Research Article |
Corresponding author: Carole C. Baldwin ( baldwinc@si.edu ) Academic editor: Devin Bloom
© 2018 Carole C. Baldwin, Luke Tornabene, D. Ross Robertson, Ai Nonaka, Grant Gilmore.
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:
Baldwin CC, Tornabene L, Robertson RD, Nonaka A, Gilmore GR (2018) More new deep-reef basslets (Teleostei, Grammatidae, Lipogramma), with updates on the eco-evolutionary relationships within the genus. ZooKeys 729: 129-161. https://doi.org/10.3897/zookeys.729.21842
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Two new Lipogramma basslets are described, L. barrettorum and L. schrieri, captured during submersible diving to 300 m depth off Curaçao, southern Caribbean. Superficially resembling L. robinsi in having 11–12 bars of pigment on the trunk, L. barrettorum is distinct from L. robinsi in having a stripe of blue-white pigment along the dorsal midline of the head (vs. a cap of yellow pigment), in patterns of pigment on the median fins, and in having 8–10 gill rakers on the lower limb of the first arch (vs. 11–12). Lipogramma schrieri is distinct from all congeners in having seven or eight dark bars of pigment on the trunk and broad, irregular, whitish blue markings on the dorsal portion of the head. The new species are genetically distinct from one another and from seven other Lipogramma species for which genetic data are available. A phylogenetic hypothesis derived from mitochondrial and nuclear genes suggests that the new species belong to a clade that also comprises L. evides and L. haberi. Collectively those four species are the deepest-living members of the genus, occurring at depths predominantly below 140 m. This study thus provides further evidence of eco-evolutionary correlations between depth and phylogeny in Caribbean reef fishes. Tropical deep reefs are globally underexplored ecosystems, and further investigation of Caribbean deep reefs undoubtedly will provide samples of species for which no genetic material currently exists and reveal more cryptic species diversity in the genus.
Caribbean Sea, manned submersible, cryptic species, integrative taxonomy, phylogeny, ocean exploration, Deep Reef Observation Project (DROP)
Baldwin et al. (2016) described two new species of western Atlantic Lipogramma basslets collected during diving to 300 m by the Curasub submersible, as part of the ongoing Deep Reef Observation Project (DROP) in the southern Caribbean. That brought the total number of species in the genus to 10. However, they noted that their deep-reef collections included two additional putative new species that superficially resemble L. robinsi
Collecting and morphology. Basslets were collected using Substation Curaçao’s manned submersible Curasub (http://www.substation-curacao.com). The sub has two flexible, hydraulic arms, one of which is equipped with a quinaldine/ethanol-ejection system and the other with a suction hose. Anesthetized fish specimens were captured with the suction hose, which empties into a vented plexiglass cylinder attached to the outside of the sub. At the surface, the specimens were photographed, tissue sampled, and fixed in 10% formalin. Measurements were made weeks to months after fixation and subsequent preservation in 75% ethanol and were taken to the nearest 0.1 mm with dial calipers or an ocular micrometer fitted into a Wild stereomicroscope. Selected preserved specimens were later photographed to document preserved pigment pattern and X-rayed with a digital radiography system. Images of parasitic cysts were made using a Zeiss Axiocam on a Zeiss Discovery V12 SteREO microscope. Counts and measurements follow
Molecular analyses. Tissue samples for 98 specimens assignable to nine species of Lipogramma were used for molecular analyses (Appendix
To further corroborate the morphologically diagnosed species using our molecular data, we conducted a coalescent-based, Bayesian species-delimitation analysis (
Depth distributions. We updated the depth histogram for Lipogramma of Baldwin et al. (2017: fig. 10) with the new-species names (originally listed as “L. ‘robinsi’ sp. 1” and “L. ‘robinsi’ sp. 2”) and with new depth information for the new species and for L. regia based on submersible-caught specimens. Additionally, with resolution of the “L. robinsi” complex, we added L. robinsi based on depth information in the original description (Gilmore 1977).
Accession numbers. GenSeq nomenclature (
Curaçao, southern Caribbean.
A species of Lipogramma distinguishable from congeners by the following combination of characters: pectoral-fin rays 15–16 (modally 16); gill rakers 12–14 (modally 12, 8–10 rakers on lower limb); four supraorbital pores present along dorsal margin of orbit, a pore present between one above mid orbit and one above posterodorsal corner of orbit; caudal fin rounded; body mostly yellow in life with 11 or 12 narrow brownish bars on trunk; posterior base of soft dorsal fin with large white- or blue-rimmed black ocellus; dorsal, anal and caudal fins yellow with blue/grey (brown in preservative) wavy bars or square-shaped spots. Pelvic fins blue/grey with scattered yellow-ringed dark spots. The new species is further differentiated genetically from congeners for which molecular data are available in mitochondrial COI and nuclear Histone 3, Rhodopsin, TMO-4C4, and RAG1.
