Bodianus atrolumbus (Valenciennes 1839), a valid species
of labrid ish from the southwest Indian Ocean
JOHN E. RANDALL
Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA
E-mail: jackr@hawaii.rr.com
BENJAMIN C. VICTOR
Ocean Science Foundation, 4051 Glenwood, Irvine, CA 92604 and Guy Harvey Research Institute,
Nova Southeastern University, 8000 North Ocean Drive, Dania Beach, FL 33004, USA
E-mail: ben@coralreefish.com
Abstract
Bodianus atrolumbus (Valenciennes 1839), a labrid ish of the southwest Indian Ocean (type locality Mauritius),
was placed in the synonymy of B. perditio (Quoy & Gaimard), antitropical in the Paciic Ocean (type locality
Tonga), by Smith (1949) who reported the irst record for southern Africa. Juveniles of both species are mostly
the same in color, featuring a white bar in the middle of the body, followed dorsally by a large black area. The
white bar develops into an oval yellow area dorsally on the body in B. perditio, whereas it narrows to a spindleshaped whitish to pink mark on the upper body that extends below the lateral line in B. atrolumbus. An analysis of
the mitochondrial DNA barcode sequence (COI) from specimens collected from all quadrants of the the species’
range reveals that the two species are 3.91% different (K2P minimum interspeciic distance), while intraspeciic
variation is no more than 0.34%. A phenetic tree of barcode sequences for twenty Bodianus species is presented,
showing that pairwise species differences range from 1.97% to 21.74%, with Indian/Paciic sibling-species pairs
accounting for the lower range of divergences (1.97% to 4.64%). A modal difference in the count of gill rakers and
the distinctly shorter pectoral ins of B. atrolumbus additionally differentiate it from B. perditio.
Key words: Labridae, coral reef ishes, phylogenetics, DNA barcoding, biogeography, taxonomy, sibling species
Introduction
The labrid ish genus Bodianus is represented by at least 44 species in the tropical to warm temperate seas of
the world, making it the third largest genus of the family Labridae, following Halichoeres and Cirrhilabrus. Only
ive species of Bodianus are reported from the Atlantic Ocean and two for the eastern Paciic, while at least 37
44
occur in the rest of the Indo-Paciic. The species range in size from the diminutive Bodianus bimaculatus Allen,
of which the largest specimen measures 59 mm standard length, to B. macrognathos (Morris), reported to 595
mm SL.
The genus was ably revised and beautifully illustrated by Gomon (2006), whose experience with the genus
dates to 1979 when it was the subject of his PhD thesis at the University of Miami in Florida (the irst author was
a reviewer of the thesis). Gomon (2006) has now divided Bodianus into 10 subgenera, seven of them taking names
of generic synonyms. Eight species were described as new. Distribution maps are provided for all species.
Gomon’s Figure 54 is a distribution map that includes Bodianus perditio (Quoy & Gaimard), with a disjunct
population: KwaZulu-Natal, Mauritius, Réunion, and St. Brandon’s Shoals in southwest Indian Ocean, and antitropical in the Paciic from southern Japan in the north to southern Queensland, New South Wales and Lord
Howe Island in the south. It ranges east in the South Paciic to New Caledonia, Tonga (type locality), Rapa, and
Mangareva in the Tuamotu Archipelago. The very similar Bodianus solatus, described as new by Gomon from
Western Australia, is reported from “between the Monte Bello Islands and the Houtman Abrolhos”, hence within
the latitudes of 20° and 30°S. However, his distribution map includes one symbol for B. solatus at the northernmost point of Western Australia. This is evidently intended for the record of B. solatus from the Northwest Shelf
of Australia, misidentiied as Bodianus perditio by Gloerfelt-Tarp and Kailola (1984: 233, ig. on adjacent plate)
and by Sainsbury et al. (1984: 256, ig. on adjacent plate). An additional specimen of B. solatus was collected in
1995 in the nearby Arafura Sea at about 10 °S (AMS I.37182).
The distributions of both Bodianus perditio and its W. Indian Ocean sibling species are noteworthy in lacking
any low-latitude records. Smith (1949: 287, pl. 52, ig. 776) reported what he believed to be the irst record of B.
perditio for southern Africa at Delagoa Bay (now Maputo Bay), Mozambique (26°S). Randall (1986: 688, ig.
221.10, pl. 90) added records from Natal (KwaZulu-Natal) and St. Brandon’s Shoals in the Indian Ocean at 16–
17°S. In the Paciic, B. perditio is almost antitropical, with no record south of Taiwan at 22°N (Shao 2013), and
none north of Efate, Vanuatu at 17.8°S (collected by Grant Norton). The present distribution and DNA sequence
data indicate that B. perditio of the Paciic has been isolated from the southwestern Indian Ocean population for
a very long time.
Juveniles in both the SW Indian Ocean and Paciic populations are similar in color, with a narrow white bar
from the base of the ninth dorsal spine extending down to the front of the anal in, followed by a large black
area dorsally on the body and extending onto most of the scaly basal sheath posteriorly on the dorsal in. Adults
in the Indian Ocean replace the white bar of the juvenile on midside of body with a lanceolate, pale pink streak
extending well below the lateral line. Adults in the Paciic lose the ventral part of the white bar of the juvenile, the
upper part enlarging to an oval, pale yellow spot, mostly above the lateral line.
