.3 mm SL, cleared and stained, tissue no. CUR12159, Curasub submersible, sta. CURASUB12-15, Curaçao, off of Substation Curaçao downline, 12.083197 N, 68.899058 W, 128 m depth, 10 August 2012, A. Schrier, B. Brandt, C. Baldwin, A. Driskell & P. Mace; USNM 440229, 12.7 mm SL, Curasub submersible, sta. CURASUB14-07, Curaçao, in between Porto Marie and Daaibooi beaches, 12.202842 N, 69.089507 W, 123 m depth, 21 March 2014, C. Baldwin et al.; USNM 440230, 13.4 mm SL, Curasub submersible, sta. CUR13-18, Curaçao, Playa Forti, Westpoint, 12.3679 N, 69.1553 W, 127 m, 15 August 2013, C. Baldwin, B. Brandt, A. Schrier, K. Johnson & C. DeForest; USNM 406140, 19.5 mm SL, tissue no. CUR11140, Curasub submersible, sta. CURASUB11-02, Curaçao, 137-146 m depth, 23 May 2011, C. Baldwin, D. Robertson & B. Van Bebber. DOMINICA: USNM 440231, 17.0 mm SL, tissue no. DOM16229, Curasub submersible, off northwest Dominica, no specific collection data available, March 2016, R/V Chapman Crew.

Non-type specimens.

BONAIRE: USNM 426754, 21.2 mm SL, tissue no. CUR13184, Curasub submersible, Bonaire, Bonaire City Dock, Kralendijk, Dive 2, 12.15 N, 68.2829 W, 121–137 m depth, 30 May 2013, B. Van Bebber, A. Schrier, C. Baldwin, T. Christiaan; USNM 426802, 9.4 and 18.3 mm SL, Curasub submersible, Bonaire, Bonaire City Dock, Kralendijk, 12.15 N, 68.2829 W, 114–137 m depth, 30 May 2013, B. Van Bebber, A. Schrier, C. Baldwin, T. Christiaan. CURAÇAO: USNM 426774, 17.6 mm SL, tissue no. CUR13267, Curasub submersible, sta. CURASUB13-18, Curaçao, Playa Forti, Westpoint, 12.3679 N, 69.1553 W, 118 m depth, 15 August 2013, C. Baldwin, B. Brandt, A. Schrier, K. Johnson & C. DeForest; USNM 426730, 12.3 mm SL, tissue no. CUR13268, Curasub submersible, sta. CURASUB13-18, Curaçao, Playa Forti, Westpoint, 12.3679 N, 69.1553 W, 118 m depth, 15 August 2013, C. Baldwin, B. Brandt, A. Schrier, K. Johnson & C. DeForest; USNM 406011, 20.9 mm SL, tissue no. CUR11011, Curasub submersible, sta. CURASUB11-22, Curaçao, off of Substation Curaçao downline, 12.083197 N, 68.899058 W, no depth data available, 27 February 2011, C. Baldwin & L. Weigt; USNM 406012, 18.0 mm SL, tissue no. CUR11012, Curasub submersible, sta. CURASUB11-22, Curaçao, off of Substation Curaçao downline, 12.083197 N, 68.899058 W, no depth data available, 27 February 2011, C. Baldwin & L. Weigt; USNM 406019, 14.0 mm SL, tissue no. CUR11019, Curasub submersible, sta. CURASUB11-22, Curaçao, off of Substation Curaçao downline, 12.083197 N, 68.899058 W, no depth data available, 27 February 2011, C. Baldwin & L. Weigt; USNM 406394, 22.2 mm SL, tissue no. CUR11394, Curasub submersible, sta. CURASUB11-06, Curaçao, 132 m depth, 31 May 2011, C. Baldwin, A. Driskell, A. Schrier & B. Van Bebber. DOMINICA: USNM 438703, 19.0 mm SL, tissue no. DOM16052, Curasub submersible, sta. CURASUB16-07, Toucari Bay, Toucari, Dominica, NW corner of island, 15.608047 N, 61.471788 W, no depth data available, 2 March 2016, A. Schrier, R. Bakmeijer, B. Van Bebber & F. van der Hoeven; JAMAICA: ANSP 134332, 12.6 mm SL, Nekton Gamma dive 141, collection 151-2, Jamaica, Discovery Bay, 145 m depth, 15 August 1972, L. Land & S. Hastings.