Counts and measurements of type specimens given in Table
Counts and measurements of type specimens of Lipogramma barrettorum sp. n. Measurements are in percent SL. “Other Caudal Rays” include “i” – a slender, flexible, non-spinous, and typically non-segmented ray and “I” – a spinous procurrent ray.
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Holotype | Paratype | Paratype | Paratype (juvenile) | Paratype | Paratype | Paratype (juvenile) | |
SL | 26.5 | 24.5 | 28.0 | 13.0 | 25.2 | 27.0 | 10.2 |
Dorsal-fin Rays | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 |
Anal-fin Rays | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 |
Principal Caudal Rays | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 |
Other Caudal Rays | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII |
Pectoral-fin Rays | 16, 16 | 15, 15 | 16, 15 | 15, 16 | 16, 16 | 16, 16 | 16, 15 |
Gill Rakers | 3+9=12 | 3+10=13 | 4+10=14 | - | 3+~9=12 | 4+8=12 | - |
Head Length | 39.6 | 36.7 | 38.2 | 39.2 | 40.1 | 38.9 | 39.2 |
Eye Diameter | 13.2 | 13.1 | 12.1 | 14.6 | 12.7 | 12.6 | 14.7 |
Snout Length | 7.9 | 7.3 | 8.2 | 7.7 | 7.9 | 8.9 | 8.8 |
Depth at Caudal Peduncle | 19.2 | 18.4 | 21.8 | 17.7 | 17.5 | 20.4 | 18.6 |
Depth at Pelvic-fin Origin | 31.3 | 31.4 | 30.0 | 29.2 | 31.7 | 30.4 | 31.4 |
Length of Pectoral Fin | 24.9 | 24.5 | Broken | Broken | 22.2 | 22.6 | 19.6 |
Length of Pelvic Fin | 49.1 | 42.9 | Broken | 31.5 | 34.5 | 40.0 | 34.3 |
Length of 12th Dorsal Spine | 18.5 | 20.8 | 19.6 | 15.4 | 17.9 | 18.9 | 19.6 |
Spinous and soft dorsal fins confluent, several soft rays at rear of fin forming slightly elevated lobe that extends posteriorly beyond base of caudal fin. Pelvic fin, when depressed, extending posteriorly to point between base of second or third anal-fin spine and posterior base of anal fin, first pelvic-fin ray elongate. Dorsal profile from snout to origin of dorsal fin convex. Diameter of eye of holotype contained three 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 one paratype (
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 6 enlarged teeth in outer row near symphysis. Vomer with chevron-shaped patch of teeth. Palatine with long series of small teeth. Conspicuous pores present in infraorbital canal (2 pores), portion of supraorbital canal bordering dorsal portion of orbit (4), on top of head (1 median coronal pore), preopercle (at least 5), and lateral-line canal in the post-temporal region (3). The 4 supraorbital pores situated as illustrated by Baldwin et al. (2016: fig 4) for L. evides. Posterior nostril situated just ventral to anteriormost supraorbital pore, nostril a single large opening. Anterior nostril at apex of elongate narial tube and situated just posterior to upper lip. No lateral line present on body.
Coloration: In life or deceased but prior to preservation (Fig.
Known only from specimens collected off Curaçao, southern Caribbean.
Lives in or immediately above elevated rocky habitat with ample cracks or holes into which the fish retreated upon approach of the submersible. The holotype was collected at 161 m, which is the only discrete depth recording for the species. Depth ranges for two specimens were recorded as 123–160 m and 132–141 m, thus providing a potential total depth range of 123–161 m. Depth ranges for two additional specimens of 90–249 m and 50–246 m reflect depths visited during an entire submersible dive and provide little information relevant to establishing this species’ depth distribution.
Named Lipogramma barrettorum in recognition of the support of Craig and Barbara Barrett for the Smithsonian’s Deep Reef Observation Project (DROP).
We propose blue-spotted basslet in reference to the numerous blue/grey markings on the dorsal, anal, and caudal fins in life.
Table
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.