Only Kuiter (2010: 36) has listed Bodianus atrolumbus as a valid species in his pictorial review of labrid ishes.
We provide here the documentation to fully establish its validity based on color pattern, morphology, and mtDNA
sequence differences. The recent development of large-scale DNA sequencing programs, such as the Barcode
of Life project (www.boldsystems.org), permits comparisons between and among populations and species and
can assist in establishing the validity of species by assessing the degree to which populations are reproductively
isolated and, along with additional genetic analyses, an estimate of the time since the populations split.
Materials and Methods
Specimens have been examined at, or obtained on loan from, the Bernice P. Bishop Museum, Honolulu
(BPBM); South African Institute for Aquatic Biodiversity, Grahamstown, South Africa (SAIAB), and the United
States National Museum of Natural History (USNM).
Lengths of specimens are given as standard length (SL), measured from the median end of the upper lip or
upper canine teeth, whichever is more anterior, to the base of the caudal in (posterior end of hypural plate); head
length is measured from the same anterior point to the posterior end of the opercular lap, and upper-jaw length
from the same anterior point to the posterior end of the maxilla; body depth is the greatest depth from the base of
the dorsal spines to the ventral margin of the abdomen (correcting for any obvious malformation of preservation);
45
body width is taken just posterior to the gill opening; orbit diameter is the greatest leshy diameter; interorbital
width the least bony width; caudal-peduncle depth is the least depth, and caudal-peduncle length the horizontal
distance between verticals at the rear base of the anal in and base of the caudal in; lengths of spines and rays of
median ins are measured to their extreme bases; caudal concavity is the horizontal distance between tips of the
longest and shortest rays; pectoral-in length is the length of the longest ray to the extreme base (average taken
of the two ins when different); pelvic-in length is measured from the base of the pelvic spine to the tip of the
longest ray.
Counts of pectoral-in rays include the rudimentary upper ray. Lateral-line scale counts do not include the one
or two pored scales on the base of the caudal in. Gill-raker counts include rudiments.
DNA extractions and sequencing were performed at the Canadian Centre for DNA Barcoding (CCDB) in
Guelph, Ontario, Canada, using the NucleoSpin96 (Machery-Nagel) kit according to manufacturer speciications
under automation with a Biomek NX liquid-handling station (Beckman-Coulter) equipped with a iltration
manifold. A 652-bp segment was ampliied from the 5′ region of the mitochondrial COI gene using a variety of
primers (Ivanova et al. 2007). PCR ampliications were performed in 12.5 µl volume including 6.25 µl of 10%
trehalose, 2 µl of ultra pure water, 1.25 µl of 10× PCR buffer (10mM KCl, 10mM (NH4)2SO4, 20mM Tris-HCl
(pH8.8), 2mM MgSO4, 0.1% Triton X-100), 0.625 µl of MgCl2 (50mM), 0.125 µl of each primer (0.01mM),
0.0625 µl of each dNTP (10mM), 0.0625 µl of Taq DNA polymerase (New England Biolabs), and 2 µl of template
DNA. The PCR conditions consisted of 94°C for 2 min., 35 cycles of 94°C for 30 sec., 52°C for 40 sec., and 72°C
for 1 min., with a inal extension at 72°C for 10 min.
Specimen information, the barcode sequence data, and the phenetic tree analysis for this study were compiled
using the Barcode of Life Data Systems (BOLD, www.barcodinglife.org; Ratnasingham & Hebert 2007). The
sequence data used for statistics and the tree are publicly accessible on BOLD and GenBank. Sequence divergence
was calculated using BOLD with the Kimura 2-parameter (K2P) model and a pairwise model. The phenetic tree
was calculated from a Kimura 2-parameter (K2P) model which generated a mid-point rooted neighbor-joining
(NJ) phenogram to provide a graphic representation of the species divergence.
Bodianus atrolumbus (Valenciennes in Cuvier & Valenciennes 1839)
Figures 1–3; Table 1
Cossyphus atrolumbus Valenciennes in Cuvier & Valenciennes 1839: 123 (Mauritius); Günther 1862: 105 (Mauritius); Bauchot 1963: 27 (holotype, MNHN A.8262, 183 mm SL, dried skin).
Cossyphus nigromaculatus Gilchrist & Thompson 1908: 197 (Durban).
Chaeropsodes pictus Gilchrist & Thompson 1909: 260 (Durban).
Lepidaplois perditio Smith 1949: 287, pl. 52, ig. 776 (Delagoa Bay); Baissac 1953: 224 (Mauritius); Fourmanoir
& Guézé 1961: 7 (Mauritius, Réunion).
Bodianus perditio Baissac 1976: 213 (Mauritius); van der Elst 1981: 189, ig. (Mozambique to Durban); Randall
1982: 203 (antitropical); Gomon in Fischer & Bianchi 1984: unnumbered pages (Mauritius, Réunion, St.
