Research Article |
Corresponding author: Iria Fernandez-Silva ( iriafernandezsilva@gmail.com ) Academic editor: Devin Bloom
© 2016 Iria Fernandez-Silva, John E. Randall, Daniel Golani, Sergey V. Bogorodsky.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Fernandez-Silva I, Randall JE, Golani D, Bogorodsky SV (2016) Mulloidichthys flavolineatus flavicaudus Fernandez-Silva & Randall (Perciformes, Mullidae), a new subspecies of goatfish from the Red Sea and Arabian Sea. ZooKeys 605: 131-157. https://doi.org/10.3897/zookeys.605.8060
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The number of goatfish species has increased recently, thanks in part to the application of molecular approaches to the taxonomy of a family with conservative morphology and widespread intraspecific color variation. A new subspecies Mulloidichthys flavolineatus flavicaudus Fernandez-Silva & Randall is described from the Red Sea and Arabian Sea, including Socotra and Gulf of Oman. It is characterized by a yellow caudal fin, 25–28 gill rakers, and 37–38 lateral-line scales and it is differentiated from nominal subspecies M. flavolineatus flavolineatus by 1.7% sequence divergence at the mitochondrial cytochrome b gene. The morphometric examination of specimens of M. f. flavolineatus revealed variation in head length, eye diameter, and barbel length, in western direction from the Hawaiian Islands, South Pacific, Micronesia, and the East Indies to the Indian Ocean. The population of Mulloidichthys f. flavicaudus subsp. n. in the Gulf of Aqaba differs from that of the remaining Red Sea by shorter barbels, smaller eyes, shorter head, and shorter pelvic fins. We present a list of 26 endemic fishes from the Gulf of Aqaba and discuss the probable basis for the endemism in the light of the geological history of this region.
cytb, marine fish, glacial refugia, phylogeography, taxonomy, vicariance
The goatfish Mulloidichthys flavolineatus was described by
Mulloidichthys flavolineatus is presently regarded as the most wide-ranging species of the family Mullidae, from the northern Red Sea (
Like other goatfishes, this species uses the pair of sensory barbels on its chin to locate prey, mainly in sedimentary substrata, as seen in Fig.
We, and surely others, have noticed that the population of Mulloidichthys flavolineatus in the Red Sea has only yellow caudal fin (Fig.
The caudal fin continues to be yellow from the Red Sea into the Gulf of Aden and Socotra, as shown by Fig.
Across the Arabian Sea to the south coast of Oman aggregations of Mulloidichthys flavolineatus in Oman and Maldives include many individuals with yellowish caudal fin mixed with a few gray-tailed and yellow-tailed fish (Figs
Type specimens were deposited at the Bernice P.
Lateral-line counts begin with the first pored scale completely posterior to the upper end of the gill opening and end at the base of the caudal fin (three pored scales continue onto the caudal fin). Counts of gill rakers were made on the first gill arch; they include all rudiments.
Lengths of specimens are given as standard length (SL), measured from the median anterior point of the upper lip to the base of the caudal fin (posterior end of the hypural plate); body depth is taken vertically from the base of the first dorsal-fin spine where it emerges from the body (not the internal base); body width is the maximum width measured just posterior to the gill openings; head length (HL) from the front of the upper lip to the posterior end of the opercular membrane, and snout length from the same anterior point to the nearest fleshy edge of the orbit; orbit diameter is the greatest fleshy diameter, and interorbital width the least fleshy width; upper-jaw length is taken from the front of the upper lip to the end of the maxilla; barbel length is the maximum straight length; caudal-peduncle depth is the least depth, and caudal-peduncle length the horizontal distance between verticals at the rear base of the anal fin and the caudal-fin base; length of fin spines and rays of the dorsal and anal fins are measured from where they emerge from the body to their tip; caudal-fin length is the horizontal length from the posterior end of the hypural plate to a vertical at the tip of the longest ray; caudal concavity is the horizontal distance between verticals at the tips of the shortest and longest rays; pectoral-fin length is measured from the base of the uppermost ray; pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest soft ray. Proportional measurements in the text are rounded to the nearest 0.05.
Only meristic characters and measurements that vary between M. f. flavolineatus and M. f. flavicaudus subsp. n. were applied in the diagnoses and comparisons: the number of gill rakers, lateral-line scale counts, barbel length, eye diameter and head length. We also compared the length of the pectoral and pelvic fins, but these did not show differences between M. f. flavolineatus and M. f. flavicaudus subsp. n.
