Corresponding author: William F. Smith-Vaniz (
Academic editor: Javier Maldonado
A new species of jawfish,
Molecular phylogenetic analysis of partial cytochrome c oxidase subunit-I sequences indicates that specimens of the two allopatric pairs,
Smith-Vaniz WF, Tornabene L, Macieira RM (2018) Review of Brazilian jawfishes of the genus
Members of the family
Prior to this study, four species of
Summary of selected characters in Brazilian species of
Characters |
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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 |
1 Includes data from non-Brazilian specimens. 2 Fleshy spine tabs present only in adults.
Institutional abbreviations mostly follow
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 (
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 |
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0.195 | n/a | |||||||||||
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0.191 | 0.161 | 0.003 | ||||||||||
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0.181 | 0.141 | 0.120 | n/a | |||||||||
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0.176 | 0.135 | 0.138 | 0.132 | 0.002 | ||||||||
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0.169 | 0.152 | 0.162 | 0.132 | 0.090 | 0.009 | |||||||
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0.162 | 0.177 | 0.153 | 0.175 | 0.125 | 0.161 | 0.003 | ||||||
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0.173 | 0.159 | 0.149 | 0.159 | 0.143 | 0.165 | 0.111 | 0.000 | |||||
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0.151 | 0.170 | 0.176 | 0.170 | 0.142 | 0.173 | 0.139 | 0.165 | n/a | ||||
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0.189 | 0.163 | 0.184 | 0.167 | 0.159 | 0.176 | 0.194 | 0.186 | 0.134 | 0.002 | |||
(Caribbean Clade 2) | |||||||||||||
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0.181 | 0.171 | 0.186 | 0.178 | 0.160 | 0.180 | 0.184 | 0.193 | 0.128 | 0.032 | 0.006 | ||
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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) | |||||||||||||
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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 |
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– | 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 |
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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 |
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– | 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 |
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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 |
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– | 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 |
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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 |
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– | 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 |
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5 | Dorsal- and anal-fin rays 12 or 13; dentary without large canines; caudal vertebrae 16 |
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– | Dorsal- and anal-fin fin rays 14 or 15; dentary with large lateral canines; caudal vertebrae 17 |
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(12 specimens 23.4–53.5 mm SL) all from Brazilian Province:
A species of
Morphometric data are given in Table
Morphometric data for holotype and six paratypes of
Character | Holotype | Range | Mean | SD |
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Standard length (mm) | 45.5 | 32.5–53.5 | 40.7 | 7.5 |
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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 |
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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 |
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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
The Caribbean allopatric
Cephalic sensory pores in selected species of
Jaws and dentition (supramaxilla shaded) in selected species of
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.
Distributions of
The conservation status of
(12 specimens 17.0–47.4 mm SL) all from Brazil Province:
A species of
Morphometric data are given in Table
Morphometric data for holotype and six paratypes of
Character | Holotype | Range | Mean | SD |
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Standard length (mm) | 43.0 | 36.0–47.4 | 40.3 | 4.06 |
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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 |
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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 |
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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.
Genetic differences (see discussion in “Phylogenetic relationships of western Atlantic
From the Latin
A Brazilian endemic (Figure
The conservation status of this species [cited as
A species of
5 specimens (107.5–129 mm SL), including the holotype, cited in
A Brazilian endemic (Figure
In the diagnosis and description of
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.
A species of
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.
A species of
3 specimens (75.3–81.2 mm SL) from Brazil and 46 specimens (27–122 mm SL) from the Caribbean.
South Carolina, Gulf of Mexico, Greater Antilles and northern South America to Brazil (Figures
Distribution of
The conservation status of this species was assessed by the International Union for Conservation of Nature (IUCN) and listed as Lease Concern (
A species of
As provisionally recognized, this species is known only from the Brazilian Province including continental localities from the State of Maranhão (
33 specimens (30.4–73.7 mm SL) all from Brazilian Province.
Brazilian specimens of
Depending on the locality, adults of Caribbean
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
Bayesian inference phylogeny of western Atlantic
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
Bayesian inference phylogeny of the
Haplotype network of the
Previous studies of several populations of Caribbean
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 (
Genus | species | GenBank Accession Number | Voucher from this study |
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USNM 438666 |
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CIUFES 2158 |
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CIUFES 2306 |
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CIUFES 2306 |
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CIUFES 2306 |
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CIUFES 2550 |
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CIUFES 2551 |
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CIUFES 2997 |
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CIUFES 2997 |
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CIUFES 2361 |
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CIUFES 2054 |
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CIUFES 2341 |
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CIUFES 2341 |
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CIUFES 2341 |
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CIUFES 2394 |
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CIUFES 2394 |
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CIUFES 2421 |
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CIUFES 2426 |
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CIUFES 0236 |
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CIUFES 2362 |
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CIUFES 2395 |
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CIUFES 3361 |
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UF 183104 |
*Numbers from Barcode of Life Databse (BOLD), not GenBank.