Brandon’s Shoals, and Natal Coast, KwaZulu-Natal); Randall in Smith & Heemstra 1986: 688, pl. 90, ig.
220.10 (Mauritius, Réunion, St. Brandon’s Shoals, and northern Mozambique to KwaZulu-Natal); Debelius
1993: 219, lower ig. (his illustration is Bodianus perditio from the Paciic); Lieske & Myers 1994: 90, igs.
(juvenile and adult; Mozambique to South Africa and Mauritius); Fricke 1999: 400 (annotated checklist);
Heemstra & Heemstra 2004: 339 (Mozambique to Aliwal Shoal, South Africa); E. Heemstra et al. 2004:
3325 (Rodrigues, Mascarene Islands); Gomon 2006: 87, igs 7b, 53-54, pls 8J, 9A-B (taxonomy); Taquet &
Diringer 2012: 460, middle ig. (Indian Ocean).
Bodianus atrolumbus Kuiter 2010: 36, 4 igs. (Indian Ocean, Mozambique to Aliwal Shoal, Mauritius, and
Réunion).
46
Diagnosis. Dorsal rays XII,10; anal rays III,11; pectoral rays 17; lateral line smoothly curved, following
dorsal contour of body, the pored scales 30 or 31; scales above origin of lateral line to origin of dorsal in 8;
predorsal scales small, becoming embedded anteriorly, ending in posterior interorbital; a patch of very small
scales on cheek posterior to center of eye, except for the broad naked lange of the preopercle; no scales on lower
jaw; a broad sheath of scales basally on median ins; total gill rakers 18–21; dorsal proile of snout to above eye
straight, becoming smoothly convex on nape; snout length 2.7–3.0 in head length; orbit diameter varying from
4.7 in head length in an 88-mm specimen to 9.3 in a 430-mm specimen; mouth slightly oblique, forming an angle
of about 20° to horizontal axis of body, the maxilla reaching to below anterior half of eye; front of jaws with two
pairs of canine teeth that interdigitate when mouth closed, the middle pair of upper jaw and lateral pair of lower
jaw largest and recurved, the middle pair of lower jaw about half length of lateral pair; side of jaws with a dental
ridge of coalesced teeth bearing a row of small, stout, close-set, conical teeth, of which a series of ive or six in
middle of jaws are largest; no teeth on palate; labial laps well-developed; dorsal spines progressively longer, the
last spine 2.0–2.2 in head length, the membranes deeply incised; seventh or eighth dorsal soft ray longest, 2.0–2.1
in head length; third anal spine longest, about 2.0–2.4 in head length; caudal in of juveniles truncate, of adults
double emarginate with long pointed lobes; pectoral ins relatively short, 3.9–4.25 in SL; pelvic ins just reaching
anus in 88-mm juvenile, reaching beyond third anal spine in a 360-mm adult. Diagnostic color differences from
B. perditio include the central white bar of the juvenile narrowing to a spindle-shaped whitish to pink mark on the
upper body extending below the lateral line (vs. a dorsal oval yellow spot) and the caudal peduncle of the juvenile
with a white bar extending full width (vs. a central white patch surrounded by dark)(Figures 1–3). Reaches 57 cm.
Remarks. Bodianus atrolumbus is known from the three Mascarene Islands (type locality, Mauritius), St.
Brandon’s Shoals, KwaZulu-Natal, and southern Mozambique. It was placed in the synonymy of B. perditio
(Quoy & Gaimard) by Smith (1949), followed by Gomon in Fischer and Bianchi (1984) and by Gomon (2006)
in his revision of the genus.
Van der Elst (1981: 189) reported this species as a common ish of deeper rock and coral reefs of South Africa.
He gave the diet as mainly sea urchins, crabs, gastropod mollusks, and other hard-shelled invertebrates, which are
crushed in the powerful pharyngeal plates that are studded with large molariform teeth.
The usual common names for Bodianus perditio in the Paciic are Goldspot Wrasse, as indicated by Grant
(1982: pl. 288), Goldspot Hogish (Randall et al. 1990: 301), Golden-spot Pigish (Kuiter 1993: 269), and Golden
Spot Hogish, the FAO English name (Westneat in Carpenter & Niem 2001: 3412). Van der Elst (1981: 189) used
Goldsaddle Hogish as the common name in South Africa, followed by Randall in Smith and Heemstra (1986:
339) and Heemstra and Heemstra (2004: 339). However, this is not appropriate, because the equivalent marking
in adult B. atrolumbus is not yellow or golden but whitish to pale pink, nor does it reach the dorsal edge of the
body. We propose the common name Palebar Hogish for B. atrolumbus.
Gomon (2006: 89) wrote: “Three specimens at the MNHN are listed as syntypes of C. atrolumbus in the
collection catalogue. MNHN A.8262 is here designated lectotype to afix the name should the Indian Ocean and
Paciic Ocean populations prove to be taxonomically separate.” However, Bauchot (1963: 27) already listed
MNHN A.8262, a dried specimen 183 mm SL, as the holotype of Cossyphus atrolumbus.