Diagnosis.

A species of Lipogramma distinguishable from congeners by the following combination of characters: pectoral-fin rays 16–18 (modally 17), gill rakers 17–20 (modally 19); three supraorbital pores present along dorsal margin of orbit, no pore present between pore at mid orbit and one at posterodorsal corner of orbit; caudal fin truncate, tips of lobes rounded; body with three broad blackish bars (one on head, two on trunk) on white background, width of bar on head sufficient to encompass entire eye, width just ventral to eye averaging 26.4% head length; trunk bars sometimes hourglass shaped, with narrower and less intensely colored central regions; anterior trunk bar covering pectoral-fin base; posterior trunk bar extending onto dorsal and anal fins as large oval blotches bordered in part by white or blue pigment to form partial ocelli; dorsal and anal fins with thin orange sub-marginal stripe. The new species is further differentiated from congeners for which molecular data are available in mitochondrial COI and nuclear Histone 3, Rhodopsin, TMO-4C4, and RAG1.

Description.

Counts and measurements of type specimens given in Table 1. Frequency distributions of pectoral-fin rays and gill rakers on the first arch are given in Table 2. Twenty specimens examined, 9.4 to 28.3 mm SL. Dorsal-fin rays XII, 9 (last ray composite); anal-fin rays III, 8 (last ray composite); pectoral-fin rays 16–18, modally 17, 17 on both sides in holotype; pelvic-fin rays I,5; total caudal-fin rays 25 (13 + 12), principal rays 17 (9 + 8), spinous procurrent rays 6 (III + III), and 2 additional rays (i + i) between principal and procurrent rays that are neither spinous nor typically segmented; vertebrae 25 (10 + 15); pattern of supraneural bones, anterior dorsal-fin pterygiophores and dorsal-fin spines 0/0/0+2/1+1/1/; ribs on vertebrae 3–10; epineural bones present on vertebrae 1-16 in holotype and cleared and stained paratype (difficult to assess in radiographs of most other specimens); gill rakers on first arch 17–20 (5-6 + 12–14), modally 19 (6 + 13), 19 (6 + 13) in holotype; uppermost four and lowermost one or two rakers very small or present only as nubs, all other gill rakers elongate and slender with tooth-like secondary rakers as in Lipogramma evides (Fig. 3); pseudobranchial filaments 5–7 (7 in holotype), filaments fat and fluffy; branchiostegals 6.

Lipogramma levinsoni sp. n. A USNM 406139, holotype, 28.3 mm SL, photographed prior to preservation, photo by D. R. Robertson and C. C. Baldwin B USNM 406394, 22.2 mm SL , photographed prior to preservation, photo by D. R. Robertson and C. C. Baldwin C and D Aquarium photos, Curaçao Sea Aquarium, photos by D. Ross Robertson.

Tooth-like, secondary rakers on the first gill arch in Lipogramma evides, USNM 34771, cleared and stained paratype. Photo by L. Tornabene.

Counts and measurements of type specimens of Lipogramma levinsoni sp. n.. Measurements are in percent SL except width of bar ventral to eye, which is in percent head length. C&S = cleared and stained; CP = caudal peduncle; PFO = pelvic-fin origin; P1 = pectoral fin; P2 = pelvic fin; DXII = twelfth dorsal-fin spine. “Other Caudal” rays include “i” – a slender, flexible, non-spinous, and typically non-segmented ray and “I” – a spinous procurrent ray.

