Memoirs of Museum Victoria 76:1-111 (2017) Pubiished 2017

1447-2554 (On-iine)

http://museumvictoria.com.au/about/books-and-journais/journais/memoirs-of-museum-victoria/

DOi: http://doi.Org/10.24199/j.mmv.2017.76.01

A review of the tuskfishes, genus Choerodon (Labridae, Perciformes), with

descriptions of three new species

(http://zoobank.org/urn:lsid:zoobank.org:pub:7B3010E9-5D84-40B6-9A3E-4E7C6761BA05)

Martin F. Gomon

Abstract

Keywords

Ichthyology, Sciences Department, Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia

(mgomon@museum.vic.gov.au)

Gomon, M.E. 2017. A review of the tuskfishes, genus Choerodon (Labridae), with descriptions of three new species.

Memoirs of Museum Victoria 76: 1-111.

The Indo-West Pacific labrid genus Choerodon comprises 27 species assigned here to six subgenera based on

genetic and morphological evidence. Three of the subgenera are monotypic with distinctive morphological features,

another two comprise species with a relatively deep body and generalised form and the remaining is a complex of small

species referred to below as dwarf tuskfishes. Twenty-three species have valid scientific names, three are undescribed and

one requires a replacement name because the name in use is a junior homonym. More than half of the species of Choerodon

occur in northern Australian waters while only four are distributed in the western Indian Ocean, with two of the four

currently undescribed. Among the undescribed species, Choerodon cypselurus sp. nov. is described from two individuals

collected on the Saya de Malha Bank in the central part of the Indian Ocean and assigned to the subgenus Aspiurocheilus

on the basis of body form and merstic values. Choerodon skaiopygmaeus sp. nov. is a dwarf species described from seven

individuals collected off Somalia. The absence of life colour information for type specimens makes comparison with

described species difficult, but the species is distinguished by pigmentation and morphometric evidence. The third new

species, Choerodon aurulentus sp. nov., the largest of known dwarf tuskfishes, is described from two specimens collected

on the northern Norfolk Ridge at the north-eastern extent of the Tasman Sea between New Caledonia and New Zealand.

Choerodonoides japonicus Kamohara, 1958 is a junior homonym of Labrus japonicus Valenciennes, in Cuvier and

Valenciennes, 1839, and is replaced by Choerodon albofasciatus nom. nov. Primary synonymies, synopses, known

distributions and a dichotomous key to known species are provided.

Xiphocheilus, sp. nov., taxonomy, Indian Ocean, western Pacific, Saya de Malha Bank

Introduction

The generic name Choerodon initially appeared in Bleeker

(1847) as a nomen nudum for Labrus macrodontus Lacepede,

1802, a junior synonym of Sparus anchorago Bloch, 1790.

Although a growing number of species were found to be

referrable to what is a clearly definable labrid genus, early

doubt as to the name’s applicability to the group apparently

existed because additional names were proposed, even by

Bleeker (1858b). By the early 1900s Choerodon finally gained

widespread acceptance for most species referred to the genus,

although reference to species as Choerops Riippell, 1852,

persisted in publications until then (e.g. Choerops rubescens

Gunther, 1862, Choerops Mae under Cartier, 1874, Choerops

cephalotes Castelnau, 1875, Choerops perpulcher De Vis,

1885, Choerops albiqenaDtWis, 1885, Choerops Hodgkinsonii

Saville-Kent, 1893, Choerops nyctemblema Jordan &

Evermann, 1902, Choerops palawanensis Seale, 1910).

Gomon (1997) provided a historical perspective, an overview

of the assemblage then thought to number 25 species, and a

hypothesis of the relationship of the genus with others in the

tribe Hypsigenyini based on morphological examination of

previously described taxa and collection materials available at

that time.

Clements et al.’s (2004) genetic comparison of several

members of the tribe with species long referred to a temperate

Australasian family Odacidae provided evidence for paraphyly

within the Hypsigenyini and a sister group relationship of the

odacid species with at least one species referred to the genus

Choerodon, its type species Choerodon anchorago. Westneat

and Alfaro (2005) subsequently published an analysis of the

families Labridae, Scaridae and Odacidae based on a larger

data set corroborating the work of Clements et al. (2004) and

the monophyly of the tribe Hypsigenyini - with the inclusion

of the “Odacidae” - as the subfamily Hypsigenyinae. Westneat

and Alfaro (2005) recovered a sister group relationship of the

monotypic Xiphocheilus with the two species of Choerodon

M.F. Gomon

included in their study as proposed by Gomon (1997), which

in turn is sister to the “odacid” species, all with strong support.

Puckridge et al. (2015) compared genetic sequences for what

proved to be 20 species referrable to Choerodon, as defined by

Gomon (1997), together with sequences of Xiphocheilus typus,

four species of “odacids”, five species of other hypsigenyin

genera and two labrids. The study provided evidence for the

monophyly of Choerodon with the inclusion of X. typus within

one of four major clades. An independent analysis of sequences

of many of the same species, as part of a larger data set for an

ongoing investigation of the interrelationships of the Labridae

by Westneat (personal communication), corroborates these

findings. Morphology also supports these relationships and

provides additional evidence for the subdivision of two of

Puckeridge et al.’s clades, resulting in a cladistic tree

comprising six clades.

This contribution presents a revised description of the

genus Choerodon, primary synonymies, synopses and known

distributions of described species referrable to the genus

arranged in a taxonomic framework of six subgenera (most

taking available names), descriptions of three new species, a

new name for a previously described species (because its

current name is a junior homonym) and a dichotomous key to

the resultant 27 species. A list of proposed scientific names

and their nomenclatural status for species referrable to the

genus Choerodon is provided in Table 1.

Methods and Materials

General terminology and methodology follows Gomon (1997),

except head depth is measured on the transverse plane at the

posterior margin of the orbit and the interorbital width is the

bony interorbital width. Caudal fin dorsal lobe length is the

length of the longest ray in the dorsal lobe. Specimen lengths

reported are standard lengths (SL), unless noted as total length

(TL), and were measured to the nearest three significant digits;

those less than 100 mm and given to two significant digits are

as recorded in the respective museum registers and not

remeasured. Head length when used as a proportional

measurentmay be abbreviated as HL. As relative measurements

of body dimensions may vary ontogenetically, morphometric

comparisons provided in taxonomic treatments are based on

specimens larger than about 60 mm SL where possible.

Specimens on which species treatments are based, apart from

newly described species, are listed in the Appendix. In the

lists of type specimens for new species treatments, the values

enclosed by parentheses following registration numbers

convey the size of the specimens in mm SL. Species

distributions are based on material in museum collections,

except where stated, to cut down on potential errors associated

with misidentifications in the literature. Institution

abbreviations follow Sabaj (2016).

For discussion purposes, Westneat and Alfaro’s (2005)

classification placing the tribe Hypsigenyini (Gomon, 2006)

together with the family Odacidae (Gomon & Paxton, 1985,

and others) in the subfamily Hypsigenyinae is adopted here,

with genera and species previously placed in the family

“Odacidae” referred to collectively as “odacids”.

Species of Choerodon, like the vast majority of labrids, are

protogynous hermaphrodites with individuals maturing first as

females before transforming into males. All appear to have

morphological differences between the two mature stages, if

only in the form of colour pattern. Juveniles likewise frequently

differ in colour pattern from mature adults. The three stages are

referred to below as juvenile, initial phase and terminal phase.

A reanalysis of the interrelationships of Choerodon species

based on mitochondrial (COl and 16S) and nuclear (RAG2

and Tmo4c4) genes was undertaken by Luisa Teasdale

(personal communication) employing sequences featuring in

the original study by Puckridge et al. (2015) with the addition

of 14 COl sequences from recently acquired tissue, including

those of the problematic species Choerodon gymnogenys not

represented in the previous analysis, and 16S, RAG2 and

Tmo4c4 sequences for a specimen of Choerodon

zosterophorus, following the methodology described in that

publication. Information about the sources and taxonomic

identities of the previous sequences are available in the

supplementary information accompanying Puckridge et al.

(2015: Table SI), with details of the new sequences provided in

Table 7. A species reported in Puckridge et al. (2015: 55, 56,

figs 1 & 2) as Choerodon cf margaritiferus is re-identified

below as Choerodon albofasciatus nom. nov. As only COl

sequences were employed for 14 of the 15 additional species

and only 16S, RAG2 and Tmo4c4 sequences for the remaining

specimen in the latest analysis, their positions in the tree are

indicative only of relative relationships. The same relative

positions were recovered in maximum likelihood, maximum

parsimony and neighbour joining trees. Consequently, scales

for branch lengths and confidence values are not indicated in

fig. 1.

Choerodon Bleeker, 1847

Figure 1; table 1

The generic synonymy is a sum of the subgeneric synonymies.

Description. Dorsal fin rays XII or XIII (rarely XI or XIV), 7

or 8 (rarely 6 or 9), total rays 20 (rarely 19); anal fin rays III, 9

or 10 (rarely 11); caudal fin rays 7-10 -i- 12 -i- 7-9 (rarely 6);

pectoral fin rays ii, 13-17 (rarely 12); vertebrae 10 or 11 -i-16 or

17 = 27; pleural ribs ending on 10th or 11th vertebra; epipleural

ribs ending on 10th-14th vertebra; lateral line scales 27 (rarely

26) -I- 2; scales above lateral line 21/2-5; scales below lateral line

approximately 814-11; predorsal scales approximately 4-15;

total gill rakers 13-18.

Body moderately slender to deep; caudal peduncle

moderately slender to very slender; head of moderate depth to

very deep; snout usually rather short; head broadly pointed to

bluntly rounded; dorsal outline of forehead and snout convexly

curved to nearly straight in lateral aspect, nape convexly

curved, strongly curved in some species; interorbital

moderately broad; jaws not attenuate.

Dorsal and anal fins usually without obvious scaly basal

sheaths, usually one much smaller scale or 1-3 progressively

smaller scales at both ends of at least some oblique scale rows

adjacent to fin bases, sheath when present rarely reaching well

Review of Choerodon tuskfishes

Table 7. Species identification, tissue number, voucher number, GenBank accession numbers and collection localities for additonal sequences

used in generating the tree in Figure 1.

M.F. Gomon

[{

Choerodon gomoni

Choerodon margaritiferus

Choerodon gymnogenys

Choerodon alboguttatus

Choerodon jordani

Choerodon zosterophorus

Choerodon frenatus

Choerodon sugiUatum

Choerodon typus

Choerodon robustus

Choerodon zamboangae

Choerodon azurio

Choerodon monostigma

Choerodon rubescens

Choerodon cauteroma

— Choerodon schoenleinii

Choerodon cephalotes

Cladela (Subgenus Peaolopesia)

] Cladelb (Subgenus Xiphocheilus)

Clade2 (Subgenus Aspiurochilus)

Clade 3 (Subgenus Choerodon)

Choerodon anchorago

Choerodon oligacanthus

Choerodon cyanodus

Choerodon venustus

- Choerodonfasciatus ] Clade 4a (SubgeHus/./enorc/e//c 7 )

- Choerodonvitta ] Clade 4b (SubgepUS/.L/fyan/Vabrus)

Heteroscarus acroptilus

Figure 1. Redrafted “Fig. 1” of Puckridge et al. (2015) with two additional species of Choerodon (red) inserted following a reassessment of the

original data set plus 17 new sequences, employing the methodology of the authors. The composition of clades is the same as hypothesised by

Puckridge et al. (2015) with two clades subdivided to reflect divergence inherent in the morphology of component species.

Review of Choerodon tuskfishes

Table 1. Proposed scientific names for species referrable to the genus Choerodon with their current statuses.

Nominal species Author, Publication date

Senior synonym

Choerops albiqena De Vis, 1885

Choerodon cyanodus

Choerodon albofasciatus Gomon, 2017

Choerodon albofasciatus

Choerodon ambiguus Ogilby, 1910

Choerodon venustus

Sparus anchorago Bloch, 1791

Choerodon anchorago

Lachnolaimus arilca Richardson, 1848

Choerodon cyanodus

Choerodon aurulentus Gomon, 2017

Choerodon aurulentus

Torresia australis Castelnau, 1875

Choerodon schoenleinii

Choerops azurio Jordan & Snyder, 1901

Choerodon azurio

Choerodon balerensis Herre, 1950

Choerodon fasciatus

Choerops brenchleyi Gunther, 1872

Choerodon zosterophorus

Choerodon cauteroma Gomon & Allen, 1987

Choerodon cauteroma

Choerops cephalotes Castelnau, 1875

Choerodon cephalotes

Chaerops crassus Castelnau, 1875

Choerodon cyanodus

Labrus cyanodus Richardson, 1843

Choerodon cyanodus

Cossyphus cyanostolus Richardson, 1846

Choerodon schoenleinii

Choerodon cypselurus Gomon, 2017

Choerodon cypselurus

Choerops dodecacanthus Bleeker, 1868

Choerodon robustus

Xiphochilus fasciatus Gunther, 1867

Choerodon fasciatus

Choerodon frenatus Ogilby, 1910

Choerodon frenatus

Choerodon gomoni Allen & Randall, 2002

Choerodon gomoni

Choerops graphicus De Vis, 1885

Choerodon graphicus

Xiphochilus gymnogenys Playfair & Gunther, 1867

Choerodon gymnogenys

Choerops Hodgkinsonii Saville-Kent, 1893

Choerodon cephalotes

Choerodonoides japonicus Kamohara, 1958

Choerodon albofasciatus

Labrus japonicus Valenciennes, in Cuvier & Valenciennes, 1839

Choerodon azurio

Choerops jordani Snyder, 1908

Choerodon jordani

Crenilabrus leucozona Bleeker, 1858

Choerodon anchorago

Torresia lineata De Vis, 1885

Choerodon schoenleinii

Choerodon macleayi Ramsay & Ogliby, 1887

Choerodon cephalotes

Cossyphus macrodon Bleeker, 1849

Choerodon anchorago

Labrus macrodontus Lacepede, 1802

Choerodon anchorago

Choerops Maeander Cartier, 1874

Choerodon anchorago

Choerodon margaritiferus Fowler & Bean, 1928

Choerodon margaritiferus

Cossyphus maxillosus Guichenot, 1865

Choerodon robustus

Choerodon melanostigma Fowler & Bean, 1928

Choerodon zamboangae

Choerops meleagris'RappeW, 1852

Choerodon anchorago

Lepidaplois mirabilis Snyder, 1908

Choerodon fasciatus

Choirodon monostigma Ogilby, 1910

Choerodon monostigma

Chaerops notatus Alleyne & Macleay, 1877

Choerodon schoenleinii

Choerops nyctemblema Jordan & Evermann, 1902

Choerodon schoenleinii

Crenilabrus oligacanthus Bleeker, 1851

Choerodon oligacanthus

Choerops olivaceus De Vis, 1885

Choerodon cyanodus

Cossyphus ommopterus Richardson, 1846

Choerodon schoenleinii

Choerops palawanensis Seale, 1910

Choerodon oligacanthus

Choerodon paynei Whitley, 1945

Choerodon cyanodus

Choerops perpulcher De Vis, 1885

Choerodon cephalotes

Choerodon pescadorensis Yu, 1968

Choerodon zamboangae

Choerodon quadrifasciatus Yu, 1968

Choerodon azurio

M.F. Gomon

Table 1 (cont.). Proposed scientific names for species referrable to the genus Choerodon with their current statuses.

Nominal species Author, Publication date

Senior synonym

Xiphochilus quadrimaculatus Gunther, 1880

Choerodon typus

Xiphochilus rohustus Gunther, 1862

Choerodon rohustus

Choerops rubescens Gunther, 1862

Choerodon rubescens

Choerodon ruhidus Scott, 1959

Choerodon schoenleinii

Cossyphus Schoenleinii Valenciennes, in Cuvier & Valenciennes, 1839

Choerodon schoenleinii

Crenilabrus stejnegeri Ishikawa, 1904

Choerodon azurio

Choerodon sugillatum Gomon, 1987

Choerodon sugillatum

Choerodon transversalis Whitley, 1956

Choerodon graphicus

Xiphocheilos typus Bleeker, 1856

Choerodon typus

Choerops unimaculatus Cartier, 1874

Choerodon schoenleinii

Choerops unimaculatus De Vis, 1885

Choerodon cyanodus

Choerops venustus De Vis, 1884

Choerodon venustus

Choerodon vitta Ogilby, 1910

Choerodon vitta

Choerodon weberi Ogliby, 1911

Choerodon anchorago

Choerodon skaiopygmaeus Gomon, 2017

Choerodon skaiopygmaeus

Choerops zamboangae Seale & Bean, 1907

Choerodon zamboangae

Choerops zosterophorus Bleeker, 1868

Choerodon zosterophorus

onto proximal parts of fins. Predorsal scales reaching forward

on dorsal midline of head, barely in advance of above centre of

eye, not quite to above dorsal end of preopercle or somewhere

between. Cheek only partially covered by scales, scales

imbricate to embedded, reaching forward to corner of mouth,

barely below lower edge of orbit or somewhere between,

moderately broad to broad naked margin posteriorly and

ventrally on preopercle; subopercle partially scaled to mostly

naked, scales confined to 1-3 rows adjacent to ventral edge of

preopercle; rows extending forward at most to just short of

below anterior extent of ventral preopercular edge, confined to

dorsoposterior corner of subopercle at the least; scales in longest

row numbering approximately 1-13; lower jaw naked. Lateral

line scales each with an unbranched or branched laterosensory

canal tube, tube extremely dentritic with irregularly curved

branches in some species, best developed in large individuals;

centre of second penultimate lateral line scale usually lateral to

posterior endge of hypural. Cephalic sensory canal pores

numerous but confined to lines or short branches associated

with major canals, or extremely numerous covering much of

dorsal side of head and snout, often extending onto anterodorsal

portion of cheek. Posterior edge of preopercle serrate to smooth.

Mouth mostly horizontal, posterior corner situated below point

just anterior to forward extent of orbit, positioned slightly

posterior to centre of eye, or somewhere between; lower lip

rather narrow, often mostly obscured by skin flap on upper side

of mouth when jaws occluded; upper lip similarly narrow and

obscured by skin flap except at anterior end of jaw; posterior end

of maxilla never exposed; crease at corner of mouth curved

downward. Gill rakers on first arch simple to arborescent.

Upper jaw with 2 prominent anterior canines; a third very

small canine often present mesial to first prominent canine and

adjacent to symphysis of jaw; first prominent canine nearly

equal to or longer than second, almost four times the length of

second in some species; both canines directed mostly ventrally,

first occasionally slightly mesially or slightly laterally, second

often slightly laterally, occasionally recurved slightly

posteriorly; dental ridge smooth to rough with numerous minute

teeth along edge; posterior canine present or absent, usually

rather small when present and directed mostly ventrally. Lower

jaw with 2 prominent anterior canines; first canine approximately

1/3 to twice the length of second; first canine directed

anterodorsally to dorsally and slightly mesially, second canine

directed anterodorsally or dorsally to dorsolaterally or curved

dorsolaterally to laterally and posteriorly; dental ridge entirely

smooth to rough with numerous minute teeth along edge, or

mostly smooth or rough edged on the anterior 1/4-2/3 of jaw

with approximately 1-10 small to moderately sized caniniform

teeth posteriorly; teeth when present in one or two series of a

single row; anteriormost teeth usually small, becoming

progressively larger posteriorly; second series, when present,

uniformly short, 2-4 in number. Vomerine teeth absent.

Dorsal fin continuous, spines subequal, pungent; posterior

tips of dorsal and anal fins broadly to narrowly rounded,

posterior rays not filamentous; posterior tips of fins reaching

short of posterior edge of hypurals, almost midway between

hypural edge and posterior edge of scaly caudal fin base, or

somewhere between. Posterior edge of caudal fin rounded,

slightly pointed midposteriorly, truncate, slightly forked or

double emarginate, upper and lower lobes only slightly produced

at most. Pectoral fin with upper rays distinctly longer than lower,

dorsoposterior corner slightly pointed in some, posterior edge

often obliquely straight, lower portion sometimes broadly

rounded, but some species with ventralmost rays markedly

longer than those above forming a falcate posterior margin of

fin, lower arm of crescent distinctly shorter than upper. Pelvic

Review of Choerodon tuskfishes

fin short to long; posterior tip of fin reaching distinctly short of

anus to centre of anal fin base, or somewhere between.

Species of this genus are rather variable in size, reaching a

small (106 mm SL) to large maximum size, largest specimen

examined 530 mm SL, although the same species is reported to

reach one metre.

Pigmentation in alcohol. Juveniles often pale dusky with narrow

to broad ill-defined darker bands crossing side; prominent often

ocellated dark spot frequently present on posterior half of dorsal

fin, as well as on other fins in some species; species otherwise

variously pigmented.

Initial phase adults often mostly pale or pale dusky, with or

without some combination of dusky to dark or distinctly pale

spots, blotches, bands or stripes on body and fins.

Terminal phase adults often with distinctive pigmentation

differing from that of initial phase adults.

Colour in life. Extremely variable ontogenetically and

interspecifically in most.

Distribution. Species of the genus are confined to the Indo-west

Pacific, all but one not occurring east of Guam, Palau, the

Solomon Islands, Vanuatu and New Caledonia; the sole exception,

Choerodon jordani, is recorded from Tonga, Fiji and American

Samoa. Of the 27 species of Choerodon, well over half are present

on the Australian plate, with two-thirds of that complement

essentially confined to it, while only four are distributed in the

western Indian Ocean. Practically all species occur at rather

shallow depths, with none known to range below 100 m.

Ecologically, most, if not all, prefer back or coral reef areas, or are

associated with substrates having similarly low relief.

Etymology. Choerodon is derived from a combination of the

masculine Greek nouns choiros, “pig”, and odon, “tooth”, in

reference to the prominent anterior canines present in species of

this genus.

Comments. The 27 species of the genus Choerodon are

distinguishable from those of all other hypsigenyine genera in

having XII, 8 or XIII, 7 dorsal fin rays (XI, 9-11, XII, 9-11, XIII,

9 or XIV-XXVII, 9-11 = 23^44 in others) and 27 total vertebrae

(versus 28, 27 in Anchichoerops and rarely in Semicossyphus,

and 14-33 -i-14—22 = 31-54 in “odacids”). The rather low number

of lateral line scales (26-28 -i- 2) present in all species is found

elsewhere in the subfamily only in the genus Decodon. Choerodon

differs from that genus by dorsal fin formula (XI, 9-10 in

Decodon), in having fewer vertebrae and in having predorsal

scales reaching forward only to above centre of eyes at most (at

least to anterior nostril in Decodon). Although Gomon (1989)

recognised Xiphocheilus as a monotypic genus distinct from

Choerodon, Puckridge et al. (2015) recovered it within

Choerodon, clustering as a sister group with the six species of the

subgenus Peaolopesia they examined. The recognition of

Xiphocheilus as a distinct genus appears to have been based on

autapomorphic features without valid synapomorphies supporting

the monophyly of the remaining species of Choerodon examined.

In their study of the interrelationships of 20 species of the

genus Choerodon, Puckeridge et al. (2015) recovered five major

clades, with Choerodon typus representing a clade sister to the

largest clade of species within the genus. On the basis of the

same morphological features that supported it as a monotypic

genus, the clade is regarded here as a major clade within

Choerodon, albeit a monotypic one. To further test

interrelationships of Choerodon species, sequences of an

additional 15 individuals, including the two species C.

gymnogenys and C. zosterophorus not previously represented in

the analysis, as well as sequences for western Indian Ocean

specimens of Choerodon robustus, were obtained and together

with sequences featuring in the original study reanalysed by L.

Teasdale using the methodology employed by Puckridge et al.

(2015). It was deemed important to include material from the

true C. gymnogenys because the name had been mistakenly

applied over the years to a number of species of similar form

distributed in the western Pacific. The outcome supported the

confinement of that species to the western Indian Ocean and a

sister relationship of it with a terminal clade comprising

Choerodon gomoni and C. margaritiferus. Choerodon

zosterophorus was recovered as a close sister to C. jordani

within the larger clade (fig. 1, clade la) that includes C.

gymnogenys. The conspecificity of specimens of C. robustus

sourced in the western Indian Ocean with those collected in

Japan and Indonesia was verified without significant genetic

separation between sequences obtained from specimens

collected in the widely separated localities. Figure 1 of Puckridge

et al. (2015) has been redrawn to incorporate these observations

(fig. 1) with the newly incorporated taxa in red.

The 27 species of the genus are here referred to six subgeneric

clades with the monophyly of each supported by both genetic

and morphological evidence. To his credit, Kuiter (2010) foresaw

some of this taxonomic arrangement in his image rich treatment

of the Labridae. Five subgenera take available group names, with

only the monotypic branch comprising Choerodon vitta

requiring a new epithet.

A number of species of this genus are only poorly represented

in collections, especially at juvenile and terminal adult stage

sizes. Consequently, ontogenetic changes in colour patterns may

not be known for those species. Furthermore, among those species

in which juvenile colour patterns are known, a number have

rather similar dusky banded pigmentations. The following key,

therefore, may be somewhat inadequate for individuals of extreme

sizes where only colour characters are utilised in key couplets.

Key to species of the genus Choerodon

1. Dorsal fin rays XIII, 7.2

Dorsal fin rays XII, 8.16

2. Anal fin rays III, 9; vertebrae 11 -i- 16; a broad pale saddle

present on caudal peduncle, extending anteriorly to below

posterior end of dorsal fin base; a pale wedge-shaped band

present midlaterally on side below about 5th dorsal fin spine;

head dusky above level of mouth, covered by tiny pale spots

(orange in life). (north-eastern

Indian Ocean and western Pacific from southern Japan to

north-eastern Australia and New Caledonia) anchorago

Anal fin rays III, 10; vertebrae 10 -i- 17; colour pattern not

as above.3

M.F. Gomon

3. Predorsal scales reaching forward beyond point above

posterior extent of orbit on dorsal midline of head, scales

lateral to midline reaching to above centre of orbit.4

Predorsal scales not reaching, or barely reaching, forward

to above posterior extent of orbit on dorsal midline of

head, scales lateral to midline not reaching much forward

of point above posterior extent of orbit.5

4. Second prominent anterior canine distinctly curved

posterolaterally; body of moderate depth at dorsal fin origin,

11-1.9 into SL; caudal fin slightly forked; body with a

prominent dusky stripe on lateral midline; prominent dusky

spot present midlaterally at posterior end of caudal peduncle;

no dark spot on dorsal fin in adults.

.(northern Australia, south-eastern Indonesia) vitta

Second prominent anterior canine mostly straight, only

slightly curved in large specimens, angled dorsolaterally

and sometimes slightly posteriorly; body deep at dorsal fin

origin, 1.3-1.6 into SL; caudal fin truncate to slightly

rounded; body mostly pale with faint broad dusky bands,

especially dorsally; prominent ocellated dark spot present

on dorsal fin between last few spines.

.(northern Australia, south-eastern Indonesia) monostigma

5. Scales on subopercle reaching to or nearly to below anterior

extent of ventral preopercular edge; predorsal scales

reaching nearly to above posterior extent of orbit on dorsal

midline of head (somewhat short of this in very large

specimens); body with an anteroventrally angled narrow

dusky wedge extending from base of 10th or 11th dorsal fin

spine to posterior side of pectoral fin base; body anterior to

wedge in adults somewhat dusky, paler posteriorly; juveniles

somewhat mottled, but with a faint indication of above

pattern and an ocellated dark spot posteriorly on dorsal fin...

.(south-eastern Asia from Japan to Vietnam) azurio

Scales on subopercle not reaching near anterior end of

ventral preopercular edge, though extending forward to

below midpoint of ventral edge in some species; predorsal

scales rarely reaching forward near point above posterior

extent of orbit on dorsal midline of head; body not

pigmented, as described above.6

6. Body with a prominent anteroventrally directed dark

band (occasionally restvricted to a large black spot

dorsally on body extending onto ventral portion of dorsal

fin between last spine and about fourth segmented ray)

associated with a single prominent pale band or pale spot;

pectoral fin rays ii, 13.7

Body with or without dark or pale marks, but not as above;

pectoral fin rays ii, 13-17 (ii, 13 in 2 of 11 species).8

7. Large pale spot situated below posterior end of dorsal fin

base, bordered anteriorly by a broad dark band tapering

anteroventrally towards pectoral fin base (band extending

as dusky segment around ventral and posterior side of pale

spot); dorsal edge of caudal fin dark in adults.

.(Japan, Taiwan, southern

China; northern Australia, New Caledonia and Fiji) jordani

Broad pale band directed from below base of last few dorsal

spines to upper side of pectoral fin base; large dark spot

situated dorsally on side and on ventral edge of dorsal fin at

dorsoposterior end of pale band, usually extending

anteroventrally as a dark tapering marginal band on pale

band at least along posteroventral edge; caudal fin uniformly

pale in adults.

.(Indonesia, northern New Guinea) zosterophorus

8. Subopercle with 9-11 scales in an anteriorly tapering row;

caudal fin slightly forked; pectoral fin narrowly pointed at

dorsoposterior tip in large specimens; body pale without

distinct pale or dusky to dark markings, only a small

dusky spot on each body scale and several dusky marks

on head in freshly preserved specimens.

.(north-eastern Australia) venustus

Subopercle with 1-8 scales in an anteriorly tapering row;

caudal fin rounded, truncate or slightly double emarginate;

pectoral fin broadly pointed or rounded at dorsoposterior

tip; body variously pigmented.9

9. Pores on dorsal side of head not numerous, confined to

distinct lines corresponding to major cephalic sensory

canals.10

Pores on dorsal side of head extremely numerous, so dense

as to obscure position of major cephalic sensory canals.11

10. Body with about 5-7 complete broad dusky bands in

juveniles and adults; distinct dark dusky spot centred on

lateral line along dusky band below 5th or 6th dorsal fin

spine; adults with second prominent anterior canine

approximately half the size of the first, directed mostly

dorsally.

.(north-eastern Australia, New Caledonia) graphicus

Body with about 5 very faint broad dusky bands dorsally on

side in most freshly preserved specimens; large pale spot

often present below posterior end of caudal fin of adults; no

dark dusky spot present on side below 5th or 6th dorsal fin

spine in individuals of any size; adults with second

prominent anterior canine approximately the same size as

first, second canine slanted dorsolaterally.

.(northern Australia) cyanodus

11. Subopercle with 6-10 scales, scales extending anteriorly

at least to below midpoint on ventral preopercular edge;

pectoral fin rays ii, 13-15 (rarely 15).12

Subopercle with 1-5 scales, scales distinctly not extending

anteriorly to below midpoint on ventral preopercular

edge; pectoral fin rays ii, 15-17 (rarely 15).13

12. Second prominent anterior canine in lower jaw distinctly

larger than first, directed strongly laterally and curved

posterolaterally; dorsal outline of head in adults strongly

curved above eyes in lateral aspect, nape nearly horizontal to

just behind eyes; pectoral fin rays ii, 13-14 (rarely 14);

pigmentation in alcohol uniformly pale (colour in life rosy,

though somewhat browner above, a blue spot present on

each body scale).(north-eastern Australia) Jrenatus

Review of Choerodon tuskfishes

Second prominent anterior canine in lower jaw approximately

equal in size to first, directed mostly dorsally and slightly

anterolaterally; dorsal outline of head with gentle convex

curve from origin of dorsal fin to tip of snout in lateral aspect;

pectoral hn rays ii, 14—15 (rarely 15); pigmentation in alcohol

pale to slightly dusky, often with a distinct dusky spot on side

immediately below posterior end of dorsal fin base; an

elongated pale spot often present just above lateral line under

anterior half of dorsal fin (colour in life orange with light

grey longitudinal stripes, a large bright yellow oval spot on

side above lateral line and below anterior half of dorsal hn.

.(south-eastern Asia, Indonesia, Philippines) oligacanthus

13. Body rather uniformly dusky without distinct dusky to

dark markings; underside of mouth pale; pectoral hn pale;

head covered with extremely numerous hne pale dots

(evident only on close inspection); forehead and snout not

crossed by alternating pale and dusky stripes.

.(south-western Australia) rubescens

Body pale to slightly dusky with a dark spot or dusky

blotch on side, or head covered with many small pale

spots and forehead crossed by 4 or 5 sets of alternating

dusky and pale stripes.14

14. Pectoral hn rays ii, 14-15; a prominent dark spot or

anteroventrally angled dark slash on side, located mostly

above lateral line under about 6th dorsal hn spine.

.(north-western Australia) cauteroma

Pectoral hn rays ii, 16-17; no prominent dark spots or

slash on side below 6th dorsal hn spine.15

15. Moderately small but prominent dark spot present on body at

base of last dorsal hn spine (spot not extending onto hn

membrane); body scales (each with a blue centre in life)

forming horizontal rows of spots on side; second prominent

anterior canine in lower jaw distinctly shorter than hrst in

adults, directed mostly dorsally and slightly anteriorly; 3-5

scales on subopercle.(western

Pacihc, China and Japan to northern Australia) schoenleinii

No small prominent black spot on side; horizontally

elongated dusky blotch present below centre of hn in some

specimens; head with many small pale spots on cheek

(orange in life), forehead crossed by 4 or 5 sets of alternating

pale and dusky stripes (blue and orange in life); body scales

edged with blue giving sides uneven banded appearance;

second prominent anterior canine in lower jaw distinctly

smaller than hrst in adults, directed dorsolaterally and

curved slightly posteriorly; 1-3 scales on subopercle.

.(northern Australia) cephalotes

16. Posterior edge of pectoral hn not produced ventrally

(ventral-most rays shorter than those immediately above);

scales ventrally on cheek extending forward to or nearly

to corner of mouth..17

Posterior edge of pectoral hn produced ventrally (ventral-

most rays distinctly longer than those immediately above);

scales ventrally on cheek distinctly not extending forward

to corner of mouth..21

17. Body shallow, 24.6-33.7% SL; snout short 8.3-9.8% SL;

subopercle covered by about 5 large scales forward to

about anterior end of ventral preopercular margin.

.(north-eastern Indian Ocean;

western Pacihc, China to north-eastern Australia) typus

Body deep, 36.1-44.0% SL; snout long 12.3-16.7% SL;

much of subopercle naked with 1-3 rows of about 7-10

small scales adjacent preopercular edge extending forward

nearly to anterior end of ventral preopercular margin..18

18. Pectoral hn ii, 13; predorsal scales approximately 10-14,

reaching forward in advance of above posterior extent of

orbit on dorsal midline of head; body with about 7 or 8 pairs

of alternating prominent pale and dusky to dark bands

(bands red, blue, white and black in life).(Japan, Taiwan,

Philippines, eastern Australia, New Caledonia) fasciatus

Pectoral hn ii, 14; predorsal scales approximately 5-8,

scales often reaching forward near, but not in advance of

above posterior extent of orbit on dorsal midline of head;

body without bands or with a single broad anteroventrally

slanted dark band.19

19. Caudal hn emarginate, comers distinctly produced, posterior

margin of hn concave..

.(Saya de Malha Bank, Indian Ocean) cypselurus sp. nov.

Caudal hn truncate, corners not or very little produced,

posterior margin of hn straight..20

20. Line of demarcation between anterodorsal dusky portion

of body and posteroventral pale portion extending

between posterior end of dorsal hn base and upper side of

pectoral hn base, line slightly darker than anterodorsal

dusky pigmentation; caudal hn dusky with hne pale and

dusky vermiculations posteriorly and narrow dark dusky

dorsal and ventral margins at corners; colour in life brown

anterodorsally, white posteroventrally; each body scale

with a blue centre in small individuals, forming blue

stripes posteriorly on body in larger specimens.

.(Indian Ocean, Japan, Indonesia) robustus

Line of demarcation between anterodorsal dusky portion

of body and posteroventral pale portion extending

between base of last dorsal hn spine and upper side of

pectoral hn base, line not darker than anterodorsal dusky

pigmentation; caudal hn pale to slightly dusky with darker

periphery; broad anteroventrally tapering reddish brown

stripe covering dorsal half of side posterior to greenish

brown anterodorsal portion of body in life; reddish brown

area edged ventrally by yellow stripe; ventral portion of

side white.(Japan,

Philippines, Indonesia, western Australia) zamboangae

21. Scales dorsally on cheek extending forward to or nearly to

below anterior extent of orbit; a short dark slash present on

side immediately below lateral hne under about 3rd dorsal

hn spine; body otherwise pale..

.(northern Australia) sugillatum

M.F. Gomon

Scales dorsally on cheek not extending forward to below

centre of eye; dark slash, if present, on side, below last few

dorsal fin spines.22

22. Scales present only on dorsal part of cheek (dorsal part of

space between ventral edge of orbit and ventral edge of

preopercle); body often pale in preservative (in life with a

purplish blue stripe directed dorsoposteriorly from posterior

tip of opercular flap to lateral line below last dorsal fin

spine); often a second parallel stripe or series of spots present

on ventral side of first originating on posterior side of

pectoral fin base, consisting of a series of dots on caudal

peduncle); no stripe or horizontal series of spots directed

posteriorly from centre of eye and extending well onto side..

.(Arabian Gulf to Zanzibar) gymnogenys

Scales extending to or below centre of cheek; body mostly

pale in preservative, often with dusky or pale stripes or a

horizontal series of spots, but not as described above; one

stripe or series of spots often emanating from midpoint on

posterior edge of orbit and extending well onto side.23

23. Caudal peduncle moderately slender, least depth 8.4-8.5

into SL; width of bony interorbital 16.3-19.6% head length;

body pale in preservative with only a single dusky stripe

directed horizontally from lateral line below about 8th

dorsal fin spine to point immediately below posterior end of

dorsal fin base.

.(Somalia coast, Indian Ocean) skaiopygmaeus sp. nov.

Caudal peduncle slender to very slender, least depth 8.6-10.9

into SL; width of bony interorbital 19.9-23.9% head length;

body usually with one or more continuous or broken pale

stripes or horizontal series of spots, sometimes with

additional dusky stripes (not as described above).24

24. Caudal peduncle very slender, 9.4—10.9 into SL; head with

narrow pale stripe directed horizontally forward from lower

edge of eye extending around tip of snout adjoining stripe

from opposite side; side of body with blue to white solid or

broken midlateral stripe extending onto base of tail.25

Caudal peduncle slender, 8.6-9.4 into SL; head with pale

stripe curving from lower edge of eye to upper jaw near

posterior end of mouth, sometimes obscured by overall

white colouration below eye; side of body with blue to white

continuous or broken stripe angled from above pectoral fin

base to dorsal half of caudal peduncle just above lateral

midline, stripe distinctly broader anteriorly.26

25. Eye of moderate size, 9.1-9.8% SL, 23.1-25.0% HL;

diameter of orbit 85.1-104% snout length; terminal phase

individuals with large square dusky (black when fresh) spot

just under lateral line below last few dorsal fin spines.

.(eastern Australia, Coral Sea, New Caledonia) gomoni

Eye moderately small, 8.3-8.6% SL, 21.6-23.4% HL;

diameter of orbit 71.4—83.1% snout length; terminal phase

individuals with large square dusky (reddish brown when

fresh) spot mostly above lateral line below last few dorsal fin

spines.(Taiwan, Philippines, Indonesia) margaritiferus

26. Cheek scales reaching forward to about anterior end of

ventral preopercular margin; single curved white to blue

stripe on side of head from eye to upper jaw.(Taiwan,

Japan, Indonesia, western Australia) albofasciatus nom. nov.

Cheek scales reaching forward only to below centre of eye;

2 white to blue curved stripes below curved stripe on side of

head from eye to upper jaw.

.(Norfolk Ridge, south-western Pacific) aurulentus sp. nov.

Choerodon (Choerodon)

Tables 2 & 3

ChoerodonBleeker, 1847; 10, type species - Labrus macrodontus

Lacepede (= C. anchorago) by monotypy (published as nomen nudum

in Bleeker, 1845: 513).

Choerops Ruppell, 1852: 20, type species - Choerops meleagris

Riippell (= C. anchorago) by monotypy.

Cossyphodes Bleeker, 1858b: 408, type species - Labrus

macrodontus Lacepede (= C. anchorago) by monotypy.

Hypsigenys Gunther, 1861; 383, type species - Labrus

macrodontus Lacepede (= C. anchorago) by monotypy.

Torresia Castelnau, 1875: 36, type species - Torresia austialis

Castelnau (= C. schoenleinii) by monotypy.

Choerodon (Macrochoerodon) Fowler & Bean, 1928; 200, type

species - Crenilabrus oligacanthus Bleeker, 1851 by monotypy.

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 9 or 10;

pectoral fin rays ii, 13-16, rarely 17, dorsalmost ray of

moderate length 24.9-53.9% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular to broadly rounded; body moderately deep,

31.4-47.3% SL, head depth 23.7-41.0% SL, caudal peduncle

depth 11.8-17.8% SL; head blunt, dorsal profile of snout

moderately steep, snout length 9.0-18.6% SL; predorsal scales

approximately 4-9, reaching forward on dorsal midline, not

quite to above posterior edge of preopercle, to about midpoint

between posterior extent of orbit and posterior edge of

preopercle, or somewhere between; cheek with small partially

embedded scales in about 1-10 diagonal rows, posteriormost

with about 4-15 scales to upper extent of free preopercular

edge, reaching forward to or almost to corner of upper lip

crease above mouth, with broad naked margin posteriorly and

ventrally on preopercle; 1-3 rows (only about 2 scales in

second and third rows when present) of small to moderately

large scales on subopercle adjacent preopercular edge

extending forward to about anterior end of ventral preopercular

margin, confined to upper end of preopercular surface, or

reaching somewhere between, with about 1-8 scales in

outermost row; each lateral line scale with triple to multiple

branching laterosensory canal tube; scales above lateral line

about 21 / 2 - 41 / 2 ; cephalic sensory canal pores relatively few

confined to lines or short branches associated with major

canals or numerous; dorsal and anal fins with very low basal

sheath comprising 1-5 progressively smaller accessory scales

at deepest; posterior lobe of dorsal and anal fins not reaching

or reaching just beyond hypural crease; second pair of canines

in lower jaw directed anterodorsally and little to strongly

laterally, greatly curved in some; caudal fin truncate to

Table 2. Ranges for selected counts and proportional measurements in subgenera of Choerodon. “N” designates number of counts or measurements. Aberrant values enclosed by

parentheses.

Review of Choerodon tuskfishes

1^1

-S

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s ^

.-H*'

ii ^

ffi

.-H

(

S 1

5 ^

mean

28.7

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5ji cn

’-H

F VO

ffi

,-H

ffi

,-H

-S

ffi

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« 00

ffi

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ffi

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ffi

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II, 10

I ^

ffi

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ffi

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144

146

138

105

124

125

133

130

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£ ^

t/5

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tZJ

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t/5

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Dorsal fin

Anal fin

Pectoral fin

Vertebrae

% SL

Body depth

■cS

Head length

Head depth

Orbit diamete

■s

Snout length

J=>

-2

VC^

cfl

J=l

”2

M.F. Gomon

ii fi

■g ^

H Qh

Peaolopesia [10]

16.1-69.1

range mean N

51.5-88.5 58.1 93

17.3- 29.8 19.2 116

15.3- 30.7 19.3 92

23.1-38.6 27.1 93

2.8-18.3 7.5 85

46.1-105 61.5 93

Xiphocheilus [1]

23.5-41.8

range mean N

67.8-80.2 73.6 12

20.2- 25.9 23.8 11

15.3- 20.3 18.5 14

24.7-30.0 26.6 11

24.6-33.8 27.6 10

70.3-100 89.3 10

Lutjanilabrus [1]

23.4-61.6

range mean N

60.7-88.4 75.2 13

18.4- 27.1 22.4 12

18.4- 22.8 21.1 10

29.9-43.1 34.5 12

25.5- 41.7 32.9 13

42.6- 90.6 66.1 12

Lienardella [1]

34.2-62.3

range mean N

79.5-89.7 84.2 14

21.2-27.2 23.9 15

18.9- 25.6 23.1 15

30.9- 43.6 36.1 15

21.3-52.6 31.6 13

50.0-79.8 67.3 13

Aspiurochilus [5]

21.5-115

range mean N

72.3- 114 78.1 75

16.3- 29.9 18.7 68

13.0-31.2 19.6 56

26.3- 46.8 31.8 57

23.9-45.2 28.0 54

37.5-114 46.1 57

Choerodon [9]

21.2-152

range mean N

67.9- 116 76.3 131

12.9- 29.4 15.6 137

11.1-30.1 17.0 92

25.5-61.2 30.7 112

24.9 - 53.9 33.0 95

28.5-95.4 35.2 110

Subgenus [number of species]

MORPHOMETRIC FEATURES

Head length (mm)

%HL

Head depth

Orbit diameter

Interorbital width

Snout length

X 100%

1st pectoral fin ray/pectoral fin length

Orbit diameter/Snout length

Table 3. Ranges for selected counts and proportional measurements in species of the subgenus Choerodon. “N” designates number of counts or measurements. Aberrant values enclosed

by parentheses.

Review of Choerodon tuskfishes

Tj-

r<p

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[ORPHOMETRIC

EATURES

tandard length

nm)

(Zl

S-

audal peduncle

;pth

-o

rbit diameter

iterorbital width

orsal fin base length

orsal fin last spine

ngth

orsal fin posterior

ibe length

nal fin base length

nal fin posterior

ibe length

sctoral fin length

>t pectoral fin ray

ngth

slvic fin length

audal fin dorsal lobe

ngth

audal fin central ray

ngth

(/3

S fa

2 fa

(Zl *

u-e

Q B

Q.2

<C.2

^ B

U ii

M.F. Gomon

C. venustus

25.5-126

range mean N

67.9- 102 87.9 12

16.6-28.1 24.0 14

18.9- 30.1 21.4 11

31.0-51.1 40.8 13

24.9 - 46.8 34.6 11

32.5-82.3 60.4 13

C. schoenleinii

21.7-152

range mean N

77.0-116 96.7 25

14.7- 29.4 20.8 27

20.4-28.7 24.1 10

30.7- 46.2 38.8 14

41.5- 53.9 48.0 10

32.4-95.4 60.8 13

C. rubescens

24.3-140

range mean N

85.5- 98.2 93.4 9

14.4-26.4 20.6 9

20.2- 25.7 23.3 9

34.7- 49.6 44.0 9

26.8-50.3 39.6 9

31.3-76.0 48.6 9

C. oligacanthus

30.0-86.3

range mean N

72.6-96.8 82.4 13

12.9-23.3 19.1 12

15.0-19.7 16.1 10

30.0-46.3 38.0 10

26.1- 42.8 36.4 10

37.2- 76.3 55.6 10

C. graphicus

34.7-136

range mean N

77.7-94.9 83.1 11

14.5-23.9 19.8 12

19.9-24.0 22.0 9

34.1-43.0 38.1 10

35.4-47.9 43.9 9

34.2-69.2 55.8 9

C. cyanodus

23.1-93.9

range mean N

72.1- 107 82.7 14

15.7-24.8 20.2 18

15.4-19.7 17.2 11

26.2- 49.0 40.2 16

29.9- 44.3 38.6 12

35.9- 69.5 52.1 16

C. cephalotes

21.2-89.2

range mean N

77.9-103 91.5 15

15.4- 27.9 22.2 16

14.4- 23.0 19.3 9

35.3-61.2 44.5 15

36.4- 53.1 41.4 13

28.5- 74.5 52.9 15

C. cauteroma

30.3-67.2

range mean N

82.0-97.7 87.7 10

22.8-28.2 25.2 10

17.4-23.3 19.7 10

29.1-44.0 36.2 10

41.3 - 52.4 45.6 10

55.3-88.2 70.8 10

C. anchorago

24.6-94.4

range mean N

73.8-102 89.6 22

13.1- 28.8 19.2 19

11.1- 23.4 19.4 13

25.5-52.4 39.1 15

35.0-44.9 39.9 11

29.3 - 90.9 53.5 15

Species

MORPHOMETRIC

FEATURES

Head length (mm)

%HL

Head depth

Orbit diameter

Interorbital width

Snout length

X 100%

1st pectoral fin ray/

pectoral fin length

Orbit diameter/Snout

length

Review of Choerodon tuskfishes

distinctly rounded or with concave posterior profile. (See

Table 1 for additional meristic and morphometric ranges.)