Material of Bodianus atrolumbus. Mauritius, BPBM 20293, 195 mm. St. Brandon’s Shoals, USNM 217848,
87.5 mm. South Africa, KwaZulu-Natal, Sodwana Bay, SAIAB 7397, 238 mm; Durban, SAIAB 43687, 2: 204233 mm; Park Rynie, SAIAB 189242, 307 mm.
TABLE 1
Total gill-raker counts for specimens of Bodianus
B. atrolumbus
B. perditio
18
19
20
21
1
3
4
2
3
5
22
23
3
1
47
Figure 1. Bodianus atrolumbus, top: small juvenile, Sodwana Bay, KwaZulu-Natal, South Africa (D. King); middle: juvenile, Réunion (A. Diringer); bottom: 9 cm TL, KwaZulu-Natal, South Africa (D. King).
48
Figure 2. Bodianus atrolumbus, top: KwaZulu-Natal (D. Polack); middle: 15 cm TL, Rodrigues, Mascarenes, SAIAB
69529 (P. Heemstra); bottom: 28 cm TL, KwaZulu-Natal (D. King).
49
Figure 3. Bodianus atrolumbus, top: 30 cm TL, KwaZulu-Natal (D. King); middle: Réunion (A. Diringer); bottom: Réunion
(A. Diringer).
50
Bodianus perditio (Quoy & Gaimard 1834)
Figures 4–6; Table 1
Labrus perditio Quoy & Gaimard 1834: 702, pl. 20, ig. 4 (Tongatapu); Bauchot 1963: 93 [holotype, 7 pouces (=
189.5 mm) has been lost].
Cossyphus perditio Valenciennes in Cuvier & Valenciennes 1839: 125 (Tongatapu).
Cossyphus atrolumbus [non Valenciennes] Günther 1862: 105, in part (Tongatapu, Minerva Reef, Saumarez
Reefs, and Aneiteum); Günther 1881: 241 (same localities, except error for Zanzibar); Ogilby 1889: 66 (Lord
Howe Island).
Trochocopus sanguinolentus De Vis 1883: 287 (Hutchinson Shoal, Cape Moreton, Queensland).
Cossyphus aurifer De Vis 1884: 146 (Moreton Bay, Queensland).
Cossyphus latro De Vis 1885: 878 (Moreton Bay, Queensland).
Lepidaplois perditio Jordan & Snyder 1902: 618, ig. 2 (Saikasaki, Wakanoura, Honshu, Japan); Jordan & Seale
1906: 293 (Tonga, Saumarez Reefs, Aneiteum, and Japan); Ogilby 1916: 184 (Snapper Banks off Moreton
Bay, Queensland); Fowler & Bean 1928: 206 (Riu Kiu Islands); Chen, J.T.F. 1952: 104 (Taiwan).
Lepidaplois perdito Jordan, Tanaka & Snyder 1913: 198, ig. 144 (near Wakanoura, Japan); Fowler & Bean 1928:
328 (New Caledonia).
Bodianus perditio Matsubara 1964: 888, 892 (Japan); Yu 1968: 20, ig. 9 (Taichung, Pescadores, and Tainan,
Taiwan); Masuda, Araga & Yoshino 1975: 295, pl. 102 E, F (Sagami Bay southward, Japan); Allen et al.
1976: 418 (Lord Howe Island); Grant 1982: 561, pl. 288 (southern Queensland, Australia); Randall 1982:
203 (B. perditio antitropical in the western and south Paciic); Yamakawa in Masuda et al. 1984: 203, pl. 196
B, C (Sagami Bay southward, Japan); Randall in Smith & Heemstra 1986: 688, pl. 90, ig. 220-10; Allen &
Swainston 1988: 112, ig. 729 (Western Australia: Abrolhos northward); Randall, Allen & Steene 1990: 301,
upper ig; Francis 1991: 214, ig. 30 (Norfolk Island); Kuiter 1993: 269, upper two igs. (southeast Australia;
juveniles to rocky estuaries in New South Wales); Masuda & Kobayashi 1994: 249, igs. 6–8 (Ogasawara
Islands and Izu Peninsula, Honshu, Japan); Kulbicki & Williams 1997: 19 (Ouvéa Atoll, Loyalty Islands);
Okamura & Amaoka 1997: 470, middle column of igs. (Ogasawara Islands, Kochi Prefecture, and AmamiOshima, Japan); Laboute & Grandperrin 2000: 351, 3 lower igs. (New Caledonia); Parenti & Randall 2000:
6 (antiequatorial: islands of southern Oceania and Taiwan to southern Japan); Randall 2005: 394 (western and
southern Paciic); Bacchet, Zysman & Lefèvre 2006: 369, lower ig. (French Polynesia); Gomon 2006: 87,
igs. 7b, 53-54, plates 8J, 9A & 9B (western and southern Paciic); Chen et al. 2010: 177, igs. G, H, and I
(Kenting National Park, southern tip of Taiwan); Kuiter 2010: 37, 5 igs. (southern Japan to eastern Australia,
ranging to Central Paciic); Nishiyama & Motomura 2012: 26-27, 7 igs. (Japan).
