Research Article |
Corresponding author: William F. Smith-Vaniz ( smithvaniz@gmail.com ) Academic editor: Javier Maldonado
© 2018 William F. Smith-Vaniz, Luke Tornabene, Raphael M. Macieira.
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:
Smith-Vaniz WF, Tornabene L, Macieira RM (2018) Review of Brazilian jawfishes of the genus Opistognathus with descriptions of two new species (Teleostei, Opistognathidae). ZooKeys 794: 95-133. https://doi.org/10.3897/zookeys.794.26789
|
A new species of jawfish, Opistognathus thionyi sp. n., is described from the Vitória-Trindade Chain and Fernando de Noronha Archipelago off Brazil, a disjunct distribution of ca. 1,800 km. Opistognathus thionyi and its allopatric Caribbean sister-species, Opistognathus maxillosus, both have a wide, fan-like upper margin of the subopercular flap and mostly over-lapping meristic data. The new species differs from O. maxillosus in having the darkest spot on the spinous dorsal fin, when present, between spines 2–5, versus always present between spines 6–9, the buccal area surrounding the esophageal opening pale versus very dark and fewer oblique scale rows in longitudinal series (45–52 vs. 69–85). A second new species, Opistognathus vicinus sp. n., known from Brazil’s mainland, has completely over-lapping meristic values with its allopatric Caribbean sister-species O. whitehursti, but differs in lacking vomerine teeth and a supramaxilla and retaining the juvenile color pattern of the latter species in adults. Diagnoses, photographs, an identification key, and distributional maps are given for all Brazilian species of Opistognathus.
Molecular phylogenetic analysis of partial cytochrome c oxidase subunit-I sequences indicates that specimens of the two allopatric pairs, O. thionyi – O. maxillosus and O. vicinus – O. whitehursti, form reciprocally monophyletic groups that differ from each other on average by 9 to 11%, with less than 1% average pair-wise genetic distance within-species. Similar patterns of phylogenetic structure were observed between reciprocally monophyletic (predominately allopatric) groups within nominal species of Opistognathus aurifrons, suggesting the possibility of at least two additional undescribed species from the Brazilian Province.
Brazilian Province, Fernando de Noronha Archipelago, reef fish, Trindade-Martin Vaz insular complex, Vitória-Trindade Seamount Chain.
Members of the family Opistognathidae range in size from 2 to 40 cm standard length and occur in all tropical oceans except the eastern Atlantic. Their natural history is of special interest because they construct burrows on sandy or rubble bottoms, near reefs using small stones or coral fragments to maintain structural integrity, and the males orally brood the egg clutches (
Prior to this study, four species of Opistognathus were known from Brazil. Two species, O. cuvierii Valenciennes, 1836 and O. brasiliensis Smith-Vaniz, 1997, are endemic to Brazil, and one species, O. lonchurus Jordan & Gilbert, 1882, has both Brazilian and broad Caribbean distributions. The fourth Brazilian species is Opistognathus aff. aurifrons, and it includes two genetically-distinct allopatric Brazilian morphotypes. The only other jawfish known from Brazil is Lonchopisthus lemur (Myers, 1935), a widely distributed deep-water species (
Summary of selected characters in Brazilian species of Opistognathus. Exceptional values in parentheses.
Characters | O. thionyi | O. vicinus | O. brasiliensis | O. cuvierii | O. lonchurus 1 | O. aff. aurifrons |
---|---|---|---|---|---|---|
Dorsal fin | XI, 16 (15) | XI, 14 | XI, 16 | XI, 16 | XI, 12 (13) | XI, 14–15 |
Anal fin | III, 15 (14) | II–III, 13 (12) | III, 15–16 | III, 16 | III, 12 (13) | III, 14–15 |
Caudal vertebrae | 18 | 17 | 18 | 19 | 16 | 17 |
Supraneurals | 0 | 0 | 2 | 1–2 | 1 | 0 |
Body scale rows | 45–52 | 43–47 | 59–75 | 60–72 | 63–76 | 66–76 |
LL terminus | 3–5 | 1–3 | 3–4 | 2–3 | 2–4 | 6–9 |
Unbranched dorsal rays | 0 | 0–1 | 1 | 1 | 5–8 | 6–11 |
Vomerine teeth | 1 | 0 | 1–3 | 2 | 2–7 | 1–3 |
Nasal cirrus | yes | yes | yes | yes | no | no |
Upper jaw fimbriate | no | no | yes | yes | no | no |
Supramaxilla present | no | no | yes | yes | yes | yes |
Subopercle flap wide | yes | no | no | no | no | no |
Dorsal-fin spines stiff with fleshy tabs on tips2 | yes | yes | no | no | no | no |
Buccal pigmentation | no | no | yes | yes | no | no |
Caudal fin banded | yes | yes | no | yes | no | no |
Spinous dorsal-fin with dark spot or ocellus | yes or no | yes | yes | yes | no | no |
Institutional abbreviations mostly follow
CIUFES Coleção Ictiológica, Departamento de Oceanografia e Ecologia, Universidade Federal do Espírito, Vitória, Espírito Santo, Brazil
FSBC Florida Fish & Wildlife Conservation Commission, Fish & Wildlife Research Institute, St. Petersburg, Florida