USNM 406139 USNM 406393 USNM 406140 USNM 414877 USNM 426784 USNM 440230 USNM 440229 USNM 440231 ANSP 201863 UF 238589
HolotypeParatypeParatypeParatype (C&S)ParatypeParatypeParatypeParatypeParatypeParatype
SL 28.325.719.525.324.213.426.317.024.025.0
Dorsal-fin raysXII, 9XII, 9XII, 9XII, 9XII, 9XII, 9XII, 9XII, 9XII, 9XII, 9
Anal-fin raysIII, 8III, 8III, 8III, 8III, 8III, 8III, 8III, 8III, 8III, 8
Principal caudal9+89+89+89+89+89+89+8Broken9+89+8
Other caudalIIIi+iIIIIIIi+iIIIIIIi+iIIIIIIi+iIIIIIIi+iIIIIIIi+iIIIIIIi+iIIIBrokenIIIi+iIIIIIIi+iIII
Pectoral-fin rays17, 1717, 1617, 1717, 1717*, 1717, 1717, 1816, -17, 1717, 17
Gill rakers19201919191918181918
Head length33.236.235.4-38.037.334.639.435.837.0
Eye diameter12.012.114.9-11.615.711.413.512.113.0
Snout length7.17.46.2-7.46.75.75.97.15.5
Depth at CP20.118.717.4-19.818.716.317.617.117.2
Depth at PFO33.936.634.4-35.535.135.431.240.033.6
Length P125.824.523.1-21.926.124.722.428.825.6
Length P272.445.5Broken-47.142.562.7Broken83.3Broken
Length DXII18.720.220.5-16.517.219.015.922.221.0
Width of bar ventral to eye25.528.021.5-26.126.023.125.425.628.4

6th and 7th rays (counting from dorsalmost ray) separate proximally but joined distally within same sheath and appearing as a single fat ray.

Frequency distributions of gill rakers on first arch and left and right pectoral-fin rays in Lipogramma levinsoni sp. n., Lipogramma evides, and Lipogramma haberi sp. n. Counts for the holotype and three paratypes of Lipogramma evides are included from Robins and Colins (1979). Counts of gill rakers and pectoral-fin rays for a fourth paratype of Lipogramma evides, ANSP 134330, were not given in the original description. The fifth and smallest paratype, ANSP 134332, is a specimen of Lipogramma levinsoni, and counts of that specimen made in this study are included. An asterisk indicates count of gill rakers or left pectoral-fin rays in holotype.

Gill RakersPectoral-fin Rays
15161718192021221516171819
Lipogramma levinsoni 159*1526*3
Lipogramma evides 314*111145*9
Lipogramma haberi 1*215*

Spinous and soft dorsal fins confluent, several soft rays at rear of fin forming elevated lobe that extends posteriorly beyond base of caudal fin. Pelvic fin, when depressed, extending posteriorly to point between anterior base of anal fin and beyond base of caudal fin, elongate first pelvic-fin ray broken in most preserved specimens. Dorsal profile from snout to origin of dorsal fin convex. Diameter of eye of holotype contained 2.8 times in head length. Pupil slightly tear shaped, with small aphakic space anteriorly. Scales extending anteriorly onto posterior portion of head, ending short of coronal pore. Scales present on cheeks, opercle, preopercle, interopercle, and isthmus. Scales lacking on top of head, snout, jaws, and branchiostegals. Scales large and deciduous, too many scales missing in most specimens to make accurate scale counts. In holotype, approximately 23 lateral scales between shoulder and base of caudal fin, approximately 4 scale rows on cheek, and approximately 9 scale rows across body above anal-fin origin. Scales on head and nape without cteni, scales on rest of body ctenoid. Fins naked except small scales present at bases of soft dorsal and anal fins.

