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Review of Brazilian jawfishes of the genus Opistognathus with descriptions of two new species (Teleostei, Opistognathidae)
expand article infoWilliam F. Smith-Vaniz, Luke Tornabene§, Raphael M. Macieira|
‡ University of Florida, Gainesville, United States of America
§ University of Washington, Seattle, United States of America
| Universidade Vila Velha, Vila Velha, Brazil
¶ Universidade Federal do Espírito Santo, Vitória, Brazil
Open Access

Abstract

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. thionyiO. maxillosus and O. vicinusO. 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.

Keywords

Brazilian Province, Fernando de Noronha Archipelago, reef fish, Trindade-Martin Vaz insular complex, Vitória-Trindade Seamount Chain.

Introduction

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 (Hess 1993), which are occasionally aerated and cleaned. They are strongly territorial, and a single fish occupies each burrow. Jawfishes of the genus Opistognathus Cuvier, 1816 are widely distributed in tropical waters of the western Atlantic Ocean, where 14 valid species occur in depths of 0.3 to at least 375 m (Smith-Vaniz 1997; Smith-Vaniz 2017). An additional eleven species occur in the eastern Pacific (Bussing and Lavenberg 2003) but no species are shared by both oceans.

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 (Smith-Vaniz and Walsh 2017). Here, we (1) describe two new species of Opistognathus from Brazil, O. thionyi sp. n., and O. vicinus sp. n., and compare them with their Caribbean sister species, bringing the total number of species in Brazil to six, (2) present abbreviated descriptions, photographs or illustrations and distribution maps of all the other species of Opistognathus known from the Brazilian Province, (3) provide a partial molecular phylogeny based on sequence data from mitochondrial cytochrome oxidase I for available material of Atlantic species of Opistognathus, and (4) compare and discuss O. aff. aurifrons Brazilian populations. For the latter populations, we do not assign new scientific names to them because additional analysis of their complex phylogenetic relationships with Caribbean populations of the O. aurifrons complex is required to confidently determine their taxonomic status. Comparison of the six species of Opistognathus known from the Brazilian Province is given in Table 1.

Summary of selected characters in Brazilian species of Opistognathus. Exceptional values in parentheses.

Characters O. thionyi O. vicinus O. brasiliensis O. cuvierii O. lonchurus1 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

Materials and methods

Institutional abbreviations mostly follow Fricke and Eschmeyer (2018) and include the following collection depositories:

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania

CAS California Academy of Sciences, San Francisco, California

CIUFES Coleção Ictiológica, Departamento de Oceanografia e Ecologia, Universidade Federal do Espírito, Vitória, Espírito Santo, Brazil

FMNH Division of Fishes, Department of Zoology, Field Museum of Natural History, Chicago, Illinois

FSBC Florida Fish & Wildlife Conservation Commission, Fish & Wildlife Research Institute, St. Petersburg, Florida

MCZ Museum of Comparative Zoology, Harvard University, Ichthyology Department, Cambridge, Massachusetts

MNHN Museum National d’Histoire Naturelle, Systématique et Évolution, Laboratoire d’Ichyologie Générale et Appliquée, Paris

MNRJ Museu Nacional, Departmento de Vertebrados, Setor de Ictiologia, Universidade Federal do Rio de Janeiro, Brazil

MZUSP Universidade de São Paulo, Museu de Zoologia, São Paulo, Brazil

NPM Núcleo em Ecologia e Desenvolvimento Socio-Ambiental (NUPEM), Universidade Federal do Rio de Janeiro (UFRJ), Macaé, Brazil

SIO Scripps Institution of Oceanography, Marine Vertebrate Collection, La Jolla, California

SU Stanford University, collection transferred to CAS

UF University of Florida, Museum of Natural History, Gainesville, Florida

UFPB Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, Paraíba, Brazil

USNM Smithsonian Institution National Museum of Natural History, Department of Vertebrate Zoology, Division of Fishes, Washington, D.C.

ZUEC Universidade Estadual de Campinas, Museu de História Natural “Prof. Dr. Adão José Cardoso”, Museu de Zoologia, São Paulo, Brazil

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 Page et al. (2013).

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 (Shubart et al. 2007) or GobyL6468 and GobyH7696 (Thacker 2003). Sequences were aligned with several other opistognathid sequences from GenBank and BOLD (Barcode of Life Database, www.boldsystems.org), plus a sequence from Lonchopisthus micrognathus (Poey, 1860) as an outgroup, for a total of 103 sequences. Sequences were assembled and aligned in Geneious v.10.0.9. The best fitting substitution model was chosen using PartitionFinder (Lanfear et al. 2016). Phylogenetic analyses were conducted using maximum likelihood and Bayesian methods in RAxML version 8 (Stamatakis 2014) and MrBayes ver. 3.2.6 (Ronquist et al. 2012), respectively. A distance matrix was created using mean between-group pairwise genetic distances (Table 2); and a haplotype network was created using Haploviewer (Salzburger et al. 2011). GenBank or BOLD accession numbers are listed in the Appendix.

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)

Key to Brazilian Opistognathus

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 thionyi sp. n.