NPM Núcleo em Ecologia e Desenvolvimento Socio-Ambiental (NUPEM), Universidade Federal do Rio de Janeiro (UFRJ), Macaé, Brazil
SU Stanford University, collection transferred to
Median fin-ray counts, and characters associated with the vertebral column were usually taken from radiographs. The last two elements in the dorsal and anal fins have their bases in close approximation (“split to base” condition) and were counted as one ray in accord with the general practice of most authors, although the ultimate element has a separate rudimentary pterygiophore or stay. Pectoral-fin ray counts are reported for one side only and include the uppermost rudimentary ray. Caudal-fin ray counts separated by a plus indicate rays associated with the dorsal and then the ventral hypural plate. Vertebral counts are presented as a formula: precaudal + caudal. The lateral-line terminus refers to the base of the posteriomost segmented dorsal-fin ray below which the lateral line ends. The number of oblique body scale rows is only an approximation due to the irregular size and arrangement of individual scale rows. Included in this count are all anteroventrally aligned scale rows in a longitudinal series from above the tip of the opercular flap to the base of the caudal fin (counts of posteroventrally aligned scale rows will result in lower values). The gill raker at the junction of the upper and lower limbs of the first gill arch is included in the lower-limb count; care was taken not to overlook rakers (often very small) at the ends of the gill arch. Counts of gill rakers were usually made only on the right side of specimens. English common names of species, if available, are those of
Specimen sizes in material examined are given as mm SL (standard length) rounded to the nearest 0.1 mm, with number of specimens and size range given in parentheses. Measurements of paratypes indicated by an asterisk were compared with those of the holotypes. Cleared and stained specimens are indicated as “C&S”. All measurements were made with needle-point digital calipers and recorded to the nearest 0.1 mm. Measurements of paratypes indicated by an asterisk were compared with holotypes of the new species. Head length is the distance from the middle of the upper lip to the posterodorsal tip of the opercular flap. Postorbital-jaw length is a straight-line measurement from the posterior orbital margin at its junction with the rigid sphenotic bone to a vertical from the posterior end of the upper jaw. Postorbital-jaw ratio is the postorbital jaw length divided by the orbit diameter. Orbit diameter is a diagonal (posterodorsal to anteroventral) measurement of the bony orbit; the posterodorsal point of origin is the rigid sphenotic margin. Body depth is a vertical measurement from the origin of the anal fin. Caudal-peduncle depth is a vertical measurement from the narrowest part of the caudal peduncle. In the color pattern descriptions, stripes refer to markings aligned with the longitudinal axis of the body and bands or bars refer to markings aligned with the vertical axis of the body.
We sequenced a segment of the mitochondrial gene cytochrome c oxidase subunit-I (COI) for 22 samples of Opistognathus, including one specimen of O. whitehursti from St. Croix and three of O. vicinus from Brazil, eight specimens of O. thionyi and eight specimens of O. aff. aurifrons, and one specimen each of O. lonchurus and O. brasiliensis. Whole genomic DNA was extracted using a Qiagen DNEasy Blood and Tissue kit per manufacturers’ protocols. Primers for PCR and sequencing reactions were COH6 and COL6 (
Mean between-group p-distances. Shaded values on the diagonal are mean within-group p-distances for groups with more than one sequence. The number of base differences per site from averaging over all sequence pairs between groups are shown. The analysis involved 103 nucleotide sequences. Codon positions included 1st+2nd+3rd+noncoding. All positions containing gaps and missing data were eliminated. There are a total of 103 positions in the final dataset. Evolutionary analyses were conducted in MEGA7 [1]. Caribbean Clade 1 = Florida, Bahamas, Caribbean; Caribbean Clade 2 = Aruba and Curacao
Species | L. micrognathus | O. robinsi | O. macrognathus | O. brasiliensis | O. thionyi | O. maxillosus | O. whitehursti | O. vicinus | O. lonchurus | O. aurifrons – Caribbean Clade 1 | O. aurifrons – Caribbean Clade 2 | O. aurifrons – Fernando de Noronha | O. aurifrons – Brazil mainland |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Lonchopisthus micrognathus | n/a | ||||||||||||
Opistognathus robinsi | 0.195 | n/a | |||||||||||
Opistognathus macrognathus | 0.191 | 0.161 | 0.003 | ||||||||||
Opistognathus brasiliensis | 0.181 | 0.141 | 0.120 | n/a | |||||||||
Opistognathus thionyi | 0.176 | 0.135 | 0.138 | 0.132 | 0.002 | ||||||||