Diagnosis. The morphological diagnosis for Bodianus perditio is essentially the same as B. atrolumbus, and
need not be repeated here. Diagnostic color differences from B. atrolumbus include the central white bar of the
juvenile diminishing to an oval yellow spot above the lateral line (vs. a spindled whitish to pink mark extending
below the lateral line) and the caudal peduncle of the juvenile with a central white patch surrounded by dark (vs.
a white bar extending full width)(Figures 4–6). We have found one meristic difference to partially separate the
two species: the number of gill rakers is 20–23 in B. perditio vs. 18–21 in B. atrolumbus (Table 1). Van der Elst
(1981: 189) gave the gill-raker counts of South African material of B. atrolumbus as 17–20. Evidently he did not
include the small rudiment at each end of the series of gill rakers. Furthermore, we have found a difference in the
length of the pectoral ins of the two species. Nine specimens of B. perditio from the South Paciic, 159–320 mm
SL, have a pectoral-in length 3.4–3.8 in the standard length, the in length proportionately longer with growth.
In comparison, four specimens of B. atrolumbus within the same size range from South Africa have a pectoralin length 3.9–4.25 in the standard length. The pectoral-in length of B. atrolumbus in Fig. 220.10 of Smith and
Heemstra (1986: 688) also falls within this range. Reaches 53 cm (Grant 1982: 561).
51
Figure 4. Bodianus perditio, top: 3 cm, New Caledonia (J.E. Randall); middle: 4.5 cm, New Caledonia (J.E. Randall); bottom: 7.5 cm, New Caledonia (J.E. Randall).
52
Figure 5. Bodianus perditio, top: 25 cm, Mangareva (J.E. Randall); middle: 30 cm, Mangareva (J.E. Randall); bottom: 36
cm, New Caledonia (J.E. Randall).
53
Figure 6. Bodianus perditio, The bright yellow juvenile sold in the aquarium trade, frequently shipped out of Vanuatu (photo
by Milan Kořínek, http://www.biolib.cz/en).
Genetics. DNA sequences were obtained from all quadrants of the range for the two sibling species: from
South Africa for Bodianus atrolumbus and, for B. perditio, from Taiwan in the northern-Paciic antitropical range
and in the southern Paciic from the western edge of the range in Queensland, Australia to the central population
in New Caledonia and the easternmost population in Mangareva, in the Gambier Islands of French Polynesia. The
barcode mitochondrial DNA sequences (COI) for the two species are 3.91% divergent (K2P minimum interspeciic
distance; 3.78% pairwise), compared to a maximum intraspeciic distance of 0.34%. B. solatus Gomon, the W.
Australian sibling species recently separated from B. perditio, is much more distant genetically, despite occupying
the intervening geographic range between the two species (11.96% divergent from B. perditio (10.94% pairwise)).
The interspeciic distance is well within the range of divergences found between other species of Bodianus.
The neighbor-joining phenetic tree based on 85 COI mtDNA sequences of twenty species in the genus (about half
of the known species), following the Kimura two-parameter model (K2P) generated by BOLD (Barcode of Life
Database), shows deep divergences between species and, in most cases, only minimal variation within species
(Fig. 7). The exception to the low variation within species is the 2.86% different sequence for Indian vs. Paciic
Ocean specimens of B. anthioides (Bennett) (2.74% pairwise); Gomon (2006) found a meristic difference but a
closer look at the phylogeography of the species is certainly warranted. The differences between pairs of Bodianus
species range from 1.87% to 21.74%, with a mean of 17.3% (minimum interspeciic distances by K2P; pairwise
values are 1.84% to 18.59%, mean of 15.19%). Interestingly, the lower range of interspeciic distances occur
between Indian/Paciic Ocean sibling species and Hawaiian endemic vs. widespread Indo-Paciic sibling species:
the former ranging from 1.87% between B. rubrisos Gomon and B. trilineatus (Fowler)(W. Indian/W. Paciic) to
4.64% between B. diana (Lacepède) and B. dictynna Gomon (Indian/W. Paciic) and 3.01% between the Hawaiian
endemic B. albotaeniatus (Valenciennes) and Indo-Paciic B. bilunulatus (Lacepède) (1.64%, 4.45%, and 2.92%
pairwise distances, respectively). These results conirm that the splitting of sibling species from the Indian and
Paciic Ocean basins is based on evidence of long-standing reproductive isolation and, for the genus Bodianus
at least, there is no evidence for pan-Indo-Paciic species with uninterrupted gene low. COI divergences of 3%
indicate more than a million years of separation by most estimates of the molecular clock (Bermingham et al.
1997).
Remarks. The color igures of Bodianus atrolumbus (Figs. 1–3) and B. perditio (Figs. 4 & 5) are arranged
in the order of the total length (measured for specimens, but only estimated for ish photographed underwater).
The relative size of the eye is helpful in arranging photographs of ishes according to size when the length is not
known, the smaller individuals having a relatively larger eye. However, a surprising exception was found in the
collection of specimens of B. perditio by the irst author at the island of Mangareva in the Tuamotu Archipelago.
