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. References Allen, G.R. & Swainston, R. (1988) The Marine Fishes of North-Western Australia. Western Australian Museum, Perth, Australia, 6 + 201 pp. 56 Allen, G.R., Hoese, D.F., Paxton, J.R., Randall, J.E., Russell, B.C., Starck II, W.A., Talbot, F.H. & Whitley, G.P. (1976) Annotated checklist of the ishes of Lord Howe Island. Records of the Australian Museum, 30, 365–454. Bacchet, P., Zysman, T. & Lefèvre, Y. (2006) Au Vent des Iles. Tahiti, 609 pp. Baissac, J. de B. (1953) Contribution à l’étude des poissons de l’Ile Maurice V. Proceedings of the Royal Society of Arts and Sciences of Mauritius, 1(3), 185–240. Baissac, J. de B. (1976) Poissons de mer des eaux de l’Ile Maurice. Proceedings of the Royal Society of Arts and Sciences of Mauritius, 3(2), 191–226. Bauchot, M.L. (1963) Catalogue Critique des Types de Poissons du Muséum National d’Histoire Naturelle. Publications du Museum National d’Histoire Naturelle, 20, 1–195. Bermingham, E., McCafferty, S.S. & Martin, A.P. (1997) Fish biogeography and molecular clocks: perspectives from the Panamanian isthmus. In: Kocher, T.D. & Stepien, C.A. (eds.) Molecular systematics of ishes. Academic Press, NewYork. pp. 113–128. Chen, J.-P., Shao, K.-T., Jan, R.-Q., Kuo, J.-W. & Chen, J.-Y. (2010) Marine Fishes in Kenting National Park. Kenting National Park HQ, Henchun, ROC, 650 pp. (in Chinese). Chen, J.T.F. (1952) Check-list of the species of ishes known from Taiwan (Formosa). Quarterly Journal of the Taiwan Museum, 6(2), 102–140. Cuvier, G. & Valenciennes, M.A. (1839) Histoire Naturelle des Poissons. Chez Pitois-Levrault et Cie, vol. 13, xix + 505 pp. Debelius, H. (1993) Indian Ocean–Tropical Fish Guide. Aquaprint Verag GmbH, Neu Isenburg, 321 pp. De Vis, C.W. (1883) Descriptions of (2) new genera and (11) species of Australian ishes. Proceedings of the Zoological Society of New South Wales, 8, 283–289. De Vis, C.W. (1884) On new ish from Moreton Bay. Proceedings of the Royal Society of Queensland, 1, 144–147. De Vis, C.W. (1885) New ishes in the Queensland Museum. Proceedings of the Zoological Society of New South Wales, 9, 869–887. Fourmanoir, P. & Guézé, P. (1961) Les poissons de la Réunion. Publications de l’Institute de Recherche Scientiique de Madagascar, 1–17. Fowler, H.W. & Bean, B.A. (1928) The ishes of the families Pomacentridae, Labridae, and Callyodontidae, collected by the United States Bureau of Fisheries steamer “Albatross”, chiely in Philippine Seas and adjacent waters. Bulletin of the United States National Museum, 100, 7, viii + 525 pp. Francis, M.P. (1991) Additions to the ish faunas of Lord Howe, Norfolk, and Kermadec Islands, Southwest Paciic Ocean. Paciic Science, 45(2), 204–220. Fricke, R. (1999) Fishes of the Mascarene Islands (Réunion, Mauritius, Rodriguez). An Annotated checklist with descriptions of new species. Koeltz Scientiic Books, Koenigstein, Germany, i-viii + 759 pp. Gilchrist, J.D.F. & Thompson, W.W. (1908) Descriptions of ishes from the coast of Natal. Annals of the South African Museum, 6(2), 145–206. Gilchrist, J.D.F. & Thompson, W.W. (1909) Descriptions of ishes from the coast of Natal. Pt. II. Annals of the South African Museum, 6(3), 213–279. Gloerfelt-Tarp, T. & Kailola, P.T. (1984) Trawled Fishes of Southern Indonesia and Northwestern Australia. The Australian Development Assistance Bureau, Canberra; The Directorate General of Fisheries, Indonesia, Jakarta; and The German Agency for Technical Cooperation, Eschborn, xvi + 406 pp. Gomon, M.F. in Fischer, W. & Bianchi, G. (1984) Labridae, FAO species identiication sheets for ishery purposes. Western Indian Ocean ishing area 51. Vol. 1. [pag. var.]. FAO, Rome. Gomon, M.F. (2006) A revision of the labrid ish genus Bodianus with descriptions of eight new species. Records of the Australian Museum, Supplement, 30, 1–133. Grant, E.M. (1982) Guide to ishes. Dept. of Harbours & Marine, Brisbane, Australia, 896 pp. Günther, A.C.L.G. (1862) Catalogue of the Acanthopterygii, Pharyngognathi and Anacanthini in the Collection of the British Museum. Vol. 4, Catalogue of the Fishes in the British Museum. London, xxii and 534 pp. Günther, A. (1873–1910). Andrew Garrett’s Fische der Südsee. Journal des Muséum Godeffroy, parts 3, 6, 9, 11, 13, 15, 16, and 17 in vols. 2, 4, and 6, Hamburg, Germany. Heemstra, E., Heemstra, P., Smale, M., Hooper, T. & Pelicier, D. (2004) Preliminary checklist of coastal ishes from the Mauritian island of Rodrigues. Journal of Natural History, 38, 3315–3344. 