Because goatfishes present allometric changes in body form (
During a previous phylogeographic survey of M. flavolineatus we obtained cytb sequences from 217 specimens sampled at nineteen sites throughout the Red Sea, the Arabian Sea, the Indian Ocean and the Pacific Ocean. To elucidate phylogenetic relationships we sequenced an additional fragment of the mitochondrial genome, the ATP synthetase 8 and ATP synthetase 6 (ATPase-8 and ATPase-6) regions, from individuals representative of the cytb diversity. We also sequenced an individual of M. vanicolensis and one of M. pfluegeri to use as outgroups. Briefly, DNA was extracted from fin clips and Polymerase Chain Reactions (PCR) were carried out using the primers L8331 (5'-AAA GCR TYR GCC TTT TAA GC-3') and H9236 (5'-GTT AGT GGT CAK GGG CTT GGR TC-3') (
The data underpinning the analysis reported in this paper are deposited in the Dryad Data Repository at https://doi.org/10.5061/dryad.f54m5
Mulloides
flavolineatus
(non Lacepède, 1801):
Mulloidichthys
flavolineatus
(non Lacepède, 1801):
Gulf of Suez:
Body elongate, the depth at first dorsal-fin origin 4.1–4.5 in SL; head moderately compressed, the length 3.0–3.3 in SL; snout long, slightly blunt. Barbels usually not reaching a vertical at posterior margin of preopercle, their length 4.1-5.0 in SL. Eye diameter 10.3–13.5 in SL. Pectoral-fin rays 16–18. Gill-raker counts 25–28 (usually 26 or 27); lateral-line scales 37–38. Caudal fin yellowish to yellow. [Diagnosis based on the Red Sea proper population, i.e. excluding the Gulf of Aqaba, see remarks].
Meristics are provided in Tables
Comparison of head length, eye diameter, and barbel length in SL among regions and against SL in Mulloidichthys flavolineatus flavolineatus and M. f. flavicaudus subsp. n. Below, measurements against SL. These colors identify the region of origin of each individual following the scheme in the upper panel. These are the same data as in Table
Lateral-line scale counts of M. flavolineatus subspp. In bold, counts for the holotype of M. f. flavicaudus subsp. n.
37 | 38 | 39 | 40 | mean | ||
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M. f. flavicaudus subsp. n. | Gulf of Aqaba | 12 | 10 | 1 | 37.5 | |
Red Sea off Sudan & Saudi Arabia | 18 | 5 | 37.2 | |||
† | Maldives | 2 | 4 | 37.7 | ||
M. f. flavolineatus | Islands of Western Indian Ocean ‡ | 5 | 5 | 4 | 37.9 | |
East Indies § | 4 | 24 | 5 | 38.0 | ||
Islands of Oceania (except Hawai‘i) | | 3 | 26 | 7 | 38.1 | ||
Hawaiian Islands & Johnston Atoll | 2 | 16 | 6 | 1 | 38.2 |
Total gill-raker counts of M. flavolineatus. In bold, counts for the holotype of M. f. flavicaudus subsp. n.
25 | 26 | 27 | 28 | 29 | 30 | mean | ||
---|---|---|---|---|---|---|---|---|
M. f. flavicaudus subsp. n. | Gulf of Aqaba | 3 | 6 | 8 | 2 | 4 | 26.9 | |
Red Sea off Sudan & Saudi Arabia | 4 | 5 | 11 | 2 | 26.5 | |||
† | Maldives | 1 | 3 | 2 | 0 | 27.2 | ||
M. f. flavolineatus | Islands of Western Indian Ocean ‡ | 4 | 2 | 2 | 27.8 | |||
East Indies § | 3 | 11 | 12 | 7 | 27.7 | |||
Islands of Oceania (except Hawai‘i) | | 12 | 18 | 11 | 3 | 28.4 | |||
Hawaiian Islands & Johnston Atoll | 2 | 3 | 10 | 3 | 28.8 |
Body elongate, its depth at first dorsal-fin origin 4.1 (4.2–4.5), and maximum width 6.7 (6.5–7.3), head length 3.2 (3.0–3.3, n=27), snout length 6.9 (6.8–7.7), orbit diameter 13.0 (10.3–13.5, n=27), barbel length 4.5 (4.2–5.1, n=27), caudal-peduncle length 4.8 (4.6–5.1), caudal-peduncle depth 11.6 (11.4–12.0), pelvic-fin length 4.9 (4.7–5.3), pectoral-fin length 4.8 (4.7–5.1), longest anal ray 7.5 (7.2–7.7), longest dorsal spine 4.8 (4.6–5.1), longest dorsal ray 7.2 (7.2–7.8).