L. barretorum | L. regia | L. evides | L. schrieri | L. levinsoni | L. haberi | L. anabantoides | L. trilineata | L. klayi | |
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L. barrettorum (n=7) | 0.003 | ||||||||
L. regia (n=1) | 0.182 | – | |||||||
L. evides (n=30) | 0.099 | 0.201 | 0.002 | ||||||
L. schrieri (n=7) | 0.117 | 0.197 | 0.123 | 0.002 | |||||
L. levinsoni (n=15) | 0.158 | 0.152 | 0.167 | 0.165 | 0.001 | ||||
L. haberi (n=3) | 0.107 | 0.185 | 0.108 | 0.131 | 0.182 | 0.002 | |||
L. anabantoides (n=2) | 0.185 | 0.188 | 0.210 | 0.179 | 0.153 | 0.193 | 0.005 | ||
L. trilineata (n=12) | 0.214 | 0.207 | 0.242 | 0.246 | 0.227 | 0.235 | 0.254 | 0.005 | |
L. klayi (n=21) | 0.253 | 0.243 | 0.253 | 0.253 | 0.248 | 0.264 | 0.239 | 0.240 | 0.003 |
The holotype has two cysts, one at the base of the uppermost left pectoral-fin ray and one about mid-way along the length of the elongate first left pelvic-fin ray (Fig.
Curaçao, southern Caribbean.
Holotype.
Paratypes.
A species of Lipogramma distinguishable from congeners by the following combination of characters: pectoral-fin rays 16-17 (modally 16), gill rakers 11–13 (modally 12, 8–9 rakers on lower limb); four supraorbital pores present along dorsal margin of orbit, a pore present between one above mid orbit and one above posterodorsal corner of orbit; caudal fin rounded; body mostly tan to brown in life with 7 or 8 narrow darker brown bars on trunk; head with broad, irregular, whitish blue markings along dorsal midline from lower lip across upper lip and snout to nape; dark bar through eye bordered anteriorly and posteriorly by bluish-white bars; posterior base of soft dorsal fin with large white- or blue-rimmed black ocellus; dorsal and anal fins blue-grey with yellow spots or bars. Caudal fin mostly yellow with wide blue-grey margin and several bars comprising blue-grey mostly square-shaped spots. Pelvic fins grey/blue with scattered yellow-ringed dark spots. Juveniles with irregular white blotches of pigment on trunk and two triangular white blotches on caudal-fin base. The new species is further differentiated genetically from congeners for which molecular data are available in mitochondrial COI and nuclear Histone 3, Rhodopsin, TMO-4C4, and RAG1.
Counts and measurements of type specimens given in Table
Counts and measurements of type specimens of Lipogramma schrieri sp. n. Measurements are in percent SL. “Other Caudal Rays” include “i” – a slender, flexible, non-spinous, and typically non-segmented ray and “I” – a spinous procurrent ray.
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Holotype | Paratype (juvenile) | Paratype | Paratype | Paratype | Paratype | Paratype | |
SL | 49.7 | 17.2 | 56.0 | 61.9 | 46.6 | 26.0 | 32.8 |
Dorsal-fin Rays | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 | XII, 9 |
Anal-fin Rays | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 | III, 8 |
Principal Caudal Rays | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 | 9+8 |
Other Caudal Rays | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII | IIIi+iIII |
Pectoral-fin Rays | 16, 16 | 16, 16 | 16, - | 16, 16 | 16, 16 | 17, 17 | 16, 16 |
Gill Rakers | 3+8=11 | - | 3+9=12 | 4+9=13 | 3+9=12 | 4+9=13 | 3+9=12 |
Head Length | 38.6 | 36.1 | 33.9 | 36.0 | 37.6 | 37.3 | 38.7 |
Eye Diameter | 11.7 | 15.1 | 11.4 | 12.1 | 10.9 | 13.5 | 14.0 |
Snout Length | 10.1 | 6.4 | 8.8 | 9.1 | 9.0 | 8.5 | 7.9 |
Depth at Caudal Peduncle | 19.7 | 19.8 | 20.4 | 21.0 | 20.2 | 20.0 | 19.2 |
Depth at Pelvic-fin Origin | 32.8 | 31.4 | 31.8 | 31.8 | 32.2 | 32.3 | 31.4 |
Length of Pectoral Fin | 20.9 | 21.5 | 20.9 | 22.9 | Broken | 22.7 | 21.7 |
Length of Pelvic Fin | Broken | 39.5 | 40.4 | 40.6 | Broken | 39.6 | 39.0 |
Length of 12th Dorsal Spine | 15.7 | 19.2 | 23.6 | 18.9 | 19.7 | 22.3 | 20.7 |
Spinous and soft dorsal fins confluent, several soft rays in posterior portion of fin forming slightly elevated lobe that extends posteriorly beyond base of caudal fin. Pelvic fin extending posteriorly to base of third anal-fin spine in preserved holotype when depressed, to middle or posterior portion of anal fin in aquarium photos (e.g., Fig.