Margins of bones of opercular series smooth, opercle without spines. Single row of teeth on premaxilla posteriorly, broadening to 2-3 rows anteriorly, teeth in innermost row smallest, some teeth in outer row enlarged into small canines. Dentary similar, holotype with 3 enlarged teeth in outer row near symphysis. Vomer with chevron-shaped patch of teeth, palatine with long series of small teeth. Several canals and pores visible on head, but most pores inconspicuous. Conspicuous pores present in infraorbital canal (2 pores) and portion of supraorbital canal bordering dorsal portion of orbit (3); less conspicuous pores present on top of head (1 median coronal pore), preopercle (7), and lateral-line canal in the posttemporal region (3). Anteriormost of the 3 supraorbital pores situated at anterodorsal corner of orbit, middle supraorbital pore situated above mid orbit, and posteriormost supraorbital pore situated at posterodorsal corner of orbit (Fig. 4). This pore with fleshy rim in holotype, and mid-orbit supraorbital pore with smaller fleshy rim. Posterior nostril situated just ventral to anteriormost supraorbital pore, nostril a single large opening with ventral portion of rim slightly elevated. Anterior nostril in tube with anterior flap and situated just posterior to upper lip. No lateral line present on body.

Supraorbital pore patterns in Lipogramma evides, UF 238591, 34.5 mm SL (left) and Lipogramma levinsoni sp. n., UF 238589, 25.0 mm SL (right). Arrows point to pores, which have been outlined with tiny dots for emphasis. PN – posterior nostril.

Coloration: In life (Fig. 2), ground color of head and trunk white to tan dorsally grading to white below. Head: dark brown to black bar encompassing orbit and extending ventrally to ventral midline; above orbit, bar narrowing across dorsal midline; eye with dark brown outer ring, yellowish to bluish iris. Trunk: two broad, dark brown to blackish bars present beneath dorsal fin, bars sometimes hourglass shaped, with narrower and less intensely colored central regions (central regions losing almost all dark color in some freshly dead specimens); anterior bar extending ventrally from anterior third of spinous dorsal fin to ventral midline, its anterior border extending forward to encompass base of pectoral fin; posterior bar extending ventrally from base of soft dorsal fin to posterior half of anal fin. Dorsal fin: dark trunk bars extending onto base of fin as two blotches, anterior blotch short, low, less conspicuous (than posterior blotch) and sometimes with faint orange upper border. Posterior blotch an intense, dark, longitudinal oval spanning lower half of soft dorsal and bordered posteriorly by white to bluish-white pigment. Base of fin between trunk bars whitish, central portion of fin brown to grey, and distal third of fin with bluish tint and thin, orange, submarginal stripe; this stripe breaking into spots along the rear third of fin. Anal fin: posterior trunk bar extending onto proximal portion of posterior half of fin as a strong, horizontally elongate, black blotch edged distally with bluish white line; base of fin with thin, white stripe, fin color grading into blackish to bluish-black distally. A thin, orange, sub-marginal stripe breaks into spots along posterior portion of fin. Caudal fin: basal half translucent pale orange, grading into translucent bluish distally, sometimes with indistinct, very narrow, submarginal orange band around entire edge. Pectoral fins: base blackish, fin translucent, rays translucent or tinted with orange. Pelvic fins: translucent white to bluish white, with orange tint medially on basal half of fin. In preservative (Fig. 5A), barred color pattern retained, but orange, yellow, and bluish pigments absent.

Preserved specimens of A Lipogramma levinsoni sp. n., holotype, USNM 406139, 28.3 mm SL B Lipogramma haberi sp. n., holotype, USNM 422679, 40.1 mm SL C Lipogramma evides, paratype, ANSP 134330, 30.5 mm SL Photos A and B by Sandra Raredon, C by Mark Sabaj.

Distribution.

Known from specimens collected from the Bahamas, Bonaire, Curaçao, Dominica, and Jamaica. This species was also clearly observed in October 2016 by DRR and LT from the mini-submarine “Idabel” at 140 m depth adjacent to Half Moon Bay, Roatan, Honduras.

Habitat.

Lives in or hovers above small rocky rubble on gradual slopes at depths of 108-154 m. When approached by the submersible, Lipogramma levinsoni disappears into the rubble. We observed them often in pairs.