Mouth small, maxilla not reaching a vertical at front of orbit, upper-jaw length 12.3 (12.2–13.9) in SL; jaws with small conical teeth, in two rows with teeth more irregularly placed between both rows; no teeth on the vomer and palatines; anterior nostril small, elliptical, two-thirds eye diameter in front of eye; posterior nostril small, elliptical, at dorsoanterior corner of orbit; opercular spine flat, at mid-eye height.
Scales very finely ctenoid; head fully scaled; scales on the base of caudal fin, other fins without scales; dorsal fin behind the vertical at fourth lateral line scale, origin of second dorsal above 18th (17th in some paratypes) scale. Pored scales on lateral line with many branching tubules.
Color in life silvery white to yellowish, slightly darker over lateral line; margin of each scale on upper half of body darker than scale. Yellow stripe on side of body at level of eye, from posterior margin of orbit to caudal-fin base, bordered by a narrow whitish stripe (stripe sometimes slightly blue); the stripe usually containing a black spot above posterior part of pectoral fins (under the first dorsal fin), sometimes faint due to fading, stripe anterior to spot occasionally indistinct; barbels white; dorsal fins usually transparent, sometimes first dorsal fin with yellowish tinge; pectoral, anal, and pelvic fins whitish, translucent; caudal fin yellowish or yellow. Color when fresh often pink and all fins yellow. Uniformly creamy white in preservative.
Mulloidichthys f. flavicaudus subsp. n. is named in reference to the yellow color of the caudal fin, in contrast to the whitish gray color of the caudal fin of M. f. flavolineatus.
Mulloidichthys f. flavicaudus subsp. n. is restricted to the NW Indian Ocean biogeographic province, where it ranges from various locations in the Red Sea (including the Gulf of Aqaba), the Gulf of Tadjoura, the Gulf of Aden, and Socotra (Fig.
The population of M. f. flavicaudus subsp. n. in the Gulf of Aqaba differs from that in the Red Sea proper by having smaller eyes (11.0–15.8 in SL) and shorter head (3.0–3.6) (Tables
Comparison of Head Length, Eye Diameter, and Barbel Length in subspecies of M. flavolineatus. Ranges and mean values (in brackets) are given for each ratio.
Locality | Standard length (mm) and number of specimens | Head length in standard length | Eye diameter in standard length | Barbel length in standard length | |
---|---|---|---|---|---|
M. f. flavicaudus subsp. n. | Gulf of Aqaba | 107–252 (n=23) | 3.0–3.6 (3.3) | 11.0–15.8 (13.4) | 4.1–5.2 (4.7) |
Red Sea ‡ | 97.5–203 (n=28) | 3.0–3.2 (3.1) | 10.2–13.5 (11.3) | 4.2–4.8 (4.5) | |
† | Maldives | 85.5–144 (n=6) | 2.8–3.3 (3.2) | 10.1–11.8 (10.7) | 3.7–4.8 (4.4) |
M. f. flavolineatus | Indian Ocean § | 120–192 (n=12) | 3.0–3.3 (3.1) | 10.5–11.7 (11.0) | 4.2–5.1 (4.5) |
East Indies | | 98–255 (n=34) | 3.1–3.5 (3.3) | 10.3–14.0 (12.5) | 4.3–5.6 (4.9) | |
Micronesia ¶ | 75–230 (n=26) | 3.1–3.8 (3.3) | 10.4–14.4 (11.9) | 4.2–5.9 (5.0) | |
South Pacific # | 81–198 (n=26) | 2.9–3.4 (3.2) | 10.2–12.8 (11.9) | 4.3–5.2 (4.7) | |
Hawaiian Is. †† | 83–288 (n=16) | 3.1–3.7 (3.3) | 10.4–15.6 (12.9) | 4.0–6.0 (5.1) |
Proportional measurements of type specimens of Mulloidichthys flavolineatus flavicaudus subsp. n. and of comparative material of M. f. flavolineatus as percentages of the standard length.