Seven specimens, 217–320 mm SL, were caught by hook and line from 92 m, and one of 189 mm SL was
collected at the same island from less than one meter depth. The 189-mm ish (25 cm TL) would be expected to
54
French Polynesia
Taiwan
New Caledonia
Australia
Australia
New Caledonia
New Caledonia
French Polynesia
Australia
0.02
}
}
South Africa
B. perditio
B. atrolumbus
}
NW. Australia
B. solatus
Madagascar
South Africa
South Africa
Bali, Indonesia
B. bilunulatus
South Africa
Lombok, Indonesia
South Africa
South Africa
Hawai’i B. albotaeniatus
Galápagos Islands B. eclancheri
Cape Verde, Atlantic
B. speciosus
Philippines
Madagascar
Madagascar
B. anthioides
Mozambique
Réunion
}
}
}
Madagascar
}South Africa
Madagascar
}
}
B. diana
Bali, Indonesia
B. dictynna
Réunion
Mozambique
Madagascar
Madagascar
B. axillaris
Réunion
Réunion
Réunion
Madagascar
Bali, Indonesia B. mesothorax
Portugal B. scrofa
Lombok, Indonesia
B. rubrisos
South China Sea
}
}
South Africa
B. trilineatus
Costa Rica
Costa Rica
Galápagos Islands
Costa Rica
Panama
Galápagos Islands
Cuba
Yucatan, Mexico
Belize
}
}
B. diplotaenia
B. pulchellus
}
Yucatan, Mexico
}
Cuba
Bahamas
Hainan, China
}
}
South Africa
B. rufus
B. oxycephalus
B. bimaculatus
Prionotus carolinus
Figure 7. The neighbor-joining phenetic tree based on 85 COI mtDNA sequences of 20 species of Bodianus following the
Kimura two-parameter model (K2P) generated by BOLD (Barcode of Life Database). The scale bar at left represents a 2%
sequence difference. Collection locations for specimens are indicated, and Prionotus carolinus is used as an outgroup. GenBank accession numbers and collection data for the sequences in the tree are listed in Appendix 1.
55
have a relatively larger eye than the 250-mm specimen (30 cm TL). Instead it has a distinctly smaller eye than
the ish from 92 m (compare Fig. 5 top and middle). The same unexpected larger eye was found in photographs
of B. atrolumbus dominated by red, inferring capture from relatively deep water. Like many reef ishes that are
found from shallow water to moderate depths, the species of Bodianus have more red color in the deeper water
(presumed to make them less visible, as the red end of the spectrum is lost irst with increasing depth). Note the
larger eye of the mainly red B. atrolumbus of the middle image of Fig. 2 placed in the order of total length, but
now out of order on eye size. The irst author also found a larger eye in a specimen of the Hawaiian goby Opua
nephodes Jordan collected in 130 m, compared to ones from about 15 m. We postulate that prejuvenile ishes that
recruit to deeper water may develop a relatively larger eye than those that settle out in the shallows.
Material of Bodianus perditio examined. Tuamotu Archipelago, Gambier Group, Mangareva, BPBM 14275,
189 mm; BPBM 14293, 7: 217–320 mm. Rapa, Hiri Bay, BPBM 12968, 268 mm. New Caledonia, Bulari Pass,
BPBM 27157, 35 mm. Australia, N.S.W., Sydney, BPBM 14961, 2: 30–47 mm. Vanuatu, Erromango, USNM
360098, 159 mm. Japan, Wakanoura, USNM 71782, 230 mm; USNM 71783, 132 mm.
Acknowledgments
We thank Arnold Y. Suzumoto and Loreen R. O’Hara of the Bishop Museum for curatorial assistance and
x-rays, and Shirleen Smith of the U. S. National Museum Natural History for the loan of specimens. We are also
grateful to Alain Diringer, Phillip C. Heemstra, Dennis King, Milan Kořínek, Grant Norton, Dennis Polack, and
Rudy van der Elst for providing photographs. Special thanks are due Ofer Gon and Roger Bills of the South
African Institute for Biological Diversity for making requested counts and measurements of specimens of
Bodianus atrolumbus and Jeff Williams and Serge Planes for their help with specimens from the Gambier Islands.
