Research Article |
Corresponding author: Luke Tornabene ( luke.tornabene@gmail.com ) Academic editor: Devin Bloom
© 2018 Luke Tornabene, D. Ross Robertson, Carole C. Baldwin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tornabene L, Robertson DR, Baldwin CC (2018) A new species of Lipogramma (Teleostei: Grammatidae) from deep reefs of Roatan, Honduras. ZooKeys 809: 79-95. https://doi.org/10.3897/zookeys.809.29280
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A new species of Lipogramma is described from submersible collections at 122–165 m depth off the coast of Roatan, Honduras, in the western Caribbean. The new species is distinguished from all other species in the genus by its bright blue coloration on the head, nape, and dorsal portion of the trunk beneath the spinous dorsal fin, a prominent round black blotch below the origin of the spinous dorsal fin, and a high number of gill rakers. A molecular phylogeny based on mitochondrial and nuclear genes shows that the new species belongs to a clade containing L. levinsoni, L. regia, and L. anabantoides. At Roatan, submersible observations of this and other Lipogramma species indicate clear, interspecific habitat partitioning by depth and substrate.
systematics, phylogeny, Caribbean, basslet, species delimitation, submersible
Manned submersibles have proven to be highly effective for collecting fishes from deep-reef habitats (
In 2017, DROP operations expanded to Roatan, Honduras, where the Idabel manned submersible is located. On the first dive capturing specimens with Idabel off Halfmoon Bay, West End, Roatan, the authors collected a specimen of an undescribed species of Lipogramma (Grammatidae) at 165 m depth. Subsequent dives around this depth revealed that the species is relatively common, despite never having been collected before nor observed in more than 150 submersible dives throughout the Caribbean by us or others using Curasub, Idabel, or the Johnson Sea-Link subs. Four additional specimens, including one juvenile, were subsequently collected on later dives off Roatan.
The description of the new species from Roatan brings the total number of species in Lipogramma to 13, all of which occur in the tropical western Atlantic Ocean. Lipogramma and Gramma are currently classified in the family Grammatidae based on a single synapomorphy in the arrangement of cheek musculature (
Specimens were collected using the Idabel submersible. The Idabel can accommodate a pilot and two scientists and is capable of diving to ~700 m. This sub was recently outfitted with a fish-catching system capable of delivering an anesthetic solution (5% quinaldine sulphate in seawater) and capturing specimens with a suction system powered by one of the submersible’s vertical thrusters (Fig.
Idabel submersible outfitted with fish-catching system. A acrylic holding tank B housing for HD video camera C quinaldine sulphate delivery hose D suction for the system is powered by a PVC hose connecting to one of the submersible’s vertical thrusters E two-way valve to allow for differential suction/blowing of water and or anesthetic F carbon-fiber compensator holding up to 2.5 gallons of anesthetic solution, powered by pressurized air from a SCUBA cylinder (not shown, lower tank in image is oxygen for life support systems) G housing for solenoid valve, enabling scientists to control the flow of anesthetic from a switch inside the submersible.
DNA was extracted from tissue samples using a Qiagen DNeasy Blood and Tissue kit. Four loci were sequenced for three specimens of the new species for phylogenetic analysis. A partial segment of the mitochondrial gene cytochrome c oxidase subunit I (COI), and three nuclear genes (TMO-4C4, Rag1, Rhodopsin) were amplified via PCR and sequenced using primers and PCR conditions from
A coalescent-based Bayesian species-delimitation analysis was also conducted (
Roatan, Honduras, western Caribbean.
A species of Lipogramma with pectoral-fin rays 15–16 (modally 16); gill rakers 18–20 total (10–11 elongate rakers plus 2–4 short, stout rudiments on lower limb, 3–4 elongate rakers plus 1–3 rudiments on upper limb); in life, body mostly yellow to tan with bright iridescent blue coloration on eye, dorsal portion of head, nape and dorsal portion of trunk beneath spinous-dorsal fin, oblique yellow bar from tip of snout to orbit and below eye, large, round, black blotch outlined with blue below anterior origin of dorsal fin, and dark ocellus outlined in blue with yellow or dark center at rear insertion of dorsal fin that extends onto body.