Figures 1, 2, 3, 4A, 5A, 7; Tables 1, 2, 3
Common name: Thiony’s jawfish
Nome comum (Portuguese): Peixe Bocão do Thiony

Opistognathus sp.: Simon et al. 2013a: 2116, 2120 (undescribed species; listed and photograph); Pinheiro et al. 2015: 15, color fig. S. 38 (Dogaressa Seamount); Pinheiro et al. 2018: 10, Southwestern Atlantic (SWA) Endemic reef fishes – Annotated Checklist: 31.

Holotype

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.

Paratypes

(12 specimens 23.4–53.5 mm SL) all from Brazilian Province: MNRJ 51283 (1, 39.9*) and ZUEC 16914 (1, 44.9*), 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 2344 (4, 23.4-32.7 & 32.5 C&S) and UF 239658 (1, 32.9), sandy rubble bottom at Praia do Lixo, Trindade Island, Brazil, 20°19'30"S, 29°19'20"W, 21 m, 21 February 2012, T. Simon and L.B.C. Xavier; AMNH 267140 (1, 32.5*), sandy rubble bottom at Praia do Lixo, Trindade Island, Brazil, 20°30'S, 29°20'W, 20 m, 20 February 2012, T. Simon and L.B.C. Xavier; CIUFES 2393 (1, 53.5), male, sandy rubble bottom at Portinho, Fernando de Noronha Archipelago, Brazil, 03°50'S, 32°24'W, 15 m, 11 July 2012, R.M. Macieira and H.T. Pinheiro; NPM 5029 (1, 38.4*), male, sandy rubble bottom at Enseada dos Portugueses, Trindade Island, Brazil, 20°31'30"S, 29°19'20"W, 25m, 8 August 2012, T. Simon and E.F. Mazzei; MZUSP 123868 (1, 30.9) and USNM 440401 (1, 35.3*), sandy rubble bottom at Praia do Lixo, Trindade Island, 20°31'30"S, 29°19'20"W, 17 m, 7 August 2012, T. Simon and L.B.C. Xavier.

Non-type material

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.

Diagnosis

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).

Description

Morphometric data are given in Table 3 for the holotype and specimens indicated above by an asterisk; other comparative features are presented in Table 1. Where counts differ, those of the holotype are given first, followed in parentheses by those of the paratypes. Dorsal fin XI, 15 (15–16). Anal fin III, 15 (14–15, usually 15). Pectoral-fin rays 20 (19–20). Vertebrae: 10+18, last pleural rib on vertebra 10, epineurals 13–15. Supraneurals absent. Caudal fin: procurrent rays 5+5 (4-6+4-5); segmented rays 8+8, middle 12 branched, total elements 26 (24–26); hypural 5 absent. Gill rakers (number not increasing with increase in SL in adults) 9+19 (8–11+17–20=25–31).

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 4A).

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 4A) and dorsalmost spine of opercle not noticeably elongate; posterior margin of preopercle distinct, with a well-developed groove dorsally. No papillae on inner surface of lips. Fifth cranial nerve passes over A1β section of adductor mandibulae muscle.

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 5A). Premaxilla anteriorly with an outer row of stout teeth and an inner row of smaller, backward slanting teeth, some nearly horizontal; laterally teeth uniserial and becoming progressively smaller and more closely spaced. Dentary anteriorly with an outer row of stout teeth and an inner row of smaller, backward slanting teeth; laterally teeth uniserial and smaller but not progressively so. Vomer with 1 large tooth. Infraorbital bones tubular, with numerous openings for sensory canals; third infraorbital with a wide suborbital shelf. Postcleithra consisting of two well separated bones; dorsal postcleithrum an irregular elongate oval, narrowest ventrally, ventral postcleithrum rod-shaped with pointed ends.

Color in life (Figures 1, 2). Body coloration chestnut brown to dark brown with five very irregular and poor defined dark bands that extend onto base of dorsal fin; head sometimes with pale speckles on posterior half; upper jaw with a wide white band near posterior end; eyes reddish brown sometimes with narrow pale radiating bands; lips with alternating dark and pale bands; branchiostegal membranes dark, especially in mature males; dorsal fin light yellow sometimes with diagonal rows of pale spots and a black blotch, when present, between spines 2–5; anal fin with small pale spots; pelvic fins dark; pectoral fins speckled and a large white spot on pectoral-fin base; caudal fin with pair of pale basicaudal spots and fin rays with prominent black speckles or narrow bands.

Figure 1. 

Opistognathus thionyi, CIUFES 2054, 27.5 mm SL, Dogaressa Seamount, Vitória-Trindade Chain. Photograph by Raphael M. Macieira.

Figure 2. 

Opistognathus thionyi, NPM 5029, 38.4 mm SL, Enseada dos Portugueses, Trindade Island, Brazil. Photograph by Thiony Simon.

Preserved color (Figure 3). As above except with white, brown, and black markings. Inner margin of maxilla posteriorly and adjacent membranes with dusky blotch. Buccal area surrounding esophageal opening pale.

Figure 3. 

Opistognathus thionyi, CIUFES 2347, preserved holotype, 45.4 mm SL, male, Trindade Island, Brazil. Photograph by Zackary S. Randall.