Opistognathus maxillosus | 0.169 | 0.152 | 0.162 | 0.132 | 0.090 | 0.009 | |||||||
Opistognathus whitehursti | 0.162 | 0.177 | 0.153 | 0.175 | 0.125 | 0.161 | 0.003 | ||||||
Opistognathus vicinus | 0.173 | 0.159 | 0.149 | 0.159 | 0.143 | 0.165 | 0.111 | 0.000 | |||||
Opistognathus lonchurus | 0.151 | 0.170 | 0.176 | 0.170 | 0.142 | 0.173 | 0.139 | 0.165 | n/a | ||||
Opistognathus aurifrons | 0.189 | 0.163 | 0.184 | 0.167 | 0.159 | 0.176 | 0.194 | 0.186 | 0.134 | 0.002 | |||
(Caribbean Clade 2) | |||||||||||||
Opistognathus aurifrons | 0.181 | 0.171 | 0.186 | 0.178 | 0.160 | 0.180 | 0.184 | 0.193 | 0.128 | 0.032 | 0.006 | ||
(Caribbean Clade 1) | |||||||||||||
Opistognathus aurifrons | 0.181 | 0.173 | 0.178 | 0.165 | 0.148 | 0.174 | 0.178 | 0.186 | 0.132 | 0.035 | 0.043 | 0.000 | |
(Fernando de Noronha) | |||||||||||||
Opistognathus aurifrons | 0.181 | 0.178 | 0.176 | 0.173 | 0.159 | 0.185 | 0.181 | 0.184 | 0.132 | 0.040 | 0.045 | 0.016 | 0.000 |
(Brazil mainland) |
1 | Anterior nostril with a simple cirrus on posterior margin; dorsal fin without a narrow dark margin; dorsal and anal fins with 0–1 anterior segmented rays unbranched distally | 2 |
– | Anterior nostril a simple tube without a cirrus on posterior margin; dorsal fin with a narrow dark margin (blue in life); dorsal and anal fins with 6–10 anterior segmented rays unbranched distally | 5 |
2 | Adults with posterior end of maxilla rigid, not ending as thin, flexible lamina; dorsal-fin spines stiff, straight, the skin-covered tips usually pale and slightly swollen fleshy tabs; supramaxilla absent | 3 |
– | Adults with posterior end of maxilla ending as thin, flexible lamina (slightly elongate in mature females and very elongate in males); dorsal-fin spines thin, flexible, usually curved distally, and tips without pale, slightly swollen tabs; supramaxilla present | 4 |
3 | Upper margin of subopercle a broad, fan-like flap; vomer with 1 tooth; premaxilla with two or more rows of teeth anteriorly; dorsal-fin segmented rays 15 or 16; caudal vertebrae 18 | O. thionyi sp. n. |
– | Upper margin of subopercle not a broad, fan-like flap; vomer without teeth; premaxilla with one row of teeth anteriorly; dorsal-fin segmented rays 14; caudal vertebrae 17 | O. vicinus sp. n. |
4 | Dorsum with 5–6 dark blotches some extending on to base of dorsal fin; underside of upper jaw and adjacent membranes with two elongate dark stripes (males) or one smaller stripe (females); caudal fin without pale bands; caudal vertebrae 18 | O. brasiliensis |
– | Dorsum without dark blotches along base of dorsal fin; under side of upper jaw and adjacent membranes in adults with two dark blotches, the innermost one poorly developed (males) or dark blotches absent (females); caudal fin with two pale bands; caudal vertebrae 19 | O. cuvierii |
5 | Dorsal- and anal-fin rays 12 or 13; dentary without large canines; caudal vertebrae 16 | O. lonchurus |
– | Dorsal- and anal-fin fin rays 14 or 15; dentary with large lateral canines; caudal vertebrae 17 | O. aff. aurifrons |
Opistognathus
sp.:
CIUFES 2347, 45.4 mm SL, male, sandy rubble bottom at Praia do Lixo, Trindade Island, Brazil, 20°31'30"S, 29°19'20"W, 20 m, 20 February 2012, Thiony Simon and L.B.C. Xavier.
(12 specimens 23.4–53.5 mm SL) all from Brazilian Province:
CIUFES 2054 (1, 27.5), sandy rubble bottom at Dogaressa seamount, Vitória-Trindade Chain, Brazil, 20°51'S, 33°40'W, 65 m, 12 April 2011, Expedição Cadeia Vitória-Trindade; CIUFES 2341-1 (1, 25.1), CIUFES 2341-2 (23.8) and CIUFES 2341-3 (1, 21.3), sandy rubble bottom at Praia do Lixo, Trindade Island, Brazil, 20°31'30"S, 29°19'20"W, 15 m, 18 February 2012, T. Simon and E.F. Mazzei; CIUFES 2346-3 (1, 29.2), sandy rubble bottom at Praia do Lixo, Trindade Island, Brazil, 20°31'30"S, 29°19'20"W, 20 m, 20 February 2012, T. Simon and L.B.C. Xavier.
A species of Opistognathus with the following combination of characters: anterior nostril a short tube with simple cirrus on posterior rim; maxilla rigid, not produced as a thin flexible lamina posteriorly; supramaxilla absent; subopercle with a broad, fan-like flap; vomer with 1 tooth; buccal area surrounding esophageal opening pale; body with 45–52 oblique body scale rows in longitudinal series; vertebrae 10+18; spinous dorsal fin with black blotch, when present, between spines 2–5. Body with five poorly defined irregular bands and sides sometimes with diagonal rows of pale spots smaller than eye diameter; when present, black blotch in spinous dorsal fin between spines 2–5; buccal area surrounding esophageal opening pale. This species is also easily distinguished from congeners by divergence in the mitochondrial gene COI, as specimens form a monophyletic group that differs from its closest relative (O. maxillosus) by an average of 9% (654 bp analyzed).