57 Heemstra, P.C. & Heemstra, E. (2004) Coastal ishes of Southern Africa. National Inquiry Service Centre (NISC) and South African Institute for Aquatic Biodiversity (SAIAB), Grahamstown, 488 pp. Ivanova, N.V., Zemlak, T.S., Hanner, R.H. & Hebert, P.D.N. (2007) Universal primer cocktails for ish DNA barcoding. Molecular Ecology Notes, 7, 544–548. Jordan, D.S. & Snyder, J.O. (1902) A review of the labroid ishes and related forms found in the waters of Japan. Proceedings of the U.S. National Museum, 24, 1266, 595–662. Jordan, D.S. & Seale, A. (1906) The ishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the ishes of Oceania. Bulletin of the United States Bureau of Fisheries, 25: 173–455. Jordan, D.S., Tanaka, S. & Snyder, J.O. (1913). A catalogue of the ishes of Japan. Journal of the College of Science, Tokyo Imperial University, 23(1), 1–407. Kuiter, R.H. (1993) Coastal Fishes of South-Eastern Australia. University of Hawaiʻi Press, Honolulu, HI, xxxi + 437 pp. Kuiter, R.H. (2010) Labridae Fishes: Wrasses. Aquatic Photographics, Seaford, Victoria, 390 pp. Kulbicki, M. & Williams, J.T. (1997) Checklist of the shoreishes of Ouvéa Atoll, New Caledonia. Atoll Research Bulletin, 444, 1-26. Laboute, P. & Grandperrin, R. (2000) Poissons de Nouvelle-Calédonie. Editions Catherine Ledru, Nouméa, New Caledonia, 520 pp. Lieske, E. & Myers, R. (1994) Coral Reef Fishes: Caribbean, Indian Ocean and Paciic Ocean Including the Red Sea. Harper Collin, Scranton, PA, 400 pp. Masuda, H. & Kobayashi, Y. (1994) Grand Atlas of Fish Life Modes. Tokai University Press, Tokyo, Japan, 465 pp. (in Japanese). Masuda, H., Araga, C. & Yoshino, T. (1975) Coastal Fishes of Southern Japan. Tokai University Press, Tokyo, Japan, 382 pp. Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.) (1984) The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, Japan, Vol. l (text: xxii + 437 pp.) and vol. 2 (plates). Matsubara, K. (1964) Fish Morphology and Hierarchy. Ishizaki-Shoten, Tokyo, Part II: 291–1605 (in Japanese). Nishiyama, K. & Motomura, H. (2012) A Photographic Guide to Wrasses of Japan. Toho Press, Osaka, Japan, 302 pp. (in Japanese). Ogilby, J.D. (1889) The reptiles and ishes of Lord Howe Island. Memoirs of the Australian Museum, 2, 51–74. Ogilby, J.D. (1916) Ichthyological notes (no. 3). Memoirs of the Queensland Museum, 5, 181–185. Okamura, O. & Amaoka, K. (1997) Sea Fishes of Japan. Yama-Kei Publishers, Tokyo, 783 pp. (in Japanese) Parenti, P. & Randall, J.E. (2000) An annotated checklist of the species of the labroid ish families Labridae and Scaridae. Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology, 68, 1–97. Quoy, J.R.C. & Gaimard, J.P. (1834) Poissons. In: Voyage de découvertes de “l’Astrolabe,” exécuté par ordre du Roi, pendant les années 1826-29, sous le commandement de M. J. Dumont d’Urville. Paris, Zoologie, 3(2), 645–720. Randall, J.E. (1982) Examples of antitropical and antiequatorial distribution of Indo-West Paciic ishes. Paciic Science, 1981, 35(3): 197–209. Randall, J.E., Allen, G.R. & Steene, R.C. (1990) Fishes of the Great Barrier Reef and Coral Sea. University of Hawai‘i Press, Honolulu, HI, xx + 557 pp. Randall, J.E. (2005) Reef and Shore Fishes of the South Paciic. New Caledonia to Tahiti and the Pitcairn Islands. University of Hawaiʻi Press, Honolulu, HI, xii + 707 pp. Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: The Barcode of Life Data System (www.barcodinglife.org). Molecular Ecology Notes, 7(3), 355–364. Sainsbury, K.J., Kailola, P.J. & Leyland, G.G. (1984) Continental Shelf Fishes of Northern and North-Western Australia. Clouston & Hall and Peter Pownall Fisheries Information Service, Canberra, viii + 375 pp. Shao, K.T. (2013) The Fish Datebase of Taiwan. www electronic publication; available from http://ishdb.sinica. edu.tw, (accessed 1 Oct. 2013). Smith, J.L.B. (1949) The Sea Fishes of Southern Africa. Central News Agency, South Africa, xvi + 550 pp. Smith, M.M. & Heemstra, P.C. (1986) Smiths’ Sea Fishes. Macmillan South Africa, Johannesburg, xx + 1047 pp., 144 pls. 58 Taquet, M. & Diringer, A. (2012) Poisson de l’ocean Indien et de la mer Rouge, ed. 2. Editions Quae, Versailles, France, 679 pp. van der Elst, R. (1981) A Guide to the Common Sea Fishes of Southern Africa. C. Struik, Capetown, South Africa, 367 pp. Westneat, M.W. in Carpenter, K.E. & Niem, V.H. (2001) Labridae, The Living Marine Resources of the Western Central Paciic, vol. 6, Bony ishes part 4 (Labridae to Latimeriidae). Food and Agriculture Organization of the United Nations, Rome, pp. 3381–3467. Yu, M.-J. (1968) The labrid ishes of Taiwan. Biological Bulletin Tunghai University, Ichthyological Series, 4, 1–136. *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