Comparisons

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 Smith-Vaniz (1997). The only other species of Opistognathus with a broad, fan-like subopercle are the eastern Pacific O. galapagensis Allen & Robertson, 1991 and O. fossoris Bussing & Lavenberg, 2003. In addition to other characters discussed by Bussing and Lavenberg (2003), both species differ from O. thionyi most notably in having the posterior end of maxilla ending as thin, flexible lamina (vs. end of maxilla rigid), more body longitudinal scale rows, 83–105 (vs. 45–52), most of nape immediately in front of dorsal-fin origin without cephalic pores (vs. almost completely covered with pores), and in having very different color patterns. Comparison of the six species of Opistognathus known from the Brazilian Province is given in Table 1.

Figure 4. 

Cephalic sensory pores in selected species of Opistognathus. A O. thionyi, 45.4 mm SL, holotype B O. vicinus, NPM 5030, 47.4 mm SL, Brazil C O. lonchurus, CIUFES 1426, 75.3 mm SL, Brazil D O. aff. aurifrons, CIUFES 1450, 57.0 mm SL, Brazil.

Figure 5. 

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, ANSP 126637, 75.0 mm SL, Gulf of Mexico D O. aff. aurifrons, ANSP 188905, 63.8 mm SL, Brazil.

Etymology

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.

Distribution, habitat, and natural history

Opistognathus thionyi is known only from three oceanic sites, Trindade Island, Dogaressa Seamount, and Fernando de Noronha Archipelago (Figure 6), and is an endemic species of the Brazilian Province (sensu Briggs and Bowen 2012 and Pinheiro et al. 2018). This species is possibly broadly distributed along the coast on the outer shelf, an area that is virtually unsampled. It has been recorded from 10–65 m, and found solitarily, always in small constructed burrows on sandy rubble bottoms (Figure 7). It feeds on small benthic organisms that live near the bottom (e.g., small shrimps).

Figure 6. 

Distributions of Opistognathus species in Brazilian Province; light shaded areas indicate continental shelf.

Figure 7. 

Opistognathus thionyi, in typical habitat, sandy rubble bottom in 13 m at Enseada dos Portugueses, Trindade Island, Brazil. Photograph by Thiony Simon.

Conservation

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 Brasil 2018; Giglio et al. 2018; Magris and Pressey 2018), could threat the existence of Thiony’s jawfish in that part of its range in the near future.

Opistognathus vicinus sp. n.

Figures 4B, 5B, 8, 9; Tables 1, 2, 4
Common name: Brazilian dusky jawfish
Nome comum (Portuguese): Peixe Bocão escuro

Opistognathus whitehursti (Longley, 1927): Simon et al. 2013b (listed).

Holotype

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.

Paratypes

(12 specimens 17.0–47.4 mm SL) all from Brazil Province: UF 239659 (2, 27.9–31.0), taken with the holotype; CIUFES 0868 (1, 38.3*), Ilha Rasa de Dentro, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 10 m, 29 January 2008, V.C. Brilhante; USNM 440402 (1, 38.6*), Ilha das Garças, Vila Velha, Espírito Santo, 20°36'S, 40°22'W, 30 March 2000, J.L. Gasparini; MZUSP 123869 (1, 38.9), MNRJ 51284 (1, 17.0), ZUEC 16915 (1, 42.5*) and AMNH 267141 (2, 21.7–38.9*), Ilha Escalvada, Guarapari, Espírito Santo, 20°42'S, 40°24'W, 21 m, 23 February 2010, R.M. Macieira, T. Simon and C.R. Pimentel; NPM 5030 (1, 47.4*), Ilha Rasa de Dentro, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 31 October 2005, R.M. Macieira and J.-C. Joyeux; CIUFES 0467 (1, 26.2), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 17 m, 5 December 2005, R.M. Macieira and J.-C. Joyeux; CIUFES 131797 (1, 36.0), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°21'W, 14 August 1989, J.L. Gasparini.

Diagnosis

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).

Description

Morphometric data are given in Table 4 for the holotype and specimens indicated above by an asterisk; other comparative features are presented in Table 1. Where counts differ, those of the holotype are given first, followed in parentheses by those of the paratypes. Dorsal fin XI, 14. Anal fin II, 13 (II or III [anterior spine minute if III], 12–13, usually II, 13). Pectoral-fin rays 18 (17–18). Vertebrae: 10+17, last pleural rib on vertebra 10, epineurals 12–14. Supraneurals absent. Caudal fin: procurrent rays 4+4 (5–4+3–4); segmented rays 8+8, middle 12 branched, total elements 24 (23–25); hypural 5 absent. Gill rakers (number not increasing with increase in SL in adults) 7+16 (7–8+15–17=23–25).

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 4B).

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 4B) and dorsalmost spine of opercle moderately elongate; posterior margin of preopercle distinct, with a well-developed groove dorsally. No papillae on inner surface of lips. Fifth cranial nerve passes over A1β section of adductor mandibulae muscle.

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 8). Background color of head and body brownish to reddish brown. Body with six vertically irregular and evenly spaced dark bands, widest on mid-side, that extend onto base of dorsal fin; two rows of six pale spots on sides, one along dorsal-fin base and the other along anal-fin base, each spot approximately diameter of eye; upper jaw with a wide dark band behind which is a white band at posterior end; the eyes are red; the lips with dark and pale bands; branchiostegal membranes dark; dorsal fin with dark stripe, widest anteriorly, along middle of fin, and a dark blue blotch between the second and fourth spine; pectoral fins are translucent; pelvic fins pale blue to dark or entirely pale; caudal fin with pair basicaudal spots bordered posteriorly by dark continuous band and remainder of fin vertical rows of dark spots or narrow bands.