Lipogramma schrieri sp. n. A
Margins of bones of opercular series smooth, opercle without spines. Premaxilla with band of small conical teeth, band widest at symphysis, outer row with largest teeth, 3 or 4 (4 in holotype) near symphysis enlarged. Dentary similar except 8 anterior teeth enlarge. Vomer with chevron-shaped patch of teeth, palatine with long series of small teeth. Conspicuous pores present in infraorbital canal (2 pores), portion of supraorbital canal bordering dorsal portion of orbit (4), on top of head (1 median coronal pore), preopercle (at least 5), and lateral-line canal in the posttemporal region (3). The 4 supraorbital pores situated as illustrated by Baldwin et al. (2016: fig 4) for L. evides. Posterior nostril situated just ventral to anteriormost supraorbital pore, nostril a single large opening. Anterior nostril at apex of elongate narial tube and situated just posterior to upper lip. No lateral line present on body.
Coloration: In life or deceased but prior to preservation (Fig.
Distribution. Known only from specimens collected off Curaçao, southern Caribbean.
Elevated rocky habitat with ample cracks or holes into which the fish retreated upon approach of the submersible. The holotype was collected at 197 m, and three paratypes were collected at 173–207 m. The range of 156–290 m recorded for another paratype reflects all depths visited on the submersible dive during which the specimen was collected and provides little relevant depth information.
Named Lipogramma schrieri in honor of Adriaan (Dutch) Schrier, owner of Substation Curaçao. Although the Curasub submersible was not built originally for scientific research, Dutch’s enthusiastic support of research use of his sub has exponentially expanded our understanding of fish and invertebrate faunas of Caribbean mesophotic and deeper reefs.
We propose Maori Basslet, in reference to the similarity of the markings on the dorsal midline of the forehead to the beautiful facial tattoo of the Maoris, indigenous Polynesian people of New Zealand.
Table
A 52.5 mm SL Lipogramma (RGG uncataloged), collected at 296 m in 1997 by one of us (RGG) and Richard Robins off Cuba (Fig.
Morphological comparisons. Lipogramma barrettorum, L. robinsi, and L. schrieri differ from all congeners in having at least seven dark trunk bars. A comparison of major morphological and pigmentation differences among those three species is provided in Table
Summary of major morphological and pigmentation differences among Lipogramma robinsi, L. barrettorum sp. n., and L. schrieri sp. n.
ADULT | L. robinsi | L. barrettorum | L. schrieri |
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SL in preservative | To 22 mm | To 25 mm | To 62 mm |
Gill Rakers | 14–16, 11–12 on lower limb | 12 (12–14), 8–10 on lower limb | 12 (11–13), 8–9 on lower limb |
Pectoral-fin rays | 15 | 16 (15–16) | 16 (16–17) |
Body ground color | Translucent green to flesh | Yellow to yellowish brown | Tan/brown |
Head coloration in life | Grey-brown; top of head bright yellow without blue-white marksDark bar through eye to mouth absent | Yellow-brown; top of head with median blue-white stripeDark bar through eye to mouth indistinct | Pale brown; top of head with blue-white “tattoo”marksDark bar through eye to mouth strong |
Trunk in life | 10–12 narrow yellow bars, narrow interspaces | 11–12 narrow brown bars, narrow interspaces | 7–8 narrow brown bars, wide interspaces |
Dorsal Fin in life | Transparent with yellow spots; margin whiteSoft dorsal: black ocellus with white front & rear edges | Yellow with blue-grey wavy bars and spots; margin grey-blueSoft dorsal: black ocellus ringed with blue-white | Blue-grey with yellow spots and short bars; margin blue-greySoft dorsal: black ocellus ringed with blue-white |
Anal fin in life | Translucent; base white; rows yellow spots | Yellow with blue-grey spots and wavy lines; margin blue-grey | Blue-grey with yellow spots and bars; margin blue-grey |
Caudal Fin in life | Translucent; base yellow, center with yellow spots, margin white | Yellow with bars of blue-grey spots; blue-grey margin. | Yellow with bars of blue-grey spots; margin blue-grey |
Pectoral fins in life | Translucent | Translucent | Translucent; base dark |
Pelvic fins in life | White; rows black spots | Blue-grey with yellow-ringed dark spots | Pale blue-grey to blue with yellow-ringed dark spots |
JUVENILE | Not known | Known from 10–13 mm (preserved SL) specimens | Known from 17–33 mm (preserved SL) specimens |
Trunk | Not known | Similar to adult | Scattered white spots and blotches on trunk and base of caudal fin, blotches roughly in two rows of four in smallest juvenile (17 mm SL); anterior trunk bars first evident in 26-mm SL juvenile paratype |
Anal fin | Not known | Similar to adult | Similar to adult except more yellow distally |
Caudal fin | Not known | Similar to adult | Mostly yellow with blue-grey margin and 2 large triangular white blotches on base |
Species delimitation and phylogeny. Comparative morphological analysis supports the recognition of L. barrettorum and L. schrieri as distinct species, and combinations of diagnostic morphological features that distinguish them from all other Lipogramma species are provided in the species descriptions above. Molecular data for the nine Lipogramma species for which genetic data existed prior to this study (Baldwin et al. 2016) or was generated in this study (L. anabantoides, L. barrettorum, L. evides, L. haberi, L. klayi, L. levinsoni, L. regia, L. schrieri, L. trilineata) unequivocally support the presence of nine species (molecular data not available for L. flavescens, L. roseum and L. robinsi). The neighbor-joining network (Suppl. Material 1) derived from COI data shows nine distinct lineages with very high genetic distances between lineages (10–27%, mean = 19%), which are at least 20 times greater than variation in COI within lineages (range 0.1–0.5%, mean = 0.3%). The molecular phylogeny from the Bayesian analysis of the concatenated dataset (Fig.