DNA studies were greatly facilitated by contributions of tissues of B. perditio from New Caledonia by Eric Clua
of the French Ministry of Agriculture and Fisheries in Noumea, New Caledonia and from Gneering Shoals,
off Queensland, Australia by Arie DeJong of DeJong Marinelife of the Netherlands. Comparison sequences on
the Barcode of Life Database (BOLD) were graciously provided by Nicolas Hubert of the Université de La
Réunion, Serge Planes of the Centre National de la Recherche Scientiique, and Jeff Williams of the U.S. National
Museum of Natural History (via CRIOBE (Centre de Recherches Insulaires et Observatoire de l’Environnement
CNRS-EPHE), BIOCODE (Moore Foundation), CORALSPOT (MEDDE, ANR, Polynésie), and the LABEX
“CORAIL”); Bob Ward of CSIRO Marine and Atmospheric Research, Hobart, Australia; Dirk Steinke of the
Canadian Centre for DNA Barcoding, University of Guelph; Lourdes Vásquez-Yeomans and Martha ValdezMoreno of ECOSUR, Unidad Chetumal, in Quintana Roo, Mexico; Jacob Lowenstein of the American Museum
of Natural History; Kwang-Tsao Shao of the Biodiversity Research Center Academia Sinica, Taipei, Taiwan;
Junbin Zhang of the College of Fisheries and Life Science, Shanghai Ocean University, Shanghai; Conan Nolan
of the Marine Institute, Ireland; and David Carlon of the Bowdoin Marine Laboratory, Brunswick, Maine and
the University of Hawaiʻi and Anuschka Faucci of the University of Hawaiʻi (Appendix 1). Sally Reader and
Mark McGrouther assisted with information from the Australian Museum. The manuscript was reviewed by
Sergey V. Bogorodsky, Martin F. Gomon, and Helen A. Randall. The DNA barcoding was performed at the
Biodiversity Institute of Ontario with the support of Bob Hanner and the team at BOLD. DNA barcoding was
supported by the International Barcode of Life Project (iBOL.org) with funding from the Government of Canada
via the Canadian Centre for DNA Barcoding as well as from the Ontario Genomics Institute (2008-OGI-ICI-03),
Genome Canada, the Ontario Ministry of Economic Development and Innovation, and the Natural Sciences and
Engineering Research Council of Canada.
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*note text in blue were revisions for inal publication.
59
Appendix 1. Specimen data and GenBank accession numbers for the mtDNA COI barcode sequences used to generate the
phenogram in Fig. 7.
60
Genus
species
Collection site
Voucher
GenBank # Collector/Source
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
perditio
perditio
perditio
perditio
perditio
perditio
perditio
Gambier Islands, Fr. Polynesia
Chenggong, Taiwan
New Caledonia
Gneering Shoals, Australia
Gneering Shoals, Australia
Gambier Islands, Fr. Polynesia
New Caledonia
GAM-556
ASIZP0800729
ecbp123
djbp124
djbp122
GAM-586
ecbp122
KC684990
KC684993
KC684989
KC684996
KC684988
KC684991
KC684994
J. Williams et al.
P.F. Lee/K.T. Shao
E. Clua
A. DeJong
A. DeJong
S. Planes & P. Sasal
E. Clua
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
perditio
perditio
atrolumbus
atrolumbus
atrolumbus
solatus
solatus
solatus
solatus
solatus
solatus
bilunulatus
bilunulatus
bilunulatus
bilunulatus
bilunulatus
bilunulatus
bilunulatus
bilunulatus
albotaeniatus
eclancheri
speciosus
anthioides
anthioides
anthioides
anthioides
anthioides
diana
diana
diana
diana
diana
diana
diana
dictynna
New Caledonia
Gneering Shoals, Australia
Park Rynie, South Africa
Park Rynie, South Africa
Park Rynie, South Africa
N.W. Australia
N.W. Australia
N.W. Australia
N.W. Australia
N.W. Australia
N.W. Australia
Nosy Be, Madagascar
Park Rynie, South Africa
Park Rynie, South Africa
Amed, Bali, Indonesia
Park Rynie, South Africa
Lombok, Indonesia
Park Rynie, South Africa
Park Rynie, South Africa
Honolulu, Hawaii
Isla Isabela, Galápagos
Sao Vicente, Cape Verde
Aquarium trade, Philippines
Nosy Be, Madagascar
Nosy Be, Madagascar
Pomene, Mozambique, Africa
St. Gilles, Réunion, France
Nosy Be, Madagascar
Nosy Be, Madagascar
Nosy Be, Madagascar
Nosy Be, Madagascar
Aliwal Shoal, South Africa
Aliwal Shoal, South Africa
Nosy Be, Madagascar
Amed, Bali, Indonesia
ecbp121
djbp121
ADC220.10-3
ADC220.10-1
ADC220.10-2
BW-A1221
NMV A 29676-006
CSIRO H 4041-03
BW-A1224
BW-A1220
BW-A1222
NBE1140
ADC220.7-6
ADC220.7-3
bal11700bl280
ADC220.7-5
BW-A10667
ADC220.7-1
ADC220.7-2
08COIFishC57
GA490be270
KV4
HLC-13187
NBE1113
NBE1113
ADC09-220.5-1
REU0734
NBE1117
NBE1118
NBE1116
NBE1115
ADC220.8-3
ADC220.8-2
NBE0365
bal11700bd171
KC684995
KC684992
JF492967
DQ884974
DQ884973
DQ885070
KC684997
DQ885069
DQ885068
DQ885067
DQ885066
JF434760
JF492960
JF492958
JQ839402
JF492962
JN311757
JF492961
JF492959
KC684979
JQ839404
GQ341587
FJ582899
JF434747
JF434746
JF492957
JQ349800
JF434756
JF434755
JF434757
JF434758
JF492965
JF492966
JF434759
KC684981
E. Clua
A. DeJong
A. Connell
A. Connell
A. Connell
A. Graham/R. Ward
M. Gomon & D. Bray
A. Graham/R. Ward
A. Graham/R. Ward
A. Graham/R. Ward
A. Graham/R. Ward
N. Hubert et al.
A. Connell
A. Connell
B. Victor
A. Connell
W. White/R. Ward
A. Connell
A. Connell
D. Carlon/A. Faucci
B. Victor
R. Hanel et al.
D. Yanke/D. Steinke
N. Hubert et al.
N. Hubert et al.
A. Connell
S. Planes et al.
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
P. Heemstra/A. Connell
P. Heemstra/A. Connell
N. Hubert et al.
B. Victor
Appendix 1. cont.