Usually separate juvenile, initial phase and terminal phase

colour patterns.

Reaches moderately large maximum size, one or more

species reported to attain a metre SL.

Comments. As the second largest subgenus of Choerodon, this

complex comprises nine species, most of which reach a larger

size than those of other subgenera. Two are widely distributed in

the western Pacific, one also occurring in the north-eastern Indian

Ocean. The evolutionary hypothesis presented by Puckridge et

al. (2015) implies clades within the major (subgeneric) clades,

the largest within this subgenus comprising at least four species,

C. cephalotes, C. cauteroma, C. rubescens and C. schoenleinii

(fig. 1, clade 3). Morphological support for this clade includes a

reduction in subopercular squamation and the highest number of

branched pectoral fin rays, regularly 15-17. A species of

Choerodon not included in the genetic analysis, C. graphicus, is

of similar body form to others in the subgenus, shares the reduced

subopercular squamation and is likely to be a member of this

clade. A second clade within the subgenus comprises

C. anchorago, C. cyanodus and Choerodon oligocanthus.

The four earliest group names proposed for the subgenus

were all based on the type species, C. anchorago, albeit in

reference to two of its junior synonyms. Early confusion

undoubtedly arose from limited communication during the 14

years between the publication of the first and last of the four

names, although a justification for Bleeker’s second name

Cossyphodes was not provided (Gill, 1908; Gomon, 1997).

Castelnau (1875) was certainly familiar with the genus

Choerodon because he treated three species assigned to the

genus (as Chaerops) in the publication in which he presented

his new Torresia, but probably failed to make the generic link

because of the juvenile form of his type specimen. Torresia

australis appears to be a junior synonym of C. schoenleinii.

Macrochoerodon was proposed as a subgenus by Fowler and

Bean (1928) to distinguish C. oligacanthus from other species

of Choerodon by its elongate pelvic fins, which extend past the

anal fin origin in adults. In all other regards, the species is

consistent with those of the subgenus.

Choerodon anchorago (Bloch, 1791)

Anchor Tuskfish

Figures 2, 3; table 3; appendix.

Sparus anchorago Bloch, 1791: 108, pi. 276, locality unknown.

Labrus macrodontus Lacepede, 1802: 451,522, locality unknown,

specimen sent to France from Holland.

Cossyphus macrodon Bleeker, 1847: 10, emendation of Labrus

macrodontus Lacepede.

Choerops meleagris Riippell, 1852: 20, Mare javanicum (Java

Sea).

Crenilabrus leucozona Bleeker, 1858a: 20, Biliton (Indonesia).

Choerops Maeander Cartier, 1874: 102, Cebu (Philippines).

Choerodon weberi Ogliby, 1911: 52, (listed as Chaerodon weberi

on p. 36), Dobo, Aru Islands (Indonesia).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 9; pectoral fin

rays ii, 13, rarely 14, dorsalmost ray of moderate length 35.0-

44.9% pectoral fin length, ventralmost rays shorter than those

above, posterior edge of fin obliquely straight, dorsoposterior

corner bluntly pointed, posteroventral corner angular; body

deep, 37.0-45.1% SL, head depth 28.5-37.9% SL, caudal

peduncle depth 12.3-17.3% SL; head bluntly pointed, dorsal

profile of snout moderately steep, snout length 12.0-18.3% SL;

predorsal scales approximately 7-9, reaching forward on dorsal

midline to about midpoint between posterior extent of orbit and

posterior edge of preopercle; cheek with small partially

embedded scales in about 7-10 diagonal rows, posteriormost

with about 9-15 scales to upper extent of free preopercular edge,

reaching forward to or almost to corner of upper lip crease above

mouth, with broad naked margin posteriorly and ventrally on

preopercle; 1 or 2 rows (only about 2 scales in second row when

present) of small scales on subopercle adjacent preopercular

edge extending forward to about anterior end of ventral

preopercular margin, with about 8 scales in outermost row; each

lateral line scale with multiple branching laterosensory canal

tube; scales above lateral line about 314; cephalic sensory canal

pores relatively few confined to lines or short branches associated

with major canals; second pair of canines in lower jaw directed

anterodorsally and slightly laterally; dorsal and anal fins with

very low basal sheath comprising 2 or 3 progressively smaller

accessory scales at deepest; posterior lobe of dorsal and anal fins

reaching to or beyond hypural crease; caudal fin truncate, corners

rounded to square; pelvic fin reaching to or just short of anus,

length 21.7-28.0% SL. (See Table 3 for additional meristic and

morphometric ranges.) Adults with upper third of body and head

above mouth dark grey, white below, with dorsally tapering

vertical white or yellow band below central dorsal fin spines and

white saddle-like blotch covering top of caudal pecuncle forward

to bases of middle segmented dorsal fin rays; head peppered with

fine orange spots in life.

Reaches moderately large maximum size, largest specimen

examined 273 mm SL.

Pigmentation in alcohol. Juveniles dark dusky with 5 vertical

narrow pale bands crossing side and 2 horizontal narrow pale

stripes extending from behind head to dorsal and ventral sides

of caudal peduncle, anterior two pale bands crossing nape;

head similarly dusky dark with pale dorsal margin to eye, mid¬

dorsal pale stripe covering top of snout to above eye, horizontal

pale stripe along underside of eye and pale marks on ventral

surface below eye, low on operculum and on chest; small dark

spots on dorsal and ventral profiles midway along and at

posterior corners of caudal peduncle and at centre of caudal fin

base, on leading edge, midway along and at posterior base of

dorsal fin, on leading edge and at posterior base of anal fin, and

extending from pectoral fin base along leading edge of pelvic

fin. Larger juveniles dusky with pale underside and narrow

remnants of 5 pale bands, 3rd band just posterior to pectoral fin

and 5th band across anterior end of caudal peduncle most

prominent; base of pectoral fin with large dark spot. Initial

phase adults dusky above and pale below upper end of pectoral

fin base, distinct demarcation on head at horizontal from corner

of mouth; side with pale underside extending upward as

M.F. Gomon

Figure 2. Choerodon anchorago. A, Juvenile, Amami Oshima Island, Ryukyu Islands, Japan, photo by S. Sato; B, Initial phase adult, BPBM

15809, 123 mm SL, Madang, New Guinea Island, Papua New Guinea, photo by J. Randall, BPBM; C, Terminal phase adult, Greek aquarium,

photo by Dr G. J. Reclos.

Review of Choerodon tuskfishes

0 Choerodon anchorago (examined)

O Choerodon anchorago (record)

D Choerodon cauteroma (examined)

D Choerodon cauteroma (record)

20 °

0 °

20 °

Figure 3. Distributions of Choerodon anchorago (circles) and C. cauteroma (squares) based on specimens examined (coloured) and collection

registration records (white).

dorsally tapering wedge-shaped mark behind pectoral fin with

apex reaching lateral line; caudal peduncle pale except for

midlateral extension of dusky pigmentation reaching to or

almost to caudal fin base; dusky side of head speckled with fine

pale spots; pectoral fin base covered by prominent dark spot;

dorsal fin dusky above dusky area on side; remainder of dorsal

fin and other fins pale. Terminal phase adults similar to initial

phase but with pale area on side covering ventral 2/3 of side

between front of pale wedge and vertical at middle segmented

dorsal fin rays; dusky and pale areas on head and anterior end

of side separated by distinctly dark margin.

Fresh colours. Juveniles olive brown to brown with 5 narrow

white bands, first two crossing predorsal, third below 4th or 5th

dorsal fin spines, 4th below last couple of dorsal fin spines and

5th at posterior end of dorsal fin base (fig. 2A); 2 narrow white

stripes originating above and below pectoral fin base and

terminating on base of caudal fin; head brown dorsally with

broad white stripe under eye and narrow white stripe above eye.

Dorsal, anal and pelvic fins same colour as side adjacent with

continuation of white bands on side, white ocellated black spot

near middle of segmented ray portion of dorsal and anal fins and

dorsally and ventrally on caudal fin base, ocelli with red

surround in small individuals; caudal and pectoral fins hyaline

(Okamura & Amaoka, 1997: 465, left second from top; Chen et

al, 2010: 380 & 381, figs D & E; Kuiter, 2010: 51, fig. D; Allen

& Erdmann, 2012: 645, left side fig. in middle of page).

Initial phase adults purplish grey dorsally with ventral half

white (fig. 2B); prominent white wedge-shaped bar on side just

posterior to pectoral fin and white rectangular blotch covering

dorsal half of caudal peduncle and side below last 2 dorsal fin

rays; pectoral fin base with black-orange-black ocellated spot;

head with numerous fine orange spots, orange horizontal line

from corner of mouth, several orange lines on chin. Dorsal fin

purplish grey with 3 horizontal orange stripes anteriorly, one

submarginal posteriorly, yellowish white basoposteriorly.

Anal fin yellow with light purple margin and orange

submarginal stripe. Caudal and pectoral fins yellowish orange.

M.F. Gomon

Pelvic fin white with lengthwise stripe just posterior to leading

edge (Okamura & Amaoka, 1997: 465, left bottom; Chen et

al, 2010: 381, fig. F; Kuiter, 2010: 51, figs B & F; Allen &

Erdmann, 2012: 645).

Terminal phase adults blacker posterodorsally (fig. 2C);

wedge-shaped bar orange with broad orange streak extending

posteriorly on lateral midline; pectoral fin base covered with

large black spot preceded by orange marks; very broad black

stripe on upper lip to opercular edge. Dorsal fin black anteriorly

on soft portion, white posteriorly with narrow orange lines. Anal

fin white with several narrow orange longitudinal lines breaking

up into spots posteriorly. Caudal fin black, orange somewhat

distally with narrow blue marginal line. Pelvic fin with 1 or 2

additional orange lines (Shen, 1993: pi. 143, fig. 10; Okamura &

Amaoka, 1997: 465, left second from bottom; Shibukawa,

Peristiwady & Suharti, in Kimura & Matsuura, 2003: 147;

Kuiter, 2010: 51, figs C & G; White etal, 2013: 265, fig. 89.13).

Etymology. The name anchorago may refer to an anchor-like

colour pattern, as perceived by its author.

Distribution. One of the most widely distributed species in the

genus (fig. 3), recorded from Sri Lanka (Munro 1955:185),

Nicobars and Andaman Islands in the north-eastern Indian

Ocean, throughout the tropical western Pacific from

southernmost Japan and the Ogasawara Islands (Randall et al.,

1997: 45) to the northern Great Barrier Reef in north-eastern

Australia, eastward to Palau and Yap in Micronesia (Myers,

1999: 189), Solomons, Vanuatu and New Caledonia (Fricke and

Kulbicki, 2006: 386). Occurs in coastal seagrass and sandy

areas with mixed coral and rubble to depths of about 25 m.

Comments. The specimens on which Bloch (1791) and Lacepede

(1802) based their descriptions of Spams anchorago and

Labrus macrodontus, respectively, were acquired from the

Netherlands, presumably from auctions, and lacked provenance.

Although Bloch’s accompanying figure (1791, pi. 276) did not

accurately portray the distinctive colour pattern of this species,

the morphological description, including dorsal, anal and

pectoral fin ray counts, is diagnostic for this species. Paepke

(1999: 92) identified two specimens in the Berlin Zoological

Museum collection as types, one of which has apparently been

lost, and designated the extant specimen (ZMB 2476) as

lectotype. The type of Lacepede’s description is likewise

identifiable as this species by the same characters, as is the type

specimen (MNHN A.8208). Bleeker (1849) emended

Lacepede’s name to Cossyphus macrodon without justification

and based his detailed description on three specimens from

Batavia (Java, Indonesia).

Choerops meleagris, the type species of Rtippell’s (1852)

new genus, was presented only with a description of

morphological features, which are common to species of the

genus. The Senckenberg Museum type (SMF 2759, 263 mm

SL) is a dried specimen clearly identifiable as C. anchorago.

The type of Bleeker’s (1858) Crenilabrus leucozona, now in

the British Museum (BMNH 1864.5.15.18), is a 34.3 mm SL

(42 mm TL) specimen with the characteristic meristic values

and juvenile colouration of C. anchorago. Choerops Maeander

Cartier, 1874 was based on juveniles according to the lengths

of his type series 3.9-6.7 cm from Cebu in the Philippines.

Although the types were not located, Cartier’s account

includes a detailed description that matches the juvenile

colouration of C. anchorago, and his anal fin count of “3/9”

(III, 9) is unique for that species within the genus. Ogilby’s C.

weberi was based on seven specimens, 121-197 mm, from

Dobo, Aru Islands, three of which (QMB 1.20, 1.1532, and

1.10133) are still present in the Queensland Museum collection.

The species is synonymous with C. anchorago.

Choerdon anchorago is the most often encountered

member of the genus, being found in shallow waters of the

central western Pacific, and one of the few ranging westward

in the northern part of the Indian Ocean as far as India and Sri

Lanka. Its nine segmented dorsal fin rays and vertebral count

of 11 -I- 16 are distinctive for the subgenus and in combination

for the family.

Material examined. 163 specimens, 12-273 mm SL; see appendix.

Choerodon cauteroma Gomon & Allen, 1987

Bluespotted Tuskfish

Figures 3, 4; table 3; appendix.

Choerodon sp. 1 Gloerfelt-Tarp and Kailola, 1983: 235, colour

figure on opposing page; Sainsbury & Kailola, 1984: 260, colour

figure on opposing page.

Choerodon sp. Allen, 1985: 2406, figs 330 & 331.

Choerodon cauteroma Gomon and Allen, 1987: 25. Western

Australia, Exmouth Gulf.

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 15, rarely 16, dorsalmost ray of moderate length

41.3-52.4% pectoral fin length, ventralmost rays shorter than

those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body deep, 36.3-44.0% SL, head depth 27.7-33.7%

SL, caudal peduncle depth 14.0-17.0% SL; head blunt, dorsal

profile of snout steep, snout length 9.9-15.7% SL; predorsal

scales approximately 5-8, reaching forward on dorsal midline

to above posterior edge of preopercle; cheek with small

partially embedded scales in about 4 or 5 diagonal rows,

posteriormost with about 9 or 10 scales to upper extent of free

preopercular edge, reaching forward to corner of upper lip

crease above mouth, with very broad naked margin posteriorly

and ventrally on preopercle; single row of about 5 small scales

on subopercle adjacent preopercular edge extending forward

about half way to anterior end of ventral preopercular margin;

each lateral line scale with multiple branching laterosensory

canal tube; scales above lateral line about 4; cephalic sensory

canal pores numerous confined to lines or short branches

associated with major canals; second pair of canines in lower

jaw strongly curved laterally; dorsal and anal fins with very low

basal sheath comprising 1-5 progressively smaller accessory

scales at deepest; posterior lobe of dorsal and anal fins reaching

hypural crease; caudal fin truncate to slightly rounded medially,

upper and lower corners slightly produced; pelvic fin reaching

to or just short of anus, length 21.9-24.1% SL. (See Table 3 for

additional meristic and morphometric ranges.) Tan to green

above with white underside and prominent black slash or spot

Review of Choerodon tuskfishes

Figure 4. Choerodon cauteroma. A, Juvenile, WAM P. 30087-009, 44 mm SL, Useless Loop, Shark Bay, Western Australia, 26° 09’ S, 113° 26’

E, photo by J. Hutchins, WAM; B, Initial phase adult, Mary Anne Island sanctuary. Shark Bay, Western Australia, photo by J. Shuttleworth;

C, Terminal phase adult, approximately 300 mm SL, off Gnaraloo Station, Western Australia, 23.44° S, 113.27° E, photo by L. Malton.

M.F. Gomon

between base of middle dorsal fin spines and lateral line; head

with black streak directed posteroventrally from eye and

second running posteriorly from above rear of mouth;

additional blue lines radiating from eye and blue spot or vertical

mark on most body scales, those posteriorly coalescing into

horizontal lines; prominent black spot at front of dorsal fin.

Reaches moderately large maximum size, largest specimen

examined 259 mm SL.

Pigmentation in alcohol. Juveniles pale with 4 faint dusky

bands or vertically elongate blotches on side; darker spot about

half diameter of eye on lateral line below 7th dorsal fin spine;

fins pale, dusky blotches basally on dorsal and anal fins opposite

ends of bands crossing side and dark spot in dorsal fin above 3rd

band in very small individuals. Initial phase adults pale with

faint dusky stripes posteriorly on side following horizontal scale

rows; broad dark slash on side between base of 7th dorsal fin

spine and lateral line; head with pair of narrow dusky lines

crossing snout in front of eyes, similar pair of dusky lines

directed posteriorly from eye, 3rd pair curved posteroventrally

from eye across operculum, 4th pair curved anteroventrally

from eye to mouth, pair of short dusky marks directed ventrally

from mouth and faint dusky streak angled posteroventrally from

cheek; dorsal fin with broad horizontal dark streak distally

between first 3 spines; dorsal and anal fins with 2 horizontal

series of dusky spots, inner series in each fin conjoined to form

stripe anteriorly; caudal fin occasionally with 1 or 2 vertical

series of dusky spots. Terminal phase adults similar to initial

phase but with dark mark dorsally on side reduced to spot of

pupil size; lines on head, except for those across snout, directed

posteroventrally from eye with streak across cheek less

prominent; spot anteriorly on dorsal fin intense.

Fresh colours. Juveniles yellow with blue lines on head and on

dorsal and anal fins as in initial phase (fig. 4A); large black spot

in position of black mark on side of initial phase (Allen, 1985:

2406, fig. 330, as Choerodon species).

Initial phase adults olivaceous above, yellow to white

below, centres of scales anteriorly on side bright blue, spots

coalescing into blue stripes posteriorly on side following

horizontal scale rows (fig. 4B); broad black slash with blue to

white anterior margin between base of 7th dorsal fin spine and

lateral line; head with pair of narrow blue lines crossing snout

in front of eyes, second pair curving posteroventrally from eye

and third pair curving posteroventrally from eye across

operculum, blue line directed dorsoposteriorly from eye; pair

of short blue marks directed ventrally from mouth on each

side; brown to black streak angled posteroventrally from cheek

across operculum. Fins bright yellow; dorsal fin with broad

black streak distally between first 3 spines; dorsal and anal

fins with narrow blue margins and blue spots posteriorly;

caudal fin spotted with blue. Pectoral fin rays with broad blue

stripes basally; pelvic fin with one or more lengthwise blues

stripes (Sainsbury & Kailola, 1984: 261, top figure, as

Choerodon sp. 1; Allen, 1985: 2406, fig. 331, as Choerodon

species; Kuiter, 2010: 57, fig. A).

Terminal phase adults similar to initial phase but with blue

to grey cast above and black slashdike mark reduced to spot

(fig. 4C); head with oblique white patch below eye continuing

posteriorly on lower half of side. Dorsal, anal, caudal and

pelvic fin rays blue; caudal fin green with yellow periphery

(Kuiter, 2010: 57, figs C & D).

Etymology. From the Latin cauteroma for “brand” in reference

to the characteristic brand-like marking dorsally on the side of

the body in this species.

Distribution. Confined to coastal waters of tropical Western

Australia (fig. 3) from Shark Bay to the Arafura Sea off the

Northern Territory (Russell & Houston, 1989: 83; Larson et al.,

2013: 165). Found on inner reefs with large boulders and in

open areas with algal covered rock, sponges and other attached

invertebrates at depths of 1.5-150 m.

Comments. This species shares a greatly reduced number of

scales on the subopercle with C. cephalotes, C. rubescens and

C. schoenleinii, and to a lesser extent C. graphicus, which has

slightly larger scales in this region. It is easily distinguished

from the four by colour pattern, the obvious black slash-like

marking on the side above the lateral line and below the middle

dorsal fin spines a diagnostic feature. Despite its relatively

recent recognition as a distinct species, C. cauteroma is

particularly common throughout much of its greatly restricted

range in Western Australia.

Material examined. 46 specimens examined, 20-259 mm SL;

see appendix.

Choerodon cephalotes (Castelnau, 1875)

Purple Tuskfish

Figures 5, 6; table 3; appendix.

Choerops cephalotes Castelnau, 1875: 39, Cape York

(Queensland).

Choerops perpulcher De Vis, 1885: 877, Moreton Bay

(Queensland).

Choerodon macleayi Ramsay & Ogliby, 1887a: 241, Port Jackson

(New South Wales).

Choerops Hodgkinsonii Saville-Kent, 1893: 296, 370, pi. 15, fig.

2, Port Denison (Queensland).

Diagnosis. Dorsal fin rays XIII, 7, rarely XVI, 6; anal fin rays

III, 10; pectoral fin rays ii, 17, rarely 16, dorsalmost ray of

moderate length 36.4-53.1% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body moderately deep, 37.0-42.2% SL, head

depth 27.9-36.8% SL, caudal peduncle depth 12.1-17.8% SL;

head blunt, dorsal profile of snout steep, snout length 12.8-

18.3% SL; predorsal scales approximately 5 or 6, reaching

forward on dorsal midline to above posterior edge of preopercle;

cheek with small mostly embedded scales in about 1-5 diagonal

rows, posteriormost with about 4 or 5 scales to upper extent of

free preopercular edge, variably reaching forward from below

eye posteriorly to nearly corner of upper lip crease above

mouth, with very broad naked margin posteriorly and ventrally

on preopercle; tiny patch of about 1-3 scales in one or two rows

on dorsal end of subopercle adjacent preopercular edge; each

lateral line scale with multiple branching laterosensory canal

tube; scales above lateral line about 31 / 2 ; cephalic sensory canal

Review of Choerodon tuskfishes

Figure 5. Choerodon cephalotes. A, Juvenile, CSIRO H 7676-04, 82 mm SL, Great Barrier Reef, Queensland, Australia, photo by D. Gledhill,

CSIRO; B, Initial phase adult, CSIRO H 5958-08, 104 mm SL, Cape Flattery, 14° 48.5’ S, 145° 15.4’ E, photo by T. Carter, CSIRO; C, Terminal

phase adult, 209 mm SL, north-west of Port Hedland, Western Australia, photo compliments CSIRO.

M.F. Gomon

Figure 6. Distribution of Choerodon cephalotes based on specimens examined (coloured) and collection registration records (white).

pores extremely numerous especially anteriorly and posteriorly

in front of predorsal scales; second pair of canines in lower jaw

directed anterodorsally and curved slightly to strongly laterally;

dorsal and anal fins with low basal sheath comprising 1-3

progressively smaller accessory scales at deepest; posterior

lobe of dorsal and anal fins reaching well past hypural crease in

large individuals; caudal fin rounded to broadly pointed

centrally; pelvic fin reaching to base of second anal fin ray in

large individuals, length 22.0-33.2% SL. (See Table 3 for

additional meristic and morphometric ranges.) Adults

olivaceous above, creamy white below with horizontal dark

patch on upper half of side from below middle dorsal fin spines

to dorsal side of caudal peduncle; snout and forehead crossed

by 6-8 transverse orange lines.

Reaches moderately large maximum size, largest specimen

examined 253 mm SL.

Pigmentation in alcohol. Juveniles pale dusky above with

about 7 obscure darker bands dorsally, first broad above eye,

second narrower on nape, third below anterior part of dorsal

fin, 4th broad with darker anterior and posterior margins below

middle dorsal fin spines, 5th broad below first few dorsal fin

rays, 6th noticeably darker on top of caudal peduncle and 7th

narrow traversing caudal fin base; side ventrally pale;

sometimes with midlateral row of about 4 large dusky

rectangular blotches on posterior half of body and faint

remnants of banding dorsally adjacent to dorsal fin; dark dusky

spot dorsally on pectoral fin base and axilla; head with several

dusky bands across snout and in front of eye, 6 or 7 slender

transverse bands developing above and in front of eye with

growth; narrow dusky bands radiating from front of eye

towards rear of upper jaw, ventrally from eye behind corner of

mouth, and posteriorly from eye towards pectoral fin base;

dorsal fin pale with scattered small dusky patches; anal fin pale

with small dusky patches basally and fine dusky vermiculations

distally; caudal fin pale with narrow dusky bands darkest

distally (central part of fin appearing dark when fin not spread);

pectoral and pelvic fins pale. Adults dusky above and pale

below with posteriorly tapering dark dusky blotch below lateral

line from below middle dorsal fin spines variably to below soft

dorsal fin rays and in some onto dorsal half of caudal peduncle;

Review of Choerodon tuskfishes

dorsal surface of head dark dusky crossed by up to nine narrow

pale lines from about centre of eye gradually slanting anteriorly,

last to snout tip; short dusky bar directed dorsoposteriorly from

eye and second anteroventrally from eye; curved dusky mark

posteroventral to eye; cheeks dusky with fine pale spots, opercle

darker with spots ventrally; teeth blue; small dark spot on top

of pectoral fin base; dorsal fin pale basally, distal half dusky;

anal fin pale with narrow pale stripes distally and posteriorly;

caudal fin dusky with numerous narrow pale cross bands;

pectoral and pelvic fins pale to slightly dusky.

Fresh colours. Juveniles grey above, white ventrally with

obscure broad orange brown stripe above lateral midline on

head and body superimposed with broad black to dark brown

stripe posteriorly from below middle dorsal fin spines (fig. 5A);

head with blue lines as in adults. Dorsal fin of larger juveniles

with horizontal rows of blue spots or narrow stripes (Kuiter,

20i0: 53, figs A, C & E).

Initial phase adults greenish grey to purple above, white to

yellow below; variable horizontally elongate dark brown to

black patch on side dorsoposterior to pectoral fin to upper side

of caudal peduncle (fig. 5B & C); black line in axial of pectoral

fin; body scales edged with blue; snout and forehead blue to

violet crossed by 6-8 transverse orange lines; underside of

lower jaw white crossed by 3 transverse blue bands; orbit

rimmed with blue with broader blue mark directed

dorsoposteriorly from eye and short blue horizontal mark

posteroventral to eye; numerous fine orange to yellow spots on

cheeks and operculum; teeth blue. Alternating blue and orange

bands across caudal fin and distally on dorsal and anal fins;

pelvic fin green with lengthwise blue lines (Marshall, 1964:

colour plate 43, fig. 293; Sainsbury & Kailola, 1984: 257,

bottom; Kuiter, 2010: 53, figs B, D, F and G).

Terminal phase adults similar to initial phase but with more

vivid colours; blue lines on scales joined to form closely packed

fine blue lines crossing sides; horizontal black stripe on side,

sometimes extending to caudal fin base; dorsal midline of head

and snout intensely blue crossed by numerous orange lines.

Etymology. The name cephalotes appears to be a Latinisation

of the Greek kephalotos meaning “headed”, perhaps in

reference to the large colourful head of this species.

Distribution. Occurs around the northern coast of Australia

from Shark Bay, Western Australia to at least Southport,

Queensland, with only the holotype of C. macleayi reported to

have been collected south of that in Sydney Harbour (fig. 6).

Found in areas with rather open rubble or weed bottom at

depths of 0.3-80 m, with juveniles sometimes inhabiting algal

covered reefs in estuaries (Kuiter, 2010: 53).

Comments. Castelnau’s (1875) description of Choerops

cephalotes generally matches both C. anchorago and the

species treated here, but the omission of any mention of a

prominent white wedge-shaped spot behind the pectoral fin

and rectangular blotch on the caudal peduncle characteristic of

the former favours its identity as the latter. The type was not

found by Bauchot (1963: 22) or Eschmeyer (2015) and is

presumed lost. Based on museum collection records, C.

anchorago appears to be rather uncommon in the Cape York

region, whereas this species is abundant at this locality. The

colour description of De Vis’s (1885) Choerops perpulcher is

diagnostic for the species treated here and the name is clearly a

junior synonym. Two specimens in the Queensland Museum

collection are regarded as syntypes of that species (QMB I 945

and QMB I 9920). Although De Vis provided only a maximum

size of 14 inches (350 mm) for the species, he does refer to both

wet and dry material. QMB I 945 is currently in alcohol but

was originally prepared as a dry specimen. It currently has a

TL of about 335 mm with a badly frayed tail and may be the

basis for the maximum length given by De Vis. As pointed out

by Ramsay and Douglas-Ogilby (1857), their C. macleayi is a

rare example of a species of this genus reaching as far south as

the type locality Port Jackson (Sydney Harbour). Of the three

species known to occur at this locality, C. cephalotes, C.

schoenleinii and C. venustus, the authors’ description roughly

matches the first two. The extremely high number of branched

pectoral fin rays 17 and conversely very few scales on the

subopercle identifies it as a junior synonym of C. cephalotes.

Choerodon schoenleinii is the only other member of the genus

that regularly has as many as 16 branched pectoral fin rays and

similarly greatly reduced subopercular squamation.

Choerodon cephalotes and the very similar C. cyanodus have

nearly identical distributions and are the most frequently

encountered members of the genus in the shallow waters of

Australia’s northern coast. The series of narrow orange and blue

bands crossing the forehead in adults is distinctive for this species.

Material examined. 198 specimens, 29.1-253 mm SL; see

appendix.

Choerodon cyanodus (Richardson, 1843)

Blue Tuskfish

Figures 7, 8; table 3; appendix.

Labrus cyanodus Richardson, 1843: 355, Black Point (Port

Essington, Northern Territory).

Lachnolaimus arilca Richardson, 1848: 131, Endeavour Straits,

Bramble Island (Queensland).

Chaerops crassus Castelnau, 1875: 39, Dampier Archipelago

Islands (Western Australia).

Choerops albiqena De Vis, 1885: 876, Cape York (Quensland).

Choerops olivaceus De Vis, 1885: 876, Barrier Reef (Cardwell).

CapeYork (Queensland).

Choerops unimaculatus De Vis, 1885: 877, Barrier Reef

(Queensland).

Choerodonpaynei Whitley, 1945: 29, Dirk Hartog Island, Western

Australia.

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 14, rarely 12,13 or 15, dorsalmost ray dorsalmost ray

of moderate length 29.9-44.3% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body deep, 33.2-47.3% SL, head depth 27.4-

41.0% SL, caudal peduncle depth 12.7-16.8% SL; head bluntly

pointed, dorsal profile of snout moderately steep, snout length

10.9-18.0% SL; predorsal scales approximately 5-8, reaching

forward on dorsal midline to or not quite to above posterior

edge of preopercle; cheek with small mostly embedded scales

M.F. Gomon

Figure 7. Choerodon cyanodus. A, Juvenile, USNM 174385, 26.5 mm SL, Little Lagoon, Groote Eylandt, Northern Territory, Australia (grey

scale); B, Initial phase adult, CSIRO H 7673-03, 127 mm SL, Torres Strait, east of Banks Island, Queensland, photo by D. Gledhill, CSIRO; C,

Terminal phase adult, Dampier, Western Australia, photo by G. Edgar.

Review of Choerodon tuskfishes

Figure 8. Distribution of Choerodon cyanodus based on specimens examined (coloured) and collection registration records (white).

in about 5-7 diagonal rows, posteriormost with about 9 or 10

scales to upper extent of free preopercular edge, reaching

forward to or almost to corner of upper lip crease above mouth,

with very broad naked margin posteriorly and ventrally on

preopercle; about 2 rows of 5-8 small scales (only about 2

scales in second row) on subopercle adjacent preopercular edge

extending forward about midway along ventral preopercular

margin; each lateral line scale with multiple branching

laterosensory canal tube; scales above lateral line about 3;

relatively few cephalic sensory canal pores associated with

major canals; second pair of canines in lower jaw directed

dorsolaterally and curved slightly posteriorly; dorsal and anal

fins with very low basal sheath comprising 1-3 progressively

smaller accessory scales at deepest; posterior lobe of dorsal

and anal fins reaching hypural crease; caudal fin truncate to

slightly rounded, upper and lower corners only barely produced

at most in large individuals; pelvic fin reaching to base of

second anal fin spine in large individuals, length 19.0-24.7%

SL. (See Table 3 for additional meristic and morphometric

ranges.) Adults green above, white below, usually with white

oval patch between bases of first few segmented dorsal fin rays

and lateral line, faint broad and slightly darker bands across

dorsum; large individuals often with posteriorly tapering dark

patch on side from behind pectoral fin to upper half of caudal

peduncle.

Reaches moderately large maximum size, largest specimen

examined 300 mm SL, but reported to 337 mm SL (AMS

1.18219-001).

Pigmentation in alcohol. Juveniles pale with about 8 irregularly

angled narrow dusky bands on side; 3 or 4 bands on body

continuing onto dorsal fin at least basally, band at bases of last 2

or 3 spines quite dark; about 3 bands also continuing onto anal

fin, at least basally; bands on side quickly fading with growth,

persisting longest just below dorsal fin base; pelvic fin broadly

dark adjacent to leading edge; fins otherwise unpigmented (fig.

7A). Initial phase adults dusky above and pale on underside,

especially on lower jaw and ventral side of head, with a large,

distinctly pale oval spot dorsally on side between bases of first

few segmented dorsal fin rays and lateral line. Dorsal and caudal

M.F. Gomon

fins slightly dusky, posterior corners of caudal fin discrete and

slightly darker, faint vermiculations sometimes visible on

caudal fin other than corners. Terminal phase adults similar to

initial phase but often lacking pale spot between dorsal fin and

lateral line and usually with posteriorly tapering pale area on

dorsal half of side posterior to pectoral fin base.

Fresh colours. Juveniles greenish yellow, darker dorsally with

brown horizontal lines following scale rows and white oval

spot as in initial phase adults (Kuiter, 2010: 58, fig. C).

Initial phase individuals green to greenish blue above,

yellowish white below (fig. 7B); head deep green with blue bar

before eye and blue bars on snout; chalky blue under lips, chin

blackish green; teeth blue to blue-green; usually with large

bright white oval spot between bases of first few segmented

dorsal fin rays and lateral line; 3 short, broad, brownish-olive

bands crossing back from just behind dorsal fin origin to

middle of dorsal fin base, bands variably faint to dark brown

or black depending on locality, pale spot missing from

individuals with dark banding; large individuals often with

blackish-green stripe from pectoral fin base to caudal fin base;

orange to rust coloured rectangular patch behind pectoral fin

base with horizontal orange stripes posteriorly. Distal margins

of dorsal and anal fins, posterior corners of caudal fin, and

leading edges of pectoral and pelvic fins deep to bright blue;

red-orange wavy markings on blue background covering

remainder of dorsal, anal and caudal fins; remaining pectoral

fin rays orange with yellow interradial membranes; pelvic fin

with orange submarginal line adjacent to blue margin (Kuiter,

2010: 58, figs A & E).

Terminal phase individuals similar to initial phase with

dark blue, brown or black dorsum having vertical anterior

margin just behind pectoral fin base reaching ventrally to near

level of lower part of pectoral fin base, tapering to lateral

midline and continuing to caudal tin base in some (fig. 7C);

smaller individuals with horizontal orange lines following

scale rows on caudal peduncle; white oval spot more or less

obscured in large individuals; dark area frequently replaced by

horizontally elongate copper to orange patch (Marshall, 1964:

colour pi. 44, fig. 295; Allen, 1985: fig. 327; Kuiter, 2010: 58,

figs B, D and F).

Etymology. The name cyanodus is from the Greek kyanos,

“blue” and odontos “tooth” in reference to the blue teeth

characteristic of this and other species of the genus Choerodon.

Distribution. Confined to inshore waters of Australia’s tropical

north from Shark Bay, Western Australia to Sydney Harbour,

New South Wales, recorded in Papua New Guinea at Daru

Island in the Torres Strait (fig. 8). Usually found in open algal

covered rubble and soft substrate habitats at depths of 0.2-20 m.

Comments. The type of Richardson’s (1843) Labrus cyanodus

is a dry skin (BMNH 1843.6.15.46) prepared from a specimen

collected at Black Point, Port Essington, Northern Territory.

Although faded, the skin retains the remnants of colour pattern

provided in the original account, which is consistent with the

above colour description. The name L. arilca appears in a

description of “Lachnolaimus, vel cyanodus” by Richardson

(1848), perhaps as an available name should the tentative

identification prove to be incorrect. A dried skin in the collection

of the British Museum (BMNH 47.6.17.57,179 mm SL) collected

by Richardson and cited by McCulloch (1929-30: 319) under

the name Labrus arilca is likely to be the type. The description

of C. crassus by Castelnau (1875) differs markedly from other

species in the genus by the numbers of dorsal and anal fin rays,

XIII, 12 and III, 13, respectively. Presumably, Castelnau erred

in counting branches of posterior rays as individual rays, since

an assumed syntype in the Paris collection (MNHN A.8890,

207 mm SL, 256 mm TL) has the typical counts of XIII, 7 and

III, 10. As well as being registered from the same collection

locality, the specimen has a TL approximating that given by

Castelnau of about a foot. The specimen was initially split open,

as it would have been if salted as Castelnau implied, and is

clearly identifiable as C. cyanodus.

De Vis (1885) provided names and descriptions of six new

species of Choerodon (as Choerops) from Queensland waters,

four of which have been synonymised with C. cyanodus,

although one only questionably (Parent! & Randall, 2000: 10).

Judging from sizes provided with the descriptions, C. concolor,

C. olivaceus and C. unimaculatus are likely to have been

based on juveniles, while the size of the type of C. albigena

was not given. The morphological descriptions of all four are

inconclusive for identification, but colour accounts are more

helpful. The Queensland Museum has specimens that have

been regarded as types of most of these, although the lengths

of some do not match those provided by De Vis. The length of

a mounted specimen (QMB 1.946, 140 mm TL) registered as a

syntype of C. concolor from north-east Queensland is

consistent with the SVi inches given by De Vis. The specimen

is likely to be the type and is identifiable as C. cyanodus. Two

specimens labelled as ''Choerops unimaculatus Type?” (QMB

1.95, 150 mm TL and QMB unregistered, 134 mm TL) are

distinctly longer than the type of that species (4 inches) and

therefore not the holotype, but may be others examined as

implied in the original description and therefore paratypes in

current terminology. De Vis’s colour description of C.

concolor more or less matches the pattern in juveniles of C.

cyanodus. The only other species occurring on the Great

Barrier Reef with a distinctive “bright oval spot below the end

of the soft dorsal” in juveniles is C. schoenleinii, which has a

prominent black spot anterodorsal to it that would certainly

have been mentioned by De Vis. Choerops concolor is

therefore regarded as a synonym of C. cyanodus. No specimen

was found in the Queensland Museum collection that is

conclusively identifiable as the type of C. olivaceus from

Barrier Reef (Cardwell), Cape York, although the

abovementioned specimen registered as QMB 1.95 is recorded

as coming from that locality. That specimen cannot be the

type because it is much longer than the “2 inches” stated by De

Vis. The original description of C. olivaceus refers to “a pale

blotch beneath the posterior half of the dorsal fin”, as

mentioned above for C. unimaculatus, and is likewise

considered to be synonym of C. cyanodus. Based on the

original description of C. albigena and a registered type

(QMB 1.110, 285 mm TL), the name is regarded as another

synonym of C. cyanodus. Collection details for the type are

identical to those provided by De Vis in his description. Still,

Review of Choerodon tuskfishes

the reference to “a dark blotch (sometimes obsolete) on the

back beneath the ninth dorsal spine” in that description is

difficult to reconcile unless it refers to one of the bands

crossing the back of many individuals. These bands are

intensely dark in fish occurring in the Kimberley Region of

northern Western Australia.

Choerodon cyanodus is geographically variable in both

colouration and genetic configuration, but the degree of

genetic separation appears to be sufficiently minor as to regard

the populations as infraspecific. Different colour forms

apparently occur within the same areas, at least in the central

western and Kimberley coasts of Western Australian

(Fairclough, 2005: 26 and personal communication). Whitley’s

(1945) Choerodon paynei was based on a terminal phase adult

form prevalent in the Shark Bay area of Western Australia,

which lacks the oval white blotch dorsally on the side below

the posterior half of the dorsal fin typical of individuals in

northern and eastern Australia. A second form that is well

known in the Kimberley Region of Western Australia is

marked by prominent broad dark bands on the back that are at

the most faint in individuals in north-eastern Australia.

Choerodon cyanodus shares with its close congener C.

cephalotes, and to a slightly lesser extent C. schoenleinii, a

particularly high pectoral fin count and few scales on the

subopercle, the combination of which are unique for the genus.

The species is distinguishable from the two by the sparse

sensory pores on the top of the head restricted to the main

sensory canals.

Choerodon cyanodus is one of the most abundant

tuskfishes in shallow coastal areas of northern Australia

having a sand and rubble substrate.

Material examined. 150 specimens, 19.8-300 mm SL; see

appendix.

Choerodon graphicus (De Vis, 1885)

Graphic Tuskfish

Figures 9, 10; table 3; appendix.

Choerops graphicus De Vis, 1885: 878, Queensland Coast

(Cardwell).

Choerodon transversalis Whitley, 1956: 258, fig. 7, Heron Island,

Great Barrier Reef, Queensland.

Diagnosis. Dorsal fin rays XIII, rarely XII, 7; anal fin rays III, 10;

pectoral fin rays ii, 14, dorsalmost ray dorsalmost ray of moderate

length 35.4-47.9% pectoral fin length, ventralmost rays shorter

than those above, posterior edge of fin obliquely straight,

dorsoposterior comer bluntly pointed, posteroventral comer

angular; body deep, 35.3^14.2% SL, head depth 27.4—35.1% SL,

caudal peduncle depth 14.5-17.2% SL; head bluntly pointed,

dorsal profile of snout moderately steep, snout length 12.2-

16.3% SL; predorsal scales approximately 5-7, reaching forward

on dorsal midline to above posterior edge of preopercle; cheek

with small partially embedded scales in about 7-9 diagonal rows,

poster!ormost with about 15 scales to upper extent of free

preopercular edge, reaching forward almost to comer of upper lip

crease above mouth, with very broad naked margin posteriorly

and ventrally on preopercle; 1 or 2 rows of about 4—7 small scales

(only 1 or 2 scales in second row when present) on subopercle

adjacent preopercular edge extending forward to about middle of

ventral preopercular margin; each lateral line scale with multiple

branching laterosensory canal tube; scales above lateral line

about IVi or 3; numerous cephalic sensory canal pores on top of

head behind eye and on cheek anteroventral to eye, fewer

between eyes and on snout; second pair of canines in lower jaw

directed anterodorsally, very little laterally; dorsal and anal fins

with very low basal sheath comprising 1-3 progressively smaller

accessory scales at deepest; posterior lobe of dorsal and anal fins

reaching beyond hypural crease; caudal fin tmncate to slightly

rounded, upper and lower comers rounded; pelvic fin reaching to

anal fin origin in large individuals, length 21.2-25.5% SL. (See

Table 3 for additional meristic and morphometric ranges.)

Juveniles and adults olive yellow with broad irregular, angled

darker bands on side and broad darker lines radiating from eye;

dark spot or blotch on lateral line below middle dorsal fin spines.

Reaches moderately large maximum size, largest specimen

examined 368 mm SL, but reported to about 500 mm.

Pigmentation in alcohol. Juveniles pale with 6 broad dusky

bands, anterior 2 across nape, subsequent 3 below dorsal fin and

last across posterior half of caudal peduncle and base of caudal

fin; bands below dorsal fin and on caudal peduncle subdividing

to form more numerous narrower dusky bands with broader

pale interspaces, areas near middle of several darkened as dark

midlateral spots; narrow dusky bands on head radiating from

eye, narrow band or pair of bands directed dorsally across top of

head, pair of bands directed anterodorsally across snout, band

directed anteroventrally across mouth midlaterally traversing

underside of lower jaw, band directed ventrally across underside

of head behind mouth, and 2 bands directed posteriorly from

eye, one towards upper end of pectoral fin base and other

towards origin of lateral line; about 4 dusky bands on side

extending onto dorsal fin and 3 extending onto anal fin; caudal

fin base with small darker spot on dorsal and ventral corners

and pair of smaller spots separated by pale space midlaterally;

dorsal, anal and caudal fins otherwise pale to transparent;

pectoral fin transparent; pelvic fin broadly dusky to dark along

leading edges, pale to transparent along posterior edges. Initial

phase adults retaining juvenile bands, although those on body

more uniformly broad and often fainter and those on head

narrower; band directed posteroventrally from eye towards

pectoral fin base usually darker; usually with large rather dark

spot on lateral line below central dorsal fin spines and second on

dorsal profile of side at base of last segmented dorsal fin ray.

Terminal phase adults with faint bands on body if at all visible

and band directed posteroventrally from eye rather dark.

Fresh colours. Juveniles and initial phase individuals white

with 5 or 6 prominent irregular broad brown to black vertical

bands from nape to base of caudal fin, band below central

dorsal fin spines split vertically, anterior section superimposed

with black spot on and below lateral line (fig. 9A & B); bands

continuing on head as bars radiating from eye, 3 crossing

forehead, 2 directed ventrally and 2 posteriorly, additional

vertical bar crossing operculum; dorsal, anal and caudal fins

crossed by blue lines or rows of fine blue spots. (Kuiter, 2010:

50, bottom of page figs B and C)

M.F. Gomon

Figure 9. Choerodon graphicus. A, Juvenile. Red Rock Estuary, Coffs Harbour, New South Wales, photo by 1. Shaw; B, Initial phase adult,

BPBM 11413, 287 mm SL, New Caledonia, photo by J. Randall, BPBM; C, Terminal phase adult, approximately 500 mm SL, Hot Canard,

Noumea, New Caledonia, photo by J. Dubose.

Review of Choerodon tuskfishes

Figure 10. Distributions of Choerodon graphicus (circles), C. oligacanthus (squares) and C. rubescens (triangles) based on specimens examined

(coloured) and collection registration records (white).

Terminal phase individuals olive yellow with slate grey to

brownish grey anastomosing transverse bands with green

flecks, most body scales behind pectoral fin with vertical blue

streak (fig. 9C); head dull orange to yellow on cheeks and

operculum, chin bright green, bars radiating posteriorly from

eye black; fins mostly dark slate grey, dorsal fin with blue

margin, reddish orange submarginal line, and scattered blue

and orange markings; anal fin blue with numerous fine orange

markings; pectoral fin brown, although more grey basally, with

green and brown base (Kuiter, 2010: 50, bottom of page fig. A).