Morphometric data are given in Table
Morphometric data for holotype and six paratypes of Opistognathus thionyi.
Character | Holotype | Range | Mean | SD |
---|---|---|---|---|
Standard length (mm) | 45.5 | 32.5–53.5 | 40.7 | 7.5 |
Percentage of SL | ||||
Head length | 38.6 | 34.8–37.6 | 36.5 | 0.99 |
Postorbital head length | 22.6 | 19.6–22.0 | 21.0 | 0.91 |
Jaw length | 22.3 | 21.2–22.1 | 21.7 | 0.41 |
Postorbital jaw length | 6.2 | 3.9–7.7 | 5.7 | 1.40 |
Orbit diameter | 13.8 | 11.8–14.2 | 12.9 | 0.80 |
Pelvic-fin length | 23.7 | 24.2–25.3 | 24.7 | 0.50 |
Caudal-fin length | 24.8 | 23.4–26.4 | 24.7 | 1.10 |
Body depth | 20.1 | 19.0–21.8 | 19.9 | 1.13 |
Caudal peduncle depth | 10.0 | 9.3–10.6 | 10.1 | 0.53 |
Predorsal length | 36.0 | 32.2–35.4 | 33.9 | 1.24 |
Preanal length | 56.9 | 58.1–61.2 | 59.7 | 1.13 |
Dorsal-fin length | 62.6 | 62.6–67.4 | 64.6 | 1.79 |
Anal-fin length | 31.9 | 29.3–34.9 | 32.8 | 2.16 |
Percentage of HL | ||||
Postorbital head length | 58.7 | 54.5–59.7 | 57.4 | 1.70 |
Jaw length | 57.7 | 56.5–61.3 | 59.4 | 2.10 |
Postorbital jaw length | 16.1 | 10.9–21.3 | 15.6 | 4.00 |
Orbit diameter | 35.9 | 32.1–37.9 | 35.5 | 2.20 |
Ratio | ||||
POJaw length/orbit diameter | 0.45 | 0.30–0.61 | 0.44 | 0.13 |
Scales absent from head, nape, pectoral-fin base and breast; belly completely scaled, and sides fully scaled except for area above lateral line anteriorly. Body with 48 (45–52) oblique scale rows in longitudinal series. Lateral-line terminus below verticals between segmented dorsal-fin ray 3 (3–5). Anterior lateral-line pores relatively numerous and arranged in branched series along lateral-line tubes, all of which are embedded in skin. Mandibulo-preopercular pore positions all consisting of multiple pore series, except first two mandibular pore positions occupied by simple pores. Infraorbital pore positions consisting of multiple series that extend onto cheeks. Nape nearly to completely covered by sensory pores except for V-shaped naked area immediately in front of dorsal-fin origin (Figure
Anterior nostril positioned closer to posterior nostril than to dorsal margin of upper lip, and adults with a rounded cirrus that usually reaches anterior margin of orbit when depressed; height of cirrus 2.0–3.0 times maximum diameter of posterior nostril. Dorsal fin moderately low anteriorly, with posterior rays slightly longer; profile relatively uniform without noticeable change in fin height at junction of spinous and segmented rays. Dorsal-fin spines stiff and straight with pungent tips and in larger specimens the skin covered tips usually with pale, slightly swollen fleshy tabs. Segmented dorsal- and anal-fin rays all typically branched distally. Outermost segmented pelvic-fin ray not tightly bound to adjacent ray and interradial membrane strongly incised distally; tip of depressed pelvic fin in front of anal-fin origin. Upper margin of subopercle consisting of a broad, truncated flap (Figure
Upper jaw not sexually dimorphic, extending 0.45 (0.3–0.6) eye diameters behind orbit in specimens 32.5–53.5 mm SL; posterior end of maxilla rigid and truncate, without a thin flexible lamina; supramaxilla absent. Coronoid (ascending) process of articular slightly tilted backward and somewhat club-shaped with anterodorsal end bluntly pointed and posteroventral end bluntly rounded (Figure
Color in life (Figures
Preserved color (Figure
The Caribbean allopatric Opistognathus maxillosus Poey, 1860 shares with O. thionyi the same subopercle shape but in addition to having more longitudinal body scale rows (69–85 vs. 45–52), the dark spot in the dorsal fin is always between spines 6–9 (vs. when present between spines 2–5), and the buccal area immediately surrounding the esophageal opening very dark (vs. pale). An updated description of Opistognathus maxillosus is given in
Jaws and dentition (supramaxilla shaded) in selected species of Opistognathus. A O. thionyi, CIUFES 2393, 53.5 mm SL, Fernando de Noronha Archipelago, Brazil B O. vicinus, NPM 5030, 47.4 mm SL, Brazil C O. lonchurus,
The specific name honors our colleague and dear friend Thiony Simon (1985–2016), who passed away during preparation of this article. He collected most of the type material of the new species and dedicated his life to study and conservation of Brazilian reef ecosystems.