Figure 8. 

Opistognathus vicinus, CIUFES 0796, holotype, 43.0 mm SL, male, Ilha Rasa de Dentro, Guarapari, Espírito Santo, Brazil. Photograph by Raphael M. Macieira.

Preserved color (Figure 9). Body with dark bands and large pale spots as above; other makings various shades of brown. Inner margin of maxilla posteriorly and adjacent membranes with brownish blotch, best developed in males. Buccal area surrounding esophageal opening pale.

Figure 9. 

Opistognathus vicinus, NPM 5030, preserved paratype, 47.4 mm SL, Ilha Rasa, Guarapari, Espírito Santo, Brazil. Photograph by Zackary S. Randall.

Figure 10. 

Opistognathus whitehursti, USNM 396062, 22.0 mm SL, Saba Bank. Photograph by Jeffrey T. Williams.

Figure 11. 

Opistognathus whitehursti, UF 183105, 43.0 mm SL, St. Croix. Photograph by Howard L. Jelks.

Comparisons

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 Smith-Vaniz 1997: fig. 33). Initially we considered both species to be only “genovariants” sensu Victor (2015). However, O. vicinus lacks the small supramaxilla (see Smith-Vaniz 1997: fig. 32a) present in O. whitehursti and vomer without teeth (typically one or two teeth present in O. whitehursti). The color pattern of juveniles of O. whitehursti (Figure 10) is virtually identical to those of adult Brazilian fish. Unlike Opistognathus vicinus, adults of O. whitehursti usually have more reduced body bands (Figure 11) and the spinous dorsal-fin spot is often absent. As noted by Böhlke and Chaplin (1968: 486) for Bahamas fish, “The spot on the spinous dorsal fin is … blue in color in young, frequently missing or poorly defined in adults.” Thus, the combined differences in COI, adult coloration, and the lack of a supramaxilla and vomerine teeth support the recognition of O. vicinus as a distinct species. Comparison of the six species of Opistognathus known from the Brazilian Province is given in Table 1.

Etymology

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.

Distribution and Habitat

A Brazilian endemic (Figure 6), known from Ceará to Espírito Santo State but absent from oceanic islands. Common in coastal regions, in depths of 10–25 meters, associated with gravel-sand bottoms, near coral reefs and rocky areas. Feeds on small benthic organisms near the bottom (e.g., small shrimps, crabs, and isopods).

Conservation

The conservation status of this species [cited as Opistognathus whitehursti (Longley 1927) – unpublished data] 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.

Opistognathus brasiliensis Smith-Vaniz, 1997

Figures 12, 13; Tables 1, 2
Common name: Darkfin jawfish
Nome comum (Portuguese): Peixe Bocão-da-Nadadeira-Negra

Opistognathus brasiliensis Smith-Vaniz, 1997: 1104, fig. 20 (original description; Alcatraces [misspelled Alcatrazes] Island: holotype MZUSP 13257); Carvalho-Filho 1999: 194 (abbreviated description); Menezes 2011: 42 (listed); Mincarone et al. 2017: 207 (listed); Pinheiro et al. 2018, Southwestern Atlantic (SWA) Endemic reef fishes – Annotated Checklist: 28.

Abbreviated description

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) (Smith-Vaniz 1997, Figure 9c).

Figure 12. 

Opistognathus brasiliensis, CIUFES 3361, 60.6 mm SL, female, Ilhas Escalvada, Guarapari, Espírito Santo, Brazil. Photograph by Raphael M. Macieira.

Figure 13. 

Opistognathus brasiliensis, ANSP 141861, 122.5 mm SL, male, southeastern Brazil. Drawn by Jack R. Schroeder; after Smith-Vaniz (1997).

Material examined

5 specimens (107.5–129 mm SL), including the holotype, cited in Smith-Vaniz (1997) plus the following additional material: CIUFES 1459 (1, 64.0), Ilhas das Garças, Vila Velha, Espírito Santo, 20°36'S, 40°22'W, 30 March 2000, J.L. Gasparini.

Distribution, habitat, and natural history

A Brazilian endemic (Figure 6), known from Espírito Santo to São Paulo State, but absent from oceanic islands. A rare species only known from the type series collected in São Paulo’s coastal waters (trawled in 50–69 m) and two specimens from Guarapari Islands, Espírito Santo. Occurs in depths of 15–69 meters, associated with gravel and sand or silt and sand bottoms, near coral reefs and rocky areas.

Remarks

In the diagnosis and description of Opistognathus brasiliensis, Smith-Vaniz (1997) stated, in part, “buccal pigmentation consisting of a dark area widely surrounding esophageal opening …” versus area around esophageal opening pale in O. cuvierii. This reported distinction is no longer valid because in a recently examined female specimen of Opistognathus brasiliensis (CIUFES 1459) the area around the esophageal opening is pale. Both species have black areas in front of each pharyngeal tooth patch that are separated by a pale median area.

Conservation

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.

Opistognathus cuvierii Valenciennes, 1836

Figures 14, 15; Tables 1, 2
Common name: Bartail jawfish
Nome comum (Portuguese): Peixe Bocão-de-Cauda-com-Barras

Opisthognathus [sic] cuvierii Valenciennes in Cuvier and Valenciennes 1836: 504, color pl. 343 (original description; Bahia: holotype MNHN A. 2108); Roux 1964: 413, pl. 10 (listed; original illustration of holotype reproduced).