Etymology. The name graphicus is from the Greek “graphikos”,

“of writing”, perhaps in reference to the complex markings on

the side of the head and body resembling letters.

Distribution. A species with limited range restricted to the

Queensland coast from Nymph Island off the Cape York Peninsula

and south, at least to Southport near the New South Wales border,

and New Caledonia (fig. 10). Found on inshore reefs, lagoons and

estuaries with open substrate and rubble at depths of 2-36 m.

Comments. De Vis’s (1885) brief description of Choerops

graphicus agrees with the species described above, while a

specimen registered as the type (QMB 1.944,360 mm TL) with

the same collection information and approximate length given

in the original account (14 inches) confirms the identity. Only

C. fasciatus with XII, 8 dorsal fin rays has a prominent darkly

banded colour pattern at this size within the genus. Whitley’s

(1956) Choerodon transversalis is based on a terminal phase

specimen (AMS IB.3527, 296 mm SL) collected at Heron

Island, Queensland.

The distinctive colouration of this species is unlike that of

other species except at a small juvenile size. Individuals of less

than about 40 mm SL approach those of C. anchorago and C.

azurio of similar size in overall pattern. Choerodon graphicus

is separable from the former by pectoral fin count and vertebral

formula and from the latter by distance, the two occurring on

opposite sides of the tropics.

Material examined. 25 specimens, 13.4-368 mm SL; see

appendix.

M.F. Gomon

Choerodon oligacanthus (Bleeker, 1851)

Whitepatch Tuskfish

Figures 10, 11; table 3; appendix.

Crenilabrus oligacanthus Bleeker, 1851: 489, Rio (Riau, Sumatra,

Indonesia).

Choerops palawanensis Seale, 1910: 523, Puerto Princessa,

Palawan Island (Philippines).

Diagnosis. Dorsal fin rays XIII (rarely XIV), 7; anal fin rays

III, 10; pectoral fin rays ii, 14, rarely 15, dorsalmost ray of

moderate length 26.1-42.8% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body deep, 31.4-39.3% SL, head depth 24.4-

33.4% SL, caudal peduncle depth 13.4-14.5% SL; head bluntly

pointed, dorsal profile of snout moderately steep, snout length

10.2- 16.4% SL; predorsal scales approximately 5-7, reaching

forward on dorsal midline to about midpoint between posterior

extent of orbit and posterior edge of preopercle, reaching

forward to or in advance of posterior edge of orbit more

laterally; cheek with small partially embedded scales in about

7-10 diagonal rows, posteriormost with about 13 scales to

upper extent of free preopercular edge, reaching forward to

corner of upper lip crease above mouth, with relatively broad

naked margin posteriorly and ventrally on preopercle; single

row of 6 or 7 rather large scales on subopercle adjacent

preopercular edge extending forward approaching anterior end

of ventral preopercular margin; each lateral line scale with

triple branching laterosensory canal tube; scales above lateral

line about 3 or 314; cephalic sensory canal pores moderately

numerous to numerous on snout and anteroventral to eye,

extremely numerous above and behind eyes; second pair of

canines in lower jaw directed anterodorsally and slightly

laterally; dorsal and anal fins with very low basal sheath

comprising 1-3 progressively smaller accessory scales at

deepest; posterior lobe of dorsal and anal fins not quite reaching

hypural crease; caudal fin truncate to slightly rounded, upper

corners slightly produced in large individuals; pelvic fin not

quite reaching anus in juveniles, elongate in large individuals

reaching at least to base of 4th segmented anal fin ray, length

22.3- 44.6% SL. (See Table 3 for additional meristic and

morphometric ranges.) Adults green above, white on underside,

usually with large oval white spot pach on lateral line below

bases of 6th to 8th dorsal fin spines and black spot below

anterior end of oval patch.

Reaches moderately large maximum size, largest specimen

examined 250 mm SL.

Pigmentation in alcohol. Juveniles and initial phase adults pale,

slightly dusky on dorsum, with dark blotch slightly smaller than

eye below lateral line beneath central dorsal fin spines and

second spot of similar size or smaller above lateral line anteriorly

on caudal peduncle; large elongate pale oval patch developing

below middle dorsal fin spines above lateral line in slightly

larger individuals posterior to first dark blotch, which

progressively extends posteriorly as dark dusky marginal stripe

on underside of pale patch; fins pale. Terminal phase adults as in

large initial phase adults with posterior dark spot disappearing.

Fresh colours. Juveniles similar to initial phase adults with little

red colouration and black spot above lateral line midway along

caudal peduncle (White et al., 2013: 267, fig. 89.15 juvenile).

Initial phase adults green above, reddish yellow laterally

and white on underside, with numerous blue horizontal lines

on side and 3 narrow blue bands directed anteroventrally from

eye (fig. 11 A); black spot below lateral line under 5th to 7th

dorsal fin spines; second above lateral line past dorsal fin base;

much larger elongate pale spot above lateral line under middle

dorsal fin spines. Dorsal fin red; soft portion of dorsal, anal

and caudal fins spotted with yellow to pearl; anal fin rays with

oblique pearly stripes (Kuiter, 2010: 60, top of page figs A-D).

Terminal phase adults similar to initial phase with more

prominent reddish colouration on side and fins (fig. IIB; White

et al., 2013: 267, fig. 89.15 adult).

Etymology. The name oligacanthus is from the Greek oligo for

“few” and akantha for “thorn or spine”, in reference to the

thirteen dorsal fin spines of this species, which is low for

members of the genus Crenilabrus as recognised at the time of

the description. The generic name has since been referred to a

subgenus of the temperate North Atlantic Symphodus, which

has a high number of dorsal fin spines relative to other members

of the Labridae.

Distribution. Confined to the western extreme of the Pacific

from the central Philippines and Malaysia to the Java Sea side

of southern Indonesia (fig. 10). Found in areas with mud or sand

and rubble bottom adjacent coastal reefs, often in turbulent

conditions, at depths of 2-15 m (Kuiter, 2010: 60; Allen &

Erdmann, 2012: 647).

Comments. The four specimens on which Bleeker (1851) based

the description of Crenilabrus oligacanthus are in the

Rijksmuseum collection (RMNH 6532,4: 77.0 mm SL, 96.8 mm

TL-88.6 mm SL, 110 mm TL) and are consistent with the original

account. The largest specimen in the type series is in reasonable

condition and should be regarded as lectotype. The jar with this

series also contains larger specimens, the longest 209 mm SL. All

are considered too large to be potential types. The type of Seale’s

(1910) Choerops palawanensis appears to be no longer extant.

The morphological description is not especially diagnostic and

the dorsal fin spine count of 14 is assumed to be an individual

anomaly, but the colour description mostly matches that of C.

oligacanthus. It is regarded as a junior synonym of that species.

A relatively common species confined to equatorial waters

of the western extreme of the Pacific, C. oligacanthus is most

easily distinguished from congeners having fourteen

segmented pectoral fin rays by the characteristic elongate oval

pale spot above the lateral line below the central portion of the

dorsal fin and in smaller specimens by the pair of black spots

dorsally on the side, the anterior centrered below the sixth

dorsal fin spine and the second above the lateral line just

posterior to the dorsal fin base. Choerodon oligacanthus is the

only species of the genus that develops elongate pelvic fins as

adults reaching well beyond the anal fin origin.

Material examined. 68 specimens, 50.5-250 mm SL; see

appendix.

Review of Choerodon tuskfishes

Figure 11. Choerodon oligacanthus. A, Juvenile, CSIRO H 7406-07, 107 mm SL, Tanjung Luar, Lombok, Indonesia, photo by W. White,

CSIRO; B, Terminal phase adult CSIRO H 7987-01, 229 mm SL, Kedonganan, Bali, Indonesia, photo by W. White, CSIRO.

M.F. Gomon

Choerodon rubescens (Giinther, 1862)

Baldchin Groper

Figures 10, 12; table 3; appendix.

Choerops rubescens Gunther, 1862: 97, Houtmans Abrolhos

(Western Australia).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 16, rarely 15, dorsalmost ray dorsalmost ray of moderate

length 26.8-50.3% pectoral fin length, ventralmost rays shorter

than those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body deep, 38.1-42.6% SL, head depth 29.8-36.1% SL,

caudal peduncle depth 13.4-16.5% SL; head bluntly pointed,

dorsal profile of snout steep, snout length 12.3-18.2% SL;

predorsal scales approximately 5-8, variably reaching forward

on dorsal midline not quite to or in advance of posterior edge of

preopercle; cheek with small embedded scales in about 4-6

diagonal rows, posteriormost with about 10 scales to upper extent

of free preopercular edge, reaching forward only slightly in

advance of preopercular corner, with very broad naked margin

posteriorly and ventrally on preopercle; 2 or 3 rows of 3-6 small

scales (only 1 or 2 scales in second and third rows when present)

on subopercle adjacent preopercular edge extending forward only

slightly in advance of preopercular corner; each lateral line scale

with multiple branching laterosensory canal tube; scales above

lateral line about 41 / 2 ; cephalic sensory canal pores extremely

numerous on top of head and anteroventral to eye, continuing

onto cheek; second pair of canines in lower jaw directed mostly

dorsally, little laterally; dorsal and anal fins with very low basal

sheath comprising 1-3 progressively smaller accessory scales at

deepest; posterior lobe of dorsal and anal fins reaching hypural

crease, not reaching crease in small individuals; caudal fin

truncate to slightly rounded, upper and lower corners rounded,

only barely produced at most in large individuals; pelvic fin to just

beyond or just short of anus, length 20.1-23.9% SL. (See Table 3

for additional meristic and morphometric ranges.) Adults pink to

dark blue with white underside of head below mouth and

prominent white spot covering pectoral fin base.

Reaches large maximum size, largest specimen examined

521 mm SL, but reported in the literature to 90 cm (Hutchins

& Swainston, 1986: 901).

Pigmentation in alcohol. Juveniles pale dusky, lower half of side

and anal fin slightly darker. Adults dusky; underside of head

below gape angled posteroventrally to opercular edge stark white;

large white blotch covering pectoral fin base. Fins mostly dusky.

Fresh colours. Juveniles yellow to grey, underside of trunk and

tail sometimes black (fig. 12A); pectoral fin yellow (Hutchins

& Swainston, 1986: fig. 491).

Initial phase adults pink with abruptly white underside of

head below level of upper jaw and white oval spot covering

pectoral fin base (fig. 12B); fins of similar colour with narrow

blue distal margins (Allen, 1985: fig. 328; Kuiter, 2010: 56,

bottom of page fig. A; Fairclough, personal communication).

Terminal phase adults similar to initial phase adults but

blue to grey overall (fig. 12C; Allen, 1985: fig. 329; Kuiter,

2010: 56, bottom of page fig. C).

Etymology. The name rubescens is from the Latin rubesco

for “reddish”, in reference to the reddish ground colour of the

type specimen (Kuiter, 2010: 56, bottom of page fig. B).

Distribution. This Western Australian endemic is the most

temperate member of the genus on Australia’s west coast with

the most restricted distribution confined to the region from

Shark Bay to Garden Island, south-west of Perth (fig. 10).

Smaller individuals occur on shallow sand and inshore reefs,

while large adults are usually found on deeper reefs at depths

of 4-30 m (Hutchins, 1979: 71; Kuiter, 2010: 56).

Comments. The type of Gunther’s (1867) Choerops rubescens

is a dry skin (BMNH 1844.2.15.68, 279 mm SL) with 15

branched pectoral fin rays that matches the original description.

Other specimens of this species examined in the course of the

study regularly have 16.

This species takes its vernacular name Baldchin Groper

from its stark white lower jaw that along with its white pectoral

fin base stands out against the uniform, often dark colouration

of the remaining head and body.

Material examined. 15 specimens, 54.4-521 mm SL; see

appendix.

Choerodon schoenleinii (Valenciennes, 1839)

Blackspot Tuskfish

Figures 13-15; table 3; appendix.

Cossyphus Schoenleinii Valenciennes, in Cuvier & Valenciennes,

1839: 143, Celebes (Indonesia).

Cossyphus cyanostolus Richardson, 1846: 256, Canton (China).

Cossyphus ommopterus Richardson, 1846: 257, Canton (China).

Choerops unimaculatus Cartier, 1874: 102, Cavite (Philippines).

Torresia australis Castelnau, 1875: 36, Cape York (Queensland).

Chaerops notatus Alleyne & Macleay, 1877: 344, pi. XVI, fig. 1,

Cape Grenville (Queensland).

Torresia lineata De Vis, 1885: 881, Cardwell (Queensland).

Choerops nyctemblema Jordan & Evermann, 1902: 353, fig. 21,

Formosa.

Choerodon rubidus Scott, 1959: 89, fig. 7, Shark Bay (Western

Australia).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 16, rarely 17, dorsalmost ray dorsalmost ray of

moderate length 41.5-53.9% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body deep, 32.8-43.4% SL, head depth 23.7-

38.5% SL, caudal peduncle depth 11.8-16.4% SL; head bluntly

pointed, dorsal profile of snout moderately steep, snout length

9.0-16.9% SL; predorsal scales approximately 5-7, reaching

forward on dorsal midline not quite to or just in advance of

posterior edge of preopercle; cheek with moderately large non-

imbricate scales in about 5-9 diagonal rows, posteriormost

with about 13 scales to upper extent of free preopercular edge,

reaching forward to or nearly to corner of upper lip crease

above mouth, with moderately broad naked margin posteriorly

and ventrally on preopercle; row of 2-5 small scales on

subopercle adjacent preopercular edge at or just above

preopercular corner; each lateral line scale with multiple

Review of Choerodon tuskfishes

Figure 12. Choerodon rubescens. A, Juvenile, WAM P. 27955-018, 60 mm SL, Port Denison, Western Australia, 29° 16’ S, 114° 55’ E, photo by

B. Hutchins, WAM; B, Initial phase adult, CSIRO H 4875-02, 324 mm SL, Western Australia, photo by T. Carter, CSIRO; C, Terminal phase

adult, CSIRO H 4875-03, 380 mm SL, Western Australia, photo by T. Carter, CSIRO.

M.F. Gomon

Figure 13. Choerodon schoenleinii. A, Juvenile, 15 mm SL, Okinawa-jima Island, Ryukyu Islands, Japan, photo by T. Tsuhako; B, Initial phase

adult, Okinawa, Ryukyu Islands, Japan, photo by K. Shimada; C, Terminal phase adult, CSIRO H 7899-03, 320 mm SL, Kedonganan, Bali,

Indonesia, photo by W. White, CSIRO.

Review of Choerodon tuskfishes

Figure 14. Choerodon schoenleinii. A, Cossyphus cyanostolus Richardson, 1846, BMNH, terminal phase adult, fourteen inches (280 mm SL),

Canton, China Seas; B, Cossyphus ommopterus Richardson, 1846, BMNH, initial phase adult, 614 inches (124 mm SL), Canton, Sea of China.

M.F. Gomon

20 °

Figure 15. Distribution of Choerodon schoenleinii based on specimens examined (coloured) and collection registration records (white).

branched laterosensory canal tube in large individuals; scales

above lateral line about 3 or 31 / 2 ; cephalic sensory canal pores

numerous above and behind eyes less so in front of and below

eyes; second pair of canines in lower jaw directed anterodorsally,

very little laterally; dorsal and anal fins with low basal sheath

comprising 1-3 progressively smaller accessory scales at

deepest; posterior lobe of dorsal and anal fins not quite reaching

hypural crease in Juveniles, reaching hypural crease in adults;

caudal fin truncate to slightly rounded, upper and lower corners

only barely produced at most in large individuals; pelvic fin

reaching to base of second anal fin spine in large individuals,

length 19.1-27.3% SL. (See Table 3 for additional meristic and

morphometric ranges.) Adults bluish green above with yellow

sides and vertical blue line of each body scale; black spot on

body at base of last dorsal fin spot and white patch often

between lateral line and middle segmented dorsal fin rays; head

with diagonal blue lines angled from and below eye.

Reaches large maximum size, largest specimen examined

530 mm SL. Reported in the literature to reach 1 m (Kuiter,

2010: 59).

Pigmentation in alcohol. Juveniles pale dusky above, pale

below with narrow dusky stripes following horizontal rows of

scales especially posteriorly, prominent dark spot on scaly

sheath of dorsal fin below last dorsal fin spine; head dusky

above and pale ventrally, with 2 dark dusky stripes directed

posteriorly from lower half of eye and another 2 from lower

half of eye anteroventrally towards upper jaw; fins pale, dorsal

and anal with dusky sub-basal stripe, anal with sub-distal

stripe; caudal fin pale dusky basally near middle of fin. Initial

phase adults pale with duskier dorsum; dark spot on scaly

dorsal fin sheath prominent. Terminal phase adults rather dark,

scales on side and chest with broad dark margins; lateral line

and pectoral fin base dark; caudal peduncle with about 6

horizontal rows of dark spots; dark spot on scaly base of dorsal

fin small or absent; fins dusky.

Fresh colours. Juveniles (fig. 13A) tan to pale brown with 8-10

fine white mostly vertical bars dorsally across back, those

posteriorly across underside of caudal peduncle as well; head

with narrow brown bands directed dosally, anteroventrally and

Review of Choerodon tuskfishes

ventrally from eye; white spot at base of pectoral fin, and

dorsally and ventrally on caudal fin base; fins transparent with

broad vertical tan patches covering anterior and middle of

dorsal fin, pelvic fin and anterior end of anal fin; small dark

brown to black spot at base of last dorsal fin spine.

Initial phase adults olive, washed with blue above, often

yellow below with vertical blue line on each body scale, those

posteriorly merging to form horizontal blue lines (fig. 13B);

pairs of blue lines directed anteroventrally and posteriorly

from orbit; underside of head orange to yellow, lower jaw blue;

teeth blue; blue line at base of pectoral fin; distinctive black

spot on body at base of last dorsal fin spine; white blotch often

on body below segmented dorsal fin rays in smaller individuals.

Dorsal and anal fins with narrow horizontal blue lines; caudal

fin with narrow blue bands; pelvic fin with blue leading endge

(Masuda etal., 1984: pi. 194, fig. A; Shen, 1993: pi. 144, fig. 6;

Okamura & Amaoka, 1997: 465, right second from bottom;

Chen et al., 2010: 383, fig. F; Kuiter, 2010: 59, figs B-E; Allen

& Erdamann, 2012: 648, top of page; White et al., 2013: 267,

fig. 89.17 female).

Terminal phase adults dark bluish green with little if any

indication of white spot posteroventral to black spot (fig. 13C);

lower jaw and caudal fin rays blue (Masuda et al., 1984: pi.

194, fig. B; Sainsbury & Kailola, 1984: 259, top; Okamura &

Amaoka, 1997: 465, centre bottom; Kuiter, 2010: 59, figs A &

F; Allen & Erdamann, 2012: 648, centre of page).

Etymology. The origin of the name schoenleinii is unclear,

although Valenciennes speculated that Agassiz intended it to

recognise Johann Lukas Schoenlein, an important medical

scientist of the Biedermeier Zeit, following the Napleonic wars,

1815-1858 (Pietsch, Eschmeyer and Fairclough, personal

communication).

Distribution. Occurs along the western edge of the Pacific from

Okinawa in southern Japan (Nakabo, 2000: 971) and China to

both the eastern and western coasts of northern Australia, at

least to Shark Bay in Western Australia and Sydney Harbour in

New South Wales (fig. 15); not recorded east of the Philippines,

Papua New Guinea and eastern Australia. Found on silty or

sand to rubble bottoms often with algal cover in lagoons and

near coastal reefs at depths of 0.3-46 m.

Comments. Valenciennes (in Cuvier & Valenciennes, 1839:

143) based his description of Cossyphus Schoenleinii on a

colour figure of a specimen collected in Celebes (Sulawesi,

Indonesia) and conveyed to him by Agassiz. Meristic

discrepancies involving soft anal fin and pectoral fin ray counts

between his description and the species described here are

attributed to the inaccuracy of the figure.

Both Cossyphus cyanostolus and Cossyphus ommopterus

were described by Richardson (1846: 256,BMNH 1968.3.11.15,

280 mm SL, and 257, BMNH 1968.3.11.16, 124 mm SL,

respectively) from dried specimens collected in Canton

(China) with accompanying detailed colour drawings (fig. 14A

and B, respectively). They are clearly synonyms of C.

schoenleinii. Cartier’s (1874: 102) description of Choerops

unimaculatus is based on juveniles as corroborated by the

stated lengths of his type series, 3.9-6.7 cm. Although the

types appear to have been lost (Eschmeyer, 2015), the colour

pattern is that of C. schoenleinii. The description of Castelnau’s

(1875: 36) Torresia australis is similarly brief and could apply

to more than one species, but the colour account most closely

matches that of juvenile C. schoenleinii. The disposition of the

type is unknown. Morphological details in Alleyne and

Macleay’s (1877) description of Chaerops notatus are

inconclusive as to the identity of this species, but the

colouration and type (AMS 1.16360-001,216 mm SL) specimen

identify it as a synonym of C. schoenleinii. De Vis’s (1885:

881) description of Torresia lineata is also very brief

morphologically, but the colour description is clearly that of a

juvenile C. schoenleinii. The type (QMB 1. 11/82, ^80 mm SL)

is in poor condition, but is identifiable as C. schoenleinii in

having 16 branched pectoral fin rays. This is the only species

in the Cardwell, Queensland region regularly with that number

of pectoral fin rays.

Choerops nyctemblema Jordan & Evermann (1902: 353, fig.

21) was based on a single 18 inch (72 cm) specimen collected in

Formosa (Taiwan) that no longer appears to be extant. The

description is brief, but of the seven species of Choerodon

known to occur in Taiwan, it most closely resembles C.

schoenleninii. The account is consistent with a large example,

none of the other six species attaining a length greater than 430

mm SL. Although the authors did not provide a pectoral fin

count, the published figure has ii, 16 rays, supporting the

identification, with the other six species having 14 (rarely 15) or

fewer branched rays. Choerodon rubidus described by Scott

(1959: 89, a second fig. 7 on that page, labelled Stethojulis

rubromacula sp. nov.) is based on a 167 mm SL (210 mm TL,

SAM F2985) specimen from Point Samson, Western Australia,

identifiable as C. schoenleinii. Scott gave Shark Bay as the type

locality but this appears to be an error, as is the caption for the

figure of the species, which is repeated verbatim from the

preceding figure. The description also contains a number of

contradictions that are likely to have been innaccuracies in data

capture, such as the anal fin count of “iii, 11” rather than III, 10,

and the pectoral fin count of “17” rather than ii, 16. The type has

the latter values typical of C. schoenleinii.

As discussed below. Parent! and Randall (2000: 10)

synonymised Choerodon quadrifasciatus Yu, 1968 with

Choerodon schoenleinii but that name is considered here to be

a synonym of C. azurio.

Choerodon schoenleinii is a common species that reaches

a relatively large size, explaining its frequency in fish markets

throughout the western extreme of the tropical Pacific and

consequently its representation in museum collections. The

relatively small but prominent black spot on the scaly sheath at

the base of the dorsal fin sets it apart from adults of other

species, with the possible exception of C. monostigma, which

has a larger black spot setting higher on the fin and 14 rather

than 16 or 17 segmented pectoral fin rays.

Ebisawa et al. (1995) studied reproduction and sex change

in this species.

Material examined. 113 specimens, 21-530 mm SL; see

appendix.

M.F. Gomon

Choerodon venustus (De Vis, 1884)

Venus Tuskfish

Figures 16, 17; table 3; appendix.

Choerops venustus De Vis, 1884: 147, Moreton Bay (Queensland).

Choerodon ambiguus Ogilby, 1910b: 100, 19 miles N 30° W from

Double Island Point (Queensland).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10;

pectoral fin rays ii, 14, dorsalmost ray of moderate length

24.9- 46.8% pectoral fin length, ventralmost rays shorter than

those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body deep, 36.4-41.5% SL, head depth 26.1-37.2%

SL, caudal peduncle depth 12.9-15.0% SL; head bluntly

pointed, dorsal profile of snout moderately steep, snout length

11.9- 18.6% SL; predorsal scales approximately 4-6, reaching

forward on dorsal midline to not quite to above posterior edge

of preopercle; cheek with small partially embedded scales in

about 5 or 6 diagonal rows, posteriormost with about 10-12

scales to upper extent of free preopercular edge, reaching

forward noticeably short of corner of upper lip crease above

mouth, with broad naked margin posteriorly and ventrally on

preopercle; row of about 8-11 small scales on subopercle

adjacent preopercular edge extending forward in advance of

middle of ventral preopercular margin; each lateral line scale

with multiple branched laterosensory canal tube; scales above

lateral line about 31/2 or 4; many cephalic sensory canal pores

above and behind eyes; second pair of canines in lower jaw

directed anterodorsally, very little laterally; dorsal and anal

fins with very low basal sheath comprising 1-3 progressively

smaller accessory scales at deepest; posterior lobe of dorsal

and anal fins not quite reaching hypural crease; caudal fin

double emarginate, upper and lower lobes distinctly but not

greatly produced, posterior margin of fin concave, smoothly

curved; pelvic fin reaching to or just past anus, length 21.3-

22.9% SL. (See Table 3 for additional meristic and

morphometric ranges.) Adults bluish green above, red

laterally and white below, each scale on side with blue dot;

juveniles with black spot between lateral line and bases of

middle segmented dorsal fin rays.

Reaches moderately large maximum size, largest specimen

examined 384 mm SL but reported to at least 430 mm by

De Vis (1884).

Pigmentation in alcohol. Juveniles pale with dark spot larger

than pupil above lateral line and below second and third soft

dorsal fin rays, much smaller spot on dorsal edge of caudal

peduncle midway between dorsal fin insertion and hypural

crease; dorsal and caudal fins mostly faintly dusky; other fins

pale. Adults pale, slightly duskier dorsally with dusky spot on

each scale of caudal peduncle, becoming less evident anteriorly;

head with pair of dusky marks adjacent to anteroventral margin

of eye with additional marks posteroventral to eye and

sometimes on snout near upper lip. Fins pale, dorsal and anal

fin with row of dusky spots submarginally and second above

each base; distal half of caudal fin dusky.

Fresh colours. Juveniles brown above with broad black

midlateral smudges and distinct black spot in position occupied

by smaller red spot below segmented dorsal fin rays of adults

(fig. 16A; Kuiter, 2010: 52, fig. B).

Initial phase adults bluish green above, pale crimson

midlaterally and white below; each scale on side with blue

spot (fig. 16B); red blotch midlaterally posterior to pectoral fin

base and smaller red blotch just above lateral line beneath

segmented dorsal fin rays; eye edged with blue, with yellow

patch anteriorly, forward directed mark at middle of patch;

blue lines along edges of lips, upper followed by yellow line;

chin and throat pale blue. Dorsal fin yellow with blue basal

and marginal stripes and intermediate row of blue spots; anal

fin yellow with broad blue submarginal stripe broken

posteriorly and series of blue spots near base; caudal fin

greenish yellow with some middle rays blue. Pectoral fin with

upper rays and bases of other rays blue; base of fin yellow

(Kuiter, 2010: 52, figs. A & C).

Terminal phase individuals with vivid colours and much of

side midlaterally dominated by bright pinkish red colouration

(fig. 16C; Marshall, 1964: colour pi. 44, fig. 294; Kuiter, 2010:

52, fig. D).

Etymology. The name venustus is Latin for “beautiful”, no

doubt a reference to the attractive colouration of this species.

Distribution. Confined to eastern Australia from the central

Great Barrier Reef north of Townsville to off Newcastle, New

South Wales (fig. 17). The northernmost verifiable record of

distribution is the south-western end of Wardle Reef (17° 27' S,

146° 30' E) based on a photographic image (J. Johnson, personal

communication). Occurs in shallow inshore weedy habitats,

open substrates with rubble and attached sargassum, ranging

into deeper sponge habitats (Kuiter, 2010: 52) at depths of 20-

60 m.

Comments. Although De Vis gave the length for his Choerops

venustus as “to 22 inches” (550 mm), only a single specimen

(QMB 1.4735, 234 mm SL) in the Queensland Museum

collection appears to be a type for this species. This specimen

no longer retains the life colours but matches the description

morphologically. Ogilby’s (1910b) detailed description of

Choerodon ambiguus likewise matches this species. Only

one of two specimens used for the description (AMS 1.12535,

134 mm SL) is so identified in the Australian Museum

collection, but at least seven others in the collection taken by

the Endeavour in the same general area may be some of the

13 trawled at the same time as reported in the description.

The second “co-type” is the larger of the two, measuring 181

mm TL (144 mm SL), and is likely to be AMS E.1473 with

those lengths.

Taken occasionally along Australia’s Queensland coast by

commercial trawlers at a marketable size, this attractive

species is distinguishable from most other congeners in the

area by the concave to forked posterior margin of the caudal

fin, which is truncate to rounded in others, apart from C. vitta

that is recognisable by its more symmetrically pointed head.

Material examined. 56 specimens, 48-384 mm SL; see

appendix.

Review of Choerodon tuskfishes

100 mm

Figure 16. Choerodon venustus. A, Juvenile, BPBM 14507, 70 mm SL, One Tree Island, Queensland, Australia, photo by J. Randall, BPBM; B,

Initial phase adult BPBM 15972, 241 mm SL, Heron Island, Australia, photo by J. Randall, BPBM; C, Terminal phase adult, CSIRO H 4307-13,

395 mm SL, Queensland, photo by G. Yearsley, CSIRO.

M.F. Gomon

Figure 17. Distribution of Choerodon venustus based on specimens examined.

Choerodon (Aspiurochilus) Fowler, 1956

Tables 2 & 4

Lepidaplois {Aspiurochilus) Fowler, 1956: 176, type species -

Crenilabrus stejnegeri Ishikawa (= C. azurio) by monotypy.

Diagnosis. Dorsal fin rays XII, 8 or XIII, 7, rarely XI, 9; anal

fin rays III, 10; pectoral fin rays ii, 14, rarely 15, dorsalmost ray

of moderate length 23.9-45.2% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular to broadly rounded; body moderately deep,

34.8-45.9% SL, head depth 26.6-42.2% SL, caudal peduncle

depth 12.2-16.8% SL; head blunt, dorsal profile of snout

moderately steep to steep, snout length 9.9-16.7% SL; predorsal

scales approximately 5-11, reaching forward on dorsal midline

to or just in advance of posterior edge of preopercle, to above

middle of eye or to somewhere between; cheek with small

imbricate to mostly embedded scales in about 4-9 diagonal

rows, posteriormost with about 9-17 scales to upper extent of

free preopercular edge, reaching forward to or almost to corner

of upper lip crease above mouth, with broad naked margin

posteriorly and ventrally on preopercle; 1-4 rows (only about 2

scales in second row when present) of small scales on

subopercle adjacent preopercular edge extending forward to or

just short of anterior end of ventral preopercular margin, with

about 6-11 scales in outermost row; each lateral line scale with

unbranched to multiple branching laterosensory canal tube;

scales above lateral line about IVi or 3; cephalic sensory canal

pores relatively few to moderately numerous confined to lines

or short branches associated with major canals; second pair of

canines in lower jaw directed anterodorsally and slightly

laterally, often strongly curved laterally; dorsal and anal fins

with low to very low basal sheath comprising 1-3 progressively

smaller accessory scales at deepest; posterior lobe of dorsal

and anal fins reaching at most to base of first anal spine, usually

to or just short of hypural crease; caudal fin truncate to

distinctly rounded or with concave posterior profile. (See Table

2 for additional meristic and morphometric ranges.) Bicolour

pattern in adults of 3 of 5 species, dusky anteriorly and paler

Review of Choerodon tuskfishes

posteriorly, contrasting areas separated by diagonal interface.

Reaches moderately large size, ranging in maximum

lengths of from about 225 to more than 430 mm SL.

Comments. Fowler (1956: 176) incorrectly referred Ishikawa’s

(1904) Crenilabrus stejnegeri to Lepidaplois (= Bodianus),

separating it from the other five species he recognised as

occurring in Chinese waters by a suite of characters, some of

which conflicted with his diagnosis for that genus. The same

characters were considered a justification for placing it in a new

subgenus AspiurochUus. Although clearly not a group name for

an assemblage of Bodianus, the name is available for species

within Choerodon to which C. stenjegeri (= C. azurio) belongs.

The subgenus comprises five species, one of which is described

here as new. Species of this subgenus (fig. 1, clade 2) are

most similar to those of the subgenus Choerodon (fig. 1, clade

3) in having a rather deep body without characteristic

modifications, some differing from members of that assemblage

in having XII, 8 rather than XIII, 7 dorsal fin elements and all

retaining 14 branched, segmented rays in the pectoral fin as

well as scales on the subopercle reaching to or almost to the

anterior extent of the ventral subopercular margin. Five of the

nine species of the subgenus Choerodon typically have 13 or

15-17 branched rays in the pectoral fin and only two of the nine

species have scales on the subopercle reaching in advance of

the midpoint between the corner of the preopercle and the

anterior extent of the ventral subopercular margin, with several

considerably more reduced. Most reach a moderate size

exceeding 250 mm SL, although one, C. monostigma, has only

been verifiably recorded to about 160 mm SL.

Choerodon azurio (Jordan & Snyder, 1901)

Azurio Tuskfish

Figures 18, 19; table 4; appendix.

Labrus japonicus Valenciennes, in Cuvier & Valenciennes, 1839:

99, Japan.

Choerops azurio Jordan & Snyder, 1901: 747, a replacement name

for Labrus japonicus, Valenciennes, a junior homonym of Labrus

japonicus Houttuyn (1782).

Crenilabrus stejnegeri Ishikawa, 1904: 13, pi. VI, fig. 2,

Miyakojima (Japan).

Choerodon quadrifasciatus Yu, 1968: 11, fig, 5, Tongkong

(Taiwan).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10, rarely 9;

pectoral fin rays ii, 14, rarely 15, dorsalmost ray dorsalmost ray

of moderate length 29.3-41.4% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body deep, 34.8-45.9% SL, head depth 26.6-

42.2% SL, caudal peduncle depth 14.7-16.8% SL; head blunt,

dorsal profile of snout steep, snout length 10.2-15.5% SL;

predorsal scales approximately 5-7, reaching forward on dorsal

midline to about midpoint between posterior extent of orbit and

posterior edge of preopercle; cheek with small imbricate to

partially embedded scales in about 8 diagonal rows,

posteriormost with about 10 scales to upper extent of free

preopercular edge, reaching forward to or almost to corner of

upper lip crease above mouth, with very broad naked margin

posteriorly and ventrally on preopercle; row of about 9 or 10

small scales on subopercle adjacent preopercular edge

extending forward to about anterior end of ventral preopercular

margin, about 3 rows dorsally; each lateral line scale with

mutlple branching laterosensory canal tube; scales above

lateral line about 214; cephalic sensory canal pores relatively

few confined to lines or short branches associated with major

canals, more numerous above eyes; second pair of canines in

lower jaw directed anterodorsally and slightly laterally, curved

posterolaterally; dorsal and anal fins with low basal sheath

comprising 1-3 progressively smaller accessory scales at

deepest; posterior lobe of dorsal and anal fins reaching to of

just short of hypural crease; caudal fin truncate to slightly

rounded; pelvic fin length reaching to or just short of anus,

length 17.8-23.6% SL. (See Table 4 for additional meristic and

morphometric ranges.) Adults with broad dark oblique band

angled from pectoral fin base towards centre of dorsal fin base

with broad pale band bordering it posteriorly.

Reaches moderately large maximum size, largest specimen

examined 308 mm SL.

Pigmentation in alcohol. Juveniles dark dusky with 6 or 1

somewhat obscure narrow pale vertical bands crossing side

from above and behind eye to caudal fin base, anterior 2

crossing nape and merging with pale underside of head with

broad dusky bands directed ventrally from eye and adjacent

posterior margin of operculum; dusky banddike interspaces on

sides extending onto dorsal fin near anterior, central and

posterior spines and onto front, middle and posterior end of

anal fin, with prominent ocellated dark spot posteriorly on

dorsal fin and at middle of anal fin, with smaller dark spots at

front of dorsal fin and basally on pelvic fin; caudal fin and

posterior ends of dorsal and anal fins transparent. Initial phase

with pale head and dusky dorsum anterior to darker oblique

band angled from base of pectoral fin towards bases of spines

at centre of dorsal fin, followed by broad pale band and pale

dusky area posteriorly; scales posteriorly on side with dusky

vertical bar to spot in large individuals; spinous portion of

dorsal fin dusky with darker blotch covering last spine or two,

pale posteriorly; anal fin pale with dusky blotches; caudal fin

dusky; pectoral and pelvic fins pale. Terminal phase similar to

initial phase but duskier overall. Juveniles dusky with darker

mottling; dark dusky blotch at anterior ends of dorsal and anal

fins, prominent pale-edged dark spot on dorsal fin centred on

first few rays and dusky blotches basally on anal fin posteriorly;

pelvic fin pale with dusky blotch basally.

Fresh colours. Juveniles olive brown to brown with 5 narrow

white bands, first 2 crossing predorsal, 3rd below 4th or 5th

dorsal fin spines, 4th below last couple of dorsal fin spines and

5th at posterior end of dorsal fin base (fig. 18A); 2 narrow

broken black stripes on and above lateral line; head brown

dorsally and white ventrally with brown band from eye to tip of

snout and second below eye. Dorsal, anal and pelvic fins same

colour as side adjacent with continuation of white bands on

side; white ocellated large black spot near middle of segmented

ray portion of dorsal and anal fins and less defined ocelli

dorsally and ventrally on caudal fin base, caudal and pectoral

fins hyaline (Masuda et al., 1984: pi. 193, fig. C; Okamura &

M.F. Gomon

Figure 18. Choerodon azurio. A, Juvenile, approximately 40 mm SL, Osezaki, Numazu, Shizuoka, Japan, photo by Izuzuki; B, Initial phase

adult, approximately 220 mm SL, Osezaki, Numazu, Shizuoka, Japan, photo by Izuzuki; C, Terminal phase adult, photo by K. Berlin, Creative

Commons Attribution Share Alike 2.0 Generic.

Review of Choerodon tuskfishes

Figure 19. Distributions of Choerodon azurio (circles) and C. monostigma (squares) based on specimens examined (coloured) and collection

registration records (white).

Amaoka, 1997: 465, centre second from top and right top;

Kuiter 2010: 55, fig. A).

Initial phase adults pinkish brown with white underside and

blue bar on each body scale (fig. 18B); prominent oblique

transverse dull olive green band from axilla of peetoral fin to

bases of 8th and 9th dorsal fin spines, immediately adjacent

posteriorly to broad pink or white band; head with 3 short blue

lines directed anteriorly, ventralmost longest and directed

slightly downward; 4th blue line oriented horizontally below

posterior half of eye. Dorsal and anal fins brown with narrow

blue distal margins; caudal fin brown with narrow blue dorsal

and ventral margins, broadest at posterior corners. Peetoral fin

hyaline; pelvic fin with pair of blue lengthwise lines separated by

yellow line along leading edge. (Shen, 1993: pi. 144, fig. 1;

Okamura & Amaoka, 1997: 465, centre top; Chen et al., 2010:

384, fig. A; Kuiter, 2010: 55, fig. C; Allen & Erdmann, 2012: 645)

Terminal phase with blue hue dorsally and ventrally on

head and yellow to orange patch posterior to pectoral fin base

(fig. 18C); dark transverse band on side black. Anal fin yellow

basally; caudal fin nearly black (Masuda et al., 1984: pi. 193,

fig. D; Okamura & Amaoka, 1997: 465, left top; Kuiter, 2010:

55, figs B & D).

Etymology. The name azurio is likely to be from the French

azur or Pershian lazhuward for “a blue colour”, in reference to

the blue colouration featuring on adults of this species.

Distribution. Confined to the north-western edge of the North

Pacific (fig. 19) from Tokyo, Japan and the Korean Peninsula to

at least Nho Trang, Vietnam, eastward to the Ogasawara Islands

in the north (Randall et al., 1997: 45). Oceurs on open bottom

near the edges of deep rocky reefs to depths of 20 or 30 m, with

juveniles secluded in rocky outgroups or under ledges on rocky

walls (Kuiter, 2010: 55; Allen & Erdmann, 2012: 645).

Comments. Labrus japonicus proposed by Valenciennes (in

Cuvier & Valenciennes, 1839) for a species occurring in Japanese

waters based on a specimen in the Berlin Museum, presented by

M. Langsdorff is a junior homonym of Labrus japonicus

M.F. Gomon

Houttuyn, 1782 (= Pseudolabrus japonicus) and therefore

unavailable. Jordan and Snyder (1901) recognised the problem

and provided Choerops azurio as a replacement for the name,

incorrectly attributed to Schlegel, without further comment.

Ishikawa’s (1904: 13, pi. VI, fig. 2) Crenilabrus stejnegeri

was based on a juvenile (80 mm TL) of C. azurio with a

developing adult colouration. The type is evidently no longer

extant (Eschmeyer, 2015). Parent! and Randall (2000: 10)

synonymised Choerodon quadrifasciatus Yu, 1968 with

Choerodon schoenleinii, citing Gomon (personal

communication) as authority. Based on Yu’s description, this

referal is contradicted by his pectoral fin count of “2, 14”, in

contrast to ii, 16, the usual number in C. schoenleinii. The

types were not found at the Pisces Collection of the National

Museum of Marine Biology and Aquarium, Pintung, Taiwan ,

the current repository for most of Yu’s specimens, and despite

Eschmeyer’s (2015) listing, may not have been transferred

there (Ho, personal communication). The original description

more closely matches C. anchorago (based on anal fin value)

and C. azurio (based on pectoral fin value). The type specimen

was clearly a juvenile, and judging from the figure (Yu, 1968:

11, fig. 5) more closely matches juveniles of C. azurio at the

approximate size of the type. In either case, the name is a

junior synonym.

This is the most frequently encountered member of the

genus in the coastal waters of Japan and China, easily

recognised by its steep, black and white white oblique band

angled upward on its rather deep body from behind the

pectoral fin base. Others in the region with an oblique dark

band on the side are either much more slender or have twelve

spines and eight segmented rays in the dorsal fin.

Material examined. 40 specimens, 24.1-308 mm SL; see

appendix.

Choerodon cypselurus sp. nov.

http://zoobank.org/urn;lsid:zoobank.org:act:AC4D29C3-820C-

4582-A19E-56BCDD3FEBF7

Proposed vernacular: Swallowtail Tuskfish

Figures 20, 21; table 4

Holotype. NSMT-P 110838 (234) Indian Ocean, Seychelles, Saya

de Malha Bank, 10° 35' S, 61° 35' E, 4 December 1978, FV Ryuyo-

maru 2 (orig. no. SKSK8896).

Paratype. (1 specimen, 164 mm SL.) NSMT-P 111345 (164)

Indian Ocean, Seychelles, Saya de Malha Bank, 10° 36' S, 61° 34' E,

30 November 1978, FV Ryuyo-maru 2 (orig. no. SKSK8523).

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 14, rarely 15, dorsalmost ray dorsalmost ray of

moderate length 30.7-30.9% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body deep, 39.8-40.7% SL, head depth 29.2-

31.2% SL, caudal peduncle depth 12.2-13.0% SL; head blunt,

dorsal profile of snout moderately steep, snout length 12.2-

15.8% SL; predorsal scales approximately 5, reaching forward

on dorsal midline to or just in advance of posterior edge of

preopercle; cheek partially covered by small, mostly embedded

scales in about 8 vertical rows, posteriormost with about 10

scales to upper extent of free preopercular edge, reaching

forward almost to corner of upper lip crease, with broad naked

margin posteriorly and ventrally on preopercle; subopercle

with row of about 10 large embedded scales adjacent ventral

edge of preopercle extending forward to just short of below

anterior extent of free ventral preopercular edge; each lateral

line scale with unbranched laterosensory canal tube; scales

above lateral line about 214 ; cephalic sensory canal pores

moderately numerous confined to lines or short branches

associated with major canals; second pair of canines in lower

jaw directed dorsolaterally and slightly posteriorly; dorsal and

anal fins with very low basal sheath comprising 1-3

progressively smaller accessory scales at deepest; posterior

lobe of dorsal and anal fins almost reaching hypural crease;

caudal fin double emarginate, upper and lower lobes distinctly

but not greatly produced, posterior margin of fin concave,

smoothly curved; pelvic fin reaching anus, length 21.4-22.4%

SL. Uniformly pale in preservative.

Description. Dorsal fin rays XII, 8; anal fin rays III, 10; caudal

fin rays 9-1-12-1-9; pectoral fin rays ii, 14 (14-15); vertebrae 10

-I- 17 = 27; pleural ribs ending on 10th vertebra; epipleural ribs

ending on 12th (13th) vertebra; lateral line scales 27 -i- 2; scales

above lateral line 214 ; scales below lateral line approximately

814 ; predorsal scales approximately 5 (6); total gill rakers 9.

(See Table 4 for selected measurements expressed as

percentage of SL or HL.) Body deep, greatest depth at pelvic

fin origin, 2.5 in SL; caudal peduncle moderately slender,

depth 8.2 (7.7) in SL; head large and deep, length 2.6 (2.8) in

SL, depth at posterior extent of orbit 1.2 in HL; snout of

moderate length, 2.4 (2.9) in HL; snout and head broadly

rounded; dorsal outline of forehead and snout convexly curved,

nape a gentle convex curve; eye large, orbit 5.0 (3.7) in HL;

interorbital moderately broad; jaws not attenuate.

Dorsal and anal fin without well developed scaly basal

sheath, small scales at base only. Predorsal scales reaching

forward on dorsal midline of head barely in advance of above

dorsal end of preopercle. Cheek only partially covered by scales,

scales small, mostly embedded, about 8 (9) vertical rows,

posteriormost row with about 10 (13) scales to upper extent of

free preopercular edge, scales reaching forward almost to comer

of upper lip crease, broad naked margin posteriorly and ventrally

on preopercle; opercle fully covered by large embedded scales;

subopercle with row of about 10 large embedded scales adjacent

ventral edge of preopercle extending forward to just short of

below anterior extent of free ventral preopercular edge; lower

jaw naked. Lateral line scales each with unbranched laterosensory

canal tube. Cephalic sensory canal pores moderately numerous

confined to lines or short branches associated with major canals.

Posterior edge of preopercle very finely serrate. Mouth mostly

horizontal, posterior comer of upper lip fold below point just

anterior to forward extent of orbit; lower lip moderately narrow;

upper lip narrow and mostly obscured by skin flap, except at

anterior end of jaw; posterior end of maxilla not exposed. Gill

rakers on first arch small, simple, those dorsally (about 6) and

ventrally (about 5) distinctly finer than fleshy intermediate rakers.

Review of Choerodon tuskfishes

50 mm

Figure 20. Choerodon cypselurus sp. nov. Holotype, NSMT-P 110838, 234 mm SL, Indian Ocean, Seychelles, Saya de Malha Bank, 10° 35’ S,

61° 35’ E, ethanol preserved.