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Bodianus
Prionotus
dictynna
dictynna
axillaris
axillaris
axillaris
axillaris
axillaris
axillaris
axillaris
axillaris
mesothorax
scrofa
rubrisos
rubrisos
trilineatus
trilineatus
trilineatus
trilineatus
diplotaenia
diplotaenia
diplotaenia
diplotaenia
diplotaenia
diplotaenia
pulchellus
rufus
rufus
rufus
rufus
rufus
rufus
rufus
rufus
oxycephalus
oxycephalus
oxycephalus
oxycephalus
oxycephalus
oxycephalus
oxycephalus
bimaculatus
bimaculatus
bimaculatus
carolinus
Amed, Bali, Indonesia
Amed, Bali, Indonesia
St. Leu, Réunion, France
Pomene, Mozambique, Africa
Nosy Be, Madagascar
Nosy Be, Madagascar
St. Leu, Réunion, France
Hermitage, Réunion, France
Hermitage, Réunion, France
Nosy Be, Madagascar
Amed, Bali, Indonesia
Funchal, Madeira, Portugal
Lombok, Indonesia
South China Sea
Park Rynie, South Africa
Park Rynie, South Africa
Park Rynie, South Africa
Park Rynie, South Africa
Guanacaste, Costa Rica
Guanacaste, Costa Rica
Isla Fernandina, Galápagos
Guanacaste, Costa Rica
Isla Contadora, Panama
Isla Isabela, Galápagos
Aquarium trade, Cuba
Quintana Roo, Mexico
Turneffe, Belize
Quintana Roo, Mexico
Isla Mujeres, Mexico
Quintana Roo, Mexico
Quintana Roo, Mexico
Aquarium trade, Cuba
San Salvador, Bahamas
Hainan, China
Hainan, China
Hainan, China
Hainan, China
Hainan, China
Hainan, China
Hainan, China
Park Rynie, South Africa
Park Rynie, South Africa
Sodwana Bay, South Africa
Massachusetts, USA
bal11700bd123
bal11700bd133
REU1617
ADC09-220.6-1
NBE0075
NBE0356
REU0995
REU0775
REU0735
NBE0074
bal11800bm126
LB122
BW-A10666
MBCSCHSY08337
ADC220.9-1
ADC220.9-3
ADC220.9-2
ADC220.9-5
JHLOW00156
JHLOW00230
gal98606bd108
JHLOW00340
pe821120bd600
gal98528bd101
HLC-11025
ECOCH5748-58b
cn10b99
ECOCH5748-58c
ECOCH7143-209
ECOCH5748-58d
ECOCH5748-58d
HLC-12307
bah91729br120
MBCSCHSY08518
MBCSCHSY08519
MBCSCHSY08515
MBCSCHSY08517
MBCSCHSY08336
MBCSCHSY08516
MBCSCHSY08520
ADC220.7a-2 5
ADC220.7a-1
ADC12 220.7a-3
DAL 07-097
KC684982
KC684980
JF434748
GU805092
JF434753
JF434752
JF434749
JF434750
JF434751
JF434754
JQ839406
GQ341586
JN311756
FJ237633
JF492970
JF492968
JF492971
JF492969
KC684983
KC684987
KC684984
KC684986
JQ839403
KC684985
FJ582902
GU225156
HQ987864
GU225154
JN311758
GU225155
GU225155
FJ582903
JQ839407
FJ237628
FJ237627
FJ237631
FJ237629
FJ237632
FJ237630
FJ237626
JF492963
JF492964
KF489507
KC015843
B. Victor
B. Victor
N. Hubert et al.
A. Connell
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
N. Hubert et al.
B. Victor
R. Hanel et al.
W. White/R. Ward
Junbin Zhang
A. Connell
A. Connell
A. Connell
A. Connell
J. Lowenstein/B. Victor
J. Lowenstein/B. Victor
B. Victor
J. Lowenstein/B. Victor
B. Victor
B. Victor
D. Yanke/D. Steinke
M. Valdez-Moreno
C. Nolan/B. Victor
M. Valdez-Moreno
L. Vásquez-Yeomans
M. Valdez-Moreno
M. Valdez-Moreno
D. Yanke/D. Steinke
G. Wellington/B. Victor
Junbin Zhang
Junbin Zhang
Junbin Zhang
Junbin Zhang
Junbin Zhang
Junbin Zhang
Junbin Zhang
A. Connell
A. Connell
A. Connell
P. Chase, NOAA
61