Opistognathus thionyi is known only from three oceanic sites, Trindade Island, Dogaressa Seamount, and Fernando de Noronha Archipelago (Figure
The conservation status of Opistognathus thionyi (cited as Opistognathus maxillosus Poey, 1860 – unpublished data) has been assessed by the Ministério do Meio Ambiente/Instituto Chico Mendes de Conservação da Biodiversidade (MMA/ICMBio - Brazil) and listed as Least Concern. However, anthropogenic activities on oceanic marine ecosystems (i.e., seamount mining, fisheries, marine traffic, tourism, and human occupation of the islands), and the inadequate protection from these impacts currently provided by new Brazilian marine protected areas in the Vitória-Trindade Seamounts Chain (see
Opistognathus
whitehursti
(Longley, 1927):
CIUFES 0796, 43.0 mm SL, male, Ilha Rasa de Dentro, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 15m, 11 March 2008, R. M. Macieira and T. Simon.
(12 specimens 17.0–47.4 mm SL) all from Brazil Province:
A species of Opistognathus with the following combination of characters: anterior nostril a short tube with simple cirrus on posterior rim; maxilla rigid, not produced as a thin flexible lamina posteriorly; supramaxilla absent; subopercle without a broad, fan-like flap; vomer without teeth; body with 43–47 oblique body scale rows in longitudinal series; vertebrae 10+17; sides with two rows of pale spots, each approximately diameter of eye. Body with six vertically irregular, evenly spaced bands, widest on mid-side, and two rows of six pale spots, each spot approximately diameter of eye; buccal area surrounding esophageal opening pale. This species is also easily distinguished from congeners by divergence in the mitochondrial gene COI, as specimens form a monophyletic group that differs from its closest relative (O. whitehursti) by an average of 11% (654 bp analyzed).
Morphometric data are given in Table
Morphometric data for holotype and six paratypes of Opistognathus vicinus.
Character | Holotype | Range | Mean | SD |
---|---|---|---|---|
Standard length (mm) | 43.0 | 36.0–47.4 | 40.3 | 4.06 |
Percentage of SL | ||||
Head length | 37.1 | 34.1–37.5 | 35.7 | 1.11 |
Postorbital head length | 24.0 | 21.2–24.2 | 22.8 | 1.23 |
Jaw length | 20.5 | 17.2–21.2 | 19.9 | 1.57 |
Postorbital jaw length | 7.8 | 6.3–8.9 | 7.7 | 0.88 |
Orbit diameter | 10.2 | 10.1–11.3 | 10.9 | 0.57 |
Pelvic-fin length | 23.4 | 23.1–25.6 | 24.6 | 1.04 |
Caudal-fin length | 25.1 | 23.8–28.7 | 26.4 | 1.94 |
Body depth | 22.5 | 19.5–22.2 | 20.9 | 0.94 |
Caudal peduncle depth | 12.1 | 11.1–12.8 | 11.6 | 0.64 |
Predorsal length | 34.8 | 31.3–36.6 | 34.3 | 1.80 |
Preanal length | 53.8 | 55.6–58.8 | 57.4 | 1.16 |
Dorsal-fin length | 71.0 | 60.9–68.1 | 64.5 | 2.35 |
Anal-fin length | 36.4 | 31.1–36.5 | 34.5 | 1.94 |
Percentage of HL | ||||
Postorbital head length | 64.9 | 59.6–67.5 | 64.0 | 3.27 |
Jaw length | 55.3 | 48.6–59.9 | 55.9 | 5.08 |
Postorbital jaw length | 21.0 | 16.8–26.0 | 21.6 | 3.08 |
Orbit diameter | 27.6 | 27.0–32.0 | 21.6 | 3.08 |
Ratio | ||||
POJaw length/orbit diameter | 0.76 | 0.62–0.88 | 0.71 | 0.09 |
Scales absent from head, nape, pectoral-fin base and breast; belly completely scaled, and sides fully scaled except for area above lateral line anteriorly. Body with 46 (43–47) oblique scale rows in longitudinal series. Lateral-line terminus below verticals between segmented dorsal-fin ray 1 (2–3). Anterior lateral-line pores relatively numerous and arranged in branched series along lateral-line tubes, all of which are embedded in skin. Mandibulo-preopercular pore positions all consisting of multiple pore series, except first two mandibular pore positions occupied by simple pores. Infraorbital pore positions consisting of multiple series that extend onto cheeks. Nape nearly to completely covered by sensory pores except for V-shaped naked area immediately in front of dorsal-fin origin (Figure
Anterior nostril positioned closer to posterior nostril than to dorsal margin of upper lip, and adults with a slender cirrus that reaches anterior margin of orbit when depressed; height of cirrus 2.0 times maximum diameter of posterior nostril. Dorsal fin moderately low anteriorly, with posterior rays slightly longer; profile relatively uniform without noticeable change in fin height at junction of spinous and segmented rays. Dorsal-fin spines stiff and straight and in larger specimens the skin covered tips usually with pale, slightly swollen fleshy tabs. Segmented dorsal- and anal-fin rays all typically branched distally. Outermost segmented pelvic-fin ray not tightly bound to adjacent ray and interradial membrane strongly incised distally; tip of depressed pelvic fin in front of anal-fin origin. Upper margin of subopercle oval-shaped without a broad, truncated flap (Figure
Upper jaw not sexually dimorphic, extending 0.76 (0.62–0.88) eye diameters behind orbit in specimens 36.0–47.4 mm SL; posterior end of maxilla rigid and truncate, without a thin flexible lamina; supramaxilla absent. Premaxilla with a single row of teeth, largest anteriorly becoming smaller and more closely spaced posteriorly, except in mature males posteriormost three or four teeth stouter and more strongly hooked than adjacent teeth. Dentary anteriorly with two rows of teeth, innermost smaller and slanted backwards; laterally teeth uniserial and larger than anterior teeth, posterior teeth of males larger and more strongly hooked than others. Vomer without teeth. Infraorbital bones tubular, with numerous openings for sensory canals; third infraorbital with a wide suborbital shelf. Postcleithra closely attached; dorsal postcleithrum an irregular elongate oval, narrowest ventrally where it overlaps head of ventral postcleithrum; ventral postcleithrum club-shaped, broadest dorsally and with a pointed ventral end.