Figure 21. Distributions of Choerodon cypselurus sp. nov. (circles) and C. robustus (squares) based on specimens examined (coloured) and

collection registration records (white).

M.F. Gomon

Table 4. Ranges for selected counts and proportional measurements in species of the subgenus Aspiurochilus. “N” designates number of counts

or measurements. Aberrant values enclosed by parentheses.

Species

C. azurio

C. cypselurus nsp

C. monostigma

C.robustus

C. zamboangae

MERISTIC FEATURES

Holotype Paratype

Dorsal fin

XIII, 7

XII, 8

XIII, 7

XII, 8 (XI, 9)

XII (IX), 8

Anal fin

III, 10

Pectoral fin

ii, 14(15)

ii, 14

ii, 14/15

ii, 14(15)

ii, 14

Vertebrae

10+17 =

10+17

= 27

10+17

= 27

10+17 =

= 27

10+17 =

MORPHOMETRIC FEATURES

Standard length (mm)

58.6-308

234

164

64.1-

158

146-278

95.6-244

% SL

range

mean

range

mean

range

mean

range

mean

Body depth

34.8-45.9

40.3

40.7

39.8

37.4 - 44.3

40.9

31.5-44.0

39.1

36.1-40.7

38.1

Caudal peduncle depth

14.7-16.8

15.9

12.2

13.0

14.6-15.7

15.2

12.8-15.9

14.5

13.3-15.8

14.5

Head length

33.3-37.3

35.5

38.4

35.5

34.9-39.3

36.6

31.3-41.3

36.3

32.7-38.5

36.2

Head depth

26.6-42.2

32.4

31.2

29.2

29.8-35.9

32.9

28.9-39.2

32.3

29.0-34.5

31.9

Orbit diameter

6.2-10.3

8.2

7.7

9.6

7.7-11.3

8.8

5.7-8.7

7.4

7.3-10.1

8.6

Interorbital width

4.7-11.2

8.2

10.4

10.3

7.0-10.0

7.8

7.3-10.9

8.9

6.0-10.3

8.5

Snout length

10.2-15.5

12.3

15.8

12.2

9.9-14.8

13.2

12.6-16.7

14.6

12.3-16.2

14.1

Dorsal fin base length

50.7-60.3

56.4

53.6

53.1

52.1-59.6

55.9

49.8-58.2

55.0

49.7-58.2

54.7

Dorsal fin last spine length

9.4-13.8

12.1

8.0-12.4

10.7

9.3-13.3

10.6

8.8-14.7

11.6

Dorsal fin posterior lobe length

12.6-15.5

13.8

12.7

11.4

12.9-16.8

14.9

12.0-14.6

13.1

11.1-13.6

12.2

Anal fin base length

23.3-30.1

26.4

23.0

25.2

24.9-32.8

29.5

23.6 - 28.3

25.9

24.2-27.9

25.8

Anal fin posterior lobe length

11.5-16.1

13.2

14.3

12.0

10.4-19.0

14.4

11.5-15.4

13.3

10.7-14.0

12.5

Pectoral fin length

18.2-29.0

26.3

27.7

27.6

26.3-33.3

30.5

23.1-28.3

26.0

22.2-30.1

26.8

1st pectoral fin ray length

7.5-10.2

9.2

9.3 -14.4

11.0

6.8-10.2

8.2

7.9-11.1

9.3

Pelvic fin length

17.8-23.6

21.5

22.4

21.4

18.8-24.1

21.2

21.4-24.8

22.8

21.1-24.9

22.8

Caudal fin dorsal lobe length

23.3-27.0

25.8

28.7

26.7

23.2-29.5

26.0

24.0-28.0

26.2

24.5-31.6

27.2

Caudal fin central ray length

25.0-30.2

26.7

21.9

19.9

23.7-29.2

25.4

18.8-26.6

23.4

22.1-30.3

25.7

Head length (mm)

21.5-

115

89.9

58.3

24.1-

56.7

52.7-

104

32.7-86.6

%HL

range

mean

range

mean

range

mean

range

mean

Head depth

72.3-114

91.4

81.1

82.2

75.9-100

89.3

80.0-101

89.2

81.2-96.9

88.1

Orbit diameter

16.7-27.8

23.0

20.0

27.0

21.2-29.9

24.0

16.3-25.1

20.6

19.2-27.8

23.7

Interorbital width

13.0-31.2

23.1

27.0

28.9

18.2-26.3

21.2

22.8-28.6

24.8

17.0-28.8

23.2

Snout length

28.9-43.2

34.4

41.1

34.2

26.3-40.8

36.1

35.7-46.8

40.5

33.8 - 43.2

38.5

X 100%

1st pectoral fin ray/pectoral fin length

29.3-41.4

35.3

30.9

30.7

31.6-45.2

36.0

24.7-36.8

31.3

23.9-38.6

33.4

Orbit diameter/Snout length

43.7-96.2

73.6

48.7

79.0

53.9-114

68.2

37.5-68.8

52.1

46.8-82.4

63.1

Upper jaw with 2 prominent anterior canines; third very

small canine mesial to first prominent canine and adjacent

symphysis of jaw; first prominent canine slightly (to distinctly)

longer than second; both canines directed mostly ventrally, first

slightly laterally, second slightly laterally and recurved slightly

posteriorly; dental ridge smooth to slightly rough; posterior

canine apparently absent. Lower jaw with 2 prominent anterior

canines, both as well as anterior pair of upper jaw exposed when

mouth occluded; first canine approximately equal in length to

second; first canine directed anterodorsally, second curved

dorsolaterally; dental ridge smooth anteriorly, followed by f or 2

triangular teeth of moderate size fused with dental ridge and

smaller teeth posteriorly. Vomerine teeth absent.

Dorsal fin spines subequal, pungent; pointed membrane

behind tip of each spine attached basal to and extending

distinctly beyond spine tip, posterior tip of dorsal and anal fin

Review of Choerodon tuskfishes

narrowly rounded, posterior rays not produced; posterior tip of

fins reaching to (or almost to) posterior edge of hypurals.

Caudal fin emarginate, upper and lower lobes pointed, distinctly

but not greatly produced, dorsalmost rays 1.3 times length of

rays at centre of fin, posterior margin concave. Pectoral fin with

dorsalmost ray of moderate length 30.7-30.9% pectoral fin

length, upper rays much longer than lower, posterior edge

oblique, almost straight, dorsoposterior corner pointed,

posteroventral corner angular to slightly rounded. Pelvic fin of

moderate length, posterior tip of fin reaching anus.

Reaches moderately large maximum size, largest specimen

examined 234 mm SL.

Pigmentation in alcohol. Uniformly pale (fig. 20).

Fresh colours. Unknown.

Etymology. The name cypselurus is from the Greek kypselos

for “swallow” and oura for “tail”, in reference to the characteristic

swallowtail-like caudal fin of this species that differs from

those in other members of the C. azurio clade within this genus.

Distribution. Known only from the type series collected on the

Saya de Malha Bank, of the Seychelles, in the western central

Indian Ocean (fig. 21).

Comments. Of the 25 currently described species of Choerodon,

all but eight characteristically have 13 spines and seven

segmented, branched soft rays in the dorsal fin. Five of the

eight species with 12 spines and eight soft rays are members of

the subgenus Pealopesia, distinguished by a slender body and

modified pectoral fin with produced ventral rays that afford the

fin a distinctive sickle-shaped profile. The remaining group of

three, comprising Choerodon fasciatus (Gunther, 1867),

Choerodon robustus (Gunther, 1862) and Choerodon

zamboangae (Seale & Bean, 1907), have a rather deep body

and pectoral fin with rays that are progressively shorter

ventrally. Choerodon cypselurus shares a dorsal fin count and

the latter characteristics with these three, but differs from them

by the emarginate shape of its caudal fin, the three having a

truncate caudal fin without noticeably produced corners.

Choerodon cypselurus is further separable from C. fasciatus

by its pectoral fin count of ii, 14 (rarely 15), versus ii, 13 in the

latter, and number of predorsal scales, approximately five that

extend forward only to above the preopercular margin, in

contrast with 10-14, which reach forward in advance of above

the posterior extent of the orbit on the dorsal midline of the

head. Although the faded condition of the type specimens

precludes a description of life colours, they are unlikely to be

as distinctive as the prominent banding of C. fasciatus, which

perseveres in preservation.

Choerodon monostigma (Ogilby, 1910)

Darkspot Tuskfish

Figures 19, 22; table 4; appendix.

Choirodon monostigma Ogilby, 1910b; 102, 13 miles from Pine

Peak bearing S 58° E (-21° 25 'S, 150° 08 ' E; Queensland).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 14, rarely 15, dorsalmost ray of moderate length 31.6-

45.2% pectoral fin length, ventralmost rays shorter than those

above, posterior edge of fin obliquely straight, dorsoposterior

comer bluntly pointed, posteroventral comer angular; body deep,

37.4-44.3% SL, head depth 29.8-35.9% SL, caudal peduncle

depth 14.6-22.8% SL; head bluntly pointed, dorsal profile of

snout moderately steep, snout length 9.9-14.8% SL; predorsal

scales approximately 7-11, variably reaching forward on dorsal

midline to above posterior extent of eye or to above middle of

eye; cheek with small partially embedded scales in about 5

diagonal rows, posteriormost with about 9 scales to upper extent

of free preopercular edge, reaching forward to above anterior end

of ventral preopercular margin, with very broad naked margin

posteriorly and ventrally on preopercle; about 3 or 4 rows (only

about 2 scales in second to 4th rows when present) of about 8

small scales on subopercle adjacent preopercular edge extending

forward to about anterior end of ventral preopercular margin;

each lateral line scale with mutlple branching laterosensory canal

tube; scales above lateral line about IVi or 3; about 2 rows of

superorbital cephalic sensory canal pores and about 8 vertical

rows of suborbital pores; second pair of canines in lower jaw

directed anterodorsally and slightly laterally; dorsal and anal fins

with very low basal sheath comprising 2 or 3 progressively

smaller accessory scales at deepest; posterior lobe of dorsal and

anal fins reaching to or beyond hypural crease; caudal fin tmncate

to slightly rounded; pelvic fin reaching to base of first anal fin

spine in largest individuals, length 18.8-24.1% SL. (See Table 4

for additional meristic and morphometric ranges.) Pink to green

above with faint brown bands on head and side; prominent black

spot on dorsal fin from 11th to 13th spines; yellow-edged blue

lines on head and fins.

Reaches moderately small maximum size, largest

specimen examined 225 mm SL.

Pigmentation in alcohol. Juveniles pale with 7 or 8 evenly

spaced vertical dusky bands crossing dorsum and sides; fins

pale except for prominent ocellated dark spot between last 3

spines of dorsal fin. Adults similar to juveniles with dusky

bands far less prominent or lacking.

Fresh colours. Juveniles similar to adults with more defined

banding. Initial phase adults rosy pink to pinkish brown above

with narrow brown midlateral stripe and white underside; about 8

faint brown bands most prominent dorsally, first 3 on predorsal

region between back of eye and dorsal fin origin, last at termination

of dorsal fin base (fig. 22A & B); darker brown dashes where bands

cross lateral line; pair of faint blue stripes posteriorly on body

centred on lateral midline; head with yellow-edged blue stripe

from eye to snout, second along maxilla, third short stripe below

posteroventral corner of eye, 4th circling eye dorsoposteriorly;

orange blotch covering much of opercle and yellow blotches below

eye and at corner of mouth. Spinous portion of dorsal fin yellow

with blue to violet midlateral stripe and large blue to violet

ocellated black spot between 11th and 13th spines; soft portion of

dorsal and anal fins blue to mauve with yellow vertical bands and

yellow margin; anal fin yellow with subbasal and marginal blue

stripes; caudal fin blue with narrow yellow bands and tip. Pectoral

fin hyaline; pelvic fin blue with lengthwise yellow lines (Sainsbury

& Kailola, 1984: 257, bottom; Kuiter, 2010: 60, bottom fig.; Allen

& Erdmann, 2012: 647, top right).

M.F. Gomon

Figure 22. Choerodon monostigma. A, Juvenile, CSIRO CA 2169, 114 mm SL, Forestier Island, Western Australia, photo by G. Leyland,

CSIRO; B, Initial phase adult, Fakfak Peninsula, West Papua, Indonesia, photo by G. Allen, WAM; C, Terminal phase adult, CSIRO CA 2170

(146) Port Musgrave, Gulf of Carpentaria, photo compliments of CSIRO.

Review of Choerodon tuskfishes

Terminal phase similar to initial phase but more greenish

blue dorsally and blue lines on head edged with yellow (fig. 22C).

Etymology. The name monostigma is from the Greek monos

for “single” and stigma meaning “spot”, in reference to the

characteristic violet to black spot present between the last three

spines of the dorsal fin of this species.

Distribution. Endemic to northern Australia and the West

Papua region of eastern Indonesia, reaching south to Rowley

Shoals off Western Australia and Port Curtis, Queensland on

Australia’s east coast (fig. 19). Lives on sand, silt, open rubble

and soft-bottom mixed habitats (Kuiter, 2010; Allen &

Erdmann, 2012) at depths of 1-90 m.

Comments. Ogilby’s (1910b: 102) account of Choerodon

monstigma is detailed and leaves no doubt as to the species

described. A single specimen registered as a co-type (AMS

1.12518,104 mm SL) is the smaller of his two types and matches

the description. The larger type, given as 160 mm TL, is likely

to be a Queensland Museum specimen registered as QMB

1.1563 (131 mm SL, 155 mm TL) collected at the type locality.

Ogilby implied that only a single collection locality near Pine

Peak yielded specimens of this species.

A species endemic to northern Australia and southern

New Guinea, C. monostigma appears to be common in coastal

waters at depths of 10-50 m, judging from specimens taken by

commercial fishing and survey vessels. Choerodon

monostigma is unique among relatively deep bodied species in

having a prominent black spot above the scaly basal sheath of

the dorsal fin between the last few spines.

Material examined. 116 specimens, 55.4-225 mm SL; see

appendix.

Choerodon robustus (Gunther, 1862)

Robust Tuskfish

Eigures 21, 23, 24; table 4; appendix.

Xiphochilus robustus Giinther, 1862: 98, Mauritius.

Cossyphus maxillosus Guichenot, 1865: 23, Reunion, nomen

nudum attributed to Valenciennes.

Choerops dodecacanthusBleeker, 1868: 275, Borbonia(Reunion).

Diagnosis. Dorsal fin rays XII, 8, rarely XI, 9; anal fin rays III,

10; pectoral fin rays ii, 14, rarely 15, dorsalmost ray of moderate

length 24.7-36.8% pectoral fin length, ventralmost rays shorter

than those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body deep, 31.5-44.0% SL, head depth 28.9-39.2%

SL, caudal peduncle depth 12.8-15.9% SL; head bluntly

pointed, dorsal profile of snout moderately steep, snout length

12.6-16.7% SL; predorsal scales approximately 5-7, reaching

forward on dorsal midline almost to or slightly in advance of

midpoint between posterior extent of orbit and posterior edge

of preopercle; cheek with small partially embedded scales in

about 6 or 7 diagonal rows, posteriormost with about 16-17

scales to upper extent of free preopercular edge, reaching

forward to or almost to corner of upper lip crease above mouth,

with very broad naked margin posteriorly and ventrally on

preopercle; 1 or 2 rows of about 7-10 small scales (only about

2 scales in second row when present) on subopercle adjacent

preopercular edge extending forward to about anterior end of

ventral preopercular margin; each lateral line scale with

multiple branching laterosensory canal tube; scales above

lateral line about 21 / 2 ; relatively few cephalic sensory canal

pores mostly confined to major canals dorsally, rows of pores

below eyes; second pair of canines in lower jaw directed mostly

dorsally and strongly curved laterally; dorsal and anal fins with

very low basal sheath comprising 1-3 progressively smaller

accessory scales at deepest; posterior lobe of dorsal and anal

fins reaching hypural crease; caudal fin truncafe to slightly

rounded, upper and lower corners only barely produced at most

in large individuals; pelvic fin reaching to or just short of anus,

length 21.4-24.8% SL. (See Table 4 for additional meristic and

morphometric ranges.) Tan above, white below separated by

oblique white band from inner side of pectoral fin base to

dorsal side of caudal peduncle, brown in front of band; most

scales on side with blue bar forming 6 or 7 horizontal blue lines

on caudal peduncle; additional blue lines on head, especially

adjacent eye.

Reaches moderately large maximum size, largest specimen

examined 285 mm SL.

Pigmentation in alcohol. Juveniles unknown. Initial phase

adults dusky anterodorsally, pale posteroventrally, abrupt

demarcation between areas a darker diffuse narrow band

angled from inner base of pectoral fin to posterior end of dorsal

fin base; head slightly dusky above with pale underside; dark

line directed anteriorly from middle of anterior border of eye,

second horizontal line beneath eye from anterior third of eye

nearly to preopercular edge; submarginal dark line on upper

lip; anteriorly tapering dusky horizontal stripe on lower jaw

from symphysis to anterior end of preoperculer edge; opercular

margin with broad dusky margin posteroventrally; dorsal fin

dusky with pale distal portion posteriorly and narrow dark

dusky distal edge; anal fin pale with dusky distal margin and

second narrow dusky stripe separated from margin by narrow

pale stripe; caudal fin dusky with fine pale and dusky

vermiculations posteriorly and narrow dark dusky dorsal and

ventral margins at corners; pectoral and pelvic fins pale.

Terminal phase adults similar to initial phase with more

pronounced markings on head; scales near middle of side in

pale area posteriorly with vertical dusky blotch or spot that

align to form faint dusky stripes on caudal peduncle and base

of tail.

Fresh colours. Juveniles unknown.

Adults fawn above, white below with broad oblique white

band from axilla of pectoral fin to dorsal side of caudal

peduncle (fig 23); side adjacent to dorsal edge of pale band

dark brown; each scale on side apart from those on band and

ventrally on body with pale blue vertical line; 6 or 7 horizontal

blue lines on caudal peduncle; lips cobalt blue; blue lines also

through centre of orbit, along base of orbit, below orbit, on

operculum parallel to posterior edge of preopercle; opercular

membrane blue. Dorsal fin yellow with blue basal and distal

margins and blue midlateral stripe; anal fin yellow with series

of blue spots at base, blue distal margin and distinct yellow

M.F. Gomon

Figure 23. Choerodon robustus. A, Initial phase adult, CSIRO H 7989-01,146 mm SL, Pelabuhanratu, West Java, Indonesia, photo by W. White,

CSIRO; B, Presumed initial phase adult, Oman, photo by K. Wilson; C, Terminal phase adult, MZB 23115, 220 mm SL, Pelabuhanratu, West

Java, Indonesia, photo by W White, CSIRO.

Review of Choerodon tuskfishes

Figure 24. Pigmentation of preserved specimens showing oblique separation of dusky and pale areas on the body and caudal fin pattern. A,

Choerodon robustus, CSIROH 7989-01,146 mm SL, lndonesia,West Java, Pelabuhanratu, 7° 00’ S, 106° 30’ E, 11 March 2009; B, C. zamboangae,

NMV A 31432-001, 157 mm SL, Indonesia, Lombok, Tanjung Luar, 8° 48’ S, 116° 2’ E, 26 Lebruary 2011; photo by C. Devine, CSIRO.

submarginal stripe; caudal fin with blue lines along rays.

Pectoral and pelvic fins pale fawn; pelvic with blue marginal

line and yellow submarginal line along leading edge (Bleeker,

1874: pi. 3; Masuda et al., 1984: pi. 194, fig. D; Kuiter, 2010:

54, bottom figs A-C; Allen & Erdmann, 2012: 647 bottom;

White et al., 2013: 267, fig. 89.16).

Etymology. The name robustus is a Latin word meaning “hard

and strong like oak”, perhaps to contrast this species with

another considered by Gunther as appropriately referred to the

genus Xiphochilus {=Xiphocheilus), X. typus, a much more

slender species.

Distribution. The distribution of the species is remarkably

broad for members of the genus with verified specimens known

from Eilat in the Red Sea and Persian Gulf to Mauritius and

Reunion, the central Indian Ocean, including Sri Lanka and the

Seychelles, north-eastern Indian Ocean west of the Malayan

Peninsula (Kyushin et al., 1977: 296), the Indian Ocean coasts

of Indonesia off eastern Java and Bali, and Wakayama, Japan

(Mabuchi et al., 2002: 388, fig. lA) in the north-western Pacific

(fig. 21) at depths of 20-120 m. Specimens collected off north¬

western Australia in collections initially identied as C. robustus

have proven to be large individuals of C. zamboangae.

Comments. Xiphochilus robustus, described by Gunther (1862:

98) in his catalogue of fishes in the British Museum, was based

on a mounted specimen (BMNH 1840.12.12.10, 275 mm SL)

from Dr Janvier’s collection obtained in Mauritius that retained

little of its original colour pattern. The species was subsequently

figured in Playfair and Gunther (1866: pi. XII, fig. 3) from a

specimen collected in Zanzibar (presumably BMNH

1867.3.9.20, skin, 242 mm SL). Based on collection specimens,

the species is a reasonably common example of the genus in

that area.

Guichenot’s (1865: 23, 28) Cossyphus maxillosus,

apparently so named for the form of the species’ jaws and

attributed to Valenciennes, is regarded as a nomen nudum

because it appears elsewhere only in synonymies. Three dried

and mounted specimens from Reunion identified as this

species are in the MNHN collection (A.8264, 275 mm SL;

A.8265, 245 mm SL; A.8266, 256 mm SL). All are specimens

of C. robustus.

Bleeker (1868: 275) based his Choerops dodecacanthus on

a specimen from Mauritius, distinguishing it from C. robustus

by the less obtuse profile of the head and the presence of a

violet bordered yellow spot on the opercle as figured in Bleeker

(1874: pi. 3). A specimen in the Leiden museum (RMNH 6534,

M.F. Gomon

204 mm SL, 240 mm TL) is regarded as the type (Eschmeyer,

2015) but is slightly shorter that the 260 mm given by Bleeker.

This specimen otherwise matches the description and is

identifiable as C. robustus. As discussed below, Parent! and

Randall (2000: 10) synonymised C. pescadoresis (as C.

pescadorensis) with C. robustus but the name is considered

here to be a synonym of C. zamboangae.

Choerodon robustus and C. zamboangae are among the

most often confused species in the genus in collections and the

literature because the diagnostic colour patterns are faint,

especially in preserved material. Both are bicoloured with the

demarcation between the slightly darker anterior pigmentation

obliquely angled from behind the pectoral fin base

dorsoposteriorly to the dorsal fin base. In specimens where it

is still evident, the angle of separation is shallower in the

former, ending near the posterior end of the dorsal fin base, but

terminating near the middle of the fin base in the latter (fig.

24). The caudal fin of C. robustus often retains remnants of a

reticulate pattern in advance of the darker posterior margin

posteriorly, rather than being more uniformly pale as in C.

zamboangae.

Mabuchi et al. (2002) compared mitochonial DNA of two

colour variants of specimens from Japanese waters identified

as C. robustus, concluding that they represent separate species.

Judging from the accompanying figures (Mabuchi et al., 2002:

fig. 1), their type A is C. robustus and their type B is most

likely C. zamboangae, although the image of the latter may be

that of a terminal phase individual in breeding colouration

rather than a more typical pattern.

Sequences attributed to specimens of C. robustus involved

in the analysis performed by Puckridge et al. (2015) are only

from Japanese and Indonesian material and therefore do not

represent specimens from or near the type locality of the

species in the distant western Indian Ocean. To test a

hypothesis that individuals in the two widely separated areas

represent different taxa, a sequence from a specimen collected

on the Mascarene Ridge in the western Indian Ocean was

added to the data set in the reanalysis of relationships described

above. The outcome confirmed that sequences for specimens

of C. robustus from all three localities have little or no

divergence, thus supporting the identity of the species.

Material examined. 32 specimens, 146-285 mm SL; see

appendix.

Choerodon zamboangae (Seale & Bean, 1907)

Zamboangan Tuskbsh

Figures 24-26; table 4; appendix.

Choerops zamboangae Seale and Bean, 1907: 236, fig. 6,

Zamboanga (Philippines).

Choerodon melanostigma Fowler and Bean, 1928: 199, Jolo

Market, Jolo (Philippines).

Choerodon pescadorensis Yu, 1968: 10, fig. 4, Pescadores

(Taiwan).

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 14, dorsalmost ray of moderate length 23.9-38.6%

pectoral fin length, ventralmost rays shorter than those above.

posterior edge of fin obliquely straight, dorsoposterior corner

bluntly pointed, posteroventral corner angular; body deep, 36.1-

40.7% SL, head depth 29.0-34.5% SL, caudal peduncle depth

13.3-15.8% SL; head blunt, dorsal profile of snout steep, snout

length 12.3-16.2% SL; predorsal scales approximately 7 or 8,

variably reaching forward on dorsal midline almost to or

distinctly in advance of midpoint between posterior extent of

orbit and posterior edge of preopercle; cheek with small partially

embedded scales in about 6-9 diagonal rows, posteriormost with

about 10 or 11 scales to upper extent of free preopercular edge,

reaching forward just in advance of corner of upper lip crease

above mouth, with very broad naked margin posteriorly and

ventrally on preopercle; 1 or 2 rows of about 6-11 small scales

(only 1-4 scales in second row when present) on subopercle

adjacent preopercular edge extending forward to about anterior

end of ventral preopercular margin; each lateral line scale with

multiple branching laterosensory canal tube; scales above lateral

line about IVi or 3; dorsal and anal fins with very low basal

sheath comprising 2 or 3 progressively smaller accessory scales

at deepest; posterior lobe of dorsal and anal fins reaching to or

beyond hypural crease; cephalic sensory canal pores numerous

dorsoposterior to eye but relatively few associated with major

canals elsewhere; second pair of canines in lower jaw directed

mostly dorsally and strongly curved laterally; dorsal and anal

fins with very low basal sheath comprising 1-3 progressively

smaller accessory scales at deepest; posterior lobe of dorsal and

anal fins reaching just past hypural crease; caudal fin truncate to

slightly rounded, upper and lower corners square, only barely

produced at most in large individuals; pelvic fin reaching to or

just short of anus, length 21.1-24.9% SL. (See Table 4 for

additional meristic and morphometric ranges.) Brown to green

above, white below, separated by anteriorly tapering dark brown

wedge-shaped band from behind pectoral fin base to below rear

third of dorsal fin, with nearly horizontal yellow to orange stripe

immediately below; head with blue lines above and below eye

and on lower jaw.

Reaches moderate maximum size, largest specimen

examined 249 mm SL.

Pigmentation in alcohol. Juveniles and subadults pale with

large elongate dark blotch angled anteroventrally from bases of

first few segmented dorsal fin rays, followed posteriorly by

distinctly pale smaller blotch adjacent to dorsal edge of side.

Initial phase adults pale with faint dusky to dark anteroventrally

tapering wedge-shaped blotch below bases of soft dorsal fin

rays directed towards inner side of pectoral fin base; head pale

with short, broad dark line directed anteriorly from middle of

anterior border of eye, second horizontal line beneath eye from

anterior third of eye nearly to preopercular edge; similar dark

mark on top of eye; broad dusky horizontal stripe on lower jaw

from symphysis to anterior end of preoperculer edge; opercular

margin with faint narrow dusky margin posteroventrally;

pectoral fin base with dusky band at base of dorsalmost rays;

dorsal fin pale with narrow dark distal edge becoming slightly

broader posteriorly; anal fin pale with dusky basal and margin

stripes, as well as third narrow dusky stripe separated from

marginal stripe by narrow pale stripe; caudal fin pale to slightly

dusky with darker periphery; pectoral and pelvic fins pale.

Review of Choerodon tuskfishes

Figure 25. Choerodon zamboangae. A, Juvenile, Milne Bay, Papua New Guinea, photo by G. Allen, WAM; B, Initial phase adult, CSIRO H

7219-07, 166 mm SL, Tanjung Luar, Lombok, Indonesia, photo by W. White, CSIRO; C, Terminal phase adult, NMV A 29706-001, 244 mm SL,

north-western Australia, photo by J. Pogonoski, CSIRO.

M.F. Gomon

Figure 26. Distribution of Choerodon zamboangae based on specimens examined (coloured) and collection registration records (white).

Terminal phase adults similar to initial phase with pronounced

dusky area anterodorsally and pale area posteroventrally,

abrupt demarcation between areas angled from inner base of

pectoral fin to bases of last couple of dorsal fin spines, dusky

area quite dark above lateral line; freshly preserved specimens

with immaculate anteroventrally tapering wedge-shaped area

below middle of dorsal fin base adjacent to dusky area; dorsal

fin with similar pigmentation as that on adjacent side.

Fresh colours. Juveniles and subadults with dark brown to

black slanted wedge-shaped patch instead of orange stripe or

patch obscuring stripe mesoposteriorly from midside below

central dorsal fin spines to bases of segmented dorsal fin rays

followed posteriorly by prominent yellow spot at termination of

dorsal fin base (fig. 25A); blue markings on head as in adults

(Allen & Erdmann, 649, centre left).

Initial phase adults red to reddish brown dorsally, white

ventrally with dark reddish brown to black anteriorly tapered

wedge-shaped oblique band angled from midside below

central dorsal fin spines to bases of segmented dorsal fin rays

followed posteriorly by prominent yellow spot at termination

of dorsal fin base, dark wedge overlying angled orange stripe

from axilla of pectoral fin to caudal fin base just above lateral

midline (fig. 25B); cheeks yellow, opercle orange; underside of

head and chest pale blue, extending dorsally to free margin of

opercle and preopercle; 3 short blue lines and spot adjacent to

eye. Dorsal fin pale blue to purple anteriorly, reddish purple

posteriorly with dark blue to purple basal and distal margins

and yellow subdistal stripe; anal fin blue to purple with yellow

basal stripe, spots and and ocelli; caudal fin red with blue to

purple distal margin and yellow submarginal streaks. Pectoral

fin red, base golden; pelvic fin pale blue with blue marginal

and yellow submarginal lines on leading edge. (Chen et al.,

2010: 382, fig. E; Kuiter, 2010: 54, top figs B & C; Allen &

Erdmann, 2012: 649, top right)

Terminal phase adults dark red above, paler below with

horizontal orange stripe from upper end of pectoral fin base to

middle of caudal peduncle (fig. 25C); some with broad oblique

yellow stripe angled from upper side of pectoral fin base

towards bases of last few dorsal fin spines; dorsal side of head

Review of Choerodon tuskfishes

and nape slate green; lips yellow with blue submarginal line;

head colouration otherwise as in initial phase individuals

(Masuda et al., 1984: pi. 194, fig. C, as C. robustus\ Sainsbury

& Kailola, 1984: 259, bottom; Shen, 1993: pi. 144, fig. 5, as C.

robustus’, Shibukawa, Peristiwady & Suharti, in Kimura &

Matsuura, 2003: 147, as C. robustus-, Kuiter, 2010: 54, top figs

A, D & E; Allen & Erdmann, 2012: 649, centre right; White et

al., 2013: 267, fig. 89.18 male).

Etymology. The name zamboangae acknowledges the type

locality of this species, Zamboanga in the Philippines.

Distribution. Reliable occurrences range from Wabuka in

southern Honshu, Japan (Ikeda & Nakabo, 2015: 454, pi. 177-4)

and Taiwan, through the Philippines to Lombok and Raja

Ampat, Indonesia, Timor Leste and north-western Australia

south of Rowley Shoals (fig. 25). Reported from seaward reef

slopes and rubble bottom at depths of 20-140 m (Allen &

Erdmann, 2012: 649).

Comments. Choerops zamboangae was described by Seale and

Bean (1907) from two preserved adult specimens (USNM

57846: 236, 226 mm SL, holotype, USNM 61154, 203 mm SL,

paratype) collected in Zamboanga, Mindinao, Philippines,

with the remnants of an orange lateral stripe present in fresh

material. The types of Eowler and Bean’s (1928: 199)

Choerodon melanostigma are smaller (USNM 89967, 171 mm

SL, holotype; USNM 5578, 95.6 mm SL and USNM, 5579,164

mm SL, paratypes) than the C. zamboangae types and have a

dark wedge-shaped blotch from the midside to the base of the

rear third of the dorsal fin with contrasting pale areas preceding

and following the dark area. Morphologically, the two appear

identical. Kuiter (2010: 54) and Allen and Erdmann (2012: 649)

figured some individuals with the horizontal orange stripe on

the side described by Seale and Bean (1907: 236-237) and

others with the dark marking described by Eowler and Bean

(1928: 199-200) as C. zamboangae, as well as a few with both,

stating that the dark marking is present in juveniles and females

but not in larger, presumably male, individuals. Their

interpretation appears to be justified.

Choerodon pescadoresis Yu, 1968 was synonymised (as

Choerodon pescadorensis) with C. robustus by Parenti &

Randall (2000: 10) on the authority of S. C. Shen (personal

communication). As with C. quadrifasciatus, Yu’s type of C.

pescadoresis was not found at the National Museum of Marine

Biology and Aquarium, Pintung, Taiwan and is thought to be

lost. The original description more closely matches C.

zamboangae and is considered to be a synonym of that species.

Choerodon zamboangae closely resembles C. robustus in

general colouration and in particular the markings on the head

and tail. Specimens of the former examined in Japanese and

Taiwanese collections were frequently misidentified as the

latter. Morphologically, the profile of the head and snout is

more rounded in initial phase adults of C. zamboangae and the

body is slightly shallower. The blue stripe on the lower jaw of

the former broadly wraps around the underside at the front,

while the stripe in C. robustus tapers to a narrow line across

the symphysis. The scales on the sides of C. zamboangae lack

the blue spot or vertical blue line characteristic of C. robustus.

In preservation, the demarcation between the dusky and pale

areas on the side is between the inner side of the pectoral fin

and the base of the last couple of dorsal fin spines in C.

zamboangae, except in smaller individuals that retain

remnants of the dark anteroventrally tapering wedge-shaped

blotch below the bases of the soft dorsal fin rays (fig. 24B). In

C. robustus the two areas are separated between the inner side

of the pectoral fin and the rear end of the dorsal fin base (fig.

24A). The two species appear to be mostly allopatric with C.

robustus, which is by far the more broadly distributed.

Documented overlap areas are confined to southern Japan and

Taiwan. Puckridge et al. (2015: figs 1 & 2d) and the reanalysis

provided above both recovered the two as sister species.

Material examined. 36 specimens examined, 95.6-249 mm

SL; see appendix.

Choerodon (Lienardella) (Eowler & Bean, 1928)

Table 2

Lepidaplois (Lienardella) Fowler & Bean, 1928: 202, type species

- Lepidaplois mirabilis Snyder (= C.fasciatus) by monotypy.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray of moderate length 21.3-34.5%

pectoral fin length, ventralmost rays shorter than those above,

posterior edge of fin obliquely straight, dorsoposterior corner

bluntly pointed, posteroventral corner angular to broadly

rounded; body moderately deep, 40.7-44.7% SL, caudal

peduncle depth 15.1-17.1% SL, head depth 27.8-31.9% SL;

head bluntly pointed, dorsal profile of snout moderately steep,

snout length 10.5-14.9% SL; predorsal scales approximately

10-14, reaching forward on dorsal midline to above centre of

eye; cheek with small partially embedded scales in about eight

or nine diagonal rows, posteriormost with about 10 scales to

upper extent of free preopercular edge, reaching forward

almost to corner of upper lip crease above mouth, with very

broad naked margin posteriorly and ventrally on preopercle; 1

or 2 rows of about 8 small scales (only about 2 scales in second

row when present) on subopercle adjacent preopercular edge

extending forward nearly to anterior end of ventral preopercular

margin; each lateral line scale with unbranched laterosensory

canal tube; scales above lateral line about 3!^; cephalic sensory

canal pores confined to lines or short branches associated with

major canals on top of head, scattered pores more numerous

anteroventral to eye; second pair of canines in lower jaw mostly

straight directed dorsolaterally; dorsal and anal fins with

relatively deep scaley basal sheath comprising approximately

about 2-4 enlarged scales; posterior lobe of dorsal and anal

fins reaching just short of hypural crease; caudal fin truncate,

corners pointed; pelvic fin not reaching anus, length 21.1-

25.6% SL. (See Table 2 for additional meristic and morphometric

ranges.) Head and body with prominent vertical bands, dark

grey and white in juveniles, red to black and white in adults.

Comments. Eowler and Bean (1928) erected Lienardella as a

subgenus of Lepidaplois (= Bodianus), saying they had

examined the type specimen of the type species L. mirabilh

Snyder and believed it to be allied with the subgenus

M.F. Gomon

Lepidaplois. Oddly, the statement immediately followed a

diagnosis of the genus Lepidaplois where they obviously

overlooked the discrepancy in dorsal and anal fin rays counts

between Lepidaplois and L. mirabilis. They also failed to

recognise the similarity of meristic values of L. mirabilis with

those provided in their preceding treatment of Choerodon,

despite the incomplete nature of the diagnosis in the latter. The

name was subsequently elevated to genus by Myers (1939: 88)

and then synonymised with Choerodon by Gomon (1997: 809).

This monotypic subgenus (fig. 1, clade 4a) is strikingly

patterned, the starkly contrasting black and white bands of

adults distinctly unlike colour patterns of other species.

Predorsal scales in the one species reach to above the centre of

the eyes, well in advance of that in all others, except C.

mono stigma and C. vitta, which have predorsal scales

extending almost as far or as far forwards. The scaley basal

sheaths on the dorsal and anal fins are also considerably

deeper than those of other species.

Choerodon fasciatus (Gunther, 1867)

Harlequin Tuskfish

Figures 27, 28; table 2; appendix.

Xiphochilus fasciatus Gunther, 1867: 101, pi. X, Cape York

(Queensland), Australia.

Lepidaplois mirabilis Snyder, 1908: 96, Japan and Riu Kiu

Islands.

Choerodon balerensis Herre, 1950: 149, Baler, Quezon (Tayabas)

Province, Luzon (Philippines).

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray of moderate length 21.3-52.6%

pectoral fin length, ventralmost rays shorter than those above,

posterior edge of fin obliquely straight, dorsoposterior corner

bluntly pointed, posteroventral corner angular to broadly

rounded; body moderately deep, 39.2-44.7% SL, caudal

peduncle depth 15.1-17.1% SL, head depth 27.8-31.9% SL;

head bluntly pointed, dorsal profile of snout moderately steep,

snout length 10.5-14.9% SL; predorsal scales approximately

10-14, reaching forward on dorsal midline to above centre of

eye; cheek with small partially embedded scales in about 8 or 9

diagonal rows, posteriormost with about 10 scales to upper

extent of free preopercular edge, reaching forward almost to

corner of upper lip crease above mouth, with very broad naked

margin posteriorly and ventrally on preopercle; 1 or 2 rows of

about 8 small scales (only about 2 scales in second row when

present) on subopercle adjacent preopercular edge extending

forward nearly to anterior end of ventral preopercular margin;

each lateral line scale with unbranched laterosensory canal

tube; scales above lateral line about 31 / 2 ; cephalic sensory canal

pores confined to lines or short branches associated with major

canals on top of head, scattered pores more numerous

anteroventral to eye; second pair of canines in lower jaw mostly

straight directed dorsolaterally; dorsal and anal fins with

relatively deep scaley basal sheath comprising approximately

about 2-4 enlarged scales; posterior lobe of dorsal and anal

fins reaching just short of hypural crease; caudal fin truncate,

corners pointed; pelvic fin not reaching anus, length 21.1-

25.6% SL. (See Table 2 for additional meristic and morphometric

ranges.) Head and body with prominent vertical bands, dark

grey and white in juveniles, red to black and white in adults.

Reaches moderately small maximum size, largest

specimen examined 188 mm SL but reported in the literature

to reach 300 mm TL.

Pigmentation in alcohol. Juveniles with narrow well-defined

dark-edged dusky bands and contrastingly immaculate

interspaces extending from tip of snout to base of tail, with

prominent ocellated dark spots at front and rear of dorsal fin,

midway along anal fin, and at base of pelvic fin, and much

smaller dark spots dorsally and ventrally on caudal fin base.

Adults pale with prominent broad dark-edged vertical dusky

bands on head and body becoming poorly defined ventrally on

sides and merging into broad dark area below segmented rays

of dorsal fin, continuing narrowly onto base of dorsal fin and

broadly onto base of anal fin; posterior end of caudal peduncle

and caudal fin abruptly pale; pair of dark lines crossing snout in

front of eye with broad dark edged dusky band across top of

head between eyes; anterior 2 broad dusky bands on head

contiguous with those of opposite side ventrally; dorsal, anal

and pelvic fins with narrow dark margins; dark blotch between

first 3 spines of dorsal fin; caudal fin with fine dusky edge

posteriorly; pectoral fin pale.

Fresh colours. Juveniles with similar patterns as adults but

banding brown with dark brown to black edges and with

prominent eye-sized ocellated black spots anteriorly and

posteriorly on dorsal fin, midway along anal fin and on basal

half of pelvic fin. (fig. 27A; Masuda et al., 1984: pi. 194, fig. E,

as Lienardella fasciatua-, Kuiter, 2010: 61, figs C-E)

Adults yellow to white with green dorsum and prominent

violet edged bright red transverse bands on head and side, 3

crossing head and 6 across side; anterior head band covering

snout and jaws, interrupted in front of eye by pair of horizontal

blue lines, second through eye, third encircling head at middle

of operculum (fig. 27B & C); first band on side broad, passing

through dorsal fin origin, subsequent bands narrower and evenly

spaced, last across scaly base of caudal fin, posterior bands

often obscured in life by dark blue to black pigmentation; teeth

blue. Dorsal and anal fins red with blue basal and distal margins,

dorsal fin with black spot between first 2 spines; caudal fin white

often with broad pink distal margin, sometime yellow near

centre. Pectoral fin yellow, red basally; pelvic fin red with blue

anterior and posterior margins. (Masuda et al., 1984: pi. 194,

fig. F, as Lienardella fasciata-, Shen, 1993: pi. 144, fig. 2;

Okamura & Amaoka, 1997: 465, right bottom; Chen et al., 2010:

384, fig. B; Kuiter, 2010: 61, figs A, B & F; Allen & Erdmann,

2012: 646, top; Motomura & Matsuura, 2014: 383, 3 figs)

Etymology. The name fasciatus is Latin for “enveloped with

bands”, in reference to the brilliant red body banding

characterising adults of this species.

Distribution. Anti-equatorial from Okinawa in southern Japan

to the northern Philippines in the Northern Hemisphere, and

Papua New Guinea, the Queensland coast of Australia, Lord

Howe Island and New Caledonia south of the equator (fig. 28).

Usually shelters in caves or beneath ledges in bays and inner

reefs to outer reef lagoons at depths of 4-15 m.

Review of Choerodon tuskfishes

Figure 27. Choerodon fasciatus. A, Juvenile, BPBM 27100,35 mm SL, New Caledonia, photo by J. Randall, BPBM; B, Initial phase adult, BPBM

41249, 140 mm SL, Lizard Island, photo by J. Randall, BPBM; C, MNHN 1980-0782, 162 mm SL, New Caledonia, photo by J. Randall, BPBM.

M.F. Gomon

Figure 28. Distribution of Choerodonfasciatus based on specimens examined (coloured) and collection registration records (white).

Comments. Giinther (1867: 101, pi. X) based his description of

Xiphochilus fasciatus on two dried specimens (BPBM

1867.6.24.3, lectotype, 161 mm SL, and BPBM 1867.6.24.4,

paralectotype, 143 mm SL) from Cape York, Australia. Snyder

(1908: 96) was unlikely to have been familiar with Gunther’s

species because he erred in the generic placement of his

Lepidaplois mirabilis and the two descriptions are remarkably

similar, at least with respect to colouration. The two would be

distinct only if the Northern and Southern Hemisphere

populations are found to be separate species. Genetic

comparisons remain to be done. Similarly, Herre (1950: 149)

must have been unfamiliar with the previous two descriptions

because he, like the other authors, commented on “its brilliant

colors and striking color pattern”.

The prominent colouration described by the authors of the

species and its synonyms is highly attractive to fish fanciers

making it a popular aquarium species in the tropical marine

aquarium trade. The distinctive pattern easily separates the

species from congeners, although the more obscure banding in

juveniles of a number of species and adults of C. graphicus are

no doubt ancestral precursors. Donaldson (1995: 313)

described the courtship and spawning of this species on the

Great Barrier Reef, Queensland.

Material examined. 49 specimens, 31-188 mm SL; see appendix

Choerodon (Lutjanilabrus nsubgen.)

Table 2

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 14, rarely 15, dorsalmost ray of moderate length

25.5-41.7% pectoral fin length, ventralmost rays shorter than

those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular to broadly rounded; body moderately deep, 29.8-39.6%

SL, caudal peduncle depth 12.0-14.9% SL, head depth 22.2-

31.8% SL; head broadly pointed, dorsal profile of snout oblique,

snout length 10.9-15.3% SL; predorsal scales approximately

12-15, reaching forward on dorsal midline to or almost to above

middle of eye; cheek with small partially embedded scales in

Review of Choerodon tuskfishes

about 8 diagonal rows, posteriormost with about 12 or 13 scales

to upper extent of free preopercular edge, reaching forward

almost to corner of upper lip crease above mouth, with broad

naked margin posteriorly and ventrally on preopercle; about 2

rows of 10-12 small scales (2-4 scales in second row dorsally)

on subopercle adjacent preopercular edge extending forward

nearly to anterior end of ventral preopercular margin; each

lateral line scale with multiple branching laterosensory canal

tube; scales above lateral line about 4 or 5; few cephalic sensory

canal pores confined to lines or short branches associated with

major canals; second pair of canines in lower jaw directed

dorsolaterally and curved posteriorly; dorsal and anal fins with

very low basal sheath comprising 1 or 2 slightly smaller

accessory scales at deepest; posterior lobe of dorsal and anal

fins not reaching hypural crease; caudal fin truncate to slightly

rounded. (See Table 2 for additional meristic and morphometric

ranges.) Distinctive colouration with white head and body and

prominent brown to black midlateral stripe from eye to base of

tail at all sizes, although less distinct in largest individuals;

smaller individuals with darkened circular to elongate spot

superimposed on stripe on rear half of caudal peduncle.

Etymology. The group name Lutjanilabrus nsubgen is from the

Malayan ikan lutjang, a vernacular for members of the snapper

genus Lutjanus and labrus, apparently a Greek vernacular for

wrasse, in reference to the snapper-like form of this monotypic

subgenus of labrid.

Comments. Kuiter (2010: 62) placed Choerodon vitta in a

separate unnamed subgenus based on its “unusual form,

superficially like some members of Lutjanidae”. The species

was found by Puckridge et al. (2015: 67, fig. 1) to be a sister

species of Choerodon fasciatus based on genetic evidence.

These two species differ considerably from each other in

overall form, dorsal and pectoral fin counts, caudal fin form,

colouration and apparent habitat preference.