Etymology.

Named Lipogramma levinsoni in recognition of the generous, continuing support of research on neotropical biology at the Smithsonian Tropical Research Institute (Panamá) made by Frank Levinson.

Common name.

We propose “Hourglass basslet” (Cabrilleta hierba-horaria as the Spanish equivalent) to differentiate this species from the Banded Basslet, Lipogramma evides, and the Yellow-banded Basslet, Lipogramma haberi (see description below), both of which have narrower, straight-sided bars on the trunk.

Genetic comparisons.

Table 3 shows average inter- and intraspecific divergences in COI among species of Lipogramma analyzed genetically in this study. With the exception of a single substitution in one specimen, the 15 specimens of Lipogramma levinsoni exhibit no intraspecific genetic variation at this locus and differ from other Lipogramma species by 15.4–26.0%. Lipogramma levinsoni differs from Lipogramma evides by 17.1% and Lipogramma haberi by 19.0%.

Average Kimura two-parameter distance summary for species of Lipogramma based on cytochrome c oxidase I (COI) sequences analyzed in this study. Intraspecific averages are in bold.

robinsi1robinsi2 levinsoni haberi anabantoides trilineata klayi evides
robinsi1” (n=6) 0.003
robinsi2” (n=7)0.119 0.002
levinsoni (n=15)0.1620.169 0
haberi (n=3)0.1110.1320.19 0.002
anabantoides (n=2)0.1950.1840.1540.202 0.005
trilineata (n=12)0.2170.2510.2270.2360.258 0.005
klayi (n=21)0.2660.2590.260.2790.2460.242 0.003
evides (n=30)0.1030.1280.1710.110.220.2490.263 0.001
Comments.

The smallest paratype of Lipogramma evides, ANSP 134332 (Fig. 1B), 12.6 mm SL, is a specimen of Lipogramma levinsoni. Although Robins and Colin (1979) indicated 15 pectoral-fin rays on both sides of this specimen, we count 17 on the right and find the left side too bent to make an accurate count. Lipogramma levinsoni typically has 17–18 pectoral-fin rays, modally 17. The gill-raker count of 19 given by Robins and Colin (1979) was confirmed by our examination, and is the typical count for Lipogramma levinsoni. Counts of pectoral-fin rays (15–16, usually 16) and gill rakers on the first arch (19–21, usually 20 or 21) given by Robins and Colin (1979) for the remaining paratypes of Lipogramma evides support their identification as specimens of Lipogramma evides. As noted, previous authors have mistakenly identified the broad-banded Lipogramma levinsoni as the juvenile form of the more narrow-banded Lipogramma evides. Our material includes juvenile specimens of both Lipogramma levinsoni and Lipogramma evides, which in each case have the adult configuration of dark bands (Fig. 6).

Comparison of juveniles of A Lipogramma levinsoni sp. n., USNM 440230, paratype, 13.4 mm SL and B Lipogramma evides, USNM 431410, 12.7 mm SL.

Previously published images of A Lipogramma evides, 34.4 mm SL, ANSP 134329, holotype, from Robins and Colin (1979: fig. 1) B Lipogramma levinsoni sp. n., 12.6 mm SL, ANSP 134332, as juvenile paratype of Lipogramma evides in Robins and Colin (1979: fig. 2) C Lipogramma levinsoni sp. n., 14.1 mm SL, IRCZM 107: 07660, as juvenile of Lipogramma evides in Gilmore and Jones (1988: fig. 3). Images reproduced with permission from Bulletin of Marine Science.

GilmoreRGJonesRS (1988) Lipogramma flavescens, a new grammid fish from the Bahama Islands, with descriptive and distributional notes on L. evides and L. anabantoides. Bulletin of Marine Science 42: 435445. RobinsCRColinPL (1979) Three new grammid fishes from the Caribbean Sea. Bulletin of marine Science 29: 4152.