Color in life (Figure
Preserved color (Figure
Genetic differences (see discussion in “Phylogenetic relationships of western Atlantic Opistognathus”), suggested that Opistognathus vicinus and the Caribbean O. whitehursti could be separate species despite their very similar appearance, including meristic values and sexually dimorphic premaxillary teeth (see
From the Latin vicinus (near, neighboring), referring to the allopatric distribution and sister-species phylogenetic relationship of the new species and the Caribbean Opistognathus whitehursti.
A Brazilian endemic (Figure
The conservation status of this species [cited as Opistognathus whitehursti (
Opistognathus
brasiliensis
Smith-Vaniz, 1997: 1104, fig. 20 (original description; Alcatraces [misspelled Alcatrazes] Island: holotype
A species of Opistognathus with the following combination of characters: anterior nostril a short tube with simple cirrus on posterior rim; adults with posterior end of maxilla ending as thin, flexible lamina (slightly elongate in mature females and very elongate in males); supramaxilla present; subopercle without a broad, fan-like flap; most of nape without sensory pores; dorsal-fin spines thin, flexible, usually curved distally, and tips without pale, slightly swollen tabs; dorsal fin XI, 16 with all soft rays weakly branched distally; anal fin II, 15–16; body with 59–75 oblique scale rows in longitudinal series; vertebrae 10+18; supraneurals 2; gill rakers 9–11+23–24=33–36; spinous dorsal fin with black spot encircled by a very narrow white ring between spines 4–7 and dorsum with 5 or 6 dusky bands that extend onto base of dorsal fin; pelvic fins uniformly dark; underside of upper jaw and adjacent membranes in adults with two elongate dark stripes (males) or one smaller stripe (females) (
5 specimens (107.5–129 mm SL), including the holotype, cited in
A Brazilian endemic (Figure
In the diagnosis and description of Opistognathus brasiliensis,
The conservation status of this species has been assessed by the Ministério do Meio Ambiente/Instituto Chico Mendes de Conservação da Biodiversidade (MMA/ICMBio - Brazil), and it was listed as Data Deficient.
Opisthognathus
[sic] cuvierii Valenciennes in Cuvier and Valenciennes 1836: 504, color pl. 343 (original description; Bahia: holotype
Opistognathus
cuvierii
:
Opistognathus
cuvieri
:
A species of Opistognathus with the following combination of characters: anterior nostril a short tube with simple cirrus on posterior rim; adults with posterior end of maxilla ending as thin, flexible lamina (slightly elongate in mature females and very elongate in males); supramaxilla present; subopercle without a broad, fan-like flap; most of nape without sensory pores; dorsal-fin spines thin, flexible, usually curved distally, and tips without pale, slightly swollen tabs; dorsal fin XI, 16, with all soft rays weakly branched distally; anal fin II, 16; body with 60–72 oblique scale rows in longitudinal series; vertebrae 10+19; supraneurals 1 or 2; gill rakers 9–11+20–23=30–35; spinous dorsal fin with an ocellus between spines 3–7, otherwise dorsal fin with rows of pale spots and dorsum without 5 or 6 dusky bands that extend onto base of dorsal fin; pelvic fins uniformly dark; caudal fin dark with two pale bands; underside of upper jaw and adjacent membranes in adults with two dark blotches, the innermost one poorly developed (males) (
5 specimens (80.5–11.5 mm SL), including the holotype, cited in
A Brazilian endemic (Figure
The conservation status of this species has been assessed by the Ministério do Meio Ambiente/Instituto Chico Mendes de Conservação da Biodiversidade (MMA/ICMBio - Brazil) and it was listed as Least Concern.
Opisthognathus
[sic] lonchurus Jordan & Gilbert, 1882: 290 (original description; snapper banks off Pensacola, Florida: holotype
Opistognathus
lonchurus
:
A species of Opistognathus with the following combination of characters: anterior nostril a short tube without a cirrus on posterior rim; posterior end of maxilla rigid, not produced as a thin flexible lamina; supramaxilla present; subopercle without a broad, fan-like flap; most of nape without sensory pores (Figure
3 specimens (75.3–81.2 mm SL) from Brazil and 46 specimens (27–122 mm SL) from the Caribbean. Brazil: CIUFES 1426 (1, 75.3), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 26 February 2000, D.A. Jório and J.L. Gasparini; CIUFES 2361 (1, 81.2), gravid female, Ilhas Escalvada, Guarapari, Espírito Santo, 20°41'S, 40°24'W, 23 m, 30 February 2012, R.M. Macieira and J.-C. Joyeux;
South Carolina, Gulf of Mexico, Greater Antilles and northern South America to Brazil (Figures
The conservation status of this species was assessed by the International Union for Conservation of Nature (IUCN) and listed as Lease Concern (
Opistognathus
aurifrons
Jordan & Thompson, 1905: 252, fig. 4 (original description; Garden key, Dry Tortugas, Florida);
Opistognathus
sp.