Bayesian Inference molecular phylogeny of nine species of Lipogramma based on combined mitochondrial and nuclear genes. Numbers of individuals analyzed for each species are given in Appendix
Eco-evolutionary relationships. The two new species, L. barrettorum and L. schrieri, belong to a clade that includes L. evides and L. haberi. Collectively the members of this clade are the deepest-living species in our analysis, occurring at depths predominantly below 140 m (Figure
In addition to depth, we have observed distinct interspecific variation in the types of substrata with which some of these species associate and the nature of their associations with substrata. Off Curaçao, Lipogramma klayi is a very commonly observed species, especially on the upper level of vertical faces or slopes of ~30-60° that are heavily indented with small caves and overhangs and festooned with gorgonians and other growth. We have commonly seen it occupying the same habitat at Bonaire, Dominica, Roatan, and St. Eustatius. Its sister species, L. trilineata is much more cryptic than L. klayi. It is rarely observed, as it tends to stay close to ceilings of cavities, whether those are caves or small holes formed in large rock or coral heads. Within the clade comprising the two new species, L. evides and L. levinsoni are commonly associated with small patches of cobble scattered among rocky areas, whereas L. barrettorum and L. schrieri are associated with elevated rocky habitat with ample cracks or holes. Finally, two of us (LT and DRR) recently observed multiple instances of L. flavescens off Roatan sitting on coarse sand, meters away from shelter, in areas of sand and scattered small low patches of rock.
Baldwin et al. (2016) noted that members of the large clade comprising all Lipogramma species except L. trilineata and L. klayi are characterized by a dark ocellus on the posterior base of the dorsal fin. Based on this character, a relatively shallow depth range, and a modal count of 17 rays in the pectoral fin, they hypothesized that L. regia (not sampled in that study) is most closely related to L. anabantoides and L. levinsoni. The present phylogenetic analysis includes a recently collected specimen of L. regia and supports this hypothesis. Baldwin et al. (2016) also hypothesized that L. flavescens may be part of the deep-dwelling clade comprising L. evides, L. haberi, L. barrettorum, and L. schrieri (the last two as “Lipogramma ‘robinsi’” in their phylogeny) and, based on the deep depth-range of L. flavescens, a dark ocellus on the dorsal fin, bright yellow body coloration, a dark bar through the eye, and a low gill-raker count (15–16), possibly most closely related to L. haberi. Collection of fresh material of L. flavescens would provide the genetic material needed to test their hypotheses. We note that L. robinsi is likely part of this deep-dwelling clade as well, based on the presence of a dorsal-fin ocellus, a barring pattern on the body similar to that of L. schrieri and L. barrettorum, and its depth range (although the latter is based on very few specimens). Fresh material of L. robinsi for genetic analyses is also desirable. In addition, more information is needed on the depth distributions of all of the less common species, particularly to determine the extent of habitat partitioning within locations, as well as its consistency between locations.