M. f. flavicaudus subsp. n. | M. f. flavolineatus | |||||||||||
Sudan | Mauritius | Johnston Atoll | ||||||||||
Holotype | Neotype | |||||||||||
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Standard length (mm) | 142 | 125 | 120 | 118 | 111 | 108 | 106 | 181 | 162 | 120 | 135 | 166 |
Body depth | 24.3 | 23.1 | 23.2 | 24.1 | 22.8 | 22.1 | 22.2 | 22.1 | 22.2 | 22.1 | 22.0 | 25.1 |
Body width | 14.9 | 14.9 | 15.3 | 14.4 | 14.5 | 13.6 | 13.9 | 15.1 | 13.1 | 12.3 | 9.6 | 14.2 |
Head length | 31.3 | 33.2 | 32.6 | 32.3 | 31.9 | 32.1 | 30.9 | 32.1 | 30.6 | 31.5 | 32.2 | 33.2 |
Snout length | 14.4 | 14.4 | 13.9 | 13.5 | 14.7 | 13.2 | 13 | 14.4 | 14.5 | 13.1 | 13.5 | 15.2 |
Orbit diameter | 7.7 | 9.3 | 9.5 | 9.0 | 8.8 | 9.2 | 7.7 | 8.6 | 9.2 | 8.5 | 8.6 | 9.3 |
Interorbital width | 7.8 | 7.5 | 7.2 | 7.4 | 7.8 | 7.0 | 7.0 | 7.5 | 9.0 | 7.6 | 6.9 | 8.0 |
Upper-jaw length | 8.1 | 7.0 | 8.0 | 8.0 | 7.8 | 7.2 | 7.6 | 8.2 | 8.3 | 8.4 | 8.1 | 7.8 |
Barbel length | 22.3 | 23.4 | 24.0 | 22.5 | 21.8 | 22.5 | – | 21.0 | 19.8 | 20.8 | 22.2 | 20.0 |
Caudal-peduncle least depth | 8.6 | 8.6 | 8.3 | 8.6 | 8.5 | 8.4 | 8.7 | 8.8 | 9.2 | 8.8 | 9.2 | 9.2 |
Caudal-peduncle length | 24.3 | 22.4 | 20.1 | 23.0 | 23.1 | 24.9 | 21.8 | 25.2 | 23.3 | 30.5 | 20.3 | 23.4 |
Snout to origin of first dorsal fin | 41.1 | 39.2 | 39.6 | 39.3 | 40.4 | 37.3 | 27.3 | 38.6 | 40.6 | 38.9 | 40.1 | 42.0 |
Snout to origin of second dorsal fin | 68.1 | 67.0 | 66.6 | 67.3 | 66.5 | 63.8 | 65.1 | 65.5 | 65.9 | 62.8 | 63.2 | 68.8 |
Preanal-fin length | 69.4 | 65.6 | 67.5 | 68.9 | 66.5 | 66.4 | 65.7 | 67.7 | 67.0 | 64.8 | 6.7 | 67.4 |
Prepelvic-fin length | 33.2 | 33.2 | 35.2 | 34.9 | 32.8 | 34.3 | 33.0 | 33.5 | 32.9 | 31.3 | 31.8 | 32.7 |
Second dorsal-fin base | 12.8 | 12.5 | 11.7 | 12.3 | 12.2 | 11.5 | 12.1 | 13.1 | 11.8 | 10.8 | 11.1 | 12.4 |
Anal-fin base | 9.9 | 10.5 | 9.1 | 10.6 | 9.6 | 9.9 | 10.7 | 9.7 | 9.0 | 10.4 | 9.4 | 10.4 |
First dorsal-fin base | 16.7 | 18.8 | 19.9 | 21.8 | 17.6 | 16.6 | 17.4 | 17.2 | 15.3 | 17.3 | 16.0 | 19.0 |
Pectoral-fin base | 4.9 | 5.1 | 5.2 | 5.2 | 5.7 | 4.7 | 4.9 | 5.6 | 5.0 | 5.2 | 4.8 | 5.4 |
Longest dorsal spine | 20.8 | 19.9 | 21.3 | 21.8 | 19.7 | 18.7 | 19.0 | 21.3 | 20.6 | 20.2 | 21.2 | 20.1 |
Longest dorsal ray | 14.6 | 14.1 | 14.1 | 13.4 | 14.3 | 14.5 | 13.2 | 13.4 | 14.3 | 15.4 | 15.2 | 14.3 |
Longest anal ray | 13.9 | 13.4 | 13.5 | 13.7 | 13.2 | 12.9 | 13.6 | 13.8 | 13.3 | 14.7 | 14.4 | 13.7 |
Caudal-fin length | 28.1 | 27.1 | 26.8 | – | 26.6 | 24.9 | 25.0 | – | 26.3 | 27.5 | 28.6 | 25.9 |
Caudal concavity | 19.1 | 18.1 | 19.1 | – | 16.4 | 16.9 | 16.5 | – | 16.7 | 19.7 | 20.6 | 16.8 |
Pectoral-fin length | 20.8 | 20.3 | 20.6 | 21.5 | 21.2 | 19.9 | 20.7 | 19.8 | 20.0 | 21.3 | – | 21.7 |
Pelvic-fin length | 20.6 | 20.3 | 20.4 | 21.0 | 20.1 | 19.0 | 21.1 | 19.8 | 20.3 | 22.1 | 21.1 | 21.0 |