Opistognathus
sp. 1
Opistognathus
sp. 2
Opistognathus
aff.
aurifrons
Jordan & Thompson:
A species of Opistognathus with the following combination of characters: anterior nostril a short tube without a cirrus on posterior rim; posterior end of maxilla rigid, not produced as a thin flexible lamina; supramaxilla present; dorsal-fin spines thin, flexible, usually curved distally, and tips without pale, slightly swollen tabs; subopercle without a broad, fan-like flap; most of nape without sensory pores (Figure
As provisionally recognized, this species is known only from the Brazilian Province including continental localities from the State of Maranhão (0°53'S, 44°17'W) south to Armação de Búzios (22°45'S, 41°59'W) in the State of Rio de Janeiro and oceanic sites of the Vitória-Trindade Seamounts Chain and Fernando de Noronha Archipelago (Figure
33 specimens (30.4–73.7 mm SL) all from Brazilian Province. Mainland localities:
Brazilian specimens of Opistognathus aff. aurifrons (n=28) differ from Caribbean O. aurifrons (n=292) in consistently having 17 vs. 16 caudal vertebrae. Brazilian fish are represented by two allopatric and slightly different genetic populations (see discussion below in “Phylogenetic relationships of western Atlantic Opistognathus”). Mainland and Vitória-Trindade Seamounts Chain specimens have long pelvic fins that when depressed extend at least to the anal-fin origin (25.7–38.2% SL, mean 30.4%, in 22 specimens 30.4–74.8 mm SL) and in fresh adult specimens the top of the head is yellow, bordered posteriorly by a narrow blue band extending from slightly behind the eye to upper jaw and across the nape; remainder of the head and body greenish-yellow to bluish-yellow (Figures
Depending on the locality, adults of Caribbean Opistognathus aurifrons may have relatively short or long pelvic fins, color patterns not found in Brazilian populations or that duplicate them.
The conservation status of Brazilian populations of this species has been assessed by the Ministério do Meio Ambiente/Instituto Chico Mendes de Conservação de Biodiversidade (MMA/ICMBio – Brazil) and listed as Least Concern.
The molecular phylogenies inferred from the COI data using ML and Bayesian inference were very similar in topology (Figure
Bayesian inference phylogeny of western Atlantic Opistognathus based on COI data. Support values are Bayesian posterior probabilities. For clarity the clade containing Opistognathus aurifrons is collapsed (see Figure
Two groups in our initial analysis showed distinct phylogenetic structure and geographic genetic variation that suggested the presence of cryptic species. In the first case, specimens initially identified as O. whitehursti from Brazil and the Caribbean each formed two reciprocally monophyletic clades with considerable genetic differentiation in COI (mean between group p-distance = 0.11 (mean within group p-distance ≤ 0.003, Table
Previous studies of several populations of Caribbean O. aurifrons based on morphology (
This study would not have been possible without the cooperation of the curators, collection managers, and support staff of the institutions cited in the Materials and methods section who loaned specimens, provided catalog numbers and other curatorial assistance. We especially thank the following individuals who provided radiographs, curatorial assistance and laboratory facilities: Jean-Christophe Joyeux (CIUFES), Marcelo Ribeiro de Britto (
Genus | species | GenBank Accession Number | Voucher from this study |
---|---|---|---|
Lonchopisthus | micrognathus | MH751526 | USNM 438666 |
Opistognathus | afer | KU176393 | n/a |
Opistognathus | aurifrons | JQ842264 | n/a |
Opistognathus | aurifrons | JQ841740 | n/a |
Opistognathus | aurifrons | JQ841739 | n/a |
Opistognathus | aurifrons | JQ841738 | n/a |
Opistognathus | aurifrons | JQ840957 | n/a |
Opistognathus | aurifrons | JQ840629 | n/a |
Opistognathus | aurifrons | JF297892 | n/a |
Opistognathus | aurifrons | JF297891 | n/a |
Opistognathus | aurifrons | JF297890 | n/a |
Opistognathus | aurifrons | JF297889 | n/a |
Opistognathus | aurifrons | JF297888 | n/a |
Opistognathus | aurifrons | JF297887 | n/a |
Opistognathus | aurifrons | JF297886 | n/a |