Counts and measurements of type specimens given in Table
Counts and measurements from type series. All measurements except SL are in % SL. Abbreviations: 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, sometimes segmented ray, and I, a spinous procurrent ray.
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MUVS-V-137 | UF 240986 |
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Holotype | Paratype | Paratype | Paratype | Paratype | |
Tissue number | ROA17003 | ROA18041 | ROA18042 | ROA17020 | ROA17026 |
SL (mm) | 26.2 | 24 | 22.5 | 26.3 | 10 |
Dorsal-fin rays | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 |
Anal-fin rays | III, 8 | III, 7 | III, 8 | III, 8 | III, 8 |
Principal caudal rays | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 |
Other caudal rays | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII |
Pectoral-fin rays | 16 | 15 | 16 | 16 | 16 |
Gill rakers - upper limb | not counted | 11+4 rudiments | 10+2 rudiments | not counted | not counted |
Gill rakers - lower limb | not counted | 4+1 rudiment | 3+3 rudiments | not counted | not counted |
Head length | 37.4 | 37.9 | 38.2 | 37.3 | 40.0 |
Eye diameter | 13.0 | 14.6 | 14.2 | 12.9 | 2.0 |
Snout length | 8.8 | 9.2 | 8.9 | 6.8 | 7.0 |
Depth at CP | 20.2 | 17.9 | 19.1 | 19.0 | 17.1 |
Depth at PFO | 36.3 | 30.8 | 35.6 | 33.4 | 35.5 |
Length P1 Fin | 24.8 | 24.2 | 23.1 | 24.7 | 22.1 |
Length P2 Fin | 50.4 | 45.4 | 44.9 | 46.4 | 35.0 |
Length DXII | 14.8 | 13.7 | 12.0 | 14.1 | 13.0 |
Spinous and soft sections of dorsal fin confluent, several soft rays in posterior portion of fin forming slightly elevated lobe that extends posteriorly beyond base of caudal fin. Pelvic fin, when depressed, extending at least to origin of first soft ray of anal fin, and to origin of penultimate anal-fin ray in holotype. Dorsal profile from snout to origin of dorsal fin convex. Diameter of eye contained 2.8–3.0 times in head length. Pupil tear-shaped, with small aphakic space anteriorly. Scales extending anteriorly onto top of head, ending at a vertical just behind posterior margin of eye. Scales present on cheeks, operculum, and isthmus, absent on snout, jaws, and branchiostegals. Scales large and deciduous, missing on anterodorsal flank of several specimens. Approximately 22–25 lateral scales between shoulder and base of caudal fin (24 in holotype), 4–6 cheek rows, 11 rows across body from above anal-fin origin. Scales on nape and along dorsal midline with reduced or absent cteni, those on cheek and opercula lacking cteni. Fins naked except base of posteriormost dorsal-fin rays, which possess 1 or 2 small embedded scales. No modified lateral-line scales present on body, but faint indication of a lateral line present superficially in fresh photographs.
Margins of bones of opercular series smooth, opercle without spines. Single row of teeth on premaxilla posteriorly, broadening to 2 or 3 rows anteriorly, teeth in innermost row smallest, some teeth in outer row enlarged into small canines. Dentary similar with 3 or 4 enlarged teeth in outer row near symphysis. Vomer with chevron-shaped patch of teeth that extends posterolaterally nearly length of premaxilla, palatine with long series of small teeth.
Cephalic head pores arranged as in Fig.
Coloration in fresh specimens (Fig.
Juvenile coloration: Coloration of the single small juvenile is essentially the same as in the adult, except that the posterior portion of the body is more noticeably yellow, the entire center of the anal fin, and, apparently, much of the soft dorsal and caudal fins are yellow, and the dark center of the ocellus at the lower rear corner of the dorsal fin is solid black.