Mulloidichthys f. flavicaudus subsp. n. differs from its nominal subspecies M. f. flavolineatus in having 25–28 (usually 26 or 27) gill-raker counts (26–30, usually 27–29, in M. f. flavolineatus), usually 37–38 lateral-line scales (37–40 in M. f. flavolineatus) and a yellow caudal fin (white to light gray in M. f. flavolineatus). Also, the eyes are smaller in M. f. flavicaudus subsp. n. (10.3–13.5 in SL) than in M. f. flavolineatus (9.8–15.6 in SL).
Mullus flavolineatus Lacepède, 1801: 384, 406 (locality unknown, no types known).
Mulloidichthys
flavolineatus
(
Hawaiian Islands:
Body elongate, the depth at first dorsal-fin origin 4.0–4.6 in SL; head moderately compressed, the length 2.9–3.8 in SL; snout long, slightly blunt anteriorly. Barbels usually not reaching a vertical at posterior margin of preopercle, their length 3.7–6.0 in SL. Eye diameter 10.1–15.6 in SL. Pectoral-fin rays 16–18. Gill-raker counts 27–29 (rarely 26 or 30); lateral-line scales 37–40 (usually 38). Caudal fin varying from usually white or light gray to occasionally yellowish or yellow.
Silvery white to yellowish, slightly darker over lateral line, margins of each scale on upper half of body darker than scale. Yellow stripe on side of body at level of eye, beginning from posterior margin of orbit and ending at caudal-fin base, bordered by two whitish narrow stripes (sometimes slightly blue); the stripe usually containing a black spot above posterior part of pectoral fins (under the first dorsal fin), sometimes faint due to fading, stripe anterior to spot occasionally indistinct; barbels white; dorsal fins usually transparent, sometimes first dorsal fin with yellowish tinge; pectoral, anal, and pelvic fins whitish, translucent; caudal fin varying from usually white or light gray to occasionally yellowish or yellow. Sometimes body color pattern of broad irregular red-brown bars, especially at night. When fresh, body color can turn pink and all fins yellow. Uniformly creamy white in preservative.
Mulloidichthys f. flavolineatus is wide-ranging from East Africa north to the Maldives and Chagos Archipelago and east to the Hawaiian, Marquesas and Pitcairn Islands, north to the Ryukyu and Bonin Islands and south to Lord Howe Island, New Caledonia and Rapa Island (
Distribution map of Mulloidichthys flavolineatus surveyed in this study. Red symbols denote locations of specimens of M. f. flavicaudus subsp. n. and blue symbols denote locations of specimens of M. f. flavolineatus. Squares indicate locations included in the genetic surveys. Circles indicate locations of specimens for which only morphological analyses were carried out.
The parsimony-based haplotype networks constructed with mtDNA cytb sequences from 217 M. flavolineatus specimens revealed a separation between individuals from the NW Indian Ocean (including the Red Sea, the Gulf of Aden and Oman) and individuals in the rest of the Indian Ocean and the Pacific Ocean (Fig.