Opistognathus | aurifrons | JF297885 | n/a |
Opistognathus | aurifrons | JF297884 | n/a |
Opistognathus | aurifrons | JF297883 | n/a |
Opistognathus | aurifrons | JF297882 | n/a |
Opistognathus | aurifrons | JF297881 | n/a |
Opistognathus | aurifrons | JF297880 | n/a |
Opistognathus | aurifrons | JF297879 | n/a |
Opistognathus | aurifrons | JF297878 | n/a |
Opistognathus | aurifrons | JF297877 | n/a |
Opistognathus | aurifrons | JF297876 | n/a |
Opistognathus | aurifrons | JF297875 | n/a |
Opistognathus | aurifrons | JF297874 | n/a |
Opistognathus | aurifrons | JF297873 | n/a |
Opistognathus | aurifrons | JF297872 | n/a |
Opistognathus | aurifrons | JF297871 | n/a |
Opistognathus | aurifrons | JF297870 | n/a |
Opistognathus | aurifrons | JF297869 | n/a |
Opistognathus | aurifrons | JF297868 | n/a |
Opistognathus | aurifrons | JF297867 | n/a |
Opistognathus | aurifrons | JF297866 | n/a |
Opistognathus | aurifrons | JF297865 | n/a |
Opistognathus | aurifrons | JF297864 | n/a |
Opistognathus | aurifrons | JF297863 | n/a |
Opistognathus | aurifrons | JF297862 | n/a |
Opistognathus | aurifrons | JF297861 | n/a |
Opistognathus | aurifrons | JF297860 | n/a |
Opistognathus | aurifrons | JF297859 | n/a |
Opistognathus | aurifrons | JF297858 | n/a |
Opistognathus | aurifrons | JF297857 | n/a |
Opistognathus | aurifrons | JF297856 | n/a |
Opistognathus | aurifrons | JF297855 | n/a |
Opistognathus | aurifrons | JF297851 | n/a |
Opistognathus | aurifrons | JF297854 | n/a |
Opistognathus | aurifrons | JF297853 | n/a |
Opistognathus | aurifrons | JF297852 | n/a |
Opistognathus | aurifrons | JF297850 | n/a |
Opistognathus | aurifrons | JF297849 | n/a |
Opistognathus | aurifrons | JF297848 | n/a |
Opistognathus | aurifrons | JF297847 | n/a |
Opistognathus | aurifrons | JF297846 | n/a |
Opistognathus | aurifrons | JF297845 | n/a |
Opistognathus | aurifrons | JF297844 | n/a |
Opistognathus | aurifrons | JF297843 | n/a |
Opistognathus | aurifrons | JF297842 | n/a |
Opistognathus | aurifrons | JF297841 | n/a |
Opistognathus | aurifrons | JF297840 | n/a |
Opistognathus | aurifrons | FJ583762 | n/a |
Opistognathus | aurifrons | FJ583761 | n/a |
Opistognathus | aurifrons | FJ583760 | n/a |
Opistognathus | aurifrons | FJ583759 | n/a |
Opistognathus | aurifrons | FJ583758 | n/a |
Opistognathus | aurifrons | FJ583757 | n/a |
Opistognathus | aurifrons | FJ583756 | n/a |
Opistognathus | aff. aurifrons | MH751545 | CIUFES 2158 |
Opistognathus | aff. aurifrons | MH751543 | CIUFES 2306 |
Opistognathus | aff. aurifrons | MH751544 | CIUFES 2306 |
Opistognathus | aff. aurifrons | MH751542 | CIUFES 2306 |
Opistognathus | aff. aurifrons | MH751538 | CIUFES 2550 |
Opistognathus | aff. aurifrons | MH751539 | CIUFES 2551 |
Opistognathus | aff. aurifrons | MH751541 | CIUFES 2997 |
Opistognathus | aff. aurifrons | MH751540 | CIUFES 2997 |
Opistognathus | aff. jacksoniensis | JF911713 | n/a |
Opistognathus | lonchurus | MH751546 | CIUFES 2361 |
Opistognathus | macrognathus | JQ841946 | n/a |
Opistognathus | macrognathus | JN193388 | n/a |
Opistognathus | maxillosus | JQ840631 | n/a |
Opistognathus | maxillosus | JQ840630 | n/a |
Opistognathus | muscatensis | KU176461 | n/a |
Opistognathus | muscatensis | KU176379 | n/a |
Opistognathus | robinsi | JN193389 | n/a |
Opistognathus | thionyi | MH751535 | CIUFES 2054 |
Opistognathus | thionyi | MH751530 | CIUFES 2341 |
Opistognathus | thionyi | MH751531 | CIUFES 2341 |
Opistognathus | thionyi | MH751532 | CIUFES 2341 |
Opistognathus | thionyi | MH751536 | CIUFES 2394 |
Opistognathus | thionyi | MH751537 | CIUFES 2394 |
Opistognathus | thionyi | MH751534 | CIUFES 2421 |
Opistognathus | thionyi | MH751533 | CIUFES 2426 |
Opistognathus | vicinus | MH751528 | CIUFES 0236 |
Opistognathus | vicinus | MH751527 | CIUFES 2362 |
Opistognathus | vicinus | MH751529 | CIUFES 2395 |
Opistognathus | brasiliensis | MH751525 | CIUFES 3361 |
Opistognathus | maxillosus | LIDM1299-8* | n/a |
Opistognathus | whitehursti | LIDMA332-10* | n/a |
Opistognathus | whitehursti | LIDMA349-10* | n/a |
Opistognathus | whitehursti | MFLII323-7* | n/a |
Opistognathus | whitehursti | MFLII324-7* | n/a |
Opistognathus | whitehursti | MH751547 | UF 183104 |