Comments about live coloration: As can be seen in the video of the holotype being captured (https://doi.org/10.5281/zenodo.1334518, or https://zenodo.org/record/1334518#.W89WsSPMyFU), live fish have the upper third of the head and body bright blue, the lower 2/3 yellow, and a prominent large round black blotch on the shoulder.
Color in preservation (Fig.
The species was frequently observed in the mid-to-upper rariphotic zone between 122–165 m depth, in or around small rock crevices, rock piles, or caves situated on steep limestone walls covered with coarse sediment and fine rubble composed of dead sections of the green macroalga Halimeda (Fig.
The specific epithet idabeli refers to the Idabel submersible, which was used to collect the type series, and recognizes of the efforts of its owner-designer and pilot Karl Stanley and engineer Thomas Trudel, who made these and other collections of fishes possible by constructing a fish-catching system that converted Idabel from an observation-only vessel to one capable of collecting scientific specimens. The name idabeli is to be treated as a patronym (adjective) formed from the female name Idabel. The generic name Lipogramma is feminine and is formed from lipo (without) and gramma (a line of text, feminine), referring to the absence of a well-developed lateral line. The common name Cabrilleta de Dorso Azul (Blue-backed Basslet in English) refers to its distinctive coloration.
Lipogramma idabeli is easily distinguished from all other species of Lipogramma by the bright blue coloration on the head and eye, and by the pair of blue-margined ocelli below the anterior origin of the dorsal fin and at the posterior insertion of the dorsal fin. There are only two other species of Lipogramma in which the head is a markedly different color than the body; L. klayi and L. anabantoides both have rose, pink, or purple heads with tan or yellow bodies. No other known species has bright blue coloration on the dorsal portion of the head, nape, and dorsal portion of the trunk. All species of Lipogramma except L. klayi, L. rosea, and L. trilineata possess an ocellus on the posterior portion of the dorsal fin, but the pattern of barring and shading on the head and body differ among all of those species (Fig.
Coloration unambiguously diagnoses L. idabeli and supports its recognition as a distinct species. Molecular data from the ten species of Lipogramma for which tissue samples were available also support this distinction. The molecular phylogenies from the Bayesian and Maximum Likelihood analyses (Fig.
Our analyses show L. idabeli as part of a well-supported clade containing L. regia, L. levinsoni, and L. anabantoides, with the relationships within this clade being less resolved. All of these species occur from the mid-to-upper rariphotic zone and shallower (20–165 m). Similar to our observations of species of Lipogramma occurring off Curacao and other localities in the Caribbean (
We thank Karl Stanley and Thomas Trudel of the Roatan Institute of Deep-sea Exploration for their support with piloting Idabel and designing, building, and maintaining the fish-catching system and other research tools for Idabel. We thank Rachel Manning for her assistance in the field, and Sam Ghods for his help with molecular work. We are grateful for the help of Katherine Maslenikov, who assisted in the field and with cataloging specimens. Funding for the Deep Reef Observation Project was provided by the Herbert R. and Evelyn Axelrod Endowment Funding for systematic ichthyology to CCB, STRI funds to DRR, and funds from the University of Washington School of Aquatic and Fishery Sciences to LT. Rob Robins at UF, and Diane Pitassy and Jeff Williams at
DNA voucher specimens, GenBank numbers, and GenSeq nomenclature for new sequences generated in this study.
Catalog number | Tissue number | Type | Species | GenBank COI | GenBank TMO-4c4 | GenBank Rag1 | GenBank Rhodopsin | GenSeq Designation |
MUVS-V-137 | ROA18041 | paratype | Lipogramma idabeli | MK227831 | MK227821 | MK227827 | MK227824 | genseq-2 |
UF 240986 | ROA18042 | paratype | MK227830 | MK227822 | MK227826 | MK227825 | genseq-2 | |
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ROA17026 | paratype | MK227829 | MK227820 | MK227828 | MK227823 | genseq-2 |
Figure S1
Data type: statistical data
Explanation note: Maximum likelihood inference molecular phylogeny of Lipogramma. Numbers at nodes are bootstrap support values.