Median-joining haplotype network based on mitochondrial cytochrome b sequence data (715 bp) from 217 Mulloidichthys flavolineatus individuals sampled across the Red Sea, Arabian Sea, Indian Ocean and Pacific Ocean. Each circle represents a haplotype, with size proportional to its total frequency. Branches separated by black crossbars represent a single nucleotide change, whereas open circles indicate unsampled haplotypes; colors indicate collection location as in the embedded key. The network depicts two distinct clades separated by seven mutational steps (corrected sequence divergence, d = 1.7%;
We obtained a concatenated alignment of a 715-bp segment of the cytb gene and a 731-bp segment of the ATPase-8 and ATPase-6 genes of the mitochondrial genome from seven individuals from the Red Sea (Jeddah) and five from the Pacific (Hawai‘i and Okinawa). Phylogenetic reconstructions based on Bayesian inference (Fig.
Bayesian-inference based phylogenetic tree showing relationships among mtDNA concatenated haplotypes of segments of the cytb and ATPase-8 & ATPase-6 genes from seven individuals of Mulloidichthys flavolineatus flavicaudus subsp. n. from the Red Sea (Jeddah), five individuals of M. f. flavolineatus from the Pacific (O‘ahu in Hawai‘i and Okinawa) and two Mulloidichthys spp. as outgroups. The nodes show posterior probabilities. Branch lengths are according to estimated divergence time (note that the branch leading to M. pfluegeri was reduced by 50%).
Higher gill-raker and lateral-line counts, smaller eyes and stable yellow coloration of the caudal fin in M. flavolineatus from the Red Sea are characters in alignment with the genetic isolation of a mitochondrial lineage in the NW Indian Ocean biogeographic province (as per
Some ichthyologists, notably
Notably, the age of split of the Mulloidichthys flavolineatus subspecies is older than the radiation that gave rise to M. vanicolensis, M. mimicus, M. dentatus (Gill, 1862) and M. martinicus (Cuvier, 1829) less than 350,000 years ago (unpublished results).
It is remarkable that individuals of Mulloidichthys f. flavicaudus subsp. n. from the Gulf of Aqaba have consistently smaller eyes, longer head, and longer barbels than fish from the Red Sea proper (Fig.
Endemic fishes of the Gulf of Aqaba | Remarks |
---|---|
Amblyeleotris neglecta Jaafar & Randall, 2009 | |
Cabillus nigrostigmus Kovačić & Bogorodsky, 2013 | Known from Sharm el Moya, close to the entrance of the Gulf of Aqaba |
Callionymus profundus Fricke & Golani, 2013 | Deep-water species |
Chromis pelloura Randall & Allen, 1982 | |
Cirrhilabrus blatteus Springer & Randall, 1974 | |
Evoxymetapon moricheni Fricke, Golani & Appelbaum–Golani, 2014 | |
Gymnapogon melanogaster Gon & Golani, 2002 | |
Gymnothorax baranesi Smith, Brokovich & Einbinder, 2008 | |
Hetereleotris psammophila Kovačić & Bogorodsky, 2014 | Recently photographed at Safaga |
Heteronarce bentuviai (Baranes & Randall, 1989) | |
Limnichthys marisrubri Fricke & Golani, 2012 | |
Myxomyrophis longirostris Hibino, Kimura & Golani, 2014 | |
Paragunnellichthys springeri Dawson, 1970 | Formally endemic to Gulf of Aqaba, known from Sharm el Moya, close to the entrance |
Parascolopsis baranesi Russell & |
|
Pseudogramma megamyctera Randall & Baldwin, 1997 | Reported from West Papua ( |
Scorpaenodes steinitzi Klausewitz & Fröiland, 1970 | A specimen identified as S. steinitzi collected from Djibouti, but no voucher available for confirmation |
Stalix davidsheni Klausewitz, 1985 | |
Suculentophichthus nasus Fricke, Golani & Appelbaum–Golani, 2015 | |
Symphysanodon disii Khalaf & Krupp, 2008 | |
Syngnathus safina Paulus, 1992 | |
Thamnaconus erythraeensis Bauchot & Maugé, 1978 | |
Tomiyamichthys dorsostigma Bogorodsky, Kovačić & Randall, 2011 | |
Upeneus davidaromi Golani, 2001 | |
Uropterygius genie Randall & Golani, 1995 | Known at Ras Mohammed, close to the entrance of the Gulf of Aqaba |
Uropterygius golanii McCosker & Smith, 1997 | Known at Ras Mohammed, close to the entrance of the Gulf of Aqaba |
Vanderhorstia opercularis Randall, 2007 |
Our range-wide phylogeographic survey of Mulloidichthys flavolineatus (
We are grateful to Loreen O’Hara and Arnold Suzumoto from the Bernice P. Bishop Museum, Honolulu, HI, U.S.A. (