Opistognathus cuvierii: Menezes and Figueiredo 1985: 42, fig. 47 (description); Smith-Vaniz 1997: 1106, fig. 21 (description); Pinheiro et al. 2018, Southwestern Atlantic (SWA) Endemic reef fishes – Annotated Checklist: 28–29, color fig. 18.

Opistognathus cuvieri: Roux 1973: 151 (description); Carvalho-Filho 1999: 194 (abbreviated description and occurrence to São Paulo); Mincarone et al. 2017: 207 (listed).

Abbreviated description

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) (Smith-Vaniz 1997: fig. 9d), or these blotches absent (females); buccal pigmentation consisting of a dark blotch on either side of esophageal opening widely separated by pale median area that continues between upper pharyngeal tooth patches (Smith-Vaniz 1997: fig. 13c).

Figure 14. 

Opistognathus cuvierii, 80 mm TL estimated, Ilha dos Frades, Baía de Todos os Santos, Bahia, Brazil. Photograph by Cláudio L. S. Sampaio.

Figure 15. 

Opistognathus cuvierii, SU 52354, 102 mm SL, male, Bahia, Brazil. Drawn by Jack R. Schroeder; after Smith-Vaniz (1997).

Material examined

5 specimens (80.5–11.5 mm SL), including the holotype, cited in Smith-Vaniz (1997).

Distribution, habitat, and natural history

A Brazilian endemic (Figure 6), known from Bahia to São Paulo but absent from oceanic islands. Distributed in coastal regions, in depths between 1–35 meters, associated with gravel and sand or silt and sand bottoms, near coral reefs and rocky areas. This species has been observed resting on the bottom (Figure 14) or in burrows (Pinto 1970).

Conservation

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.

Opistognathus lonchurus Jordan & Gilbert, 1882

Figures 4C, 5C, 16; Tables 1, 2
Common name: Moustache jawfish
Nome comum (Portuguese): Peixe Bocão-de-Bigode

Opisthognathus [sic] lonchurus Jordan & Gilbert, 1882: 290 (original description; snapper banks off Pensacola, Florida: holotype USNM 30864); Böhlke and Thomas 1961: 514–515, Table 2.

Opistognathus lonchurus: Floeter and Gasparini 1999: 58 (Brazil occurrence); Carvalho-Filho 1999: 194 (abbreviated description).

Abbreviated description

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 4C); dorsal fin XI, 12–13, with spines thin and flexible, usually curved distally, tips without pale, slightly swollen tabs, and anterior 5–8 soft rays unbranched; dorsal fin sexually dimorphic, fin rays distinctly higher in large males; anal fin III, 12, with 7–10 anterior rays unbranched; outermost segmented pelvic-fin ray tightly bound to adjacent ray and interradial membrane not incised distally; body with 63–87 oblique scales in longitudinal series; vertebrae 10+16; supraneurals 1; gill rakers 13–17+22–28=35–45; dentary without large canines (Figure 5C). Head and body brown to greenish-tan; upper lip blue; body with two narrow blue stripes on side; distal margins of dorsal and anal fins with narrow blue stripe; outer margin of caudal fin and outer ray of pelvic fin blue.

Figure 16. 

Opistognathus lonchurus, CIUFES 2361, 81.2 mm SL, female, Ilhas Escalvada, Guarapari, Espírito Santo, Brazil. Photograph by Raphael M. Macieira.

Material examined

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; ZUEC 3160 (1), Mar do Bom Nome, Espírito Santo, 21°09'S, 40°30'W, 30 m, 8 December 1996, J.L. Gasparini. Caribbean, only abbreviated localities, standard lengths and depths given: ANSP 83680 (1, 88), Haiti; ANSP 126637 (1, 75 C&S), off Mobile, Alabama, 35 m; ANSP 134237 (1, 96), Puerto Rico, 49 m; ANSP 138138 (1, 86), Puerto Rico, 52 m; ANSP 138362 (1, 93), Puerto Rico, 52 m; ANSP 140955 (5, 77-97), Puerto Rico, 50 m; ANSP 142700 (1, 107), Puerto Rico; 14 m; ANSP 174157 (1, 83), Gulf of Mexico, 74 m; ANSP 177885 (2, 82-97), Gulf of Mexico, 63 m; ANSP 136545 (1, 92), off South Carolina, 60 m; CAS 1863 (1, 84), CAS 29250 (1, 84) and CAS 36683 (1, 95), Pensacola, Florida; FSBC 1706 (2, 85.5-94), 32°01'N, 79°24'W, R/V Silver Bay sta. 1788, 64–82 m; FMNH 79582 (1, 76), off Guyana, 49–55 m; FSBC 3314 (1, 107), off St. Petersburg, Florida, 47 m; FSBC 3324 (1, 93), Gulf of Mexico, 37 m; FSBC 12575 (1, 99), Florida Keys, 76–80 m; MCZ 52103 (2, 112-122), 55 m; Dry Tortugas, 55 m; SIO 70–186 (2, 77–97), Florida Keys, 40 m; SIO 70–186 (2, 76–96), Florida Keys, 40 m; SIO 70–224 (1, 47), Florida Keys, 38 m; UF 186218 (1, 80), Gulf of Mexico; UF 186226 (1, 89), Gulf of Mexico, 61 m; UF 186239 (1, 102), Gulf of Mexico; UF 218742 (1, 28), Florida, 38 m; UF 203991 (1, 90), off Guyana, 55–60 m; UF 219018 (1, 35), Florida, 40 m; UF 238298 (1, 196), Gulf of Mexico; USNM 31903 (1, 64), Pensacola, Florida; USNM 34976 (1, 69), holotype of Gnathypops mystacinus), Pensacola, Florida; USNM 117035 (1, 27), Dry Tortugas; USNM 217802 (1, 96), off Colombia, 73 m; USNM 217083 (1, 106) off Colombia, 84 m; USNM 30712 (1, 104), Pensacola, Florida; USNM 358160 (1, 68), Gulf of Mexico, 62 m; USNM 358161 (1, 73), Gulf of Mexico, 67 m.