Submersible exploration. Effective capture of fish specimens during deep-sea submersible dives has only been realized since 1982 with the Johnson Sea-Link submersibles and, much more recently, with the Curasub. Fresh and often living specimens are brought to the surface, providing quality material for color photography and genetic analyses to investigate phylogenetic relationships and evolutionary trends. Capture of cryptobenthic species, including gobiids (
Photographs or illustrations by C. C. Baldwin, R. G. Gilmore,
1 | Posterior base of soft dorsal fin with prominent dark spot, ocellus, or elongate blotch | 2 |
– | Soft dorsal fin without prominent markings | 10 |
2 | With prominent black bar through eye | 3 |
– | Without prominent black bar through eye | 7 |
3 | Trunk without bars, body yellow above, white below | flavescens |
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– | Trunk with bars | 4 |
4 | Trunk with 2 bars | 5 |
– | Trunk with 7–8 bars | schrieri |
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5 | Bar through eye wide, encompassing entire eye; trunk bars of equal intensity, often hourglass shaped; pectoral-fin rays modally 17, gill rakers modally 19 | levinsoni |
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– | Bar through eye narrow, encompassing only pupil; anterior trunk bar more pronounced than posterior bar, neither bar hourglass shaped; pectoral-fin rays modally 16, gill rakers 15–16 or 20–21 | 6 |
6 | Posterior trunk bar a broad, yellowish inverted triangle; gill rakers modally 15–16 | haberi |
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– | Posterior trunk bar a black rectangle; gill rakers modally 20–21 | evides |
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7 | Trunk without bars; head reddish, trunk grey-brown | anabantoides |
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– | Trunk with bars; head and trunk not colored as above | 8 |
8 | Trunk with 6 yellow bars, anterior 5 extending onto dorsal fin and 3rd-5th extending onto anal fin; head with two prominent broad yellow stripes behind eye; top of head without yellow cap | regia |
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– | Trunk with 10 to 12 bars, none extending onto dorsal or anal fins; head without broad yellow stripes behind eye, and top of head with or without yellow cap | 9 |
9 | Median fins transparent with yellow spots; top of head with yellow cap, without median blue-white stripe; lower-limb gill rakers 11–12 | robinsi |
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– | Median fins yellow with blue-grey spots; top of head without yellow cap, with median blue-white stripe; lower-limb gill rakers 8–10 | barrettorum |
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10 | Dorsal fin XI, 6–7; circum-peduncular scales 16; head and body yellow to rose colored, caudal fin yellow with dark spots, dorsal and anal fins red with pale spots | rosea |
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– | Dorsal fin XII-XIII, 8–10; circum-peduncular scales 18 to 21; not colored as above | 11 |
11 | Strongly bicolored, purplish red anteriorly, yellow posteriorly; no stripes on head; scales in lateral series 29 to 35; gill rakers 20 to 21; anal-fin soft rays 8; upper caudal-fin spines 4 or 5 | klayi |
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– | Uniformly yellowish, 3 blue stripes on head: one along dorsal midline from snout to dorsal-fin, one from top of each eye to shoulder and anterior portion of trunk; lateral scales 25 to 29; gill rakers 13 to 18; anal-fin soft rays 7; upper caudal-fin spines 3 | trilineata |
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We thank Bruce Brandt, Barry Brown, Cristina Castillo, Loretta Cooper, Tico Christiaan, Tommy Devine, Brian Horne, Dave Johnson, Rob Loendersloot, Dan Mulcahy, Diane Pitassy, Laureen Schenk, Adriaan “Dutch” Schrier, and Barbara van Bebber for assistance in various ways with this study. Portions of the laboratory and/or computer work were conducted in and with the support of the L.A.B. facilities of the National Museum of Natural History. Funding for the Smithsonian Institution’s Deep Reef Observation Project was provided internally by the Consortium for Understanding and Sustaining a Biodiverse Planet to CCB, the Competitive Grants for the Promotion of Science program to CCB and DRR, the Herbert R. and Evelyn Axelrod Endowment Fund for systematic ichthyology to CCB, and STRI funds to DRR. Externally the research was funded by National Geographic Society’s Committee for Research and Exploration to CCB (Grant #9102-12) and the Prince Albert II of Monaco Foundation grant to CCB. This is Ocean Heritage Foundation/Curacao Sea Aquarium/Substation Curacao (OHF/SCA/SC) contribution number 32.
Links between DNA voucher specimens, GenBank accession numbers, and DNA sequences of Lipogramma derived for use in this study. BAH = Bahamas, BLZ = Belize, CUR = Curaçao, DOM = Dominica, EUS = St. Eustatius, T1K = Curaçao.