The body form of the sole representative of this subgenus

(fig. 1, clade 4b) is far more symmetrical with a distinctly

pointed head than in species of other subgenera, apart from

the dwarf members of the subgenus Peaolopesia, which have

a more slender appearance and blunter head.

Choerodon vitta Ogilby, 1910

Blackstripe Tuskfish

Figures 29, 30; table 2; appendix.

Choerodon vitta Ogilby, 1910a; 13, Aru Islands (Indonesia).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 14, rarely 15, dorsalmost ray of moderate length

25.5-41.7% pectoral fin length, ventralmost rays shorter than

those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body moderately deep, 29.8-39.6%SL, head depth

22.2-31.8% SL, caudal peduncle depth 12.0-14.9% SL; head

broadly pointed, dorsal profile of snout oblique, snout length

10.9-15.3% SL; predorsal scales approximately 12-15,

reaching forward on dorsal midline to or almost to above

middle of eye; cheek with small partially embedded scales in

about 8 diagonal rows, posteriormost with about 12 or 13 scales

to upper extent of free preopercular edge, reaching forward

almost to corner of upper lip crease above mouth, with broad

naked margin posteriorly and ventrally on preopercle; about 2

rows of 10-12 small scales (2-4 scales in second row dorsally)

on subopercle adjacent preopercular edge extending forward

nearly to anterior end of ventral preopercular margin; each

lateral line scale with mutlple branching laterosensory canal

tube; scales above lateral line about 4 or 5; few cephalic sensory

canal pores confined to lines or short branches associated with

major canals; second pair of canines in lower jaw directed

dorsolaterally and curved posteriorly; dorsal and anal fins with

very low basal sheath comprising 1 or 2 slightly smaller

accessory scales at deepest; posterior lobe of dorsal and anal

fins not reaching hypural crease; caudal fin double emarginate,

upper and lower corners rounded, posterior margin of fin

concave, smoothly curved; pelvic fin reaching to or just short of

anus, length 20.4-25.0% SL. (See Table 2 for additional

meristic and morphometric ranges.) Brown to grey above,

white below with dark midlateral stripe from eye to base of tail,

usually followed by prominent darker spot.

Reaches moderately small maximum size, largest

specimen examined 174 mm SL.

Pigmentation in alcohol. Juveniles pale with broad dark dusky

lateral stripe from snout across lower half of eye to caudal fin

base, intensified as elongate dark spot on rear half of caudal

peduncle and caudal fin base (fig. 29A). Adults pale, slightly

duskier dorsally with distinct dusky stripe on lateral midline

continuing on head beneath eye and crossing snout in front of

eye, ending posteriorly in distinct horizontally elongate spot on

posterior half of caudal peduncle; fins mostly pale, dorsal fin

slightly dusky anteriorly and basally.

Fresh colours. Juveniles white with grey dorsum and black

midlateral stripe.

Adults grey dorsally, white ventrally with broad dark

edged reddish brown stripe midlaterally from ventral half of

eye to caudal peduncle followed by black elongate oval spot on

caudal fin base (fig. 28B & C); head greenish grey above with

yellow on cheeks and above midlateral stripe behind eye, with

2-4 anastomosing, silvery blue lines on cheek and preorbital.

Dorsal and caudal fins grey; anal fin white. (Sainsbury &

Kailola, 1984: 259, centre; Allen, 1985: 2407, fig. 333)

Etymology. The name vitta is Latin for “ribbon” or “stripe”, in

reference to the characteristic dark midlateral stripe on the side

of this species.

Distribution. Confined to northern Australia and the Aru

Islands in south-eastern Indonesia, reaching south to the

Exmouth Gulf in Western Australia and to the southern part

of the Great Barrier Reef on the east coast of Australia (fig.

30). Found on open sand or rubble bottom adjacent coastal

reefs at depths of 4-75 m (Kuiter, 2010: 62; Allen & Erdmann,

2012: 648).

Comments. Ogilby (1910a: 13) described Choerodon vitta from

a 190 mm TL specimen (QMB 1.1555, 148 mm SL) collected at

Dobo in the Aru Islands (Indonesia). The symmetrical, snapper-

M.F. Gomon

Figure 29. Choerodon vitta. A, Juvenile, AMS 1. 18767-009, 33.9 mm SL, Queensland, Great Barrier Reef, Linnet Reef, 14° 47’ S, 145° 20’ E,

preserved, photo by D. Paul, NMV; B, Initial phase adult, CSIRO H 6558-04, 107 mm SL, Torres Strait, north-west of Prince of Wales Island,

Queensland, photo by D. Gledhill, CSIRO; C, Terminal phase adult, CSIRO CA 2163, 147 mm SL, Eighty Mile Beach, Western Australia, photo

compliments of CSIRO.

Review of Choerodon tuskfishes

Figure 30. Distribution of Choerodon vitta based on collection examined (coloured) and collection registration records (white).

like form and dark brown to black midlateral stripe with black

spot on the base of the caudal fin are diagnostic for the species.

Material examined. 40 specimens, 29.5-174 mm SL; see

appendix.

Choerodon (Xiphocheilus) Bleeker, 1856

Table 2

Xiphocheilus Bleeker, 1856: 223 - Xiphocheilos typus Bleeker,

1856, by monotypy.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10, rarely

9 or 11; pectoral fin rays ii, 14, rarely 12, 13 or 16, dorsalmost

ray of moderate length 24.6-33.8% pectoral fin length,

ventralmost rays shorter than those above, posterior edge of

fin obliquely straight, dorsoposterior corner bluntly pointed,

posteroventral corner angular; body shallow, 24.6-33.7% SL,

head depth 21.8-25.5% SL, caudal peduncle depth 12.8-

14.2% SL; head blunt, dorsal profile of snout steep, snout

snout length 8.3-9.8% SL; predorsal scales approximately

7-10, reaching forward on dorsal midline to or slightly in

advance of above centre of eye; cheek covered by large

imbricate scales in about 3 nearly vertical rows, posteriormost

with about 6 scales to upper extent of free preopercular edge,

reaching forward in advance of corner of upper lip crease

above mouth, below posterior half of eye, with narrow naked

margin posteriorly and ventrally on preopercle; about 5 large

scales covering subopercle forward to about anterior end of

ventral preopercular margin; each lateral line scale with

multiple branching laterosensory canal tube; scales above

lateral line about 2 or 1 V 2 \ cephalic sensory canal pores

numerous but confined to lines or short branches associated

with major canals; second pair of canines in lower jaw

directed anterodorsally and slightly laterally; dorsal and anal

fins without basal sheath, 1-3 progressively smaller accessory

scales adjacent to fin base; posterior lobe of dorsal and anal

fins barely reaching hypural crease at most; caudal fin

truncate to slightly rounded, upper corner slightly produced

in largest individuals; pelvic fin reaching well short of anus in

small individuals, to anus in largest, length 18.5-26.1% SL.

M.F. Gomon

(See Table 2 for additional meristic and morphometric

ranges.) Olive above, somewhat orange to pink below, with

black bar between lateral line and base of middle dorsal fin

spine; numerous posteroventrally angled oblique yellow-

edged blue lines on side; additional, anteroverntrally angled

lines on head.

Reaches small maximum size, largest specimen examined

110 mm SL.

Comments. The name Xiphocheilus has appeared under

several spellings, including within Bleeker’s (1856: 223)

initial description of the genus and the accompanying

description of the type species Xiphoceilos typus Bleeker

(1856: 224). The latter is regarded as a typographical error.

Subsequent referral of species to the nominal genus by

Gunther introduced the spelling Xiphochilus (Gunther, 1861,

1862, 1867, 1880; Playfair & Gunther, 1866). In more recent

years, regarded as a valid genus, the generic boundary of the

taxon was much less clear in the years following its proposal,

as Gunther alone referred species in four of the six subgenera

of Choerodon recognised here to it. Gomon (1997: 861)

acknowledged that Xiphocheilus has characters consistent

with species of Choerodon but hypothesised it diverged prior

to species within that genus primarily based on patterns of

head squamation. Puckeridge et al. (2015: fig. 1) found it to

be sister to a clade within Choerodon corresponding to the

subgenus Choerodon (Pealopesid). Based on genetic

evidence inferring interrelationships, reduction or expansion

of head squamation is likely not to have been unidirectional

within the complex or reduction has occurred independently

on more than one occasion. The sister relationship of C.

typus with the complex of species constituting C.

(Peaolopesia) hypothesised is supported by the shared small

maximum size attained but larger first pectoral fin ray

present in C. typus that approaches the condition in other

subgenera. Choerodon (Xiphocheilus) is therefore recognised

as a monotypic subgenus (fig. 1, clade lb).

Choerodon typus (Bleeker, 1856)

Bluetooth Tuskfish

Figures 31,32; table 2; appendix.

Xiphocheilos typus Bleeker, 1856: 224, Nias.

Xiphochilus quadrimaculatus Gunther, 1880: 45, pi. XX, fig. c,

Arafura Sea.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10, rarely 9 or

11; pectoral fin rays ii, 14, rarely 12, 13 or 16, dorsalmost ray of

moderate length 24.6-33.8% pectoral fin length, ventralmost

rays shorter than those above, posterior edge of fin obliquely

straight, dorsoposterior corner bluntly pointed, posteroventral

corner angular; body shallow, 24.6-31.5% SL, head depth 22.1-

26.9% SL, caudal peduncle depth 12.7-15.2% SL; head blunt,

dorsal profile of snout steep, snout snout length 7.9-9.8% SL;

predorsal scales approximately 7-10, reaching forward on dorsal

midline to or slightly in advance of above centre of eye; cheek

covered by large imbricate scales in about 3 nearly vertical rows,

poster!ormost with about 6 scales to upper extent of free

preopercular edge, reaching forward in advance of corner of

upper lip crease above mouth, below posterior half of eye, with

narrow naked margin posteriorly and ventrally on preopercle;

about 5 large scales covering subopercle forward to about

anterior end of ventral preopercular margin; each lateral line

scale with multiple branching laterosensory canal tube; scales

above lateral line about 2 or 21 / 2 ; cephalic sensory canal pores

numerous but confined to lines or short branches associated with

major canals; second pair of canines in lower jaw directed

anterodorsally and slightly laterally; dorsal and anal fins without

basal sheath, 1-3 progressively smaller accessory scales adjacent

to fin base; posterior lobe of dorsal and anal fins barely reaching

hypural crease at most; caudal fin truncate to slightly rounded,

upper corner slightly produced in largest individuals; pelvic fin

reaching well short of anus in small individuals, just beyond anus

in largest, length 18.5-26.5% SL. (See Table 2 for additional

meristic and morphometric ranges.) Olive above, somewhat

orange to pink below, with black bar between lateral line and

base of middle dorsal fin spine; numerous posteroventrally

angled oblique yellow-edged blue lines on side; additional,

anteroverntrally angled lines on head.

Reaches small maximum size, largest specimen examined

123 mm SL.

Pigmentation in alcohol. Juvenile patterns unclear. Adults

pale, slightly duskier above with dusky horizontal stripe above

lateral midline from above pectoral fin base to hypural crease

and vertical dusky mark above lateral line below central dorsal

fin spines; pectoral fin base dusky darkest along proximal ends

of rays dorsally; head with diffuse dusky blotch on opercle and

underlying blue bones faintly evident. Dorsal, anal, pectoral

and pelvic fins pale, anal with broad dusky distal stripe; caudal

fin pale, broadly blue basally and with broad dark dusky

marginal band on lower half of fin.

Fresh colours. Olivaceous above, somewhat orange or pink

below (fig. 31); broad black short band dorsally on side below

about 6th dorsal fin spine; numerous narrow evenly spaced

posteroventrally angled, oblique pale blue bands outlined with

yellowish orange on side; head with additional bands, first 2

directed anteroventrally from eye, third anoteroventrally from

origin of lateral line across cheek to corner of mouth, remaining

1 or 2 on head vertically adjacent to posterior edge of preopercle.

Dorsal fin blue with 2 or 3 narrow yellowish orange or pink

stripes; anal fin yellowish orange with numerous narrow blue

bands basally; caudal fin yellowish orange with about 5-8

narrow blue bands; fin suffused with black centrally. Pectoral

fin transparent to orange with blue band edged with black

posteriorly on fleshy fin base; pelvic fin yellowish orange with

blue leading edge (Sainsbury & Kailola, 1984: 261, bottom;

Kuiter, 2010: 66, figs A-C; Allen & Erdmann, 2012: 726).

Etymology. The name typus is from the Greek typos for “figure

or mark”, most likely a reference to the black band dorsally on

the side below the central dorsal fin spines.

Distribution. Occurs in the tropical western Pacific Ocean

from at least Taiwan to the Gulf of Carpentaria, extending into

the north-eastern Indian Ocean westward to about Madras,

India (fig. 32). Lives over sand or rubble bottom at depths of

15-60 m, occasionally near reefs (Allen & Erdmann, 2012).

Review of Choerodon tuskfishes

Figure 31. Choerodon typus. Initial phase adult, USNM 424685, 105 mm SL, Pasil market, Cebu, Philippines, photo by J. Williams, USNM.

20 °

0 °

20 °

0 °

20 °

Figure 32. Distribution of Choerodon typus based on specimens examined.

M.F. Gomon

Comments. Bleeker (1856: 223-224) described his new genus

and species X. typus in considerable detail from a 128”’ (102

mm SL) specimen collected off the island of Nias on the

western coast of Sumatra, Indonesia, and now lodged in the

Natural History Museum (BMNH 1864.5.15.35). Gunther

(1880: 45) presented his description of his Xiphochilus

quadrimaculatus in much less detail and failed to compare it

with previous species referred to the genus. His type specimen

(BMNH 1879.5.14.34) from the Arafura Sea is smaller than

Bleeker’s type but clearly an example of the same species.

As discussed above, C. typus most closedly resembles

slender species of the subgenus Choerodon (Pealopesia) (fig. 1,

clade la), differing from them in having a longer first pectoral

fin ray equal to 24.6-33.8% (versus 2.8-17.5%) of the pectoral

fin length, 7-10 predorsal scales reaching forward on the dorsal

midline to or slightly in advance of above the centre of the eye,

in contrast to 5-8 scales reaching forward no farther than above

the posterior extent of the eye and its cheek covered by large

imbricate scales rather than small, partially or almost completely

embedded scales extending over far less of the cheek. Choerodon

typus is recognised here as a monotypic subgenus (fig. 1, clade

lb) based on these considerable morphological differences.

Material examined. 45 specimens, 38.6-123 mm SL; see

appendix.

Choerodon (Peaolopesia) Smith, 1953

Tables 2, 5, 6

Peaolopesia Smith, 1953: 520, type species - Xiphochilus

gymnogenys Playfair & Gunther, by monotypy.

Choerodonoides Kamohara, 1958: 2, type species -

Choerodonoides japonicus Kamohara (= C. albofasciatus), by

monotypy.

Diagnosis. Dorsal fin rays XII, 8 or XIII, 7; anal fin rays III, 10;

pectoral fin rays ii, 13-14, rarely 14, dorsalmost ray short 2.8-

17.5% pectoral fin length; ventralmost pectoral fin ray distinctly

longer than those immediately above, posterior edge of fin falcate,

dorsoposterior corner bluntly pointed and posteroventral comer

sharply pointed or ventralmost rays shorter than those above,

posterior edge of fin obliquely straight, dorsoposterior comer

bluntly pointed and posteroventral comer angular to broadly

rounded; body moderately shallow, 24.4—37.7% SL, caudal

peduncle depth 9.2-13.3% SL, head depth 19.4—29.22% SL;

head blunt, dorsal profile of snout steep, snout length 8.0-15.5%

SL; predorsal scales approximately 5-8, reaching forward on

dorsal midline almost to above posterior edge of preopercle, to

above posterior extent of eye or somewhere between; cheek with

small partially or almost completely embedded scales in about

2-10 diagonal rows, posteriormost with about 2-10 scales to

upper extent of free preopercular edge, reaching forward

approaching upper lip crease above mouth, not reaching much

below lower margin of eye and posterior extent of orbit, or

reaching somewhere between, with broad naked margin

posteriorly and ventrally on preopercle or almost completely

naked cheek; 1-3 rows (only about 2 scales in second row when

present) of small scales on subopercle adjacent preopercular edge

extending forward to about anterior end of ventral preopercular

margin, to just in advance of posteroventral comer of preopercular

margin, or somewhere between, with about 5-10 scales in

outermost row; each lateral line scale with unbranched to multiple

branching laterosensory canal tube; scales above lateral line

about 21 / 2 - 31 / 2 ; cephalic sensory canal pores relatively few

confined to lines or short branches associated with major canals,

extremely numerous on top of head, in front of and below eye and

on lower jaw, or arranged somewhere between; second pair of

canines in lower jaw directed anterodorsally and slightly laterally,

usually curved slightly posteriorly; dorsal and anal fins with or

without very low basal sheath, comprising 1 smaller scale with

1-3 smaller or progressively smaller accessory scales adjacent to

fin base; posterior lobe of dorsal and anal fins reaching well short

of, not quite to or to hypural crease; caudal fin margin truncate to

convex. (See Table 2 for additional meristic and morphometric

ranges.) Most with prominent white to pearlescent stripe or

enlarged spot on body of adults, with or without distinctive sexual

dimorphism. Colouration of juveniles of some unknown.

Comments. Smith published Peaolopesia as a nomen nudum

first in a figure caption of a plate of illustrations (1949: pi. 101,

fig. 776a; 1950: pi. 101, fig. 776a), evidently added immediately

prior to publication, and for which no species treatments were

provided. In the third edition of Sea Fishes of Southern Africa

(1953: 520), Smith elaborated, designating Xiphochilus

gymnogenys Playfair and Giinther, 1867 as type species. Debate

continued as to whether it and closely allied species, such as

Choerodon margaritiferus Fowler and Bean, 1928, formed a

natural grouping deserving generic recognition distinct from

Choerodon. Kamohara (1958: 2) faced the same dilemma when

describing his Choerodonoides japonicus and was undoubtedly

unaware of Smith’s name. Gomon (1997: 812) placed the two in

synonymy with Choerodon but acknowledged that they

represented a natural grouping that might be recognised in the

future at subgeneric level.

The subgenus (fig. 1, clade la) comprises ten species with

a relatively slender profile and very short first pectoral fin ray,

seven of which have a modified falcate pectoral fin with an

extended ventralmost fin ray. The rather wide range of

proportional lengths for the first pectoral fin ray provided in

the above description and in Tables 2, 6 and 7 is more reflective

of the difficulty in determining where the distal tip is

positioned beneath overlying epidermal tissue than any true

variations in length. All species reach a small maximum size

as adults; the smallest C. gomoni is so far known from

individuals measuring 106 mm SL or less, and the largest C.

aurulentus sp. nov. attains only just more than 180 mm SL.

Elsewhere within the genus, only C. typus of the monotypic

subgenus Xiphocheilus has a maximum length falling within

this range. The subgenus may be legitimately regarded as

dwarf tuskfishes.

Choerodon albofasciatus nom. nov.

Ira-modoki, Whitestripe Tuskfish

Figures 33, 34; table 5; appendix.

Choerodonoides japonicus Kamohara, 1958: 2, pi. 1, fig. 1,

Mimase Market, Kochi City (Japan), junior homonym of Labrus

japonicus Valenciennes (= C. azurio).

Table 5. Ranges for selected counts and proporional measurements in eight species of the subgenus Peaolopesia. “N” designates number of counts or measurements. Aberrant values

enclosed by parentheses.

Review of Choerodon tuskfishes

X a

m fN

o >o

o o

00 o

X 3

00 o

X 3

0 ^ ^

00 o

X B

2 ^

X «•' -

X a

O ON

2 ^

^ ^ -H

o c

Review of Choerodon tuskfishes

Table 6. Ranges for selected counts and proportional measurements for types of two new species of the subgenus Peaolopesia. “N” designates

number of counts or measurements.

Species

C. aurulentus nsp

C. skaiopygmaeus nsp

MERISTIC FEATURES

Holotype

Paratype

Holotype

Paratypes

Dorsal fin

XII, 8

XIII, 7

Anal fin

III, 10

Pectoral fin

ii, 13

Vertebrae

10+ 17 = 27

10+ 17 =

MORPHOMETRIC FEATURES

range

mean

Standard length (mm)

181

177

138

75.6-133

100.4

% SL

Body depth

29.3

29.5

34.3

31.0-33.1

31.6

Caudal peduncle depth

11.4

10.6

12.1

11.7-13.0

12.2

Head length

38.2

36.6

42.0

38.0-41.4

39.9

Head depth

25.0

24.6

26.2

24.3-28.5

25.9

Orbit diameter

7.1

7.4

7.6-9.0

8.4

Interorbital width

8.4

7.9

6.6-8.1

7.2

Snout length

12.0

12.2

13.1

10.3-13.0

11.5

Dorsal fin base length

53.6

56.4

55.0

50.7-53.9

52.1

Dorsal fin last spine length

9.5

8.8

7.7

7.1-9.4

8.0

Dorsal fin posterior lobe length

11.8

11.4

9.5

7.6-10.1

9.1

Anal fin base length

26.7

26.9

25.7-29.3

27.1

Anal fin posterior lobe length

12.4

12.6

8.6

9.2- 13.0

10.3

Pectoral fin length

23.3

23.4

24.6

23.8-26.3

25.1

1st pectoral fin ray length

1.6

2.1

2.7

2.4-3.8

3.1

Pelvic fin length

24.6

25.8

22.8

19.6-22.8

21.5

Caudal fin dorsal lobe length

21.8

21.5

20.4

18.7-20.8

19.6

Caudal fin central ray length

17.1

18.9

17.7

17.4-23.0

20.7

Head length (mm)

69.1

64.7

57.9

31.3-53.3

40.0

%HL

Head depth

65.6

67.2

62.3

58.6-70.9

65.0

Orbit diameter

19.3

19.4

17.7

19.0-22.6

21.1

Interorbital width

22.0

22.9

18.9

16.3- 19.6

18.2

Snout length

31.4

33.3

31.2

27.2-32.2

28.9

X 100%

1st pectoral fin ray/pectoral fin length

7.0

9.1

11.2

9.5-14.1

12.0

Orbit diameter/Snout length

61.4

58.2

56.6

58.9-83.1

73.6

M.F. Gomon

Peaolopesia gymnogenys (not Gunther). — Masuda et al., 1975:

294; Masuda et al., 1984: 202, Okamura and Amaoka, 1997: 519;

Nakabo, 2000: 969.

Choerodon gymnogenys (not Gunther). — Shen, 1993: 452; Chen,

2010: 384.

Choerodon japoncus. — Kuiter, 2010: 63; Nakabo, 2013: 1088.

Choerodon cf margaritiferus. — White et al., 2013: 265;

Puckridge et al., 2015: 65, 66, figs 1 & 2.

Diagnosis. Dorsal fin rays XII, 8 (rarely XI, 9); anal fin rays

III, 10 (rarely 8); pectoral fin rays ii, 13 (rarely 12), dorsalmost

ray short 7.2-13.1% pectoral fin length, ventralmost ray

distinctly longer than those immediately above, posterior

edge of fin falcate, dorsoposterior corner bluntly pointed,

posteroventral corner sharply pointed; body shallow, 27.8-

32.5% SL, head depth 21.2-25.5% SL, caudal peduncle depth

10.8-11.6% SL; head blunt, dorsal profile of snout steep, snout

length 9.9-13.1% SL; predorsal scales approximately 7,

reaching forward on dorsal midline to or just in advance of

posterior edge of preopercle; cheek with small partially

embedded scales in about 4 or 5 diagonal rows, posteriormost

with about 8-10 scales to upper extent of free preopercular

edge, reaching forward almost to corner of upper lip crease

above mouth, with very broad naked margin posteriorly and

ventrally on preopercle; row of about 8-10 small scales on

subopercle adjacent preopercular edge extending forward to

about anterior end of ventral preopercular margin; each

lateral line scale with unbranched laterosensory canal tube;

cephalic sensory canal pores moderately numerous confined

to lines or short branches associated with major canals; scales

above lateral line about IVi or 3; second pair of canines in

lower jaw directed laterally, recurved posteriorly; dorsal and

anal fins without basal sheath, additional small scale at top of

some oblique rows; posterior lobe of dorsal and anal fins

reaching well short of hypural crease; caudal fin truncate,

with posterior margin slightly convex medially, upper and

lower corners slightly pointed; pelvic fin reaching to anus,

length 21.4-24.8% SL. (See Table 5 for additional meristic

and morphometric ranges.) Red to green above, white below

with with pearly blue midlateral stripe angled from underside

of eye to base of caudal fin, stripe deflected downward to

upper jaw below front of eye; terminal phase with second

narrower stripe from back of eye along lateral line to top of

caudal peduncle; scales posteroventrally with vertical pearly

blue line.

Reaches moderately small maximum size, largest

specimen examined 152 mm SL.

Pigmentation in alcohol. Juveniles unknown. Adults pale with

moderately narrow tapering dark stripe directed posteriorly

and slightly dorsally from centre of posterior margin of orbit

towards posterior end of dorsal fin base, terminating slightly

below last few rays; similar stripe directed from lower extent of

orbit to just above centre of caudal fin base; space between

stripes dark dusky tapering to point adjacent termination of

ventral stripe; fins pale.

Fresh colours. Juveniles unknown. Initial phase adults reddish

orange above, white below, with broad white stripe running

from tip of snout under eye and angled slightly dorsally to

caudal fin base above lateral midline, stripe with distinct red

margin from lower edge of eye to caudal base dorsally and

from opercle to caudal base ventrally (fig. 33A); scales below

stripe on sides blotched with red to level of lower end of

pectoral fin base; head with white cheeks and red lips. Dorsal

fin yellow with hyaline submarginal stripe; anal fin hyaline

with broad yellow midlateral stripe; caudal fin mostly yellow.

Pectoral and pelvic fins hyaline; pelvic fin with lengthwise

yellow submarginal line near leading edge (Masuda et al.,

1984: pi. 193, fig. B; Okamura & Amaoka, 1997: 519, fig. 1, as

Paeolopesia gymnogenys-. White et al., 2013: 265, fig. 89.14, as

Choerodon cf. margaritiferus).

Terminal phase adults green above, yellow reticulation

midlaterally and white ventrally, each scale on side with

vertical pearly line (fig. 33B); broad pale blue stripe from

lower margin of eye above pectoral fin base to just above

centre of caudal fin base; second narrower blue stripe from

posterior margin of eye curving ventrally to join first below

middle of soft portion of dorsal fin; head with yellow stripe

from upper jaw to anteroventral margin of eye; second

curving from posterior end of upper jaw across operculum

and along pectoral fin base; space between yellow stripes

pale blue. Dorsal fin pale pinkish brown with narrow yellow

submarginal stripe; anal fin white with narrow yellow medial

stripe; caudal fin blue basally, posterior half red. Pectoral fin

red; pelvic fin white with lengthwise yellow submarginal line

near leading edge (Masuda et al., 1984: pi. 193, fig. A; Shen,

1993: pi. 144, fig. 3, as C. gymnogenys-, Chen et al., 2010:

384, fig. C).

Etymology. The name albofasciata is from the Latin albo for

“white” and fasciatus meaning “envelope with bands”, in

reference to the prominent white stripe on the side as part of the

initial phase colouration of this species.

Distribution. Occurs along the western edge of the Pacific

from Inami, on the south-eastern coast of Honshu Island in

Japan to Shark Bay, Western Australia (fig. 34) at depths of at

least 95-120 m.

Comments. Kamohara’s Choerodonoides japonicus (1958: 2,

pi. 1, fig. 1) is clearly a species of Choerodon (Gomon, 1997:

812) and therefore a junior homonym of Labrus japonicus

Valenciennes, in Cuvier & Valenciennes, 1839 (= Choeodon

japonicus), itself a homonym of Labrus japonicus Houttuyn,

1782. The species therefore requires a new name and C.

albofasciatus is proposed as a replacement.

The species has been treated variously in the literature, as

cited in the synonomy above, and therefore confused with C.

gymnogenys and C. margaritiferus, with which it shares the

white to pearly lateral stripe or pair of stripes. The same basic

pattern occurs in C. gomoni as well as C. auruilentus described

below. Choerodon albofasciatus differs from all four in

having its cheek scales extending forward to the anterior end

of the exposed ventral edge of the preopercle versus to or less

than to the middle of the preopercular edge.

Material examined. 11 specimens, 96.7-152 mm SL; see

appendix.

Review of Choerodon tuskfishes

Figure 33. Choerodon albofasciatus nom. nov. A, Initial phase adult, CSIRO H 7220-04, 136 mm SL, Lombok, Tanjung Luar, photo by W.

White, CSIRO; B, Terminal phase adult, north of Bonaparte Archipelago, Western Australia, photo compliments of CSIRO.

Choerodon aurulentus sp. nov.

http://zoobank.org/urn:lsid;zoobank.org:act:CB22CDB8-2FC7-

4417-8FA9-ABAD417033D6

Proposed vernacular: Gilded Tuskfish

Figures 34, 35; table 6

Holotype. NSMT-P 122933 (181) western South Pacific, New

Zealand, Norfolk Ridge, 29° 28.8' S, 168° 10.6' E, 90 m, 17 January

1976, RV Kaiyo-maru (orig. no. ED166).

Paratype. (1 specimen, 177 mm SL). NSMT-P 122932 (177)

western South Pacific, New Zealand, Norfolk Ridge, 28° 44.4' S, 167°

55' E, 79 m, 18 January 1976, RV Kaiyo-maru (orig. no. EC301).

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray very short 7.0-9.1% pectoral fin

length, ventralmost ray distinctly longer than those immediately

above, posterior edge of fin falcate, dorsoposterior corner bluntly

pointed, posteroventral corner sharply pointed; body shallow,

10.6-11.4% SL, head depth 24.6-25.0% SL, caudal peduncle

depth 10.6-11.4% SL; head blunt, dorsal profile of snout steeply

rounded, snout length 12.0-12.2% SL; predorsal scales

approximately 7, reaching forward on dorsal midline just in

advance of posterior edge of preopercle; cheek with small

partially embedded scales in about four diagonal rows,

posteriormost with about 7 scales to upper extent of free

preopercular edge, scales not reaching forward quite to below

centre of eye, lower third or more of cheek above ventral edge of

preopercle naked; 1 or 2 rows of about 7 or 8 small scales on

subopercle adjacent preopercular edge extending forward to just

more than half way to anterior extent of free ventral preopercular

edge; each lateral line scale with unbranched laterosensory canal

tube; cephalic sensory canal pores moderately numerous

M.F. Gomon

20 °

0 °

20 °

Cr.

O Choerodon albofasciatus (examined)

O Choerodon albofasciatus (record)

EH Choerodon aurulentus sp. nov, (examined)

A Choerodon frenatus (examined)

A Choerodon frenatus (record)

20 °

0 °

20 °

Figure 34. Distributions of Choerodon albofasciatus nom. nov. (circles), C. aurulentus sp. nov. (squares) and C. frenatus (triangles) based on

specimens examined (coloured) and collection registration records (white).

confined to lines or short branches associated with major canals

between and behind eye, less numerous in front of and below

eye; scales above lateral line about IVi-, second pair of canines in

lower jaw directed laterally, recurved posteriorly dorsal and anal

fins without basal sheath, additional small scale at top of some

oblique rows; posterior lobe of dorsal and anal fins just reaching

hypural crease; caudal fin truncate, with posterior margin mostly

straight, upper and lower corners slightly pointed; pelvic fin to

anus or anal fin origin, length 24.6-25.8% SL. Green above,

white below with with pearly blue midlateral stripe angled from

underside of eye to base of caudal fin; additional blue stripes

curving from front of eye to upper jaw and across lower part of

head from underside of lower jaw to preopercular margin;

terminal phase golden midlaterally, scales with vertical pearly

bar and second pearly blue stripe from back of eye along lateral

line to top of caudal peduncle; scales posteroventrally with

vertical pearly blue line; lines on head with gold margins.

Reaches moderately small maximum size, largest

specimen examined 181 mm SL.

Description. Dorsal fin rays XII, 8; anal fin rays III, 10; caudal

fin rays 9 + 12 + 9 (8+12 +8); pectoral fin rays ii, 13; vertebrae

10 + 17 = 27; pleural ribs ending on 10th vertebra; epipleural

ribs ending on 13th vertebra; lateral line scales 27 + 2; scales

above lateral line 21 / 2 ; scales below lateral line approximately

81 / 2 ; predorsal scales approximately 7; total gill rakers 13.

(See Table 6 for selected measurements expressed as

percent of SL or HL.) Body shallow, greatest depth at dorsal

fin origin, 3.4 in SL; caudal peduncle moderately slender,

depth 8.8 (9.5) in SL; head large and shallow, length 2.6 (2.7)

in SL, depth at posterior extent of orbit 1.5 in HL; snout of

moderate length, 3.2 (3.0) in HL; head bluntly pointed; dorsal

outline of forehead and snout convexly curved, nape a gentle

convex curve; eye large, orbit 5.3 in HL; interorbital moderately

broad; jaws not attenuate.

Predorsal scales reaching forward on dorsal midline of

head barely in advance of above dorsal end of preopercle.

Cheek only partially covered by scales, scales small, mostly

embedded, about 4 vertical rows, posteriormost row with about

Review of Choerodon tuskfishes

Figure 35. Choerodon aurulentus sp. nov. Paratype, NSMT-P 122932, 111 mm SL, Western South Pacific, New Zealand, Norfolk Ridge, 28°

44.4’ S, 167° 55’ E, photo compliments of Far Seas Fisheries Faboratory, Japan.

7 scales to upper extent of free preopercular edge, scales not

reaching forward quite to below centre of eye, lower third or

more of cheek above ventral edge of preopercle naked; opercle

fully covered by large imbricate seales; subopercle with row (1

or 2 rows) of about 8 (7 or 8) embedded scales adjacent ventral

edge of preopercle extending forward Just more than half way

to anterior extent of free ventral preopercular edge; lower jaw

naked. Lateral line scales each with unbranched laterosensory

canal tube. Cephalic sensory canal pores moderately numerous

confined to lines or short branches associated with major canals

between and behind eye, less numerous in front of and below

eye. Posterior edge of preopercle very finely serrate. Mouth

mostly horizontal, posterior corner of upper lip fold below

point just anterior to forward extent of orbit; lower lip

moderately narrow; upper lip narrow and mostly obscured by

skin flap except at anterior end of jaw; posterior end of maxilla

not exposed. Gill rakers on first arch arborescent.

Upper jaw with 2 prominent anterior canines; first

prominent canine slightly longer than second; both canines

directed mostly ventrally, first slightly laterally; dental ridge

smooth to slightly rough; posterior canine well developed.

Lower jaw with 2 prominent anterior canines, both as well as

anterior tooth of upper jaw exposed when mouth occluded;

first canine slightly shorter than second; first canine directed

antero dors ally, second dorsolaterally recurved posteriorly;

dental ridge smooth anteriorly, followed by 1 or 2 short stout

canines and dental ridge posteriorly. Vomerine teeth absent.

Dorsal fin spines subequal, pungent; pointed membrane

behind tip of each spine attached basal to and extending

distinctly beyond spine tip, posterior tip of dorsal and anal fin

narrowly rounded, posterior rays not produced; posterior tip of

fins not reaching to posterior edge of hypurals, anal approaching

edge. Dorsal and anal fins without scaley basal sheath, at most

1/2 scale at base. Caudal fin truncate with upper and lower

corners pointed, distinctly but only slightly produced,

dorsalmost rays 1.3 (1.1) times length of rays at centre of fin,

posterior margin straight. Pectoral fin with dorsalmost ray very

short, ventralmost ray distinctly longer than those immediately

above, posterior edge of fin falcate, dorsoposterior corner

bluntly pointed, posteroventral corner sharply pointed. Pelvic

fin of moderate length, posterior tip of fin reaching anus.

Reaches moderately small maximum size, largest

specimen examined 181 mm SL.

Pigmentation in alcohol. Juveniles unknown. Adults dusky

above, pale below; head with narrow darker edged pale stripe

curving from front of eye to upper lip anteriorly, second horizontal

stripe adjacent to underside of eye extending posteriorly on head

and curving downward to third stripe anteriorly behind mouth,

third stripe curving across cheek to posterior extent of upper jaw

and across lower jaw ventrally. Pectoral fin base crossed by

narrow pale band adjacent proximal ends of rays.

Fresh colours. Juveniles unknown. Adults bluish green in front

of eye and along dorsal surface above level of eye, remainder of

head and body stark white with golden markings, including

narrow golden horizontal stripe from lower margin of eye to

dorsoposterior corner of caudal peduncle (fig. 35); scales

midlaterally behind pectoral fin ouflined with gold; head with

pair of golden stripes curving from anterior margin of eye to

upper jaw; posterior band continuing onto chin and joined on

upper lip to 3rd band directed posteriorly to preopercular

margin; 4th band parallel with 3rd from underside of lower jaw

across corner of mouth to preopercular margin; opercle with

scattered irregular golden spots posterior to preopercular edge;

golden band across pectoral base with white margin posteriorly;

gold marks on scales anterior to pectoral fin base. Dorsal fin

rosy pink; anal fin white; both fins with narrow yellow

submarginal stripe distally; caudal fin white with horizontal

gold streaks. Pectoral fin hyaline; pelvic fin white with

lengthwise yellow submarginal line near leading edge.

Etymology. The name aurulentus is Latin for “ornamented

with gold”, in reference to the prominent golden markings

characterising this species.

M.F. Gomon

Distribution. Known from only two specimens collected on the

Norfolk Ridge in the north-eastern Tasman Sea between New

Caledonia and New Zealand (fig. 34) at depths of 80-90 m.

Comments. Of the six species in the subgenus with which it

shares a crescent-shaped pectoral fin and dorsal fin for formula

of XII, 8, C. aurulentus is most similar to those with the

confirmed presence of one or two white or pearlescent narrow

stripes on the body, at least in adult males of C. albofasciatus,

C. gymnogenys and the cognates C. gomoni and C. margritiferus.

Choerodon aurulentus differs from C. albofasciatus and C.

gymnogenys in having cheek scales reaching forward to below

the centre of the eye, rather than to the anterior end of the

exposed ventral edge of the preopercle in the former and forward

barely to below the posterior extent of the orbit in the latter, as

well as in details of colour patterns. Choerodon aurulentus is

the largest of the dwarf tuskfishes, and although the two types

have proportional measurements that differ from those of C.

gomoni and C. margaritiferus, the latter species reach a

considerably smaller maximum size and differences observed

in features like body depth, head length, orbital diameter and

snout length, might be interpreted as simply representing

allometric changes. Without question, the nearly identical

colour pattern of the two C. aurulentus types differs considerably

from those of both males and females of the other two.

Choerodon frenatus Ogilby, 1910

Bridled Tuskfish

Figures 34, 36; table 5; appendix.

Choerodon frenatus Ogilby, 1910b; 99, 19 miles N 30° W from

Double Island Point (Queensland).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 13, rarely 14, dorsalmost ray short 6.7-16.8% pectoral

fin length, ventralmost rays shorter than those above, posterior

edge of fin obliquely straight, dorsoposterior corner bluntly

pointed, posteroventral corner angular; body shallow, 27.1-

30.5% SL, head depth 21.8-25.6% SL, caudal peduncle depth

11.7-13.3% SL; head blunt, dorsal profile of snout steep, snout

length 11.6-14.3% SL; predorsal scales approximately 5,

reaching forward on dorsal midline to or just in advance of

posterior edge of preopercle; cheek with small partially

embedded scales in about 6 or 7 diagonal rows, posteriormost

with about 7 or 8 scales to upper extent of free preopercular edge,

reaching forward approaching corner of upper lip crease above

mouth, with broad naked margin posteriorly and ventrally on

preopercle; 2 or 3 rows of about 8 small scales (fewer scales in

outer rows) covering much of subopercle adjacent preopercular

edge extending forward to about anterior end of ventral

preopercular margin, each lateral line scale with multiple

branched laterosensory canal tube; scales above lateral line

about IVi or 3; extremely numerous fine cephalic sensory canal

pores on top of head, in front of and below eye and on lower jaw;

second pair of canines in lower jaw directed dorsolaterally and

recurved slightly posteriorly; dorsal and anal fins with very low

basal sheath, comprising 1 smaller scale; posterior lobe of dorsal

and anal fins reaching well short of hypural crease; caudal fin

with central rays longer than those above and below, posterior

margin of fin with convex angle centrally, corners of fin slightly

produced in adults, most pronounced in large adults; pelvic fin

reaching well short of anus, approaching anus in large adults,

length 20.1-23.4% SL. (See Table 5 for additional meristic and

morphometric ranges.) Olive above, white below with elongate

reddish brown stripe-like blotch above lateral midline, at least

anteriorly, and pale blue to opalescent centres on scales

posteroventrally; head with pale blue lines angled anteroventrally

from eye and broader horizontal line on lower jaw.

Reaches moderately small maximum size, largest

specimen examined 136 mm SL but attaining 175 mm TL

(Ogilby, 1910b) and reported to 200 mm TL (Kuiter, 2010: 63).

Pigmentation in alcohol. Juveniles pale, slightly duskier above.

Adults dusky above, underside pale, distinct demarcation at

horizontal in line with posterior end of upper lip; each side of

head with parallel pair of pale lines directed anteroventally

from front of eye, 3rd pale line from eye to posterior end of

upper lip, 4th angled more horizontally below posterior half of

eye about midway up cheek; dorsal fin dusky with narrow pale

distal margin; anal fin pale; caudal fin pale with faint dusky

spot or small blotch centrally on caudal fin base above posterior

edge of hypurals; middle of caudal fin dusky tapering

posteriorly, corners pale dorsally and ventrally. Pectoral and

pelvic fins pale.

Fresh colours. Juveniles similar to adults. Adults red, strongly

washed with olive brown above lateral line anteriorly (fig. 36);

side pink, uniformly so or with pale blue to lavender spot on

each scale behind appressed pectoral fin, spots covering most

of scale in large individuals; underside of head, throat and

abdomen pearly white; predorsal and interorbital area and

blotch on opercle violet; face below and in front of eye orange;

2 dark blue-edged pale blue stripes in front of eye, upper

continuing across midline in front of eye, lower across tip of

snout, 3rd line from middle of upper jaw to and along ventral

edge of eye, and 4th from angle of mouth to preopercular edge,

bands somewhat variable between individuals; bases of canine

teeth blue. Dorsal and caudal fins green to red; dorsal with pale

blue, yellow-edged submarginal stripe; caudal fin washed with

gold basally, dorsoposterior and ventroposterior corners

broadly hyaline; anal fin white with broad yellow midlateral

stripe. Pectoral and pelvic fins hyaline; pelvic fin with

lengthwise submarginal yellow line adjacent to leading edge.

Etymology. The name frenatus is apparently from the Latin

frenum meaning “bridle” or “rein”, in reference to the halter¬

like blue lines on the head.

Distribution. Endemic to the east coast of Australia from just

south of Cairns, Queensland to about the Clarence River, New

South Wales (fig. 34). Occurs over rubble bottom at depths of

28-83 m.

Comments. Only one (AMS 1.12536, 136 mm SL) of the five

specimens examined by Ogilby (1910b: 99) for his description

appears to be in the Australian Museum collection, although a

specimen in the Queensland Museum collection (QMB 1.475,

105 mm SL) is registered as being collected at the type locality

and has approximately the same length as that given for the

Review of Choerodon tuskfishes

ZS mm

Figure 36. Choerodon frenatus. A, Initial phase adult, QMB 1.37338, 111 mm SL, 19° 20.2' S, 148° 17.8' E, photo complements of Qld DPI &

Fisheries; B Terminal phase adult, AMS 1.31472-005, 105 mm SL, Yamba, New South Wales, photo by K. Graham, NSW Fisheries.

lower end of the examined specimen range, 127 mm SL. The

length of the Australian Museum specimen is close to the upper

end of that range and that specimen is regarded as the lectotype.

Choerodon frenatus is distinctive and unlikely to be

confused with others within the genus. As well as a relatively

unique profile, the species has an extraordinarily high number

of fine pores covering the head. Unlike the majority of species

in the subgenus, C. frenatus, C.jordani and C. zosteropherus

have a pectoral fin like those of other species of Choerodon

without the extended ventralmost rays. The three also have

XIII, 7 dorsal fin elements characteristic of most species in

other subgenera, rather than XII, 8 rays as in other members

having the modified pectoral fin.

Judging from the paucity of specimens in museum

collections, C. frenatus lives in habitats infrequently trawled.

When collected, it is usually taken as groups of individuals

rather than as lone specimens.

Material examined. 47 specimens, 52-136 mm SL; see

appendix.

Choerodon gomoni Allen & Randall, 2002

Gomons Tuskfish

Figures 37, 38; table 5; appendix.

Choerodon gomoni Allen and Randall, 2002: 110, figs 1-2,

Chesterfield Bank, Coral Sea.

Diagnosis. Dorsal fin rays XII, 7 or 8; anal fin rays III, 9 or 10;

pectoral fin rays ii, 13, rarely 14, dorsalmost ray short, 8.3-16.4%

pectoral fin length, ventralmost ray distinctly longer than those

immediately above, posterior edge of fin falcate, dorsoposterior

corner bluntly pointed, posteroventral corner sharply pointed;

body shallow, 24.4-28.6% SL, head depth 20.1-24.6% SL,

caudal peduncle depth 9.2-10.3% SL; head bluntly pointed,

dorsal profile of snout shallow curve, snout length 9.4-11.9% SL;

M.F. Gomon

25 mm

Figure 37. Choerodon gomoni. A, Juvenile, CSIRO H 6442-06, 85.2 mm SL, east-north-east of Hayman Island, Queensland, Australia, 19.6653°

S, 150.0759° E, photo by D. Gledhill, CSIRO; B, Initial phase adult, CSIRO H 3436-04, 92.0 mm SL, Blackwood Channel, Cape York,

Queensland, Australia, photo by D. Gledhill, CSIRO; C, Terminal phase adult, CSIRO H 3441-01, 106 mm SL, Blackwood Channel, Cape York,

Queensland, Australia, photo by D. Gledhill, CSIRO.

Review of Choerodon tuskfishes

Figure 38. Distributions of Choerodon gomoni (circles) and C. margaritiferus (squares) based on specimens examined (coloured) and collection

registration records (white).

predorsal scales approximately 6, reaching forward on dorsal

midline just in advance of midpoint between posterior extent of

eye and posterior edge of preopercle; cheek with small partially

embedded scales in about 3 diagonal rows, posteriormost with

about 7 scales to upper extent of free preopercular edge, reaching

forward about half way between corner of upper lip crease and

posterior edge of preopercle and ventrally about half way

between lower extent of orbit and ventral edge of preopercle,

with extemely broad naked margin posteriorly and ventrally on

preopercle; about 2 rows of about 7-9 small scales (only about 1

or 2 scales in second row when present) on subopercle adjacent

preopercular edge extending forward just in advance of

posteroventral corner of preopercular margin; each lateral line

scale with unbranched laterosensory canal tube; scales above

lateral line about IVi or 3; cephalic sensory canal pores

moderately numerous confined to lines or short branches

associated with major canals; second pair of canines in lower jaw

directed anterodorsally and recurved laterally; dorsal and anal

fins without basal sheath, 1-3 progressively smaller accessory

scales adjacent to fin base; posterior lobe of dorsal and anal fins

reaching well short of hypural crease; caudal fin truncate, often

with posterior margin slightly convex medially, upper and lower

corners slightly produced; pelvic fin reaching to terminal phase

or well short of initial phase anus, length 19.1-22.9% SL. (See

Table 5 for additional meristic and morphometric ranges.) Pink,

somewhat olive above, white below with broad midlateral

reddish brown stripe from above pectoral fin base to base of tail;

broad horizontal blue line below eye; terminal phase with dark

brown to black bar on upper half of side below 9th to 11th dorsal

fin spines.