Distribution, habitat, and natural history

South Carolina, Gulf of Mexico, Greater Antilles and northern South America to Brazil (Figures 6, 17) in about 15–90 m. In the Brazilian Province known from the northern coast to Espírito Santo State but absent from oceanic islands. Occur in depths of 10–91 meters, in rubble-sand bottoms near coral reefs and rocky areas. It has been observed in burrows or resting on the bottom.

Figure 17. 

Distribution of Opistognathus lonchurus, including a few records from Robertson and Van Tassell (2015).

Conservation

The conservation status of this species was assessed by the International Union for Conservation of Nature (IUCN) and listed as Lease Concern (Smith-Vaniz et al. 2015). The conservation status of this species was also assessed by the Ministério do Meio Ambiente/Instituto Chico Mendes de Conservação de Biodiversidade (MMA/ICMBio – Brazil) and listed as Least Concern.

Opistognathus aff. aurifrons Jordan & Thompson, 1905

Figures 4D, 5D, 18, 19, 20, 21; Tables 1, 2
Common name: Yellowhead jawfish
Nome comum (Portuguese): Peixe Bocão-da-Cabeça-Amarela

Opistognathus aurifrons Jordan & Thompson, 1905: 252, fig. 4 (original description; Garden key, Dry Tortugas, Florida); Moura et al. 1999: 517 (Brazilian occurrence); Pereira-Filho et al. 2015: 69–70 (Listed, Fernando de Noronha Archipelago); Stocco and Joyeux 2015: 6 (ichthyoplankton, Trindade Island).

Opistognathus sp. Rocha and Rosa 2001: 990 (listed; Manuel Luiz Marine State Park, Brazil); Gasparini and Floeter 2001: 1644 (listed; Trindade Island); Rocha 2002: 477, unnumbered color fig. (Parcel Manuel Luiz, Brazil); Menezes et al., 2003: 78; Moura and Sazima 2000: 482 (listed); Sampaio and Nottingham 2008: 171 (abbreviated description).

Opistognathus sp. 1 Carvalho-Filho 1999: 193, color fig. 185; Pinheiro et al. 2018, Southwestern Atlantic (SWA) Endemic reef fishes – Annotated Checklist: 29–30, color fig. 19 (Fernando de Noronha Archipelago).

Opistognathus sp. 2 Smith-Vaniz 1997: 1096 (in identification key); Simon et al. 2013a: 2120 (listed); Pinheiro et al. 2018, Southwestern Atlantic (SWA) Endemic reef fishes – Annotated Checklist: 30–31, color fig. 20.

Opistognathus aff. aurifrons Jordan & Thompson: Feitoza et al. 2005: 732 (Brazilian Province in 35–54 m); Simon et al. 2013b: 63 (listed); Pinheiro et al. 2015: 15, color fig. S.37 (Trinidade Island and Dogaressa Seamount).

Abbreviated description (Brazilian specimens only)

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 4D); dorsal fin XI, 14–15, with 6–11 anterior rays unbranched distally; anal fin III, 14–15, with 7–10 anterior rays unbranched; outermost segmented pelvic-fin ray tightly bound to adjacent ray and interradial membrane not incised distally; scales in longitudinal series 66–76; vertebrae 10+17; supraneurals absent; gill rakers 15–20+26–32=41–51; dentary with large lateral canines (Figure 5D). Life color of adults of the two different color morphs as in Figs 1820 and discussed below in Remarks.

Figure 18. 

Opistognathus aff. aurifrons, CIUFES 0795-1, 61.8 mm SL, female, Ilhas Escalvada, Guarapari, Espírito Santo, Brazil. Photograph by Raphael M. Macieira.

Figure 19. 

Opistognathus aff. aurifrons, adult, Bellucia shipwreck, Guarapari, Espírito Santo, Brazil. Photograph by Raphael M. Macieira.

Figure 20. 

Opistognathus aff. aurifrons, CIUFES 2551, 69.2 mm SL, Fernando de Noronha Archipelago, Brazil. Photograph by Raphael M. Macieira.

Distribution

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 6). It is common in coastal regions, in depths of 10–30 meters, and in oceanic sites of 10–65 meters, associated with rubble and sand bottoms, near coral reefs and rocky areas. Its behavior and life history are similar of that of the Caribbean O. aurifrons (Colin 1971; Colin 1973). In this species the burrows are constructed on sandy or rubble bottoms, near reefs, using small stones, shell or coral fragments. They feed on zooplankton while hovering at a small distance over the substrate during quick incursions and generally retreat tail first into the burrow.