CatalogNumber | TissueNumber | Species | GenBank COI | GenBank H3 | GenBank TMO-4C4 | GenBank Rag1 | GenBank Rhodopsin | GenSeq designation |
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Photo Voucher Only | BAH10150 | Lipogramma anabantoides | KX713732 | KX713823 | KX713880 | KX713842 | KX713862 | genseq-5 |
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BAH9160 | Lipogramma anabantoides | KX713824 | KX713881 | KX713843 | KX713863 | genseq-4 | |
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BLZ5340 | Lipogramma anabantoides | KX713733 | - | - | - | - | genseq-4 |
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CUR12013 | Lipogramma evides | KX713750 | - | - | - | - | genseq-4 |
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CUR12031 | Lipogramma evides | KX713751 | KX713834 | KX713891 | KX713852 | KX713872 | genseq-4 |
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CUR12044 | Lipogramma evides | KX713752 | - | - | - | - | genseq-4 |
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CUR12050 | Lipogramma evides | KX713753 | - | - | - | - | genseq-4 |
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CUR12078 | Lipogramma evides | KX713754 | - | - | - | - | genseq-4 |
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CUR12084 | Lipogramma evides | KX713755 | - | - | - | - | genseq-4 |
|
CUR12116 | Lipogramma evides | KX713757 | KX713835 | KX713892 | KX713853 | KX713873 | genseq-4 |
|
CUR12118 | Lipogramma evides | KX713758 | - | - | - | - | genseq-4 |
|
CUR12276 | Lipogramma evides | KX713760 | - | - | - | - | genseq-4 |
|
CUR12280 | Lipogramma evides | KX713761 | - | - | - | - | genseq-4 |
|
CUR12281 | Lipogramma evides | KX713762 | KX713837 | KX713894 | KX713855 | KX713875 | genseq-4 |
|
CUR12288 | Lipogramma evides | KX713763 | - | - | - | - | genseq-4 |
|
CUR12353 | Lipogramma evides | KX713767 | - | - | - | - | genseq-4 |
|
CUR13100 | Lipogramma evides | KX713771 | - | - | - | - | genseq-4 |
|
CUR13233 | Lipogramma evides | KX713779 | - | - | - | - | genseq-4 |
|
CUR13234 | Lipogramma evides | KX713780 | - | - | - | - | genseq-4 |
|
CUR13265 | Lipogramma evides | KX713781 | - | - | - | - | genseq-4 |
|
CUR13266 | Lipogramma evides | KX713782 | - | - | - | - | genseq-4 |
|
CUR13279 | Lipogramma evides | KX713785 | - | - | - | - | genseq-4 |
|
CUR13286 | Lipogramma evides | KX713786 | - | - | - | - | genseq-4 |
|
CUR13294 | Lipogramma evides | KX713787 | - | - | - | - | genseq-4 |
Photo Voucher Only | CUR15032 | Lipogramma evides | KX713793 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15055 | Lipogramma evides | KX713795 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15057 | Lipogramma evides | KX713796 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15060 | Lipogramma evides | KX713798 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15061 | Lipogramma evides | KX713799 | - | - | - | - | genseq-5 |
|
CUR15091 | Lipogramma evides | KX713811 | - | - | - | - | genseq-4 |
|
CUR15103 | Lipogramma evides | KX713813 | - | - | - | - | genseq-4 |
|
CUR15104 | Lipogramma evides | KX713814 | - | - | - | - | genseq-4 |
|
TIK003 | Lipogramma evides | KX713822 | - | - | - | - | genseq-4 |
|
CUR13158 | Lipogramma haberi | KX713775 | - | - | KX713860 | - | genseq-2 |
|
CUR13171 | Lipogramma haberi | KX713776 | - | - | KX713861 | - | genseq-1 |
|
CUR15092 | Lipogramma haberi | KX713812 | - | - | - | - | genseq-2 |
|
CUR11013 | Lipogramma klayi | KX713737 | KX713826 | KX713883 | KX713845 | KX713865 | genseq-3 |
|
CUR11133 | Lipogramma klayi | KX713740 | KX713828 | KX713885 | KX713847 | KX713867 | genseq-3 |
|
CUR11134 | Lipogramma klayi | KX713741 | - | - | - | - | genseq-3 |
|
CUR11375 | Lipogramma klayi | KX713744 | - | - | - | - | genseq-3 |
|
CUR11376 | Lipogramma klayi | KX713745 | KX713830 | KX713887 | KX713849 | KX713869 | genseq-3 |
|
CUR13112 | Lipogramma klayi | KX713772 | - | - | - | - | genseq-3 |
|
CUR13113 | Lipogramma klayi | KX713773 | - | - | - | - | genseq-3 |
|
CUR13114 | Lipogramma klayi | KX713774 | - | - | - | - | genseq-3 |