Reaches small maximum size, largest specimen examined

106 mm SL.

Pigmentation in alcohol. Juveniles uniformly pale. Inital phase

adult pale with narrow dusky midlateral stripe posteriorly on

caudal peduncle. Terminal phase adults dusky dorsally, lower

2/3 of side pale with narrow dusky stripe posteriorly onto base

of caudal fin; darker dusky blotch crossing lateral line below

M.F. Gomon

last few dorsal fin spines; head dusky dorsally with darker

stripe crossing snout at level of ventral margin of eye; fins pale.

Fresh colours. Juveniles unknown.

Initial phase washed with pink, rosy dorsally, paler pink to

white ventrally, separated by faint yellow to orange brown stripe

from upper part of pectoral fin base to middle of upper half of

caudal fin base; fins hyaline to white (fig. 37A & B); caudal fin

with bright red margin posteriorly (Allen & Randall, 2002: fig. 2).

Terminal phase greenish grey dorsally, mostly cream to

yellowish white ventrally with pale mauve tinge above lateral

line; head slightly pink (fig. 37C); broad white stripe crossing

snout, extending under eye and continuing along lateral

midline to centre of caudal fin base; faint yellow to dark red

lateral stripe above white stripe from upper pectoral base to

upper portion of caudal fin base; large, diffuse, reddish brown

spot on back below base of 9th to 11th dorsal fin spines, its

anterior edge bordered with white; second much smaller,

fainter red spot above pectoral fin. Fins mainly pale bluish

white to hyaline; dorsal and anal fins with pale yellow stripe

across middle of fin, distal third of dorsal pink. (Allen &

Randall, 2002: fig. 1; Allen & Erdmann, 2012: 646, middle).

Etymology. The name gomoni recognises the contributions

made to the understanding of labrid fishes by Martin F. Gomon

(NMV).

Distribution. Apparently confined to the south-western Pacific

off the north-east coast of Australia, New Caledonia and the

intervening Coral Sea (fig. 38). Occurs above rubble bottom at

depths of 24-82 m.

Comments. This species is very similar to C. margaritiferus

known from the Philippines, Taiwan and southern Indonesia.

Both have nearly identical primary phase and terminal phase

colour patterns, differing only slightly in details such as the

nature and position of the prominent dark spot posteriorly on the

side relative to the lateral line in terminal phase adults. The

marking is black with a blue leading edge and is situated just

under the lateral line below the last few dorsal fin spines in C.

gomoni but orange-brown with a blue anterior margin in a

position above the lateral line in C. margaritiferus. The eye of C.

gomoni also appears to be slightly larger (8.3-8.6% SL, 21.6-

23.4% HL, 71.4-83.1% snout length) than that of C. margaritiferus

(8.3-8.6% SL, 21.6-23.4% HL, 71.4-83.1% snout length).

Genetic evidence supports the distinction of C. gomoni and C.

margaritiferus with a close sister relationship between the two

(Puckridge et al., 2015: 61, fig. 1). A paratype of C. gomoni

(WAM P. 31498-003) collected in the Bengal Islands east of

central Sulawesi, Indonesia is a specimen of C. margaritiferus.

This is one the smallest species of Choerodon, the largest

specimen examined measuring 106 mm SL. Freshly caught

specimens are strongly washed with a bright rosy hue that

rapidly fades after death. It is also one of the few Choerodon

species known to aggregate as evidenced by large collections

along the northern Queensland coast and observations by the

original authors (Allen & Randall, 2002: 113).

Material examined. 30 specimens, 71.5-106 mm SL; see

appendix.

Choerodon gymnogenys (Playfair & Gunther, 1867)

Zanzibar Tuskfish, Purplelined Wrasse

Figures 39, 40; table 5; appendix.

Xiphochilus gymnogenys Playfair & Gunther, 1867: 85, pi. XII,

fig. 4, Zanzibar.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, rarely 14, dorsalmosf ray short 5.2-13.9%

pectoral fin length, ventralmost ray distinctly longer than those

immediately above, posterior edge of fin falcate, dorsoposterior

corner bluntly pointed, posteroventral corner sharply pointed;

body shallow, 29.3-32.0% SL, head depth 24.0-27.3% SL,

caudal peduncle depth 10.0-11.4% SL; head bluntly pointed,

dorsal profile of snout oblique, snout length 12.2-15.5% SL;

predorsal scales approximately 7, reaching forward on dorsal

midline to or just in advance of midpoint between posterior

extent of eye and posterior edge of preopercle; cheek with small

partially embedded scales in about 2-5 diagonal rows,

posteriormost with about 2-3 scales extending ventrally just

below upper extent of free posterior preopercular edge and

little below ventral extent of orbit, not quite reaching forward to

posterior extent of orbit, leaving most of cheek naked; 1 or 2

rows of about 6-9 small scales (only about two scales in second

row when present) on subopercle adjacent preopercular edge

extending forward just short of anterior end of ventral

preopercular margin; each lateral line scale with unbranched

laterosensory canal tube; scales above lateral line about 21/2 or

3; few cephalic sensory canal pores confined to lines or short

branches associated with major canals; second pair of canines

in lower jaw directed dorsolaterally and curved slightly

posterolaterally; dorsal and anal fins without basal sheath,

additional small scale at top of some oblique rows; posterior

lobe of dorsal and anal fins reaching well short of hypural

crease; caudal fin truncate, often with posterior margin slightly

convex medially, upper and lower corners slightly to distinctly

pointed; pelvic fin reaching just short of anus to just short of

anal fin origin, length 21.3-26.1% SL. (See Table 5 for

additional meristic and morphometric ranges.) Red above,

white below; terminal phase with olive back and narrow blue

lines midlaterally, others associated with eye.

Reaches moderately small maximum size, largest

specimen examined 140 mm SL.

Pigmentation in alcohol. Juveniles unknown. Initial phase

adults pale. Terminal phase adults pale dusky above, underside

pale, distinct demarcation on head at horizontal in line with

posterior end of upper lip to below eye; starck white narrow

stripe along lower lip, then horizontally to posterior edge of

preopercle; starck white line directed anteroventally from eye

to midway along upper lip, short horizontal white line adjacent

lower rim of orbit; lower half of operculum below middle of

pectoral fin base with moderately broad starck white marginal

band; broad white band angled posteroventrally across fleshy

base of pectoral fin; broad stark white nearly horizontal stripe

from above pectoral fin base and angled slightly upward to

lateral line under first few segmented dorsal fin rays, stripe

distinctly tapered anteriorly and posteriorly; second much

narrower stripe of uniform width below and parallel to first

Review of Choerodon tuskfishes

Figure 39. Choerodon gymnogenys. A, Initial phase adult, SAIAB 81739, 145 mm SL, Xai-Xai, Mozambique, 26° 9.4’ S, 32° 58.6’ E, photo by P.

Heemstra, SAIAB; B, Terminal phase adult, SAIAB 81739,150 mm SL, Xai-Xai, Mozambique, 26° 9.4’ S, 32° 58.6’ E, photo by P. Heemstra, SAIAB.

from just behind lower end of pectoral fin base; third narrow

starck white stripe or series of spots on dorsal profile of side

from above posterior end of first stripe onto caudal fin base.

Fins pale.

Fresh colours. Juveniles unknown. Initial phase adults red

above, white ventrally (fig. 39A); head wifh mauve-tinted white

patch covering lower half of cheek extending posteriorly to

middle of opercular edge, red pigmentation covering snout and

head anteroventral to eye to margin of upper jaw; pair of mauve

lines directed from eye to upper jaw. Fins milky to hyaline;

dorsal fin with red broad basal stripe and narrow distal margin;

middle of anal fin with broad faint yellow stripe; caudal fin

yellowish red basally with white distal margin; pelvic fin with

faint yellow lengthwise stripe along posterior edge of first

segmented ray (Kuiter, 2010: 62, bottom B).

Terminal phase olive green dorsally, red midlaterally on

side transitioning to orange, yellow and white ventrally with

pair of pearly or mauve to blue dorsoposteriorly angled lines

or series of spots, dorsalmost from above pectoral fin base to

lateral line below bases of first few segmented dorsal fin rays

and second from axilla of pectoral fin to centre of caudal fin

base (fig. 39B); head with pair of yellow margined blue lines

directed anteroventrally from eye to upper jaw; blue line on

lower lip continuing horizontally to posterior margin of

preopercle; mauve blotch on cheek and greenish blue blotch

dorsally on opercle; pectoral fin base with mauve to blue band;

M.F. Gomon

20 °

0 °

20 °

Figure 40. Distributions of Choerodon gymnogenys (circles) and C. skaiopygmaeus sp. nov. (squares) based on specimens examined.

dorsal fin pale blue with yellow sub-basal and sub-distal

stripes; anal fin pale blue with broad yellow midlateral stripe;

caudal fin somewhat orange basally; pectoral fin yellow; pelvic

fin milky with lengthwise yellow stripe as in initial phase

adults (Smith, 1949: pi. 101, fig. 776a, as Peaolopesia

gymnogenys-, Kuiter, 2010: 62, bottom A & C).

Etymology. The name gymnogenys is from the Greek gymno

for “naked” and genys for “cheek”, in reference to the lack of

scales on all but the dorsoposterior corner of the cheek in this

species.

Distribution. Restricted to the western Indian Ocean from

Mozambique and St Brandon Shoals through the Seychelles

Bank to the Arabian Gulf, although the last locality is based on

specimens collected in 1861 and may not be reliable (fig. 40).

Occurs on open rubble bottom in relatively deep water to

depths of at least 60 m (Kuiter, 2010: 62).

Comments. This species has been a source of confusion since

its description by Playfair and Gunther (1867: 85). Although

the authors gave only a single length of seven inches, four

specimens in the British Museum collection (BPBM

1864.11.15.28, dried skin, 132 mm SL; BPBM 1866.1.19.17,140

mm SL; BPBM 1866.1.19.18, 132 mm SL; BPBM unregistered,

118 mm SL) were collected by Playfair, presumably prior to the

publication of the original description. In the absence of

contradictory information, the largest specimen (BPBM

1866.1.19.17) is here regarded as lectotype.

Smith erected the genus Peaolopesia for the species, first as

a nomen nudum (1949: pi. 101, fig. 776a), and subsequently

(1953: 520) designated Xiphochilus gymnogenys Playfair and

Gunther, 1867 formally as type species. Smith (1957: 101) then

synonymised the species with Julis matthaei Valenciennes, in

Cuvier and Valenciennes (1839:419) on the basis of Valenciennes’

description alone. The latter is a junior synonym of Julis

genivittatus Valenciennes, in Cuvier and Valenciennes, 1839 (=

Thalassoma genivittata-, Randall and Smith, 2001: 119). Fricke’s

(1999: 405) record of Choerodon matthaei in the Mascarenes is

based on Smith’s error. Randall (in Smith & Heemstra, 1986:

690, pi. 94) corrected the misperception, but inadvertently

Review of Choerodon tuskfishes

reproduced Playfair and Giinther’s (1866: pi. XII, fig. 3) colour

figure of Xiphochilus robustus to illustrate C. gymnogenys.

Choerodon gymnogenys has been applied to very similar

con-subgenerics in the western Pacific (e.g. C. albofasciatus

as Paeolopesia gymnogenys - Masuda et al., 1975: 294,

Masuda et al., 1984: 202, Okamura & Amaoka, 1997: 519,

Nakabo, 2000: 969; and as Choerodon gymnogenys - Shen,

1993: 452, Chen et al., 2010: 384), but is clearly separable

from them by the greatly reduced cheek squamation,

as thought by its original authors. It appears to share the

rosy hue in life with C. gomoni and C. mar gar itiferas (figs

36, 38 & 42).

Material examined. 20 specimens, 99.5-140 mm SL; see

appendix.

Choerodon jordani (Snyder, 1908)

Dagger Tuskfish

Figures 41, 42; table 5; appendix.

Choerops jordani Snyder, 1908: 98, Naha (Okinawa, Japan).

Diagnosis. Dorsal fin rays XIII, 7 (rarely XIV, 6); anal fin

rays III, 10; pectoral fin rays ii, 13, dorsalmost ray short

9.6-17.5% pectoral fin length, ventralmost rays shorter than

those above, posterior edge of fin obliquely straight,

dorsoposterior corner bluntly pointed, posteroventral corner

angular; body shallow, 29.8-33.0% SL, head depth 22.1-

28.1% SL, caudal peduncle depth 11.7-13.1% SL; head

bluntly pointed, dorsal profile of snout moderately steep,

snout length 9.8-12.5% SL; predorsal scales approximately

5 or 6, reaching forward on dorsal midline to above posterior

extent of eye; cheek with small partially embedded scales in

about 7-10 diagonal rows, posteriormost with about 10

scales to upper extent of free preopercular edge, reaching

forward almost to corner of upper lip crease above mouth,

with broad naked margin posteriorly and ventrally on

preopercle; 2 rows of about 8-10 small scales with only

single scale in second row on subopercle adjacent

preopercular edge extending forward nearly to anterior end

of ventral preopercular margin; each lateral line scale with

single branched laterosensory canal tube; scales above

lateral line about IVi or 3; cephalic sensory canal pores

numerous behind eyes immediately in front of predorsal

scales, far fewer confined to lines or short branches

associated with major canals between and in front of eyes;

second pair of canines in lower jaw directed dorsolaterally

and curved posteriorly; dorsal and anal fins without

prominent basal sheath, 1-3 progressively smaller accessory

scales adjacent to fin base; posterior lobe of dorsal and anal

fins not quite reaching hypural crease; caudal fin truncate,

corners slightly produced in large individuals; pelvic fin

reaching to or not quite to anal fin origin, just short of anus

in small individuals, length 19.5-27.3% SL. (See Table 5 for

additional meristic and morphometric ranges.) Grey above,

white below, separated by broad yellow midlateral stripe,

broad anteriorly tapered wedge-shaped stripe above lateral

midline and large white oval spot below posterior end of

dorsal fin and on anterior end of caudal peduncle.

Reaches moderately small maximum size, largest

specimen examined 136 mm SL but reported to 170 mm TL

(Kuiter, 2010: 64).

Pigmentation in alcohol. Juveniles pale, slightly duskier above

with dark blotch dorsally on side extending onto base of dorsal

fin at proximal ends of first few rays and second dorsally at

posterior end of caudal pecuncle, dark blotches separated by

stark white oval patch; fins otherwise colourless. Adults pale

with broad dark anteriorly tapering, oblique band from axilla

of pectoral fin to bases of first few soft rays of dorsal fin,

covering basal third of fin to posterior edge; immaculate pale

blotch covering dorsal half of side and top of caudal pecuncle

posterior to dark band to hypural crease defined ventrally by

dusky pigment; dorsal edge of caudal fin darker.

Fresh colours. Juveniles pale grey with yellow midlateral stripe

from anterior tip of mouth to middle of side followed by broad

orange patch to base of tail (fig. 41A & B); small juveniles with

bright white stripe above yellow and orange stripes, broken by

extension of orange patch to dorsal profile of side at bases of

last few dorsal fin spines; prominent white edged black spot on

dorsal fin immediately above orange patch; caudal fin base

with prominent white spot at corners of caudal base dorsally

and ventrally; fins otherwise mostly clear. With growth, black

spot developing in advance of white spot dorsally on caudal fin

base and black spot on dorsal fin enveloping orange patch

below (Chen et al., 2010: 382, fig. A & 383, figs B & C; Kuiter,

2010: 64, figs D-F).

Adults grey dorsally, white ventrally with broad yellow

midlateral stripe separating the two (fig. 41C & D); black

wedge-shaped stripe from just above axilla of pectoral fin to

caudal fin base extending as black margin on dorsal edge of

caudal fin, lower edge on lateral midline and upper edge

angled to base of last dorsal fin spine and continuing onto base

of dorsal fin posteriorly; prominent yellow to white spot

superimposed on black wedge posteriorly, appearing as saddle

on caudal peduncle. Dorsal and caudal fins otherwise grey;

anal and pelvic fins white; pectoral fin hyaline (Masuda et al.,

1984: pi. 193, fig. E; Allen, 1985: 2407, fig. 332; Shen, 1993: pi.

144, fig. 4; Okamura & Amaoka, 1997: 465, centre second

from bottom and right second from top and middle; Chen et

al, 2010: 382, fig. D; Kuiter, 2010: 64, figs A-C and G).

Terminal phase adult colouration if differing from initial

phase not prominent.

Etymology. The name jordani recognises David Starr Jordan,

the then President of Stanford University and accomplished

ichthyologist.

Distribution. Antitropical in the western Pacific (fig. 42),

confined to Japan south of Kochi (Nakabo, 2000: 970), Taiwan

and the coast of south-east Asia to Hong Kong in the north,

occurring on both coasts of Australia south to the Abrolhos

Islands in Western Australia and One Tree Island on the Great

Barrier Reef, extending eastward to New Caledonia, Vanuatu,

Tonga, Fiji (Randall, 2005: 401; Seeto & Baldwyn, 2010: 39)

and American Samoa. Lives over sand and rubble along reef

edges at depths of 5-40 m (Kuiter, 2010: 64).

M.F. Gomon

25 mm

Figure 41. Choerodon jordani. A, Juvenile, Manabe, Kii Peninsula, Japan, photo by K. Yamasaki; B, Juvenile, Mana Island, Fiji, photo by K.

Uchino; C, Initial phase adult, KPM-NI19119, 98.0 mm SL, Miyako Island, Ryukyu Islands, Japan, photo by H. Senou, KPM; D, Terminal phase

adult, KPM-NI 30612, 124 mm SL, Motobu, Okinawa Island, Ryukyu Islands, Japan, photo by H. Senou, KPM.

Review of Choerodon tuskfishes

180 °

O Choerodon jordani (examined)

■ -X y /

A J

■ ^ y rr

1 ( f ^ U'.

O Choerodon jordani (record)

□ Choerodon zosterophorus (examined)

i-— w . Ax

^ "'-x

■ ' "30

\ _/

1 1

180 °

Figure 42. Distributions of Choerodon jordani (circles) and C. zosterophorus (squares) based on specimens examined (coloured) and collection

registration records (white).

Comments. Snyder (1908: 98) based his description of

Choerops jordani on four specimens acquired from the Naha,

Okinawa fish market, designating USNM 62235 (120 mm SL)

as holotype.

The species is distinctive, resembling only the very similar

C. zosterophorus, an apparent alloptric cognate distributed in

intervening equatorial latitudes. The two species, along with

C. frenatus, which lacks a prominent black or white marking

on the side, are the only members of the subgenus with a dorsal

fin count of XIII, 7 and the more conventional pectoral fin

profile of other subgenera. Choerodon zosterophorus is

distinguishable from C. jordani in having an obvious blunt

ended oblique white band bordered by a black blotch

dorsoposteriorly in place of the black wedge-shaped band on

the side.

Material examined. 49 specimens examined, 23-136 mm SL;

see appendix.

Choerodon margaritiferus Fowler & Bean, 1928

Pearlyscale Tuskfish

Figures 38, 43; table 5; appendix.

Choerodon margaritiferus Fowler & Bean, 1928: 197, Jolo

Market, Jolo (Philippines).

Choerodon sp. A White et ah, 2013: 266, fig. 89.19 (Lombok,

Indonesia).

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray short 7.3-12.9% pectoral fin length,

ventralmost ray distinctly longer than those immediately above,

posterior edge of fin falcate, dorsoposterior corner bluntly

pointed, posteroventral corner sharply pointed ted; body shallow,

24.7-30.8% SL, head depth 19.4-25.2% SL, caudal peduncle

depth 9.4-10.5% SL; head bluntly pointed, dorsal profile of snout

shallow curve, snout length 9.4-12.1% SL; predorsal scales

approximately 6 or 7, reaching forward on dorsal midline almost

M.F. Gomon

25 mm

Figure 43. Choerodon margaritiferus. A, Initial phase adult, USNM 435387, 114 mm SL, Cebu, Kawit, Medellin, Visayan Sea, Philippines,

photo by J. Williams, USNM; B, Terminal phase adult, USNM 435389, 115 mm SL, Kawit, Medellin, Visayan Sea, Cebu, Philippines, photo by

J. Williams, USNM.

to midpoint between posterior extent of eye and posterior edge of

preopercle; cheek with small partially embedded scales in about

3 or 4 diagonal rows, posteriormost with about 5 or 6 scales to

upper extent of free preopercular edge, reaching forward to

below middle of eye with very broad naked margin posteriorly

and ventrally on preopercle; 1-3 rows of about 6-8 small scales

(only about 2 scales in second row when present) on subopercle

adjacent preopercular edge extending forward to about midpoint

of ventral preopercular margin; each lateral line scale with

unbranched or single branched laterosensory canal tube; scales

above lateral line about Idh or 3; cephalic sensory canal pores

variable, mostly confined to lines or short branches associated

with major canals to numerous scattered on top of head and

anteroventral to eye and apparently not confined to lines or short

branches associated with major canals; second pair of canines in

lower jaw directed dorsolaterally and curved posterolaterally;

dorsal and anal fins without basal sheath, additional small scale

at top of some oblique rows; posterior lobe of dorsal and anal fins

not quite reaching hypural crease; caudal fin truncate, often with

posterior margin slightly convex medially; pelvic fin not quite

reaching anus, length 19.0-21.3% SL. (See Table 5 for additional

meristic and morphometric ranges.) Initial phase pink overall

with white underside and broad orange stripe just above lateral

midline; caudal fin with bright reed posterior margin. Terminal

phase green above, white below, with interrupted blue stripe

midlaterally, horizontal row of blue scales on lateral line

anteriorly, row of blue scales adjacent base of dorsal fin

posteriorly and horizontal blue stripe below eye from tip of snout

to preopercular edge; blue-edged, reddish brown blotch crossing

lateral line below 9th to 11th dorsal fin spines.

Reaches moderately small maximum size, largest

specimen examined 115 mm SL.

Pigmentation in alcohol. Juveniles uniformly pale. Initial phase

adults pale with narrow dusky to dark posterior margin on caudal

fin. Terminal phase dusky above, paler below, with narrow pale

stripe from tip of snout to lower edge of eye, continuing as broken

row of pale spots to pale irregular narrow blotch in pectoral fin

axilla continuing as pale spots to above middle of anal fin base;

Review of Choerodon tuskfishes

second narrow pale band in form of spots from behind eye across

opercle to side posterior to just above pectoral fin tip; pale band

from middle of opercular margin across pectoral fin base; narrow

pale stripe below dorsal fin base anteriorly to dorsal caudal

peduncle nearly to caudal fin base; additional narrow pale stripe

midlaterally underlying lateral line along length of caudal

peduncle, becoming dusky to dark with fading of overlying pale

pigment. Dorsal and anal fins pale.

Fresh colours. Juveniles unknown.

Initial phase adults pink overall, darker on dorsal half of

side and head, breast and belly white (fig. 43A); slightly darker

lengthwise stripe above lateral midline to dorsal half of caudal

peduncle. Dosal fin pale pink, darker near tips of spines; anal

and pelvic fins white; caudal fin yellow with pink basal third

and dark red posterior margin. Pectoral fin hyaline. (White et

al, 2013: 267, fig. 89.19).

Terminal phase adults greenish grey dorsally, white

ventrally with pink hue midlaterally (fig. 43B); terminal phase

with large red blotch centred just above lateral line below 9th

to 11th dorsal fin spines with oblique grey bar on anterior

edge; greyish white stripe originating on posterior edge of

operculum just above pectoral fin base breaking into scale¬

sized spots centrally on side, again becoming prominent stripe

on lateral midline below posterior end of dorsal fin, terminating

in point anteriorly on caudal fin; second horizontal series of

greyish white spots along dorsal profile of side below

segmented portion of dorsal fin and on caudal peduncle;

additional greyish white spots scattered centrally on side;

downward opening crescentic greyish white band on pectoral

fin base; head with horizontal greyish white stripe extending

from snout tip, below eye to preopercular edge; vertical greyish

white mark on dorsoposterior edge of eye; scattered pink

blotches, especially on operculum; eye red with yellow dorsal

margin on iris and greyish white margin to eye dorsally.

Dorsal fin with broad yellow basal stripe slightly narrower

pink stripe distally and narrow yellow distal margin; colours

less defined posteriorly; anal fin white with broad yellow

midlateral stripe and fine yellow distal margin; caudal fin

hyaline with pink hue. Pectoral fin hyaline with greyish white

rays dorsally; pelvic fin hyaline with pink leading edge

bordered by similarly narrow yellow longitudinal stripe

(Kuiter, 2010: 63, top; Allen & Erdmann, 2012: 246, bottom;

Miyamoto et al., 2015: 84, fig. lA and B).

Etymology. The name margaretiferus is Latin for “pearl

bearer”, in reference to the series of pearlescent spots on the

side of the body.

Distribution. Known conclusively only from Okinawa in

southernmost Japan, Taiwan, Philippines, and Indonesia off

eastern Sulawesi and Lombok (fig. 38) at depths to at least 30 m.

Comments. Choerodon margaritiferus was described by

Fowler and Bean (1928) from a single specimen (USNM 13558,

115 mm SL) purchased at a market in Jolo, Philippines. The

apparent absence of other specimens in collections until

recently and the overall similarity of members of this complex

to one another has caused confusion over the identity of the

species to which the name was affixed.

The name C. margaritiferus was appropriately applied by

Puckridge et al. (2015: 5, fig. 1, as Choerodon margaritiferus)

in their investigation of the interrelationships of species of the

genus employing genetic markers. The study recovered C.

margaritiferus as sister to C. gomoni, a species so similar that

Randall and Allen (2002: 110) designated a specimen of the

former from the Bangai Islands off eastern Sulawesi, Indonesia

as a paratype of the latter. The two appear to be among the

smallest species of Choerodon. They are extremely similar

and difficult to separate at initial phase and juvenile sizes.

The specimen from which the tissue was taken for

Puckridge et al.’s (2015) sequences of C. margaritiferus was

treated and figured by White et al. (2014: 266, fig. 89.19) as

Choerodon sp. A “Redtip Tuskfish”. Like C. gomoni, freshly

caught individuals have an overall bright rosy hue that obscures

underlying colouration and rapidly fades after death. The

specimen featuring in the above two studies is a female that

differs markedly in colouration from the male shown by

Miyamoto et al. (2015: 84, fig. lA & B). The last study also

recovered the taxonomic relationships found by Puckridge et

al. (2015). The species in all three studies tentatively identified

as C. cf. margaritiferus is C. albofasciatus, a new name for

Choerodonoides japonicus Kamohara, proposed above.

Material examined. 11 specimens, 82.8-115 mm SL; see

appendix.

Choerodon skaiopygmaeus sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:200B0A56-10B2-

47E4-A21D-61236AE91E88

Proposed vernacular: Western Pygmy Tuskfish

Figures 40, 44; table 6

Holotype. USNM 437323 (138) Indian Ocean, Somali coast, 11°

18' S, 51° 08' E, 25-29 m, 17 December 1964, RV Anton Bruun, Cr. 9,

Sta. 459, International Indian Ocean Expedition, trawl.

Paratypes. (6 specimens, 75.6-133 mm SL.) BMNH 2017.1.19.1

(106), MNHN-IC-2016-0605 (102), and USNM 437324 (133), same

data as holotype; SAIAB 203921 (93.1) Indian Ocean, Somali coast,

10° 00' S, 51° 15' E, 59-61 m, 16 December 1964, RV Anton Bruun,

Cr. 9, Sta. 447, International Indian Ocean Expedition, trawl; USNM

437325 (94.7). same data as SAIAB 203921; USNM 437326 (75.6)

Indian Ocean, Somali coast, 09° 41' S, 51° 03' E, 60-70 m, 16

December 1964, RV Anton Bruun, Cr. 9, Sta. 445, International Indian

Ocean Expedition, trawl.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray very short 9.5-14.1% pectoral fin

length, ventralmost ray distinctly longer than those

immediately above, posterior edge of fin falcate, dorsoposterior

corner bluntly pointed, posteroventral corner sharply pointed;

body shallow, 31.0 -34.3% SL, head depth 24.3-28.5% SL,

caudal peduncle depth 11.7-13.0% SL; head bluntly pointed,

dorsal profile of snout moderately steep, snout length 10.3-

13.1% SL; predorsal scales approximately 6, reaching forward

on dorsal midline almost to above posterior edge of preopercle;

cheek with small partially embedded scales in about 3 or 4

diagonal rows, posteriormost with about 7 or 8 scales to upper

M.F. Gomon

25 mm

Figure 44. Choerodon skaiopygmaeus sp. nov. Holotype, USNM 437323, 138 mm SL, Indian Ocean, Somali coast, 11° 18’ S, 51° 08’ E, ethanol

preserved (grey scale image).

extent of free preopercular edge, reaching ventrally to level of

posterior corner of upper jaw and forward little if at all beyond

posterior extent of orbit, with very broad naked margin

ventrally on preopercle; 1-3 rows of about 6-8 small scales

(only 1 or 2 scales in second of 2 rows when present) on

subopercle adjacent preopercular edge extending forward to

about midpoint of ventral preopercular margin; each lateral

line scale with single pore on laterosensory canal tube; scales

above lateral line about IVi or 3; numerous cephalic sensory

canal pores scattered on top of head and anteroventral to eye

apparently not confined to lines or short branches associated

with major canals; second pair of canines in lower jaw directed

dorsolaterally and curved laterally; dorsal and anal fins

without basal sheath, additional small scale at top of some

oblique rows; posterior lobe of dorsal and anal fins not

reaching hypural crease; caudal fin truncate, often with

posterior margin slightly convex medially; pelvic fin reaching

to or just short of anus, length 19.6-22.8% SL. Uniformly pale

in preservation except for dusky horizontal stripe from just

below lateral line under 8th dorsal fin spine to just beyond

posterior end of dorsal fin near top of caudal peduncle.

Reaches moderately small maximum size, largest

specimen examined 139 mm SL.

Description. Dorsal fin rays XII, 8; anal fin rays III, 10; caudal

fin rays 8 + 12 + 8 (8 or 9 + 12 + 8); pectoral fin rays ii, 13;

vertebrae 10 + 17 = 27; pleural ribs ending on 10th vertebra;

epipleural ribs ending on 13th (12th or 13th) vertebra; lateral

line scales 27 + 2; scales above lateral line IVr, scales below

lateral line approximately 8U; predorsal scales approximately

6; total gill rakers 17 (16 or 17, rarely 17).

(See Table 6 for selected measurements expressed as

percent of SL or HL.) Body shallow, greatest depth at dorsal

fin origin, 2.9 (3.0-3.2) in SL; caudal peduncle moderately

slender, depth 8.2 (7.7-8.6) in SL; head large and shallow,

length 2.4 (2.4-2.6) in SL, depth at posterior extent of orbit 1.6

(1.4-1.7) in HL; snout of moderate length, 3.2 (3.1-3.7) in HL;

head bluntly pointed; dorsal outline of forehead and snout

nearly straight, nape a gentle convex curve; eye of moderate

size, orbit 5.7 (4.4-5.3) in HL; interorbital moderately broad;

jaws not attenuate.

Predorsal scales reaching forward on dorsal midline

almost to above posterior edge of preopercle. Cheek only

partially covered with small partly embedded scales in about

5 or 6 (3-6) diagonal rows, posteriormost with about eight

scales to upper extent of free preopercular edge, reaching

ventrally to level of posterior corner of upper jaw and forward

little if at all beyond posterior extent of orbit, with very broad

naked margin ventrally on preopercle; opercle fully covered

by large embedded scales; subopercle with 1-3 rows of about

6-8 small scales (only 1 or 2 scales in second or third rows

when present) adjacent preopercular edge extending forward

to about midpoint of ventral preopercular margin; lower jaw

naked. Lateral line scales each with unbranched laterosensory

canal tube. Cephalic sensory canal pores numerous scattered

on top of head and anteroventral to eye apparently not

confined to lines or short branches associated with major

canals. Posterior edge of preopercle very finely serrate. Mouth

mostly horizontal, posterior corner of upper lip fold below

forward extent of orbit; lower lip moderately narrow; upper

lip narrow and mostly obscured by skin flap except at anterior

end of jaw; posterior end of maxilla not exposed. Gill rakers

on first arch arborescent.

Upper jaw with 2 prominent anterior canines; first

prominent canine noticeably longer than second; both canines

directed mostly ventrally, first slightly laterally, second

slightly laterally and recurved slightly posteriorly; dental

ridge smooth; posterior canine well developed. Lower jaw

with 2 prominent anterior canines, both as well as anterior

canine of upper jaw exposed when mouth occluded; first

canine noticeably shorter than second; first canine directed

Review of Choerodon tuskfishes

antero dors ally, second canine directed laterally and curved

posterolaterally; dental ridge smooth anteriorly, followed by 7

or 8 stout canines progressively increasing in length posteriorly.

Vomerine teeth absent.

Dorsal fin spines subequal, pungent; pointed membrane

behind tip of each spine attached just basal to and extending

distinctly beyond spine tip, posterior tip of dorsal and anal fin

narrowly rounded, posterior rays not produced; posterior tip of

fins not reaching to posterior edge of hypurals. Dorsal and

anal fins without basal sheath, 1-3 progressively smaller

accessory scales adjacent to fin base. Caudal fin truncate,

often with posterior margin slightly convex medially,

dorsalmost rays 1.2 (0.9-1.1) times length of rays at centre of

fin. Pectoral fin with dorsalmost ray very short, ventralmost

ray distinctly longer than those immediately above, posterior

edge of fin falcate, dorsoposterior corner bluntly pointed,

posteroventral corner sharply pointed. Pelvic fin of moderate

length, posterior tip of fin reaching to (to or nearly to) anus.

Reaches moderately small maximum size, largest

specimen examined 138 mm SL.

Pigmentation in alcohol. Juveniles unknown. Adults uniformly

pale except for dusky horizontal stripe about 1 scale wide from

just below lateral line under 8th dorsal fin spine to just beyond

posterior end of dorsal fin near top of caudal peduncle (fig. 44).

Fresh colours. Unknown.

Etymology. The new name skaiopygmaeus is a conjunction of

the Greek adjectives skaios for “western” and pygmaios for

“pygmy”, in reference to the distribution of this dwarf tuskfish

in the western Indian Ocean off Somalia.

Distribution. Known from only seven specimens acquired in

two collections made off the Somali coast in the western Indian

Ocean (fig. 40) at depths of about 25-60 m.

Comments. This species is clearly a member of the seven

species complex characterised by a tiny first pectoral fin ray

and falcate pectoral fin profile. Of the seven, it appears to be

closest to C. albofasciatus and C. sugillatum in having a

slightly deepter body and caudal peduncle. As the type series,

comprising seven mostly faded specimens collected off

Somalia in two tows on consecutive days in 1964, are

apparantly the only extant collection specimens, nothing is

known of life colours. Still, all lack persistent patterns

characteristic of the two described species as well as others in

the complex, like C. gymnogenys also known from the western

Indian Ocean. Choerodon skaiopygmaeus is further

distinguishable from the last by the more expansive scaled

area on the cheek that reaches distinctly below the ventral

margin of the eye. It is unclear if the slightly dusky stripe

laterally on the side is homologous with similar markings on

the side of C. margaretiferus specimens that are pearly in life,

a colour feature of some species of the complex. Long

preserved specimens of C. albofasciatus retain a similar

marking that is pearly in life. The species lacks the dark slash¬

like marking of C. sugillatum and dark spot dorsally on the

side of terminal phase C. gomoni and C. margaritiferus.

Choerodon sugillatum Gomon, 1987

Wedgetail Tuskfish

Figures 45, 46; table 5; appendix.

Choerodon sugillatum Gomon, 1987: 19, Queensland, Cape

Bedford (Australia), 15° 13.59'- 15° 15.00' S, 145° 23.36'-145° 27.87' E.

Diagnosis. Dorsal fin rays XII, 8; anal fin rays III, 10; pectoral

fin rays ii, 13, rarely 14, dorsalmost ray short 2.8-11.7% pectoral

fin length, ventralmost ray distinctly longer than those

immediately above, posterior edge of fin falcate, dorsoposterior

corner bluntly pointed, posteroventral corner sharply pointed;

body shallow, 29.3-37.7% SL, head depth 21.5-29.2% SL,

caudal peduncle depth 10.0-12.8% SL; head blunt, dorsal

profile of snout moderately steeply curved, snout length 8.0-

10.6% SL; predorsal scales approximately 6 or 7, reaching

forward on dorsal midline to midpoint between posterior extent

of eye and posterior edge of preopercle; cheek with small

partially embedded scales in about 6 diagonal rows,

posteriormost with about 6 or 7 scales to upper extent of free

preopercular edge, reaching forward to corner of upper lip

crease above mouth, with very broad naked margin posteriorly

and ventrally on preopercle; 2 rows of about 5-9 small scales

(only 1 or 2 scales in second row when present) on subopercle

adjacent preopercular edge extending forward to about anterior

end of ventral preopercular margin; each lateral line scale

usually with unbranched laterosensory canal tube; scales above

lateral line about IVi or 3; cephalic sensory canal pores

moderately numerous confined fo lines or short branches

associated with major canals; second pair of canines in lower

jaw directed dorsolaterally; dorsal and anal fins without basal

sheath, 1-3 progressively smaller accessory scales adjacent to

fin base; posterior lobe of dorsal and anal fins not quite reaching

hypural crease; caudal fin truncate, posterior margin convex

medially in adults, upper and lower corners pointed; pelvic fin

not quite reaching anus, length 17.3-21.3% SL. (See Table 5 for

additional meristic and morphometric ranges.) Olive above,

white below with blue-edged black bar below lateral line under

3rd or 4th dorsal fin spine; blue lines anteroventral from eye,

below and above eye and across pectoral fin base; caudal fin

with darker, posteriorly tapering wedge-shaped patch centrally.

Reaches moderately small maximum size, largest

specimen examined 149 mm SL.

Pigmentation in alcohol. Juveniles pale with dusky scale margins

dorsally and prominent dusky stripe above lateral midline from

opercular margin to distal caudal fin margin; darker mark on

stripe below bases of second and third dorsal fin spines; centre of

posterior caudal fin margin with small dark blotch; fins otherwise

hyaline .Adults pale, slightly dusky dorsally, with prominent dark

slash preceded by immaculate pale slash above and slightly

behind pectoral fin base; narrow dark margin wrapping around

dorsal edge of pectoral fin base; fins pale except for dark mark at

centre of caudal fin’s posterior margin.

Fresh colours. Juveniles generally similar to initial phase

adults as described below with additional brown blotches

dorsally; fins mostly transparent with 2 olive stripes in dorsal

fin and 2 blue stripes in anal fin (Kuiter, 2010: 56, top B & C).

M.F. Gomon

Figure 45. Choerodon sugillatum. Terminal phase adult, CSIRO H 3897-04, 135 mm SL, Pollard Channel, Great Barrier Reef, Queensland,

Australia, 11° 51.51’ S, 143° 27.83’ E, photo by T. Carter, CSIRO.

Figure 46. Distribution of Choerodon sugillatum based on specimens examined (coloured) and collection registration records (white).

Review of Choerodon tuskfishes

Initial phase adults olive dorsally, white ventrally with

narrow orange to brown stripe just above lateral midline from

snout to base of tail, stripe broader with blue dorsal edge on

head (fig. 45); scale centres blue, at least posteriorly, yellow

streaks between scale rows on caudal peduncle; distinctive

vertical black mark with blue anterior margin on midlateral

stripe above and slightly behind pectoral fin base; 2 yellow

edged blue lines directed anteroventrally from eye to upper

jaw, 2 others crossing snout in front of eye, short blue line

directed posteriorly from top and bottom of eye, lower line

curving ventrally anteroventral to eye and continuing onto

chin; blue stripe directed posteriorly from lower jaw to

posterior edge of preopercle; lower lip blue; broad blue band

with narrow yellow margins crossing pectoral fin base. Dorsal

and anal fins blue, dorsal fin with thin yellow horizontal line

proximally and distally; anal fin with thin yellow horizontal

lines proximally and midlaterally; caudal fin blue with

posteriorly converging yellow lines, lines edged with black

posteriorly at centre of rear margin of fin. Pectoral and pelvic

fins hyaline; pelvic fin with lengthwise yellow submarginal

line near leading edge (Sainsbury & Kailola, 1984: 261, middle

as Choerodon sp 2.).

Terminal phase adults similar to initial phase but pinker

overall with narrower midlateral stripe and vertical blue line

or spots on most scales on side. (Allen, 1985: fig 334, as

Choerodon species; Kuiter, 2010: 56, top A)

Etymology. The name sugUlatum is Latin for “black and blue”, a

reference to the characteristic marking on the side of this species.

Distribution. Restricted to northern Australia, extending at

least from Cape Tribulation, Queensland to the Monte Bello

Islands, Western Australia (fig. 46). Occurs on open sand with

sparse algal cover, like Caulerpa, at depths of 10-120 m

(Kuiter, 2010: 56).

Comments. Described from a series of specimens from both

coasts of northern Australia (Gomon and Allen, 1987: 19,

NMV A3126, 104 mm SL, holotype), C. sugUlatum frequently

features in the bycatch of northern Australian prawn fisheries.

As a dark slashdike marking is present on the side above the

lateral line in a number of species of this subgenus and at least

one other apparently unrelated subgenus, it cannot be relied on

alone for recognising this species. However, in combination

with the crescentic pectoral fin, having a nubdike first fin ray,

the features alone distinguish the species.

Material examined. 114 specimens, 35.3-149 mm SL; see

appendix.

Choerodon zosterophorus (Bleeker, 1868)

Blackblotch Tuskfish

Figures 42, 47; table 5; appendix.

Choerops zosterophorus Bleeker, 1868: 273, pi. XII, Kei-major

(Indonesia).

Choerops brenchleyi Gunther, 1872:424, Misol Island (Indonesia).

Diagnosis. Dorsal fin rays XIII, 7; anal fin rays III, 10; pectoral

fin rays ii, 13, dorsalmost ray short 11.3-18.3% pectoral fin

length, ventralmost rays shorter than those above, posterior

edge of fin obliquely straight, dorsoposterior corner bluntly

pointed, posteroventral corner angular; body shallow, 29.4-

32.9% SL, head depth 19.8-27.0% SL, caudal peduncle depth

11.8-13.6% SL; head bluntly pointed, dorsal profile of snout

moderately steep, snout length 9.5-11.6% SL; predorsal scales

approximately 6-8, reaching forward on dorsal midline almost

to midpoint between posterior extent of eye and posterior edge

of preopercle; cheek with small partially embedded scales in

about 8-12 diagonal rows, posteriormost with about 10 scales

to upper extent of free preopercular edge, reaching forward to

corner of upper lip crease above mouth, with broad naked

margin posteriorly and ventrally on preopercle; 1 or 2 rows of

about 8 small scales (only about 1 or 2 scales in second row

when present) on subopercle adjacent preopercular edge

extending forward near anterior end of ventral preopercular

margin; each lateral line scale with multiple branching

laterosensory canal tube; scales above lateral line about IVi or

3; cephalic sensory canal pores mostly confined to lines or

short branches associated with major canals; second pair of

canines in lower jaw directed dorsolaterally and curved

posterolaterally; dorsal and anal fins without prominent basal

sheath, 1-3 progressively smaller accessory scales adjacent to

fin base; posterior lobe of dorsal and anal fins not quite reaching

hypural crease; caudal fin truncate; pelvic fin reaching just

short of anus, length 19.8-22.4% SL. (See Table 5 for additional

meristic and morphometric ranges.) Grey above, white below

with broad yellow midlateral stripe, broad white stripe above it

angled towards base of first few segmentd dorsal fin rays and

marginal black blotch to stripe below posterior part of dorsal

fin base; smaller white blotch on side at posterior end of dorsal

fin base.

Reaches moderately small maximum size, largest

specimen examined 153 mm SL.

Pigmentation in alcohol. Juveniles pale with dark blotch

anteriorly at base of soft portion of dorsal fin. Initial phase

adults pale with stark white anteriorly tapering stripe on side

from base of last few dorsal fin spines to just above pectoral fin

base. Dark blotch on dorsal fin at base of anteriormost four or

five rays extending as tapered spot to about midside along

posterior margin of white stripe. Terminal phase adults as

initial phase with tapered end of dark blotch extending to

posterior side of pectoral fin base and blotch extending equal

distant along anterodorsal margin of white stripe.

Fresh colours. Juveniles pale grey above, white ventrally with

broad yellow to orange midlateral stripe with white stripe

immediately above interrupted below posterior part of dorsal

fin by dorsal expansion of yellow/orange stripe and black blotch

on dorsal apex at boundary between side and dorsal fin base

(fig. 47A); brown blotch at posterior end of caudal peduncle

with small black spot at centre of caudal fin base; fins hyaline

(Kuiter, 2010: 65, fig. B).

Initial phase adults grey dorsally, white ventrally with

broad yellow mostly horizontal band from cheek to below rear

end of dorsal fin; prominent pearly, silvery or white oblique

M.F. Gomon

Figure 47. Choerodon zosterophorus. A, Juvenile, USNM 436349, 43 mm SL, Oriental Mindoro, Puerto Galera, Philippines, photo by J.

Williams, USNM; B, Initial phase adult, Puerto Galera, Oriental Mindoro, Philippines, photo by P Ryan; C, Terminal phase adult, USNM

378714, 124 mm SL, Buyallao Island off SE Mindoro, Philippines, photo by J. Williams, USNM.

Review of Choerodon tuskfishes

stripe on dorsal edge of yellow band from above pectoral fin

base to below last few dorsal fin spines (fig. 47B); large black

blotch posterior to upper end of pale stripe, extending onto

bases of first few segmented dorsal fin rays, in small

individuals, black band extending anteroventrally along

posterior edge of pale band in larger adults; moderately small

white blotch posterior to black blotch below last few segmented

dorsal fin rays in smaller adults. Fins grey to hyaline; dorsal,

anal and caudal fins with narrow blue margins (Kuiter, 2010:

65, figs C-E; Allen & Erdmann, 2012: 649, bottom).

Terminal phase adults similar to initial phase but with

black blotch extending forward as anteriorly tapering stripe

towards forward end of white stripe (fig. 47C); additional

white blotch dorsoposteriorly absent (Kuiter, 2010: 65, fig. A).