Material examined

33 specimens (30.4–73.7 mm SL) all from Brazilian Province. Mainland localities: ANSP 188905 (2, 35.4–63.8 C&S), Naufrágio Bellucia (shipwreck), off Guarapari, Espírito Santo, 20°40'S, 40°21'W, 24.6 m, 29 August 2008, A. Carvalho-Filho, R.M. Macieira and C.R. Pimental; CAS 238006 (3, 53.2–57.5), Ilha Escalvada, Guarapari, Espírito Santo, 20°42'S, 40°24'W, 30 March 2012, L.A. Rocha; CAS 238007 (4, 58.8–70.3), Ilha Escalvada, Guarapari, Espírito Santo, 20°42'S, 40°24'W, 15 February 2012, L.A. Rocha; CIUFES 0795 (4, 24.0–66.9), Naufrágio Bellucia (shipwreck), off Guarapari, Espírito Santo, 20°40'S, 40°21'W, 27.0 m, 11 March 2008, J.-C. Joyeux, R.M. Macieira and V.C. Brilhante; CIUFES 1450 (2, 57.1–73.7 C&S), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°22'W, 15 m, 14 August 1999, J.L. Gasparini; MZUSP 44937 (1, 52.4), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°21'W, January 1992, J.L. Gasparini; MZUSP 46191 (1, 67.5), male, Cabo Frio, Rio de Janeiro, 22°53'S, 42°00'W, March 1991, A. Carvalho-Filho; MZUSP 46541 (1, 61.8), gravid female, Vitória, Espírito Santo, 20°19'S, 40°21'W, December 1990, A. Carvalho-Filho; MZUSP 52271 (2, 35.0–42.8), Ilha Sueste, Abrolhos Archipelago, 17°58'S, 38°41'W, 11 January 1997, I. Sazima, C. Sazima, J.L. Gasparini and R.L. Moura; MZUSP 52453 (1, 46.9) and UF 191039 (3, 30.4–51.6), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°22'W, 22 April 1992, D.A. Jório; ZUEC 3105 (1, 72.6), Ilhas Rasas, Guarapari, Espírito Santo, 20°40'S, 40°22'W, 18 m, 1 June 1996, D.A. Jório; ZUEC 2739 (1, 61.6), gravid female, Ilha Escalvada, Guarapari, Espírito Santo, 20°42'S, 40°24'W, 16 m, July 1995, D.A. Jório and J.L. Gasparini; UFPB 4047 (5, 52.4–67.5), Três Ilhas Archipelago, Guarapari, Espírito Santo, 20°36'S, 40°22'W, 1 December 1997, J.L. Gasparini. Fernando de Noronha Archipelago: CIUFES 2550 (1, 58.4) and CIUFES 2551 (1, 69.2), Cabeço Submarino, 03°52'S, 32°25'W, 19.6 m, 8 April 2013, R.M. Macieira and T. Simon.

Remarks

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 1819). Populations from the Fernando de Noronha Archipelago differ in having short pelvic fins that do not extend to the anal-fin origin (20.1–20.7% SL in 2 specimens 58.4–69.2 mm SL) and in fresh adults the head is pale tan-yellow, the body is pale grey-blue, and both are uniformly colored (Figure 20). To further complicate the situation, uncollected juveniles from Fernando de Noronha (Figure 21) and Bonaire (Figure 22) have identical life coloration consisting primarily of a white head crossed by diagonal brown-orange stripe about width of pupil extending from chin, through eye and across nape; body and fins pale grey. In preservation, adults from Tobago (USNM 317005) have the same head color pattern and long pelvic fins as those of the Brazilian mainland population but differ in having 16 caudal vertebrae. Unfortunately, life coloration was unrecorded and tissue samples were not obtained. A color photograph taken by Les Wilkes of a “Bluebar jawfish” from St. Vincent, Lesser Antilles (13°15'N, 61°12'W) also looks like mainland Brazilian adults. Overall, there are morphological characters that collectively differentiate the two Brazilian clades from the Caribbean haplotypes (i.e., vertebral counts) and from each other (i.e., pelvic fin length, but note the low sample size), but we know of no consistent phenotypic characters that differentiate the two Caribbean haplotypes from each other. The type locality for O. aurifrons is Dry Tortugas, Florida, making it likely that if the main groups here do indeed represent distinct species, the group containing specimens from Florida (green in Figure 24) represents the true O. aurifrons and the others new species. However, we refrain from making taxonomic changes pending a more thorough analysis comparing multiple genetic loci, and live coloration/morphology of vouchered specimens.

Figure 21. 

Opistognathus aff. aurifrons, juvenile, Fernando de Noronha Archipelago, Brazil. Photograph by João P. Krajewski.

Figure 22. 

Opistognathus aurifrons, juvenile, Bonaire. Photograph by Scott Michael.

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.

Conservation

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.