Photo Voucher Only | CUR15064 | Lipogramma klayi | KX713800 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15066 | Lipogramma klayi | KX713801 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15068 | Lipogramma klayi | KX713802 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15069 | Lipogramma klayi | KX713803 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15070 | Lipogramma klayi | KX713804 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15075 | Lipogramma klayi | KX713805 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15076 | Lipogramma klayi | KX713806 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15077 | Lipogramma klayi | KX713807 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15084 | Lipogramma klayi | KX713810 | - | - | - | - | genseq-5 |
|
DOM16036 | Lipogramma klayi | KX713817 | - | - | - | - | genseq-4 |
|
DOM16090 | Lipogramma klayi | KX713819 | - | - | - | - | genseq-4 |
|
DOM16091 | Lipogramma klayi | KX713820 | - | - | - | - | genseq-4 |
|
DOM16152 | Lipogramma klayi | KX713821 | - | - | - | - | genseq-4 |
|
CUR11011 | Lipogramma levinsoni | KX713735 | - | - | - | - | genseq-3 |
|
CUR11012 | Lipogramma levinsoni | KX713736 | - | - | - | - | genseq-3 |
|
CUR11018 | Lipogramma levinsoni | KX713738 | KX713827 | KX713884 | KX713846 | KX713866 | genseq-2 |
|
CUR11019 | Lipogramma levinsoni | KX713739 | - | - | - | - | genseq-3 |
|
CUR11139 | Lipogramma levinsoni | KX713742 | KX713829 | KX713886 | KX713848 | KX713868 | genseq-1 |
|
CUR11140 | Lipogramma levinsoni | KX713743 | - | - | - | - | genseq-2 |
|
CUR11393 | Lipogramma levinsoni | KX713747 | KX713832 | KX713889 | KX713851 | KX713871 | genseq-2 |
|
CUR11394 | Lipogramma levinsoni | KX713748 | - | - | - | - | genseq-3 |
|
CUR13183 | Lipogramma levinsoni | KX713777 | - | - | - | - | genseq-2 |
|
CUR13184 | Lipogramma levinsoni | KX713778 | - | - | - | - | genseq-3 |
|
CUR13267 | Lipogramma levinsoni | KX713783 | - | - | - | - | genseq-3 |
|
CUR13268 | Lipogramma levinsoni | KX713784 | - | - | - | - | genseq-3 |
Photo Voucher Only | CUR15031 | Lipogramma levinsoni | KX713792 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15058 | Lipogramma levinsoni | KX713797 | - | - | - | - | genseq-5 |
|
DOM16052 | Lipogramma levinsoni | KX713818 | - | - | - | - | genseq-4 |
|
CUR16008 | Lipogramma barrettorum | MG676227 | - | - | - | - | genseq-1 |
|
CUR11392 | Lipogramma barrettorum | KX713746 | KX713831 | KX713888 | KX713850 | KX713870 | genseq-2 |
|
CUR11426 | Lipogramma barrettorum | KX713749 | KX713833 | KX713890 | - | - | genseq-2 |
|
CUR12149 | Lipogramma barrettorum | KX713759 | KX713836 | KX713893 | KX713854 | KX713874 | genseq-2 |
|
CUR14079 | Lipogramma barrettorum | KX713789 | - | - | - | - | genseq-2 |
|
CUR15125 | Lipogramma barrettorum | KX713815 | - | - | - | - | genseq-2 |
|
CUR15139 | Lipogramma barrettorum | KX713816 | - | - | - | - | genseq-2 |
|
CUR14114 | Lipogramma schrieri | KX713790 | - | - | - | - | genseq-1 |
|
CUR12101 | Lipogramma schrieri | KX713756 | - | - | - | - | genseq-2 |
|
CUR12316 | Lipogramma schrieri | KX713765 | KX713839 | KX713896 | KX713857 | KX713877 | genseq-2 |
|
CUR12317 | Lipogramma schrieri | KX713766 | KX713840 | KX713897 | KX713858 | KX713878 | genseq-2 |
|
CUR13329 | Lipogramma schrieri | KX713788 | - | - | - | - | genseq-2 |
|
CUR15012 | Lipogramma schrieri | KX713791 | - | - | - | - | genseq-2 |
|
CUR12290 | Lipogramma schrieri | KX713764 | KX713838 | KX713895 | KX713856 | KX713876 | genseq-2 |
Photo Voucher Only | BLZ8127 | Lipogramma trilineata | JQ841643 | - | - | - | - | genseq-5 |
Photo Voucher Only | BLZ8128 | Lipogramma trilineata | JQ841642 | - | - | - | - | genseq-5 |
|
BLZ8168 | Lipogramma trilineata | JQ841645 | - | - | - | - | genseq-4 |
|
BLZ8274 | Lipogramma trilineata | JQ841646 | KX713825 | KX713882 | KX713844 | KX713864 | genseq-4 |
Photo Voucher Only | BLZ8343 | Lipogramma trilineata | JQ841644 | - | - | - | - | genseq-5 |
|
BLZWF204 | Lipogramma trilineata | KX713734 | - | - | - | - | genseq-4 |
|
CUR13082 | Lipogramma trilineata | KX713768 | - | - | - | - | genseq-3 |
|
CUR13089 | Lipogramma trilineata | KX713769 | - | - | - | - | genseq-3 |
|
CUR13090 | Lipogramma trilineata | KX713770 | KX713841 | KX713898 | KX713859 | KX713879 | genseq-3 |
Photo Voucher Only | CUR15034 | Lipogramma trilineata | KX713794 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15078 | Lipogramma trilineata | KX713808 | - | - | - | - | genseq-5 |
Photo Voucher Only | CUR15079 | Lipogramma trilineata | KX713809 | - | - | - | - | genseq-5 |
|
EUS17109 | Lipogramma regium | MG676228 | genseq-4 |
Figure
Data type: (measurement/occurence/multimedia/etc.)
Explanation note: Neighbor-joining network based on COI sequences of Lipogramma species investigated in this study. Scale-bar units are substitutions per site.