Etymology. The name zosterophorus is from the Greek zoster

for “belt” or “girdle” and phaeos, phiaros or phos for “light,

bright or shining”, in reference to the distinctive oblique bright

white stripe angled across the side of this species.

Distribution. Apparently restricted to the Philipines north to at

least Caban Island, Batangas in Luzon, eastern Indosnesia at

least as far west as Lombok and Papua New Guinea (fig. 42).

Like its cognate C. jordani, this species occurs on semi-open

sandy substrates along reefs, often in deep channels to lagoons

that are prone to strong tidal currents, particularly with soft

corals, at depths of 10-40 m (Kuiter, 2010: 65; Allen &

Erdmann, 2012: 649).

Comments. Bleeker (1868: 273) based his description of

Choerops zosterophorus on a 170 mm specimen from Kei-

major (Great Kei Island, Indonesia). A 157 mm TL specimen in

the Rijksmuseum (RMNH 6537: 131 mm SL; Eschmeyer, 2015,

gives RMNH 6533 as the type) is slightly smaller than the

given length but has been cut open, presumably for the

examination of the pharyngeals, and may be Bleeker’s

specimen. It otherwise agrees with the description. Gunther’s

Choerops brenchleyi (1872: 426, 1873: pi. 34) was based on a

specimen stated to be IV 2 inches (about 188 mm TL) from

Misol Island. Three specimens in the British Museum

collection are registered as types from this locality (Eschmeyer,

2015). The largest (BMNH 1870.8.31.27, 153 mm SL, 184 mm

TL) is close to that length and is here regarded as the lectotype,

if not holotype. The original description gives no indication

that it was based on more than one specimen. The species is

conspecific with C. zosterophorus.

As described above, C. zosterophorus is closely related to

the very similar C. jordani, but easily distinguished by colour

pattern at adult sizes. Small juveniles are much more alike

(figs 40A & 45A).

Material examined. 9 specimens examined, 44.7-153 mm SL;

see appendix.

Incertae Sedis

Labrus chlorodus Gray, 1854 (p. 80, no locality), a name

historically associated with the genus Choerodon, was based

on a description compiled by Gronow and published along with

descriptions of other species in Gronow’s collection by Gray

long after Gronow’s death. Labrus chlorodus was subsequently

referred to C. anchorago by Gunther (1862), Fowler and Bean

(1928) and Herre (1953: 647). Although the description does

match C. anchorago, it fails to mention a diagnostic character

that would distinguish it from not only other species of the

genus but a number of other labrids, such as those in the genus

Cheilinus. Since the type has not been located in the collection

of the British Museum (Natural History) or elsewhere, the

status of the name is considered unresolved.

Acknowledgements

The study was resurrected in conjunction with two workshops

investigating demersal fishes collected by the Far Seas Fisheries

Resources Survey, hosted by Gento Shinohara and colleagues.

A special thanks to Jeff Johnson (QMB) for revisiting presumed

types and registrations for specimens in his care, especially

those of De Vis that were poorly documented in his descriptions

at best. I am grateful to Mark McGrouther (AMS) for tracking

down specimens and literature. The reanalysis of genetic

sequences was ably performed by L. Teasdale (NMV). For

assistance with accommodation and access to specimens, as

well as information pertaining to them, particularly during the

initial phase of this study, I am grateful to: J.R. Paxton, D.E

Hoese, J.M. Leis, S. Reader, A. Hay and M. McGrouther

(AMS); Hsin-HuaLin (ASIZ); J. Mclean (BMNH); J.E. Randall

and A. Suzumoto (BPBM); W.F. Eschmeyer, T. Iwamoto and P.

Sonoda (CAS); A. Graham, J. Pogonoski, W. White, D. Gledhill

and P. Last (CSIRO); Y. Kai and N. Nakayama (FAKU);

Hsuan-Chin Ho (NMMB); Dianne J. Bray (NMV); M.-L.

Bauchot, M. Desoutter and R. Causse (MNHN); K. Matsuura

and G. Shinohara (NSMT); M. Hammer, H.K. Larson and B.C.

Russell (NTM); J. Rivaton (ORSTOM); J. Johnson and R.

McKay (QMB); M. Boesemann (RMNH); M.M. Smith, E.

Heemstra and PC. Heemstra (RUSI); R. Foster (SAM), C.

Araga (SMBL); W. Klausewitz (SMF); G. Von Wahlert

(SMNS); S. Jewett, E.N. Gramblin and J. Williams (USNM);

G.R. Allen, M. Allen, B. Hutchins, S. Morison and G. Moore

(WAM); and, M.-J. Yu (THUP), unfortunately a number of

whom are also deceased. Descriptive and distributional

observations were kindly provided by R.H. Kuiter (Seaford,

Australia). Access to images was facilitated by A. Graham

(CSIRO), Y. Fukui (KAUM), H. Senou (KPM) and K. Matsuura

(NSMT). The following photographers and organisations were

especially generous in allowing me to reproduce their images:

K. Wilson (Brighton, UK, image Ccnc); J.E. Randall (BPBM);

C. Devine, T. Carter, D. Gledhill, G. Leyland, J. Pogonoski, W.

White and G. Yearsley (CSIRO); Far Seas Fisheries Laboratory,

Japan; H. Senou (KPM); D. Paul (NMV); K. Graham (NSW

Fisheries); P. Heemstra (SAIAB); P. Ryan, B. Hazes (University

of Alberta); J. Williams (USNM); G.R. Allen and B. Hutchins

(WAM); K. Berlin, J. Dubose, Izuzuki, L. Malton, G.J.

Redos, S. Sato, I. Shaw, K. Shimida, J. Shuttleworth, T.

Tsuhako, K. Uchino and K. Yamasaki. Genetic sequences for

several Japanese species were provided by S. Chiba (NSMT).

Thanks to T. Darragh (NMV), who assisted with Latin, Dutch

and German translations. Assistance with photo editing was

kindly provided by D. Paul (NMV).

M.F. Gomon

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Review of Choerodon tuskfishes

Appendix

Specimens on which species treatments are based, apart from

newly described species, are listed below and arranged

alphabetically by species. Where a registration lot comprises a

single specimen, only the length is provided; where lengths were

not recorded, the numbers of specimens are presented; question

marks (?) indicate specimen lots taken from registration lists for

which the number of specimens and their lengths were not

obtained. Registration lots comprising specimens for which

morphological data featuring in descriptions and tables were

recorded are marked with an asterisk (*); those taken from

museum registration lists for which identities were not verified

but feature in distribution maps as white symbols are indicated

by a dagger sign (^). Descriptive localities in material sections are

edited to reduce the word count because most have latitude and

longitude coordinates that convey greater accuracy. Readers

seeking additional collection information are referred to the

museums housing the relevant specimens or to their web-based

resources.

Choerodon albofasciatus nom. nov. (11 specimens examined,

96.7-152 mm standard length [SL]). Japan, Okinawa: URM-P

04294* (146) Naha fish market, 7 September 1982; URM-P

18569* (152) Naha fish market, 29 December 1986; URM-P

31237* (148) Naha fish market, 24 February 1994; URM-P

31276* (146) Naha fish market, 4 March 1994; URM-P 39636*

(121) Naha fish market, 18 December 1998; Taiwan: THUP

00750* (133); THUP 00976* (141); THUP 03355* (136).

Indonesia: CSIRO H 7220-04* (136) Lombok, Tanjung Luar

fish market, 8° 48" S, 116° 29" E, 5 November 2010. Australia,

Western Australia: CSIRO H 6452-05* (96.7) W of Shark Bay,

25° 54.13" S, 112° 49.74" E-25° 54.38" S, 112° 49.62" E, 100-

95 m, 7 December 2005, FRY Southern Surveyor, beam trawl,

SS1005/116; NMV A1784* (117) Shark Bay, 27° 33" S, 113°

14" E-27° 25" S, 113° 11" E, 109-128 m, 2 March 1981.

Choerodon anchorago. (163 specimens examined, 12-273 mm

SL) MNHN A.8208* (273, holotype of Labrus macrodontus,

skin) unknown locality; RMNH 6530* (47.3) unknown locality.

INDIAN OCEAN, Nicobar Islands: AMS B.8221 (181) Day,

1865. Andaman Islands: BMNH 70.5.18.80 (104) Andaman

Islands. PACIFIC OCEAN, Japan: BPBM7253 (138) Ryukyu

Island, Ishigaki. Vietnam: MNHNA8891 (2, ~125-180) Poulo-

Condore Islands, 8° N, 107° E; MNHN A8892 (2, ^48-68)

Poulo-Condore Islands, 8° N, 107° E. Malaysia: USNM 35717

(1) Sarawak, Sadong. Singapore: ANSP83277 (1); CAS 303771

(1); MCZ 14375 (3, 13-175). Philippines: AMS 1.21901-015*

(2, 74.7-80.0) Bolinao, 16° 25" N, 119° 53" E, 16 April 1980;

AMS 1.31430.039' (1) Dumaguete, 9° 20" N, 123° 18" E, 16

April 1986; AMS 1.31430-009+ (203) Dumaguete, 9° 20" N, 123°

18" E, 16April 1986; AMS 1.31431.004+ (1) Negros, Bais, 10° N,

122° E, 22 August 1986; AMS 1.31431-002+ (112) Negros, Bais,

10° N, 122° E, 22 August 1986; AMS 1.40150-011+ (2) Mindoro

Island, San Jose, 12° 19" 40" N, 121° 05" 16" E, 31 May 2000;

ANSP 33254 (1) Luzon, Bacin; BPBM 14215 (2,106-109) Sulu

Archipelago, Tapau Island; CSIRO H 4132-10 (93 SL) Bolinao,

7 October 1995; NSMT-P 120137* (99.2) Cebu, Mactan Island,

Lapu-lapu fish market, 16 October 1988; USNM 56078 (1)

Luzon; USNM 58030 (2) Zamboanga; USNM 112253 (1)

Palawan; USNM 126414 (1) Racon; USNM 152153 (2) Tilig;

USNM 152169* (202) Luzon, Pt Jamelo; USNM 152171 (1)

Alimango; USNM 152172 (1) Alamango Bay; USNM 152173*

(221) Mindoro, Varadero Bay; USNM 152174 (1) Mindanao;

USNM 152175 (1) Jolo; USNM 152176 (1) South Tumindao;

USNM 152177* (230) Little Santa Cruz Island; USNM 152178

(2) Masbate; USNM 152179 (1) Cebu; USNM 152339 (1)

Balabac; USNM 152341 (1) Tataan; USNM 153471 (1) Cebu;

USNM 153472* (254) Romblon, Romblon Reef; USNM 153473

(1) Juanico Strait; USNM 153474 (1) Mindanao, Puiada Bay;

USNM 153478 (236) Little Santa Cruz Island; USNM 153479

(1) Tataan; USNM 153481 (1) Mindanao, Iramucan Bay; USNM

153541 (1) Masbate Reef; USNM 153542 (2) Palawan,

Endeavour Strait; USNM 153543 (1) Mindoro, Sablayan; USNM

153544 (1) Jolo; USNM 153545 (1) Pangasinan Island; USNM

153546 (1) Burias Island, Busin Harbor; USNM 153547 (1)

Luzon, Alibijaban Island, Ragay Gulf; USNM 153548 (3)

Ulugan Bay; USNM 153549 (1) Zamboanga, Little Santa Cruz

Island; USNM 153550 (1) Mindoro, GaleraBay; USNM 153551

(1) Cataingan Bay; USNM 153632 (1) Cebu; USNM 153477 (1)

Cataingan; USNM 153480 (1) Lubana Island, Tilig; USNM

153482 (219) Libani Bay, Kait Point; USNM 153534 (1) Jolo

Island; USNM 153535 (2) Caxisagan Island; USNM 153536 (3)

Tacao Island, San Miguel Harbor; USNM 153537 (5) Palawan,

Bolalo Bay; USNM 153538 (2) Jolo Island; USNM 153539 (1)

Point Palapag; USNM 153540 (1) Mindoro, Mansalay; USNM

153632 (1) Cebu; USNM 153845* (208) Luzon, Alibijaban

Island, Ragay Gulf. Palau: BPBM 7436 (-150) Ein Malk;

BPBM 9382 (2, 40^2) Arakubisan Island; BPBM 9474 (126)

Urukthayal; BPBM 9475 (147) Iwayama Bay; NSMT-P 46217*

(108) Iwayama Bay, 1 m, 16 June 1980; NSI^-P 46226* (71.1)

Urukthapal Island, 1 m, 23 June 1980. Yap: NSMT-P 46225*

(89.7) Inuf, 1 m, 30 June 1980. Guam: USNM 113627* (6,

32.8-107) Namru; USNM 152340 (1) Apra Bay. Caroline

Islands: USNM 113628 (1) Yap. Solomon Islands: AMS

1.15767-021* (165) Inia Atoll, off Baga Island, 7° 45" S, 157° E,

1970; AMS 1.22128-148* (126) New Georgia Island, 8° 18" S,

157° 15" E, 5 m, December 1980; BMNH 76.5.1.21 (191) Yap,

Duke of York Group; BPBM 1308 (178) Shortland; BPBM

15565 (2, 22-25) Guadalcanal; CSIRO C 418 (264 TL) Mioko

Island, 4° 15" S, 152° 28" E, 29 October 1949. Vanuatu: ANSP

77780 (1) Fate. Indonesia: AMS 1.12543+ (1) Aru Islands, Dobo,

1912; BMNH 1864.5.15.18* (34.3, type of Crenilabrus

leucozonaiy, CSIRO H 7853-03* (158) Lombok, Tanjung Luar,

8° 48" S, 116° 29" E, 25 October 2008; CSIRO H 7853-04*

(220) same collection information as CSIRO H 7853-03; CSIRO

H 7853-05* (221) same collection information as CSIRO H

7853-03; CSIRO H 7899-01 (137) Bali, Kedonganan, 8° 45" S,

115° 09" E, 21 October 2008; CSIRO H 7899-02* (149) same

collection information as CSIRO H 7899-01; CSIRO H 7899-

04* (213) same collection information as CSIRO H 7899-01;

NMV A 31429-001* (208) same collection information as

CSIRO H 7899-01; CSIRO H 7985-01* (146) same locality as

CSIRO H 7899-01, 2 March 2009; MCZ 30510 (138) Sulawesi,

Macasser; MCZ 14372 (9, 84.^179) Java; MZB 23111 (96.7)

same locality as CSIRO H 7899-01, 14 July 2008; MZB 23112*

(141) same collection information as CSIRO H 7853-03; NMV

46063* (196); NTM S.10741-001 (211) Bali, 8° 40" 59" S, 115°

15" 00" E, September 1981; NTM S.11163-005 (123) Bali,

Denpasar, 8° 04" 01" S, 115° 13" 59" E, 4April 1984; QMB 1.20

(105, syntype of Choerodon weberi) Aru Islands, Dobo; QMB

1.1532 (159, syntype of Choerodon weberi) Aru Islands, Dobo;

M.F. Gomon

QMB 1.10133 (92.8, syntype of Choerodon weberi) Am Islands,

Dobo; RMNH 10892 (140) Sleyer Island; SMBL 2759* (263,

holotype of Choerops meleagris) Mare Javanicum; USNM

153482* (219) Sulawesi, Libani B., Kait Point; USNM 438444

(234) Pulau Seribu, Pulau Kongsi, 5° 51" S, 106° 36" E, 5 April

1974; WAM P.31305-004^ (73) Rian Islands, Bintan Island, 1°

11" N, 104° 19" E, 13 May 1997. Timor-Leste: AMS

1.46112.044^ (1) Dili, 8° 32" 53" S, 125° 35" 15" E, 18 September

2012; AMS 1.46112-043^ (165) Dili, 8° 32" 53" S, 125° 35" 15" E,

18 September 2012; RMNH 2361 (200) Timor; SME (263,

holotype of Choerops meleagris, mount) Java Sea. Papua New

Guinea: AMS 1B.4656* (149) 1960; AMS 1B.4657* (138) 1960;

AMS 1.12543 (106) Am Islands, Dobo; ANSP 133156* (117)

Keraward Island; BPBM 15809 (123) New Guinea Island,

Madang, 14 August 1973; CSIRO C 1810 (225 TL) Port Moresby,

21 November 1948; NMVA18465* (3, 165-187) April 1881;

NTM S.13666-031 (3, 12-63) Madang, 5° 09" 00" S, 145°

48" 00" E, 7 October 1992; QMB 1.20 (105, syntype of Chorodon

weberi) Am Islands, Dobo; QMB 1.1532 (159, syntype of

Chorodon weberi) Am Islands, Dobo; QMB 1.10133 (92.8,

syntype of Chorodon weberi) Am Islands, Dobo. Admiralty

Islands: USNM 114810 (3) Los Negros. Australia, Queensland:

AMS 1.19461-017* (64.8) Decapolis Reef, 14° 51" S, 145°

17" E, 2-A m, 14 November 1975; AMS 1.20773-052* (3, 52.5-

65.2) Nymph Island, 14° 39" S, 145° 15" E, 1-2 m, 6 December

1978; BMNH 1861.6.11.11(220, skin) Hope Island, 27 October

1860; BPBM 15967 (70) Port Douglas; QMB 1. 169U (?) Pipon

Island, off Cape Melville, 14° 7" S, 144° 31" E; QMB 1. 6454' (?)

Pencil Bay, Palm Island, 18° 46" S, 146° 34" E; QMB 1. 9907^ (?)

Dunk Island, 17° 57" S, 146° 9" E; QMB 1. 11372' (?) Palm

Group, 18° 40" S, 146° 33" E; QMB 1. 25993’ (?) Green Island,

off Cairns, 16° 46" S, 145° 58" E, 24 Eebruary 1988; QMB 1.

26944^ (?) Elinders Island, Princess Charlotte Bay, 14° 10" S,

144° 15" E, 1 October 1990.

Choerodon azurio. (40 specimens examined, 24.1-308 mm SL).

Unknown locality, China: BMNH 1851.12.27.376* (234, skin)

Warwick; BMNH 1968.3.11.26* (231, skin) J. R. Reeve

Collection; BMNH 1968.3.11.27* (264, skin) #282, J. R. Reeve

collection; CAS 61193 (1) Hong Kong; NSMT-P 55131* (198) S

coast of Hainan Island, Sanya Bay fish market, 27 Eebruary

1997; NSMT-P 60849* (129) S coast of Hainan Island, Sanya

Bay fish market, 27 November 1997; USNM 153420* (93.1)

Hong Kong; MCZ 14376 (182). Japan: ANSP 26203 (1)

Nagasaki, Hizen; BPBM 5814 (268) Tokyo; BPBM 6524 (235)

Shirahama; CAS 7046 (1); NSMT-P 18960 (24.1) Honshu, Boso

Peninsula, off Chikura, 38° 58.5" N, 140° 1.3" E, 48 m, 13

November 1973; NSMT-P 19134 (83.9) Izu, 40 m, 28 November

1968; NSMT-P72357* (226) Hyogo Prefecture, Shinonsen-chou

Humasaka, off Kanaya, 24 October 2005; NSMT-P 77942* (2,

105-117) Kochi Prefecture, Kochi City, Mimase fishing port, 8

January 2000; NSMT-P 81036* (2, 122-126) Shizuoka

Prefecture, Suruga Bay, Masaki Lighthouse, 28 May 1994;

NSMT-P 95917* (146) Kochi Prefecture, Tosashimizu City,

Kubotsu fish market, 23 July 2009; USNM 22601* (308) Awa;

USNM 50240 (1) Nagasaki; USNM 57488 (1) Tsushima; USNM

57584 (1); USNM 57669* (260); USNM 59691 (1) Hamashima;

USNM 59807* (274) Kochi; USNM 61665* (218) Tokyo;

USNM 71219 (2) Shimizu; USNM 713579 (2) Shimizu; USNM

71584 (1) Kagoshima; USNM 71800 (2) Misaki. Taiwan: CAS

7980 (1); THUP 2527* (58.6); THUP 2589* (58.9); THUP

3360* (69.6). Vietnam: MNHN 1965-232 (~170) NhaTrang.

Choerodon cauteroma. (46 specimens examined, 20-259 mm

SL) Australia, Western Australia: AMS 1.21603-002 (163)

North West Shelf, 18° 25" S, 118° 52" E, 18° 25" S, 118° 49" E,

150-154 m, 17 May 1979; AMS 1.26373-001* (2, 136-205,

paratype) Shark Bay, 26° 00" S, 113° 30" E, 8 July 1962; BPBM

30944* (110, paratype of Choerodon cauteroma) same locality

as AMS 1.26373-001, May 1964; CSIRO A 1379 (62.2) Shark

Bay, Denham Sound, 25° 43" S, 113° 15" E, August 1955;

CSIRO C 507 (148) Shark Bay, Carnarvon, Cape Ronsard, 12.6-

14.4 m, 7 September 1948; CSIRO C 2423 (100) Hampton

Harbour, 5.4 m; CSIRO C 2511 (89.6) Exmouth Gulf, 22° S,

114° E, 1954; CSIRO C 2683 (130) Exmouth Gulf, 22° S, 114°

E, 1952; CSIRO C 3871 (99.1) Exmouth Gulf, Giralia Bay, 6

September 1966; CSIRO CA 256 (259) E Monte Bello Islands,

11 May 1978; CSIRO CA 2155 (221) Monte Bello Islands, 20°

17.0" S, 116° 04.0" E-20° 17.0" S, 116° 00.0" E, 51-52 m, 31

May 1980; CSIRO CA 2156 (154) Port Hedland, 19° 36.0" S,

118° 37.0" E-19° 38.0" S, 118° 40.0" E, 36 m, 4 June 1980;

CSIRO CA 2157 (206) Port Hedland, 19° 36.0" S, 118°

37.0" E-19° 38.0" S, 118° 40.0" E, 36 m, 4 June 1980; CSIRO

CA 2160 (179) Monte Bello Islands, 20° 15.0" S, 115° 56.0" E,

20° 17.0" S, 115° 55.0" E, 48-50 m, 27 May 1980; CSIRO H

659-1 (176, paratype) Dampier Archipelago, Cape Preston, 20°

48.0" S, 116° 00.0" E-20° 46.0" S, 115° 59.0" E, 19-22 m, 1

December 1979; CSIRO H 4043-0U (320) Port Hedland, 19°

45.8" S, 117° 57.7" E-19° 46.7" S, 117° 58.5" E, 46^8 m, 2

September 1995; CSIRO H 4101-0U (1) Point Samson, 20° S,

117° E, September 1995; CSIRO H659-01* (176, paratype of

Choerodon cauteroma) Exmouth Gulf, October 1974; CSIRO H

7198-31 (2, 108-128) Shark Bay, Carnarvon, 25° 20" S, 113°

30" E, 30 m, 25 May 1995; NMV A1918 (212) NW of Dampier,

20° 07" S-20° 02" S, 116° 12" E-116° 08" E, 60 m, 9 March

1981; NMV A3819* (4, 114—208, paratypes of Choerodon

cauteroma) same collection data as AMS 1.26373-001; NTM

S.10574-006 (212) Dampier Archipelago, 20° 03" 00" S, 115°

47" 60" E, 54 m, 7 May 1982; NTM S.11688-002 (150) North

West Shelf, King Sound, 16° 31" 59" S, 121° 25" 59" E, 42 m, 17

April 1985; NTM S.17058-002 (157) North West Shelf, Bedout

Island, 19° 22" 59" S, 118° 28" 01" E, 70 m, 25 January 1984;

USNM 280629* (2, 115-158, paratypes of Choerodon

cauteroma) same collection data as AMS 1.26373-001; WAM

P.21277-001' (1) Dampier Archipelago, Elphick Knob, 20°

28" S, 116° 37" E, 6 November 1971; WAM P.22960-00U (1)

Dampier Archipelago, Elphick Knob, 20° 28" S, 116° 37" E, 5

November 1971; WAM P.22959-00U (1) Dampier Archipelago,

Kendrew Island, Museum Beach, 20° 29" S, 116° 32" E, 17

Eebruary 1973; WAM P.5701-00U (1) Shark Bay, 25° 21" S,

113° 44" E, September 1960; WAM P.23092-001 (3, 20-10)

Dampier Archipelago, Elphick Knob, 20° 28" S, 116° 37" E, 26

November 1971; WAM P.24259-001* (195, paratype) Dampier

Archipelago, Kendrew Island, 20° 29" S, 116° 32" E, 24 Eebruary

1974; WAM P.25095-038* (6, 98-144, paratypes) Exmouth

Gulf, 22° 05" S, 114° 15" E, 0-12 m, October 1974; WAM

P.25372-001* (3, 90-104, paratypes) Exmouth Gulf, 21° 55" S,

114° 09" E, 4-5 m, 1 July 1975; WAM P.25508-042* (203,

holotype) Exmouth Gulf, 21° 57" S, 114° 12" E, 20-30 m, 6

December 1975; WAM P.30083-006^ (29) Shark Bay, Cape

Ransonnet, 26° 09" S, 113° 13" E, ^5 m, 30 March 1990; WAM

P.30087-009 (2, 4T49) Shark Bay, Useless Loop, 26° 09" S,

Review of Choerodon tuskfishes

113° 26" E, 0.5-1.5 m, 2 April 1990; WAM P.30239-005^ (2,

110-125) Exmouth Gulf, 22° 15" 00" S, 114° 20" 00" E, 24

March 1991; WAM P.31391-023^ (3, 12-15) Beagle Bay, 16°

59" S, 122° 40" E, 9-10 m, 28 September 1997; WAM P32317-

008 (133) Shark Bay, Cape Peron North, 25° 22.558" S, 113°

17.597" E, 11.5-11 m, 7 October 2002.

Choerodon cephalotes. (198 specimens examined, 29.1-253

mm SL). Australia, New South Wales: AMS 1.25419-001*

(107, holotype of Choerops macleayi, formerly E.860A) Port

Jackson, 33° 50" S, 151° 15" E, 1984. Queensland: AMS E.1471

(237) Double Island Point, 26° 25" S, 153° 30" E, 60 m, 28 June

1910; AMS E.1530 (139) Wide Bay, 25° 52" S, 153° 07" E,

1909; AMS E.1657 (158) Eraser Island, 25° 23" S, 153° 12" E,

27 m, 29 June 1910; AMS E.1658 (183) same collection data as

AMS 1.1657; AMS E.2601 (5, 92-190) Hervey Bay, 25° S, 152°

E, 20 m, 27 July 1910; AMS E.2602* (5, 96.^150) same

collection data as AMS 1.2601; AMS E.2603 (5, 105-168) same

collection data as AMS 1.2601; AMS E.2604 (5, 95-167) same

collection data as AMS 1.2601; AMS E.2605 (5, 85-143) same

collection data as AMS 1.2601; AMS E.2606 (4, 88-144) same

collection data as AMS 1.2601; AMS E.2885 (133) Rock Cod

Shoal, Bustard Head Light, 23° 52" S, 151° 37" E, 31 m, 9

August 1910; AMS E.2927 (165) Hervey Bay, 24° S, 152° E, 20

m, 27 July 1910; AMS E.2929 (2) same collection data as AMS

1.2927; AMS E.2930 (2) same collection data as AMS 1.2927;

AMS E.2931 (6) same collection data as AMS 1.2927; AMS

1.10934 (180) Double Island Point, 26° 00" S, 153° 00" E, 28

June 1910; AMS 1.10990 (173) Wide Bay, 25° 52" S, 153° 07" E,

1910; AMS 1.15421-023 (100) Townsville, Magnetic Island, 19°

10" S, 146° 50" E, 1968; AMS 1.15557-203 (8,84.1-162) Gulf of

Carpentaria, 17° 25" S, 140° 10" E, 10.1 m, 27 November 1963;

AMS 1.18248-001 (160) locality unknown, 1909; AMS 1.20751-

007 (5, 70.1-139) Lizard Island, 14° 38" S, 145° 24" E, 25 m, 8

Eebruary 1979; AMS 1.20752-011 (2, 108-120) Lizard Island,

14° 30" S, 145° 22" E, 20 m, 8 Eebruary 1979; AMS 1.20753-

007* (10, 48.2-78.0) Nymph Island, 14° 36" S, 145° 14" E, 14-

15 m, 8 Eebruary 1979; AMS 1.20771-067 (4, 89.1-134) Cape

York, Capt. Billy Creek, 11° 37" S, 142° 56" E, 16-18 m, 18

Eebruary 1979; AMS 1.20771-092^ (134) same collection data as

AMS 1.20771-067; AMS 1.20827-008 (2, 92.0-102) Cape York,

Hannibal Island, 11° 33" S, 142° 57" E, 22-23 m, 15 Eebruary

1979; AMS 1.20829-001 (2, 99.7-103) Lizard Island, Decapolis

Reef, 14° 50" S, 145° 15" E, 8 m, 25 Eebruary 1979; AMS

1.27745-009 (100) Silkwood, 17° 45" S, 146° 01" E, 1925; AMS

1.34301-051* (3 , 50.^85.9) Shoalwater Bay, Sabina Point, 22°

23" 45" S, 150° 18" 14" E, 0.3 m, 14 September 1993; AMS

1.34318-055 (29.1) Townshend Island, 22° 12" 14" S, 150°

28" 32" E, 0.2 m, 16 September 1993; AMS 1.9490 (105) Wide

Bay, 25° 52', 153° 07" E, 1908; AMS IA.6724* (100) Lindeman

Island, Kennedy Sound and Shaw Island, 20° 27" S, 149° 03" E,

1936; AMS IA.6783 (2, 42.5-102) Lindeman Island, 20° 27" S,

149° 02" E, 1936; AMS IB.1037 (16) Eraser Island, 24° 52" S,

152° 48" E, 27 m, 14 September 1938; AMS IB. 1156 (160) same

collection data as AMS IB. 1037; AMS IB.5875 (99.6) Townsville,

19° S, 146° E, 1962; BMNH 1871.9.13.68 (115) Point Bower;

CSIRO A 774 (47.1) Eraser Island, 26° S, 150° E, 4 September

1947; CSIRO C 3260 (137) Gulf of Carpentaria, Mornington

Island, 16° 34.0" S, 139° 56.4" E, 29 m, 29 August 1963; CSIRO

C 3280 (121) Gulf of Carpentaria, Nicholson River, 17° 22.7" S,

139° 40.4" E, 5.4 m, 20 August 1963; CSIRO C 3281 (118) Gulf

of Carpentaria, Nicholson River, 17° 22.7" S, 139° 40.4" E, 5.4

m, 20 August 1963; CSIRO C 3315 (129) Gulf of Carpentaria,

16° 52" S, 140° 55" E, 10.8 m, 5 August 1963; CSIRO C 3341

(108) Gulf of Carpentaria, 17° 02" S, 139° 55" E, 13 August

1963; CSIRO C 3368 (102) Gulf of Carpentaria, 16° 59" S, 139°

50' E, 14.4 m, 12 August 1963; CSIRO C 3369 (107) Gulf of

Carpentaria, 16° 59' S, 139° 50' E, 14.4 m, 12 August 1963;

CSIRO C 3402 (104) Gulf of Carpentaria, Wellesley Islands, 16°

38.7' S, 140° 05.1' E, 25.2 m, 28 August 1963; CSIRO C 3428

(111) Gulf of Carpentaria, 16° 29' S, 140° 03' E, 27.0 m, 2

September 1963; CSIRO C 3447 (135) Gulf of Carpentaria,

Pisiona Island, 25.2 m, 2 September 1963; CSIRO C 4044 (125)

Gulf of Carpentaria, Karumba, 16° 24.8' S, 140° 58.0' E, 12.6 m,

24 August 1963; CSIRO C 4169 (112) Gulf of Carpentaria,

between Sweers and Bountiful islands, 16° 52' S, 139° 45' E, 14

m, 5 November 1971; CSIRO C 4563 (121) Gulf of Carpentaria,

30 miles NW of Eairway Buoy, 16 m, 30 October 1972; CSIRO

CA 773^ (89 TL) Torres Strait, Thursday or Prince of Wales

islands, 10° 40' S, 142° 15' E, April 1979; CSIRO H 3305-03

(120) Shelburne Bay, 11° 48.8' S, 143° 12.8' E-ll° 47.2' S, 143°

11.0' E, 31 m, 14 January 1993; CSIRO H 3307-11 (2, 105-106)

Hannibal Island, 11° 41.7' S, 143° 00.9' E-ll° 43.7' S, 143°

01.7' E, 17 m, 14 January 1993; CSIRO H 3355-09^ (156) Prince

of Wales Island, 10° 50.6' S, 141° 12.7' E-10° 51.7' S, 141°

11.2' E, 27 m, 3 Eebruary 1993; CSIRO H 3896-01' (232) Cape

Grenville, 11° 42.06' S, 143° 38.86' E-ll° 41.85' S, 143°

37.38' E, 29 m, 28 October 1994; CSIRO H 5958-08 (104) Cape

Elattery, 14° 48.5' S, 145° 15.4' E, 18 m, 11 May 2002; CSIRO H

6756-11 (137) NE of Bundaberg, 24° 21.35' S, 152° 40.28' E, 14

April 2004; CSIRO H 6893-09 (2, 125-136) Palm Islands, 18°

32' S, 146° 34' E, 31 m, 27 November 2003; CSIRO H 7676-04

(82) W of Northumberland Islands, 21° 35.12' S, 149° 42.53' E,

28 April 2004; CSIRO H 7895-01 (149) Torres Strait, E of The

Three Sisters, 10° 15.51' S, 142° 55.60' E, 12 January 2004;

NMV A13308 (88.0) 18 km W of Saunders Island, 11°

41' 42.00" S, 143° 00' 53.99" E, 18-17 m, 14 January 1993;

NMV A13309 (104) 18 km W of Saunders Island, 11°

41' 42.00" S, 143° 00' 53.99" E, 18-17 m, 14 January 1993;

NMV A13310 (186) Gulf of Carpentaria, 114 km W of Prince of

Wales Island, 10° 50' 35.99" S, 141° 12' 42.00" E, 27 m, 3

Eebruary 1993; NMVA13311 (151) Gulf of Carpentaria, 109 km

W of Prince of Wales Island, 10° 49' 12.00" S, 141° 09' 18.00" E,

29 m, 3 Eebruary 1993; NMV A13312 (180) Gulf of Carpentaria,

70 km W of Weipa, 12° 39' 06.00" S, 141° 12' 54.00" E, 40 m, 22

January 1993; NMV A13313 (122) Gulf of Carpentaria, 114 km

W of Prince of Wales Island, 10° 50' 35.99" S, 141° 12' 42.00" E,

27 m, 3 Eebruary 1993; NMV A13314 (130) Gulf of Carpentaria,

114 km W of Prince of Wales Island, 10° 50' 35.99" S, 141°

12' 42.00" E, 27 m, 3 Eebruary 1993; NMVA13315 (2,114-141)

Gulf of Carpentaria, 111 km W of Prince of Wales Island, 10°

49' 48.00" S, 141° 08' 12.00" E, 30 m, 4 Eebruary 1993; NMV

A13316 (190) Gulf of Carpentaria, 105 km W of Prince of Wales

Island, 10° 47' 35.99" S, 141° 11' 30.00" E, 27 m, 3 Eebruary

1993; NMV A14169 (95) Gulf of Carpentaria, 70 km W of

Weipa, 12° 39' 06.00" S, 141° 12' 54.00" E, 40 m, 22 January

1993; NMVA18718 (150) off Innisfail, 17° 27' 56.99" S, 146°

07' 06.60" E, 25 January 1996; NMV A2921 (2, 120-125) Gulf

of Carpentaria, 45 km W of Cullen Point, 11° 55' 11.99" S, 141°

28' 11.99" E, 33 m, 7 September 1982; NMV A3080 (2, 145-

210) Gulf of Carpentaria, 55 km NNW of Duyfken, 12°

07' 11.99" S, 141° 26' 59.99" E, 36 m, 16 September 1982; NMV

M.F. Gomon

A3135 (3, 135-160), Gulf of Carpentaria, 60 km W of Weipa,

12° 36' 18.00" S, 141° 19' 55.80" E, 40 m, 25 February 1983;

NMV A3157 (3, 115-120) Great Barrier Reef, near Haggerstone

Island, 12° 03' 16.92" S, 143° 15' 10.08" E, 26-29 m, 27 February

1983; NMV A3182 (2,105-145) Gulf of Carpentaria, 155 km W

of Cape York, 10° 49' 43.20" S, 141° 08' 32.39" E, 32 m, 26

February 1983; NMV A3188 (4, 84.8-131) Cape York, 60 km E

of Cape Melville, 14° 40' 47.99" S, 145° 03' 48.60" E, 12.5 m, 28

February 1983; NTM S. 13272-005 (2, 73-93) Gulf of

Carpentaria, Booby Island, 10° 42' 00" S, 141° 39' 00" E, 12 m,

29 November 1991; NTM 8.13273-017(4, 60-94) Gulf of

Carpentaria, Booby Island, 10° 43' 59" S, 141° 52' 59" E, 10 m,

29 November 1991; QMB 1.289^ (?)Moreton Bay, Myora Banks,

27° 28' S, 153° 25' E; QMB+1.692+ (?)Cape York, Cape Grenville,

11° 58' S, 143° 14' E; QMB 1.945 (266, syntype of Choerops

perpulcherl, mount) Moreton Bay, IT 15' S, 153° 15' E; QMB

1.1054+ (?) Moreton Bay, Bulwer, 27° 5' S, 153° 22' E; QMB

1.1055+ (?) Moreton Bay, Bulwer, 27° 5' S, 153° 22' E; QMB

1.1056+ (?) Moreton Bay, Bulwer, 27° 5' S, 153° 22' E; QMB

1.1212+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB 1.1331+ (?)

Southport, 27° 58' S, 153° 25' E; QMB 1.2110+ (?) Moreton Bay,

27° 15' S, 153° 15' E; QMB 1.2130+ (?) Moreton Bay, 27° 15' S,

153° 15' E; QMB 1.2146+ (?) Moreton Bay, 27° 15' S, 153° 15' E;

QMB 1.3067+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB

l. 3113+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB 1.3347+ (?)

S coast of Queensland; QMB 1.9919* (165) Great Sandy Strait,

25° 20' N, 153° E; QMB 1.9920* (229, syntype of Choerops

perpulcherl) Moreton Bay, 27° 15' N, 153° 15' E; QMB 1.11408+

(?) Moreton Bay, 27° 15' S, 153° 15' E; QMB 1.11409+ (?) off

Caloundra, 26° 48' S, 153° 15' E; QMB 1.11778+ (?) Far North

Queensland coast; QMB 1.12164+ (?) off Cairns, 16° 50' S, 146°

E; QMB 1.13462+ (?) Gulf of Carpentaria, 17° 20' S, 139° 32' E,

5.9 m, 8 December 1963; QMB 1.14275+ (?) Dampier, 20° 39' S,

116° 43' E, 1977; QMB 1.15240+ (?) Torres Strait, 9° 49' S, 142°

48' E, 12.8-14.6 m, 4 April 1974; QMB 1.15625+ (?) 2 miles NW

of Nymph Island, 14° 36' S, 145° 14' E, 15 m, 8 February 1979;

QMB 1.15673+ (?) 5 miles WNW of Lizard Island, 15° 30' S,

145° 22' E, 20 m, 8 February 1979; QMB 1.15969+ (?) Princess

Charlotte Bay, 14° 20' S, 144° 7' E, 12-15 m, 23 February 1979;

QMB 1.16163+ (?)2 miles NE of Hannibal Island, 11° 33' S, 142°

57' E, 21.9 m, 15 February 1979; QMB 1.17532+ (?) Torres Strait,

9° 28' S, 142° 53' E, 7 m, 24 March 1974; QMB 1.17533+ (?)

Torres Strait, 9° 35' S, 143° 2' E, 10.1-11.9 m, 26 March 1974;

QMB 1.17534+ (?) Torres Strait, 9° 37' S, 142° 47' E, 8.2-12.8 m,

27 March 1974; QMB 1.17535+ (?) Torres Strait, 9° 39' S, 142°

47' E, 8.2-10.1 m, 28 March 1974; QMB 1.17536+ (?) Torres

Strait, 9° 40' S, 142° 56' E, 11 m, 29 March 1974; QMB 1.17537+

(?) Queensland, 9° 47' S, 142° 43' E, 13.7-14.6 m, 2 April 1974;

QMB 1.17538+ (?) Torres Strait, 9° 49' S, 142° 49' E, 14.6-18.3

m, 4 April 1974; QMB 1.17539+ (?) Torres Strait, 9° 51' S, 142°

49' E, 14.6-18.3 m, 5 April 1974; QMB 1.17540+ (?) Torres Strait,

9° 53' S, 142° 48' E, 15.5-17.4 m, 6 April 1974; QMB 1.17541+

(?) Torres Strait, 9° 47' S, 142° 43' E, 13.7-14.6 m, 2 April 1974;

QMB 1.17542+ (?) Torres Strait, 9° 58' S, 142° 43' E, 15.5-17.4

m, 8 April 1974; QMB 1.17543+ (?) Torres Strait, 10° S, 142°

42' E, 13.7-15.5 m, 8 April 1974; QMB 1.17544+ (?) Torres Strait,

9° 55' S, 141° 33' E, 12.8-14.6 m, 31 May 1974; QMB 1.17545+

(?)Torres Strait, Dalrymple Islet, 10° 2' S, 142° 41' E, 14 October

1974; QMB 1.17546+ (?) Torres Strait, Yorke Island, 9° 44' S,

143° 25' E, 15 October 1974; QMB 1.17547+ (?) Thursday Island

Harbour, 10° 35' S, 142° 13' E, 20 March 1974; QMB 1.17548+

(?) Torres Strait, Keats Island, 9° 41' S, 143° 27' E, 12 October

1974; QMB 1.17549+ (?) Torres Strait,Stephen to Bramble Cay,

9° 20' S, 143° 42' E, 32.9-34.7 m, 13 October 1974; QMB

1.18099+ (?) Princess Charlotte Bay, 14° 5' S, 144° 4' E, 25.6 m,

19 September 1979; QMB 1.18141+ (?) E of Hannah Island, 13°

52' S, 143° 51' E,31.1 m, 23 September 1979; QMB 1.18211+ (?)

NE of Lizard Island, 14° 20' S, 145° 51' E, 23.8 m, 18 September

1979; QMB 1.18249+ (?) NW of Bird Island, 11° 45' S, 143° 3' E,

20.1 m, 27 September 1979; QMB 1.18263+ (?) ESE of Wilkie

Island, 13° 45' S, 143° 47' E, 18.3 m, 28 September 1979; QMB

1.18270+ (?) Princess Charlotte Bay, 14° 10' S, 144° 9' E, 21.9 m,

29 September 1979; QMB 1.21195+ (?) Gulf of Carpentaria, 16°

51' S, 139° 52' E, 17 m, January 1983; QMB 1.21337+ (?) SE of

Lady Elliot Island, 24° 7' S, 152° 43' E, 46-55 m, 9 March 1982;

Qhffi 1.21448+ (?) mouth of Moon Creek, Fraser Island, 25°

14' S, 153° E, 1-A m, 3 October 1984; QMB 1.22724+ (?)

Grasstree Creek, Sarina Beach, 21° 22' S, 149° 18' E, 14 April

1987; QMB 1.23475+ (?) off Hinchinbrook Island, 18° 27' S, 146°

25' E, 24 m, 15 April 1985; QMB 1.25522+ (?) 40 miles NE of

Momington Island, 16° 4' S, 140° 3' E, February 1988; QMB

1.26809+ (?) Flinders Island, Princess Charlotte Bay, 14° 10' S,

144° 15' E, 1 July 1990; QMB 1.27732+ (?) Gulf of Carpentaria,

10° 20' S, 141° 9' E, 23 m, 3 December 1990; QMB 1.34458+ (?)