Phylogenetic relationships of western Atlantic Opistognathus

The molecular phylogenies inferred from the COI data using ML and Bayesian inference were very similar in topology (Figure 23); also see supporting information for ML trees. Most basal nodes were not well supported in either analysis, indicating that additional slower evolving genetic markers are needed to better resolve the phylogeny. Several clades containing western Atlantic Opistognathus were well supported, including a clade containing O. whitehursti and O. vicinus, another clade containing O. lonchurus and O. aurifrons, and a clade containing O. maxillosus, O. thionyi, O. brasiliensis, O. robinsi, and O. macrognathus. In the latter clade, the new species O. thionyi was resolved as a sister species to O. maxillosus, with that clade being sister to a clade containing O. robinsi, O. brasiliensis, and O. macrognathus.

Figure 23. 

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 24 for this clade expanded). Scale bar units are expected number of substitutions per site.

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 2). Subsequent analysis of coloration and morphology revealed subtle differences that support the recognition of these mitochondrial lineages as distinct species (see “Comparisons” in O. vicinus description). Opistognathus aurifrons formed four distinct groups (Figs 2425), with one large polyphyletic grade from the Caribbean, a clade nested with this grade from Aruba and Curacao, a clade from mainland Brazil and Vitória-Trindade Seamounts Chain, and a pair of sequences from Fernando de Noronha Archipelago. Genetic variation in COI was very low within each of these groups (mean p-distance ≤0.006), but moderate between each group (mean p-distance between groups = 0.016–0.045). A haplotype map of the Opistognathus aurifrons complex (Figure 25) confirms the four main clusters of haplotypes, which are separated from one another by at least nine mutations in the partial COI gene. The strong connection between the Trindade and mainland populations is probably related to the stepping-stone process provided by Vitória-Trindade Seamounts Chain (Floeter and Gasparini 2000; Macieira et al. 2015; Pinheiro et al. 2017, 2018).

Figure 24. 

Bayesian inference phylogeny of the Opistognathus aurifrons complex. Support values are Bayesian posterior probabilities. Nodes without support values shown have <0.90 support. Scale bar units are expected number of substitutions per site.

Figure 25. 

Haplotype network of the Opistognathus aurifrons complex. Dark solid circles represent un-sampled intermediate haplotypes, lines connecting haplotypes represent single substitutions.

Previous studies of several populations of Caribbean O. aurifrons based on morphology (Böhlke and Thomas 1961) and genetics (Ho et al. 2012) indicated that despite substantial variation in color pattern within and between populations, only a single species should be recognized. Several other western Atlantic reef fishes, especially small, cryptobenthic fishes like gobies and blennioids, show similar patterns where there is substantial mitochondrial divergence without detectable phenotypic differences (Victor et al. 2015; Victor 2015). Some examples include the multiple genetic lineages within Bathygobius soporator (Tornabene et al. 2010; Tornabene and Pezold 2011), Malactoctenus triangulatus (Victor 2015), several species of Starksia (Baldwin et al. 2011), and some species in the Tigrigobius (Elacatinus) multifasciatus/panamensis complex (Victor 2010, 2014). In these instances, nominal species comprise multiple highly-divergent mitochondrial lineages, each with very little genetic variation within lineages, and no recognizable phenotypic differences between lineages (such groups have been termed “genovariants” by Victor 2015). In these studies, and in most cases of such ‘cryptic species’ of reef fishes, authors have conservatively refrained from naming them as new species based solely on the presence reciprocally monophyletic mitochondrial lineages. Part of this is for practical reasons; taxonomists often prefer species descriptions to be operational, and thus having observable diagnostic characters is critically important. However, it is also possible that such genetic lineages don’t correspond to distinct species, and instead are the result of incomplete lineage sorting, or in the case of allopatric lineages, simply representative of geographic population-genetic structure. For these reasons, we follow convention and refrain from naming the distinct lineages of O. aurifrons, pending additional genetic and phenotypic data. However, for conservation purposes it is important to consider these four lineages separately.

Acknowledgments

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 (MNRJ), Fabio Di Dario and Michael Maia Mincarone, Coleção de Peixes do NUPEM/UFRJ (NPM), Sandra Raredon (USNM), Flávio C. T. Lima (ZUEC-PIS) and Michel Donato Gianeti, Mario de Pinna, Aléssio Datovo da Silva, and José Lima de Figueiredo (MZUSP). Important color photographs of jawfishes were generously provided by João L. Gasparini, Howard L. Jelks, J.-C. Joyeux, João P. Krajewski, Scott Michael, Zachary S. Randall, Cláudio L. Sampaio, Thiony Simon, and Jeffery T. Williams. Hudson T. Pinheiro kindly provided three specimens of the new species he had collected from Fernando de Noronha. Thanks to J.-C. Joyeux, C.R. Pimentel, L.B. Xavier, J.L. Gasparini, C.L.S. Sampaio, and A. Carvalho-Filho for collecting Opistognathus specimens with us. We thank Michael S. Netto, Lúcio Engler, Carlos Janovitch, Rebreather Clube do Brasil, Inner-Space Systems Corp., Liquivision Products, Inc., Atrasorb Absorvedores de CO2, Seasub, Águas Claras (Fernando de Noronha), Subaquática, and Piranha Dive Mfg. for diving support and logistics; the Abaeté crew for help and friendship on-board; TAMAR/ICMBio, ICMBio, IBAMA and the Brazilian Navy for logistics and permits. This study was supported in part by fellowships to R.M. Macieira (CAPES).

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Appendix

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