Sweers Island, rocky reef ca 400 m off SW tip, 17° 8' S, 139°

36' E, 0.5-2 m, 16 November 2002; QMB 1.34622+ (?) Sweers

Island, 150 m off beach at centre of E side, 17° 7' S, 139° 37' E,

0.3-0.5 m, 23 November 2002; QMB 1.35161+ (?) 15 36.9' S, 145

22.5' E-15 36.9' S, 145 22.5' E, 29 September 2003; QMB

l. 35806+ (?) 19.53.7' S, 148.12.9' E-19.53.7' S, 148.12.9' E, 12

m, 6 December 2003; QMB 1.9919+ (165) Moreton Bay, Great

Sandy Strait. Northern Territory: AMS 1.21848-004+ (200)

Arafura Sea, 11° 17' S, 134° 05' E-ll° 18' S, 134° 02' E, 40^

m, 18 November 1980; CSIRO C 3981 (159) Gulf of Carpentaria,

Robinson River, 15° 55' S, 137° 30' E, 31 October 1971; NTM

S.10050-001 (2, 100-110) Darwin Harbour, 12° 16' 01" S, 130°

54' 00" E, 4April 1977; NTM S.10939-005 (114) Groote Eylandt,

Bickerton Island, 13° 58' 01" S, 136° 18' 00" E, 17 m, 1 March

1983; NTM S. 10941-007 (131) Groote Eylandt, Bickerton

Island, 14° 04' 59" S, 136° 04' 59" E, 15 m, 2 March 1983; NTM

S. 13258-004 (217) Gulf of Carpentaria, Cape Grey, 13° 07' 59" S,

136° 35' 60" E, 19 m, 23 November 1991; NTM S. 12941-008 (3,

175-195) Arafura Sea, 11° 28' 59" S, 133° 35' 60" E, 23 m, 22

October 1990; NTM S.12958-011 (159) Arafura Sea, 11°

36' 00" S, 135° 31' 01" E, 33 m, 26 October 1990; NTM S. 12960-

002 (160) Marchinbar Island, Arafura Sea, 11° 13' 59" S, 135°

58' 01" E, 38 m, 26 October 1990; NTM S. 13333-012 (226)

Arafura Sea, 11° 19' 01" S, 134° 28' 01" E, 39 m, 23 October

1990; NTM S. 12445-069 (165) Cobourg Peninsula, Orontes

Reef, 11° 05' 60" S, 132° 04' 01" E, 20 m, 10 August 1986;

USNM 174393 (227); WAM P.14391-001+ (1) Darwin area, 12°

15' S, 130° 25' E, September 1965. Western Australia: AMS

1.22802-010* (3, 18^233) North West Shelf, Port Hedland, 19°

39' S, 117° 53' E, 52-53 m, 26 March 1982; CSIRO C 508 (141)

Shark Bay, Carnarvon, Cape Ronsard, 12.6-14.4 m, 7 September

1948; CSIRO C 541 (132) Shark Bay, Cape Levillian, Dirk

Hartog Island, 13 September 1948; CSIRO C 2315 (106) Shark

Bay, Cape Peron, 25° 30' S, 113° 30' E, 10.8 m, 27 August 1953;

CSIRO C 2339 (131) Exmouth Gulf, 22° S, 114° E, 1954; CSIRO

C 2497 (128) Exmouth Gulf, 22° S, 114° E, 1954; CSIRO C

2524 (109) Shark Bay, Denham Sound, 25° 43' S, 113° 15' E,

August 1955; CSIRO C 2678 (133) Dampier Archipelago,

Review of Choerodon tuskfishes

Lancelin; CSIRO C 3869 (124) Exmouth Gulf, Giralia Bay, 6

September 1966; CSIRO C 3870 (113) Exmouth Gulf, Giralia

Bay, 6 September 1966; CSIRO CA 255 (202) Monte Bello

Islands, 11 May 1978; CSIRO CA 1728 (149) Barrow Island, 20°

47.0' S, 115° 59.0' E-20° 49.0' S, 115° 59.0' E, 20 m, 6 December

1979; CSIRO CA 2144 (200) Eorestier Island, 19° 52.0' S, 118°

12.0' E-19° 53.0' S, 118° 14.0' E, 38 m, 3 June 1980; CSIRO CA

2154 (253) Monte Bello Islands, 20° 15.0' S, 115° 56.0' E-20°

17.0' S, 115° 55.0' E, 48-50 m, 27 May 1980; CSIRO CA 2159

(158) Admiralty Gulf, 14° 01' S, 125° 34' E-14° 03' S, 125°

33' E, 44 m, 25 June 1980; CSIRO H 2782-03 (128) North West

Shelf, Port Hedland, 19° 29.7' S, 118° 52.2' E-19° 29.2' S, 118°

52.7' E, 39 m, 24 October 1983; CSIRO H 4028-01' (266)

Dampier Archipelago, 20° 18.3' S, 116° 29.8' E-20° 18.6' S, 116°

28.4' E, 40 m, 28 August 1995; NTM S. 11924-014 (136) Dampier

Archipelago, 20° 04' 01" S, 116° 37' 01" E, 80 m, 10 June 1986;

NTM S.14362-015 (175) Holothuria Banks, West Holothuria

Reef, 13° 25' 12" S, 125° 39' 18" E, 50 m, 12 June 1996; NTM

S.17058-003 (150) North West Shelf, Bedout Island, 19°

22' 59" S, 118° 28' 01" E, 70 m, 25 January 1984; QMB 1.10234^

(?) Shark Bay, 30 miles S of Carnarvon, 25° 16' S, 113° 40' E, 3

June 1972; QMB 1.2320P (?) Exmouth Gulf, 22° S, 114° 20' E,

9-11 m, 2 May 1972; WAM P.25397-018' (8, 116-140) Rowley

Shoals, 17° 29' S, 121° 52' E, 0^2 m, 21 December 1969; WAM

P.26197-010^ (175) 20° 05' 00" S, 117° 05' 00" E, 0^ m, 17

May 1978; WAM P.25354-022+ (3, 137-162) Monte Bello

Islands, 20° 25' S, 115° 30' E, April 1975; WAM P.25354-053^

(250) Monte Bello Islands, 20° 25' S, 115° 30' E, April 1975;

WAM P.25508-04P (129) Exmouth Gulf, 21° 57' S, 114° 12' E,

20-30 m, 6 December 1975; WAM P.2489-00P (1) Exmouth

Gulf, 22° 05' S, 114° 15' E, October 1943; WAM P.23798-00P

(1) Exmouth Gulf, Sunday Island, 22° 05' S, 114° 15' E, 12.8-

18.2 m, September 1973; WAM P.25095-018^ (8, 87-176)

Exmouth Gulf, 22° 05' S, 114° 15' E, 0-12 m, October 1974;

WAM P.25095-033^ (5, 98-119) Exmouth Gulf, 22° 05' S, 114°

15' E, 0-12 m, October 1974; WAM P.30239-006^ (2, 86-190)

Exmouth Gulf, 22° 15' 00" S, 114° 20' 00" E, 24 March 1991;

WAM P.23455-00P (1) Learmonth, 22° 15' S, 114° 05' E, June

1973; WAM P.23683-00P (1) Carnarvon, 24° 53' S, 113° 40' E,

29.8 m, 19 July 1972; WAM P.13953-00P (1) Shark Bay, 25°

21' S, 113° 44' E, August 1965; WAM P.32279-002+ (97) Shark

Bay, Cape Peron North, 25° 23.122' S, 113° 25.822' E, 16.8-16.9

m, 3 October 2002; WAM P.31978-024' (85) Shark Bay, 25°

25' S, 113° 35' E, November 1998; WAM P.32267-004' (80)

Shark Bay, Cape Peron North, 25° 30.544' S, 113° 33.699' E,

13.2-13 m, 1 October 2002.

Choerodon cyanodus. (150 specimens examined, 19.8-300 mm

SL). Australia: BMNH 1876.5.11.32 (230) “Tahiti”, Godeffroy.

New South Wales: AMS I.45736-00P (1) Port Jackson, 33°

52' S, 151° 50' E, 33° 48' S, 151° 54' E. Queensland: AMS 1.391

(202) Cardwell, 18° 16' S, 146° OP E, 1886; AMS 1.3433 (275)

Thursday Island, 10° 35' S, 142° 13' E, 1895; AMS 1.3434 (110)

same collection data as AMS 1.3433; AMS 1.6076 (211) Northern

Territory, Port Darwin, 12° 27' S, 130° 48' E, 1903; AMS 1.9487

(122) Wide Bay, 25° 52' S, 153° 07' E, 1908; AMS 1.11024' (1)

Eraser Island, Boomerang Hill (5 miles NW), 25° 20' S, 153°

17' E, 27, 29 June 1910; AMS 1.11929 (189) Cape York, 10°

00' S, 142° 00' E, 1907; AMS 1.11930 (193) same collection data

as AMS 1.11929; AMS 1.11931* (176) same collection data as

AMS 1.11929; AMS 1.11932 (165) same collection data as AMS

1.11929; AMS 1.11933 (165) same collection data as AMS

1.11929; AMS 1.11934 (175) same collection data as AMS

1.11929; AMS 1.11935 (169) same collection data as AMS

1.11929; AMS 1.11936* (178) same collection data as AMS

1.11929; AMS 1.11937* (166) same collection data as AMS

1.11929; AMS 1.12542 (135) Great Sandy Strait, 23° 35' S, 152°

58' E, 1912; AMS 1.12632 (220) Masthead Island, 23° 32' S, 151°

44'E, August 1912;AMS 1.15557-207* (134) Gulf of Carpentaria,

17° 25' S, 140° 10' E, 10.1, 27 November 1963; AMS 1.15680-

018 (250) One Tree Island, 23° 30' S, 152° 05' E, 1 m, 24

November 1969; AMS I.18219-00P (337) One Tree Island, 23°

30' S, 152° 05' E, 4 December 1974; AMS 1.19356-033 (2, 3A-

116) Prince of Wales Island, 10° 41' S, 142° 07' E, 1 m, 2 July

1976; AMS 1.20464-019 (195) One Tree Island, 23° 30' S, 152°

05' E, 1 m, 8 October 1972; AMS 1.20773-025' (103) Nymph

Island, 14° 39' S, 145° 15' E, 1-2 m, 6 December 1978; AMS

l. 20776-028 (2, 50-58) Cape York, Ealse Orfordness, 11° 23' S,

142° 52' E, 3-4 m, 18 Eebruary 1979; AMS 1.22072-029 (90)

Port Douglas, 16° 29' 58" S, 145° 28' 00" E, 1 m, 24 September

1980; AMS 1.26248-014 (152) Gulf of Carpentaria, between

Sweers and Little Bountiful islands, 16° 45' S, 139° 50' E, 15 m,

January 1971; AMS 1.33759-006' (183) Bet Reef, 10° 10' 32" S,

142° 56' 01" E, 18-21 m, 30 January 1993; AMS 1.34301-048

(37) Shoalwater Bay, Sabina Point, 22° 23' 45" S, 150° 18' 14" E,

0.3 m, 14 September 1993; AMS 1.34301-052 (3, 80-95)

Shoalwater Bay, Sabina Point, 22° 23' 45" S, 150° 18' 14" E, 0.3

m, 14 September 1993; AMS 1.34302-002^ (240) Shoalwater

Bay, Sabina Point, 22° 23' 49" S, 150° 18' 15" E, 2 m, 14

September 1993; AMS 1.34311-020 (14, 43-136) Shoalwater

Bay, Collins Island, 22° 14' 47" S, 150° 19' 08" E, 3 m, 15

September 1993; AMS 1.34311-029' (11, 50-138) Shoalwater

Bay, Collins Island, 22° 14' 47" S, 150° 19' 08" E, 3 m, 15

September 1993; AMS 1.34318-023 (3, 62-106), AMS 1.34318-

024 (36.7) Townshend Island, 22° 12' 14" S, 150° 28' 32" E, 0.2

m, 16 September 1993; AMS IA.2591 (1) Capricorn Group,

North West Island, 23° 30' S, 152° 00' E, 1926; AMS IA.2725

(220) Harvey Bay, 25° S, 152° E, 1926; AMS IA.6381 (150)

Hayman Island, 20° 03' S, 148° 53' E, 1935; AMS lA. 6723 (1)

Lindeman Island, Kenney Sound and Shaw Island, 20° 27' S,

149° 03' E, 1936; AMS IB.2170 (1, head) islands off Emu Park,

23° 15' S, 150° 53' E, 1948; AMS IB.2171 (1, head) same

collection data as AMS IB.2170; AMS IB.3870 (1) Heron Island,

23° 26' S, 151° 55' E, 1957; AMS IB.8343' (260) Townsville,

Bay Rock, 19° 07' S, 146° 45' E, 1966; BMNH 1847.6.17.57

(179, type of Labrus arilcal, skin) Endeavour Straits, Bramble

Island, 13 Eebruary 1845; BMNH 1850.7.20.65* (195, skin)

Cape York, 31 October 1849, HMS Rattlesnake; BMNH

1911.4.1.30-31 (2,166-168) Northern Queensland, Saville Kent;

CSIRO C 4239 (220) Gulf of Carpentaria, Lakes Islet, 29 October

1971; CSIRO C 4479 (157) Gulf of Carpentaria, Bentnick Island,

16° 55' S, 139° 37' E, 12.5 m, 1 November 1972; CSIRO H 7673-

03 (127) Torres Strait, E of Banks Island, 10° 06.74' S, 142°

39.63' E, 12 January 2004; QMB 1.95* (121) Cardwell, 18° 16' S,

146° 1' E; QMB 1.110* (226, type of Choerops albigena) Cape

York, 10° 41' S, 142° 32' E; QMB 1.852' (?) Capricorn Group,

Masthead Island, 23° 32' S, 151° 44' E; QMB 1.853' (?) same

collection data as QMB 1.852; QMB 1.946 (117, type of Choerops

concolor, mount) North-East Queensland; QMB 1.1332' (?)

Great Sandy Strait, 25° 20' S, 153° E; QMB 1.1870' (?) Magnetic

Island, 19° 8' S, 146° 50' E; QMB 1.4734' (?) Dunk Island, 17°

57' S, 146° 9' E; QMB 1.4737' (?); QMB 1.5471' (?) Heron

M.F. Gomon

Island, 23° 27' S, 151° 55' E; QMB 1.5524^ (?) Heron Island, 23°

27' S, 151° 55' E; QMB 1.5600^ (?) Salamander Rocks, off Cape

Cleveland, 19° 11' S, 147° 4' E; QMB 1.5840^ (134) Great Palm

Island, Butler Bay, 18° 46' S, 146° 36' E; QMB 1.5841 (1) Great

Palm Island, Butler Bay, 18° 46' S, 146° 36' E; QMB 1.5876^ (?)

Whitsunday Group, Shaw Island, 20° 29' S, 149° 5' E; QMB

1.6109* (1) Townsville; QMB 1.6145^ (?) Eour Eoot Rocks, off

Cape Cleveland, 19° 11' S, 147° 3' E; QMB 1.6146^ (?) same

collection data as QMB 1.6145; QMB 1.6147^ (?) same collection

data as QMB 1.6145; QMB 1.6972+ (?) Magnetic Island, 19° 8' S,

146° 50' E; QMB 1.9902+ (?) Lindeman Island, 20° 27' S, 149°

2' E, 1936; QMB 1.11537+ (?) Mackay, 21° 9' S, 149° 12' E; QMB

1.13459+ (?) Bushy Island, 80 km ENE of Mackay, 20° 58' S,

150° 5' E; QMB 1.17550+ (?) Torres Strait, Warrior Islet, 9° 46' S,

142° 57' E, 1 April 1974; QMB 1.17551+ (?)Torres Strait, Yam

Island, 9° 53' S, 142° 45' E, 6 April 1974; QMB 1.17669+ (?)

Torres St, Aeroplane Sandbank, 10° 23' S, 143° 19' E, 8 m, 20

July 1974; QMB 1.17550+ (?) Torres Strait, Warrior Islet, 9°

46' S, 142° 57' E, 1 April 1974; QMB 1.17551+ (?) Torres Strait,

Yam Island, 9° 53' S, 142° 45' E, 6 April 1974; QMB 1.17669+ (?)

Torres Strait, Aeroplane Sandbank, 10° 23' S, 143° 19' E, 8 m, 20

July 1974; QMB 1.21128+ (?) Weipa Channel, 12° 37' S, 141°

52' E, 17 May 1984; QMB 1.21129+ (?) Weipa Channel, 12°

37' S, 141° 52' E, 17 May 1984; QMB 1.21891+ (?)Sarina,

Campwin Beach Headland, 21° 23' S, 149° 19' E, 0.5-1 m, 1

December 1985; QMB 1.22718+ (?) Sarina Beach, Victor Island,

21° 19' S, 149° 19' E, 8 April 1987; QMB 1.22757+ (?)Sarina

Beach, Victor Island, 21° 19' S, 149° 19' E, 8 April 1987; QMB

l. 22795+ (?) Sarina Inlet, 21° 24' S, 149° 19' E, 0.5 m, 10 April

1987; QMB 1.26070+ (?) Proserpine, Bait Reef, 19° 48' S, 149°

4" E, 29 March 1975; QMB 1.26836+ (?)Torres Strait, Warrior

Reefs, 9° 48' S, 142° 58' E, 27 March 1990; QMB 1.28360+ (?)

Sabina Point, 22° 24' S, 150° 18' E, 0.5 m, 14 September 1993;

QMB 1.33697+ (?) Bountiful Island, 16° 39' S, 139° 53' E, 0-3.5

m, 22 November 2002; QMB 1.33732+ (?) Bountiful Island, 16°

38' S, 139° 53' E, 2^ m, 22 November 2002; QMB 1.33773+ (?)

Sweers Island, 17° 9' S, 139° 36' E, 2-3.5 m, 15 November 2002;

QMB 1.34432+ (?) Sweers Island, 17° 8' S, 139° 36' E, 0.5-2 m,

16 November 2002; QMB 1.34503+ (?) Sweers Island, 17° 8' S,

139° 37' E, 1.5-2.5 m, 17 November 2002; QMB 1.34546+ (?)

Sweers Island, 17° 5' S, 139° 39' E, 0.5-3 m, 18 November 2002;

QMB 1.34588+ (?) Sweers Island, 17° 8' S, 139° 37' E, 0.1-1.5 m,

21 November 2002; QMB 1.34627+ (?) Sweers Island, 17° T S,

139° 37' E, 0.3-0.5 m, 23 November 2002; QMB 1.34648+ (?)

Sweers Island, 17° 10' S, 139° 38' E, 0.1-2 m, 24 November

2002; QMB 1.34696+ (?) Sweers Island, 17° 8' S, 139° 37' E,

0.1-1 m, 19 November 2002; QMB 1.34714+ (?) Sweers Island,

17° 8' S, 139° 37' E, 0.1-1 m, 19 November 2002; Northern

Territory. AMS 1.23948-018 (3, 39^8) East Arm, 12° 29' S,

130° 55' E, 11 August 1983; AMS 1.24676* (13, 19.8-61.7)

Darwin, 12° 28' S, 130° 54' E, 1 m, 29 August 1984; AMS

1.24678-056 (11, 90-218) Darwin, 12° 25' S, 130° 49' E, 1 m, 31

August 1984; AMS 1.25637-001 (158) Darwin, 12° S, 130°

48' E, 1977; AMS IA.1463* (157) Gulf of Carpentaria, Pellew

Group, 15° 33' S, 136° 59' E, June 1923; AMS IA.1670+ (1) Gulf

of Carpentaria, Pellew Group, 15° S, 136° E, 1923; AMS IA.1671

(187) Gulf of Carpentaria, Pellew Group, 15° S, 136° E, 1923;

AMS IA.2541 (129) Pellew Group, Vanderlin Island, 15° 42' S,

136° 59' E, 1926; AMS IA.3854 (116) Darwin, 12° 27' S, 130°

48' E, 1929; AMS IA.7769 (164) Darwin, Point Charles, 12°

23' S, 130° 37' E, 1938; AMS IA.7828 (178) Melville Island, 11°

37' S, 131° 26' E, 1938; BMNH 1843.6.15.46* (207, holotype of

Labrus cyanodus, skin) Black Point, Port Essington; BMNH

1847.6.17.56* (254, skin) Port Essington, Earl of Derby; BMNH

1858.12.27.36* (190, skin) “Victoria" (Northern Territory?),

HMS Herald; CSIRO CA 704 (243) Groote Eylandt, Dalumbu

Bay, 14° 11' S, 136° 44' E, 19 March 1979; CSIRO CA 1227

(187) Alyangula, 13° 50' S, 136° 24' E, 15 March 1980; CSIRO

CA 1228 (186) Alyinga Island, 13° 4' S, 136° 3' E, 1 September

1980; NMV A7991 (228) The English Company Islands, 11°

51' S, 136° 23' E, 10-20 m, 1989; NTM S. 10454-010+ (6, 12^

192) Port Essington, Orontes Reef, 11° 04' 01" S, 132° 04' 59" E,

5 May 1982; NTM S. 10415-013+ (2,30-51) Lee Point Reef, 12°

19' 59" S, 130° 54' 00" E, 13 December 1981; NTM S. 16133-

002+ (180) Bynoe Harbour, Indian Island, 12° 39' 11" S, 130°

31' 34" E, 12 m, 7 April 2002; NTM S.10472-001+ (178) Darwin

Harbour, Channel Island, 12° 33' 00" S, 130° 52' 01" E, 15 July

1982; NTM S.10040-001+ (194) Darwin Harbour, South Shell

Island, 12° 30' 00" S, 130° 54' 00" E, 8 January 1982; NTM

S.10015-011+ (2, 134-144) Cobourg Peninsula, Coral Bay, 11°

10' 59" S, 132° 04' 01" E, 17 October 1981; NTM S.10431-008+

(5, 25-165) Cobourg Peninsula, Danger Point, 11° 07' 01" S,

132° 19' 59" E, 1 May 1982; NTM S. 10436-005+ (177) Cobourg

Peninsula, Danger Point, 11° 07' 01" S, 132° 19' 59" E, 29 April

1982; NTM S. 10603-034+ (3, 35-64) North Oxley Island, 11°

00' 00" S, 132° 49' 01" E, 19 October 1982; NTM S. 10004-005+

(5, lid—165) Cobourg Peninsula, Sandy Island, 11° 07' 01" S,

132° 16' 59" E, 21 October 1981; NTM S.10005-013+ (17, 36-

100) Cobourg Peninsula, Burford Island, 11° 28' 59" S, 131°

56' 60" E, 13 October 1981; NTM S.10006-010+ (84) Cobourg

Peninsula, Burford Island, 11° 28' 59" S, 131° 56' 60" E, 13

October 1981; NTM S.10397-002+ (2, 166-185) Darwin, 1977;

NTM S.10432-012+ (105) Port Essington, Table Head, 11°

13' 01" S, 132° 10' 01" E, 3 May 1982; NTM S.10033-013+ (3,

12-58) Darwin Harbour, Dudley Point, 12° 25' 01" S, 130°

49' 01" E, 13 November 1981; NTM S.11295-016+ (210) East

Bremer Islet, 11 March 1976; NTM S. 12449-005+ (167), Darwin

Harbour, South Shelll2° 30' 00" S, 130° 52' 59" E, 18 Eebruary

1982; NTM S. 16122-031+ (188) Port Patterson, Turtle Island,

12° 37' 48" S, 130° 27' 29" E, 8 April 2002; NTM S.11199-002+

(285) Cape Arnhem, Gove, “White Island”, 27 March 1975;

NTM S. 11304-001+ (207) Gove, 15 March 1976; NTM S. 13130-

001+ (235) Groote Eylandt, NW Bluff, 13° 49' 59" S, 136°

24' 00" E, 9 Eebruary 1989; NTM S. 11578-001+ (2, 105-215)

Melville Bay, South Granite Island, 12° 13' 59" S, 136° 40' 01" E,

25 May 1975; NTM S. 11306-002+ (214) Melville Bay, Gove, 29

October 1975; NTM S. 16115-013+ (177) Port Patterson, Moira

Reef, 12° 30' 54" S, 130° 30' 40" E, 13 m, 3 May 2002; NTM

S.16116-003+ (167) Port Patterson, Simms Reef, 12° 34' 12" S,

130° 26' 60" E, 20 m, 4 May 2002; NTM S. 12811-002+ (50)

Darwin Harbour, Channel Island, 12° 31' 01" S, 130° 50' 60" E,

14 November 1985; NTM S. 13237-013+ (5, 145-195) Rimbija

Island, Cape Wessel, 11° 01' 01" S, 136° 40' 59" E, 1 m, 16

November 1990; NTM S. 13228-005+ (2, 107-152) English

Company Islands, Wigram Island, 11° 43' 59" S, 136° 37' 59" E,

12 November 1990; NTM S. 13226-003+ (2, 122-158) English

Company Islands, Wigram Island, 11° 46' 59" S, 136° 35' 17" E,

12 November 1990; NTM S. 14469-022+ (5, 125-211) Eield

Island, 12° 03' 32" S, 132° 23' 17" E, 14 June 1997; NTM

S.14471-018+ (3, 81-113) Eield Island, 12° 05' 17" S, 132°

19' 08" E, 5 June 1997; NTM S. 14665-004+ (200) Eield Island,

12° 04' 12" S, 132° 19' 19" E, 5 June 1998; NTM S. 14949-004+

Review of Choerodon tuskfishes

(145) Adam Bay, Cape Hotham, 12° 05' 10" S, 131° 15' 11" E, 9

m, 12 June 1996; NTM S. 11263-018^ (5, 105-188) Cobourg

Peninsula, Coral Bay, 11° 12' 00" S, 132° 03' 00" E, 18 May

1983; NTM S. 16160-005+ (46, 32-170) Port Patterson, Quail

Island, 12° 31' 23" S, 130° 25' 37" E, 7 December 2002; NTM

S.15221-021+ (7, 36-50) Darwin Harbour, Nightcliff, 12°

22' 59" S, 130° 50' 17" E, 1 m, 12 October 2000; NTM S. 14965-

018+ (2, 105-117) Eield Island, 12° 05' 13" S, 132° 18' 36" E, 1

m, 10 October 1999; NTM S.14966-007+ (165) Eield Island, 12°

03' 32" S, 132° 23' 20" E, 1 m, 10 October 1999; NTM S. 16155-

005+ (56) Port Patterson, Middle Reef, 12° 28' 12" S, 130°

30' 18" E, 5 December 2002; NTM S. 15916-020+ (4, 11^3)

Kakadu National Park, Eield Island, 12° 03' 25" S, 132° 24' 32" E,

1 m, 19 August 2004; NTM S.16112-001+ (2, 209-253) Groote

Eylandt,Awangmarra, 13° 42' 00" S, 136° 39' 00" E, 16 August

1979; NTM S.16037-001+ (193) Darwin Harbour, Blackmore

Point, 12° 34' 16" S, 130° 50' 53" E, 4 m, 27 January 2005; NTM

S. 11260-034+ (3, 100-184) Cobourg Peninsula, Coral Bay, 11°

10' 59" S, 132° 03' 00" E, 16 May 1983; NTM S.10443-003+ (2,

195-237) Cobourg Peninsula, Danger Point, 11° 07' 01" S, 132°

19' 59" E, 30 April 1982; NTM S.11488-006+ (3, 6^80) Darwin

Harbour, Vesteys Beach, 12° 27' 00" S, 130° 49' 59" E, 15

November 1982; NTM S.10979-004+ (7, 53-66) Darwin

Harbour, Dudley Point, 12° 12' 14" S, 130° 49' 01" E, 15

November 1982; NTM S. 17312-001+ (200) Port Bremer, Sandy

Island, 11° 07' 01" S, 132° 17' 31" E, 20 October 1981; NTM

S. 16708-064+ (42,41-182) Darwin Harbour, Vesteys Beach, 12°

26' 10" S, 130° 49' 44" E, 0.3 m, 14 November 2008; NTM

S.17390-002+ (180) Darwin Harbour, Stevens Rock, 12°

29' 10" S, 130° 47' 06" E, 5 m, 15 August 2012; USNM 174367*

(96.6); USNM 174385 (22, 23.2-117) Groote Eylandt, Little

Lagoon, 25 April 1948; USNM 174395* (200) Yirrkala; USNM

174405* (174) ; USNM 174383 (1) Groote Eylandt, 0-1 m, 19-

25 April 1948; USNM 134388* (109) East Point reef, NNW of

Darwin, 0-0.5 m, 26 March 1948; USNM 174397* (2,156-194)

Amham Land, Yirrkala, NW of Cape Arnhem, 5-6 m, 6 August

1948; USNM 174401* (247) Yirrkala, NW of Cape Arnhem,

14-16 August 1948; USNM 174406* (237), Arnhem Land, Port

Bradshaw, W from Cape Arnhem, 0^ m, 22-26 July 1948;

USNM 174409* (232) Bickerton Island, South Bay, 0 m, 2 June

1948. Western Australia: AMS 1.12950 (300), Port Hedland, 20°

18' S, 118° 35' E, 1914; AMS 1.12951 (222) Port Hedland, 20°

18' S, 118° 35' E, 1914;AMS 1.17060-013 (2,215-255) Exmouth

Gulf, Bundegi Reef, 21° 52' S, 114° 09' E, 1 m, 19 January 1972;

AMS IB.1590 (159) Onslow, 21° 38' S, 115° 07' E, 1946; MNHN

A.8890* (207, type of Chaerops crassus) Dampier Archipelago,

Castelnau; NTM S. 12587-024+ (2, 46-74) Cape Leveque, 16°

25' 01" S, 122° 25' 01" E, 1 m, 18 March 1987; WAM P.25310-

002 (2, 52-68) Houtman Abrolhos, Beacon Island, 28° 30' S,

113° 44' E, 2-15 m, 18 May 1975;WAM P.25749-001 (218)

Rottnest Island, 31° 59' S, 115° 33' E, 8-10 m, 3 March 1977;

WAM P.25759-001 (117) Rottnest Island, 31° 59' S, 115° 29' E,

2-A m, 10 March 1977;WAM P.31017-018+ (4,31-72) Exmouth

Gulf, Tent Island, 0.1-0.3 m, 18 August 1995; WAM P.31516-

011+ (2, 28-67) Dampier Archipelago, Searipple Passage, 28

October 1998; WAM P.31092-030+ (2,4^110) Cape Talbot, 13°

45' S, 126° 45' E, 0.1-0.5 m, 27 November 1995; WAM P.31097-

046^ (3, 106-121) Cape Londonderry, 13° 45' S, 126° 58' E,

0.5-1.5 m, 29 November 1995; WAM P.30307-023+ (2,72-102)

Long Reef, 13° 48' 00" S, 125° 47' 00" E, 0.1-1 m, 17 August

1991; WAM P.33432-014+ (4, 58-143) Kimberley, Long Reef,

13° 48' 35" S, 125° 49' 26" E, 0-1 m, 23 October 2010; WAM

P.33429-010+ (2,103-118) Kimberley, Long Reef, 13° 49' 48" S,

125° 49' 57" E, 0-1, 21 October 2010; WAM P.31244-022+ (2,

112-114) Cassini Island, 13° 57' S, 125° 39' E, 0.5 m, 29

November 1996; WAM P.31085-022+ (4, 56-121) Long Island,

Vansittart Bay, 13° 59' S, 126° 20' E, 0.1-0.3 m, 24 November

1995; WAM P.31249-011+ (115) Colbert Island, 14° 53' S, 124°

43' E, 1-2 m, 01 December 1996; WAM P.31250-028+ (5, 33-

102) Wildcat Reefs, 15° 17' S, 124° 10' E, 0.5 m, 2 December

1996; WAM P.33615-009+ (6, 30-141) Kimberley, Beagle Reef,

15° 19' 36" S, 123° 32' 9" E, 0.6 m, 19 October 2011; WAM

P.33605-018+ (2, 28-29) Kimberley, Champagny Island, 15°

19' 57" S, 124° 14 09" E, 1.0 m, 15 October 2011; WAM

P.30316-003+ (6, 27-124) Beagle Reef, 15° 21' 00" S, 123°

32' 00" E, 0.1-1 m, 24 August 1991; WAM P.33274-033+ (5,

55-65) Kimberley, Adele Island, 15° 29.474 S, 123° 09.798' E,

0-1 m, 15 October 2009; WAM P.33273-011+ (9, 57-92)

Kimberley, Adele Island, 15° 31.700' S, 123° 11.611' E, 0-1 m,

14 October 2009; WAM P.33281-024+ (4, 63-97) Kimberley,

Montgomery Reef, 15° 51.323' S, 124° 18.875' E, 0-1 m, 20

October 2009; WAM P.33283-015+ (4, 56-82) Kimberley,

Montgomery Reef, 15° 52.727' S, 124° 19.602' E, 0-1 m, 20

October 2009; WAM P.33291-050+ (2, 79-103) Kimberley,

Montgomery Reef, 15° 53.700' S, 124° 20.340' E, 0-0.5 m, 24

October 2009; WAM P.33278-030+ (2, 83-83) Kimberley,

Montgomery Reef, 15° 53.815' S, 124° 19.531' E, 0-1 m, 19

October 2009; WAM P.33279-026+ (8, 36-109) Kimberley,

Montgomery Reef, 15° 53.815' S, 124° 19.531' E, 0-1, 19

October 2009; WAM P.33284-030+ (84) Kimberley, Montgomery

Reef, 15° 53.895' S, 124° 10.901' E, 0-1 m, 21 October 2009;

WAM P.31251-034+ (3, 68-94) Montgomery Reef, 15° 55' S,

124° 04 E, 0.7 m, 3 December 1996; WAM P.33286-013+ (4,

48-74 m) Kimberley, Montgomery Island, 15° 56.659' S, 124°

16.004 E, 0-1 m, 22 October 2009; WAM P.33285-015+ (4, 48-

72) Kimberley, Montgomery Reef, 15° 57.528' S, 124° 16.144' E,

0-0.5 m, 22 October 2009; WAM P.33287-025+ (3, 70-98)

Kimberley, Montgomery Reef, 15° 58.089' S, 124° 16.918' E,

0-1 m, 22 October 2009; WAM P.30851-008+ (13, 20-93)

Montgomery Reef, 15° 59' 00" S, 124° 17' 00" E, 0.1-0.5 m, 28

September 1994; WAM P.33288-001+ (3, 102-113) Kimberley,

Montgomery Reef, 16° 00.865' S, 124° 10.389' E, 0-1 m, 20

October 2009; WAM P.30916-020+ (77) Admiral Island, 16°

04 00" S, 123° 24 00" E, 0.1 m, 20 November 1994; WAM

P.30320-042+ (5, 102-118) Buccaneer Archipelago, Powerful

Island, 16° 05' 00" S, 123° 27' 00" E, 0.1-1 m, 26 August 1991;

WAM P.31204-003+ (11, 47-112) Kingfisher Island, 16° 07' S,

124° 05' E, 0.1-0.5 m, 27 September 1996; WAM P.30929-019+

(103) Whirlpool Pass, 16° 16' 00" S, 123° 30' 00" E, 0.1-0.8 m,

25 November 1994; WAM P.30911-012+ (2, 63-66) Gregory

Island, 16° 19' 00" S, 123° 19' 00" E, 0.1-0.5 m, 19 November

1994; WAM P.31205-033+ (44) Swan Bay, 16° 22' S, 123° 02' E,

O. 1-0.5 m, 27 September 1996; WAM P.30902-010+ (5, 59-160)

Tallon Island, 16° 24' 00" S, 123° 07' 00" E, 0.5 m, 17 November

1994; WAM P.30898-021+ (4, 63-116) Sunday Island, 16°

25' 00" S, 123° 11' 00" E, 0.1-1 m, 15 November 1994; WAM

P. 30321-005+ (3 , 39-86, Sunday Island, 16° 26' 00" S, 123°

11' 00" E, 0.1-L5 m, 27 August 1991; WAM P.30908-011+ (43)

Mermaid Island, 16° 26' 00" S, 123° 21' 00" E, 0.1-1 m, 18

November 1994; WAM P.31391-011+ (211) Beagle Bay, 16°

59' S, 122° 40' E, 9-10 m, 28 August 1997; WAM P.28417-003+

(6, 23-122) Coulomb Point, 17° 21' S, 122° 09' E, 2 m, 1

100

M.F. Gomon

September 1981; WAM P.32171-017^ (2, 107-172) James Price

Point, 17° 26' S, 122° 10' E, 2-7 m, 31 July 1982; WAMP32169-

016^ (4, 28-16) Quondong Point, 17° 34' S, 122° 09' E, 1-3 m,

25 August 1982; WAM P30263-025+ (215) Broome, 17°

58' 00" S, 122° 14' 00" E, 28 July 1982; WAM P27274-02+ (3,

61-96) Broome, 17° 58' S, 122° 14' E, 1-2 m, 18 January 1981;

WAM P27457-00E (224) Port Hedland, 20° 18' S, 118° 35' E,

1981; WAM P.31514-034^ (1) Dampier Archipelago, Keast

Island, 20° 24' S, 116° 50' E, 0.1-0.5 m, 26 October 1998; WAM

P.25354-002+ (2, 254-280) Monte Bello Islands, 20° 25' S, 115°

30' E, April 1975; WAM P.31512-017^ (9, 39-79) Dampier

Archipelago, Collier Rocks, 20° 25' S, 116° 51' E, 0.1-0.4 m, 24

October 1998; WAM P.30298-014^ (147) Monte Bello Islands,

20° 26' 00" S, 115° 37' 00" E, 12 December 1979; WAM P.31510-

008^ (2, 21-145) Dampier Archipelago, 20° 26' S, 116° 49' E,

O. 1-0.5 m, 22 October 1998; WAM P.30672-017+ (2, 36-45)

Monte Bello Islands, 20° 28' 33" S, 115° 32' 13" E, 0.1-1.5 m, 25

August 1993; WAM P.25112-00E (3, 155-225) Dampier

Archipelago, Kendrew Island, 20° 28' S, 116° 32' E, 0.1-0.5 m, 4

November 1974; WAM P.22230-00E (1) Dampier Archipelago,

Kendrew Island, 20° 29' S, 116° 32' E, 26 October 1972; WAM

P. 22416-00E (1) Dampier Archipelago, Kendrew Island, 20°

29' S, 116° 32' E, 23 October 1972; WAM P.22848-001' (1)

Dampier Archipelago, Kendrew Island, Museum Bay, 20° 29' S,

116° 32' E, 20 Eebruary 1973; WAM P.25107-015+ (185)

Dampier Archipelago, Kendrew Island, 20° 29' S, 116° 32' E,

3-5 m, 21 October 1974; WAM P.25114-012^ (55) Dampier

Archipelago, Rosemary Island, 20° 29' S, 116° 35' E, 1-5 m, 8

November 1974; WAM P.31655-014^ (1) Dampier Archipelago,

Malus Island, 20° 30.90' S, 116° 40.22' E, 0.1-0.2 m, 27 August

1999; WAM P.4524-00E (1) Dampier Archipelago, 20° 33' S,

116° 32' E, December 1957; WAM P.31659-022+ (2, 40-43)

Dampier Archipelago, Enderby Island, 20° 35.20' S, 116°

30.91' E, 1 September 1999; WAM P.31661-016' (2, 28-33)

Dampier Archipelago, Enderby Island, 20° 36.22' S, 116°

33.06' E, 0.1 m, 2 September 1999; WAM P.2850-00E (175)

Onslow, 21° 38' S, 115° 07' E, January 1945; WAM P.31013-02r

(9, 30-62) Exmouth Gulf, Burnside Island, 22° 06' 00" S, 114°

31' 00" E, 0.1-0.5 m, 16 August 1995; WAM P.31015-032' (36)

Exmouth Gulf, Simpson Island, 22° 07' 00" S, 114° 29' 00" E,

O. 1-0.3 m, 17 August 1995; WAM P.12850-00E (1) Shark Bay,

25° 21' S, 113° 44' E, September 1960; WAM P.12851-00r (1)

Shark Bay, 25° 21' S, 113° 44' E, September 1960; WAM

P. 33864-00E (2, 206-265) Shark Bay, Peron Peninsula, 25°

32' S, 113° 30' E, 25 September 2002. Papua New Guinea:

WAM P.28154-024+ (24, 50-108) Daru Island, Daru, 9° 05' S,

143° 15' E, 0.1-0.4 m, 19 September 1983.

Choerodon fasciatus. (49 specimens examined, 31-188 mm

SL). Japan, Okinawa: URM-P 22567* (125) May-September

1989; URM-P 40874* (146) Awase fish market, 7 September

2000; URM-P 44551* (170) Henza fish market, 31 May 2008;

USNM 62234 (1) Naha. Philippines: USNM 202508 (1) Luzon.

Australia, Queensland: AMS 1.12690 (176) Dunk Island, 17°

57' S, 146° 09' E, 1912; AMS 1.15459-001 (188) Heron Island,

23° 27' S, 151° 57' E, 7 October 1964; AMS 1.15571-010 (130)

Capricorn Group, North West Island, 23° 18' S, 151° 42' E, 1969;

AMS 1.15647-075 (2, 133-182) One Tree Island, 23° 30' S, 152°

05' E, 13-14 m, 9 October 1968; AMS 1.15681-057 (138) One

Tree Island, 23° 30' S, 152° 05' E, 25 November 1969; AMS

1.17445-068^ (4, 52-155) One Tree Island, 23° 30' S, 152° 05' E,

3-f m, 19 September 1968; AMS 1.18739-026^ (148) Lizard

Island, Palfrey Island, 14° 42' S, 145° 27' E, 3-10 m, 21

November 1975; AMS 1.18755-033* (98.4) Lizard Island,

Palfrey Island, 14° 41' S, 145° 27' E, 4-8 m, 2 November 1975;

AMS 1.19108-024+ (19) Lizard Island, 14° 40' S, 145° 28' E,

1-10 m, 17 November 1975; AMS 1.19460-033+ (99) Decapolis

Reef, 14° 51' S, 145° 17' E, 10 m, 14 November 1975; AMS

1.19607-085+ (2, 30-102) Lizard Island, North Point Reef, 14°

40' S, 145° 27' E, 5-6 m, 30 January 1975; AMS 1.20762-069+ (2,

46-70) Lizard Island, 14° 41' S, 145° 27' E, 1-5 m, 1 Eebruary

1975; AMS 1.20765-009 (31) Lizard Island, Palfrey Island,

(bearing W .25 mile), 14° 42' S, 145° 27' E, 5 m, 1 January 1975;

AMS 1.20769-029+ (105) Cape York, Halfway Island, 11° 23' S,

142° 57' E, 4-9 m, 18 Eebruary 1979; AMS 1.20769-061+ (111)

Cape York, Halfway Island, 11° 23' S, 142° 57' E, 4-9 m, 18

Eebruary 1979; AMS 1.20793-015 (44.0) Cape York, Clack

Island, 14° 03' S, 144° 16' E, 3-7 m, 24 Eebruary 1979; AMS

1.20982-036* (95.8) Lizard Island, 14° 41' S, 145° 27' E, 5-6 m,

28 November 1978; AMS 1.21422-040* (130) Lizard Island, 14°

41' S, 145° 26' E, 1-8 m, 27 January 1975; AMS 1.21495-001

(103) Lady Musgrave Reef, 23° 54' S, 152° 25' E, 5 m, 20

Eebruary 1980; AMS 1.22578-006* (2, 106-157) Escape Reef,

15° 49' S, 145° 50' E, 6-10 m, 28 October 1981; AMS 1.22582-

005* (122) Escape Reef North, 15° 49' S, 145° 50' E, 1^17 m,

29 October 1981; AMS 1.23702-083 (107) Lizard Island, Palfrey

Island, 14° 42' S, 145° 27' E, 2-6 m, 29 December 1974; AMS

1.31433-002+ (1) Orpheus Island, 18° 37' S, 146° 30' E, 4 m, 11

November 1982; AMS 1.31433-001+ (164) Orpheus Island, 18°

37' S, 146° 30' E, 4 m, 11 November 1982; AMS 1.44750-008*

(112) Lizard Island, Mermaid Cove, 14° 38' 45" S, 145° 27' 17" E,

0-10 m, 17 September 2008; AMS IA.1314 (179) Bowen, 20°

or S, 148° 15' E, 1923; AMS IA.207+ (1) Palm Islands, 18° 47' S,

146° 34' E, 1921; AMS IA.2097 (1) Great Barrier Reef, Beaver

Reef, 17° 50' S, 146° 30' E, 1924; AMS IA.2098 (1) 1924; AMS

IA.2241 (169) Great Barrier Reef, North Barnard Island, 17°

40' S, 146° 10' E, 1924; AMS IA.2719+ (1) 160, Hervey Bay, 25°

S, 152° E, 1926; AMS IA.4615+ (1) Capricorn Group, North

West Island, 10° 40' S, 142° 07' E, May 1930; AMS IA.5003+ (1)

Capricorn Group, North West Island, 23° 17' S, 151° 42' 28" E,

May 1931; AMS IB.7415+ (155) Townsville, 19° 16' S, 146°

49' E, 1965; BMNH 1867.6.24.3* (161, lectotype of Xiphocheilus

fasciatus, mount) Cape York; BMNH 1867.6.24.4* (143,

paralectotype of Xiphocheilus fasciatus, mount) Cape York;

BMNH 911.4.1.28-29 (2, 150-155) northern Queensland,

Saville-Kent; BMNH 1933.1.25.92 (184) Capricorn Group;

BPBM 13666 (65) Eitzroy Island; BPBM 41249 (140) Lizard

Island, 25 November 1982; MCZ 36846 (2,105-150) Gladstone;

NMV A22006 (75) Lizard Island, 14° 40' 59.87" S, 145°

27' 45.00" E, November 1979; QMB 1.4234 (1) off Cape

Moreton, 27° 2' S, 153° 30' E; QMB 1.4301+ (?) off Cape

Moreton, 27° 2' S, 153° 30' E; QMB 1.4479+ (?) Elinders Reef, off

Cape Moreton, 26° 58' S, 153° 29' E; QMB 1.5220 (1) off Cairns;

QMB 1.5388 (1) Moreton Bay, 27° 15' S, 153° 15' E; QMB

1.5479+ (?) Heron Island, 23° 27' S, 151° 55' E; QMB 1.6750+ (?)

Magnetic Island, 19° 8'S, 146° 50'E; QMB 1.7602(1) Townsville,

Lodestone Reef, 18° 41' S, 147° 5' E; QMB 1.7619+ (?) Dent

Island, 20° 21' S, 148° 56' E; QMB 1.7628+ (?) Mooloolaba, 26°

40' S, 153° 15' E; QMB 1.7998 (1) Proserpine, 20° 25' S, 148°

55' E; QMB 1.8028 (1) Cape Moreton, Smith Rock, 27° S, 153°

29' E; QMB 1.8029+ (?) Cape Moreton, Smith Rock, 27° S, 153°

29' E; QMB 1.8192 (1) Alexandra Headland, 26° 40' S, 153° 7' E;

Review of Choerodon tuskfishes

101

QMB 1.8821 (1) Cape York, 10° 41' S, 142° 32' E; QMB 1.1020r

(?) Cairns, July 1972; QMB 1.10336^ (?)Cape Bowling Green,

Darley Reef, 19° 12' S, 148° 12' E, 23 March 1973; QMB

1.11360^ (?) Kelso Reef, 18° 24' S, 147° E; QMB 1.14866^ (?)

Lizard Island, Yonge Reef, 14° 35' S, 145° 37' E, 6 January 1975;

QMB 1.15435+ (?) Clack Reef, 14° 3' S, 144° 15' E, 1.5-7.6 m, 24

Eebruary 1979; QMB 1.19109+ (?)Caims, Arlington Reef, 16°

42' S, 146° 4' E, 1980; QMB 1.20105+ (?) Erankland Islands, 17°

11' S, 146° 4' E, 6-10 m, November 1982; QMB 1.31635+ (?)

Pompey Group, Renes Nook, 21° 17' S, 151° 31' E, 2.5-6 m, 13

March 2000; QMB 1.31697+ (?) Pompey Group, 21° 42' S, 151°

44' E, 3-5 m, 11 March 2000; QMB 1.31719+ (?) Pompey Group,

21° 2' S, 151° 28' E, 12.5-15 m, 15 March 2000; QMB 1.31953+

(?) Pompey Group, 21° 42' S, 151° 44' E, 3-6 m, 11 March 2000;

QMB 1.33355+ (?) Swain Reefs, 21° 35' S, 152° 22' E, 3-5 m, 11

Eebruary 2001; QMB 1.33430+ (?) Swain Reefs, Gannet Cay, 21°

59' S, 152° 21' E, 2.5-3.5 m, 5 Eebruary 2001; QMB 1.33485+ (?)

Swain Reefs, Gannet Cay, 21° 53' S, 152° 21' E, 4.5-7.5 m, 6

Eebruary 2001; QMB 1.33638+ (?) Erigate Reef, Swain Reefs,

21° 42' S, 152° 21' E, 4.5-6 m, 7 Eebruary 2001; QMB 1.37049+

(?) Star Reef, Swain Reefs, 21° 30' S, 152° 25' E, 4-6 m, 10

Eebruary 2001; QMB 1.37611+ (?) Shag Rock, North Stradbroke

Island, 27° 24' S, 153° 31' E, 7-10 m, 14 December 2005; USNM

176924 (1) Great Barrier Reef; USNM 368244* (1) Eairfax

Island Lagoon, 1966-1968; USNM 368246 (122) One Tree

Island; WAM P.33532-005+ (85) Great Barrier Reef, Lizard

Island, 14° 38.45' S, 145° 27.17' E, 0-10 m, 17 September 2008.

New Caledonia: AMS 1B.4467 (154) Noumea, 1959; BPBM

27100 (35) 9 January 1979; MNHN 1980-0782 (162) 10

January 1979.

Choerodon frenatus. (47 specimens examined, 52-136 mm SL).

Australia, New South Wales: AMS 1.26536-013 (52) Namba,

29° 28' S, 153° 30' E-29° 33' S, 153° 25' E, 51 m; AMS 1.31472-

002 (2, 75-92) Yamba, 29° 24' S, 153° 33' E-29° 28' S, 153°

35' E, 66-73 m, 31 August 1990; AMS 1.31472-005 (105) Yamba,

29° 24' S, 153° 33' E-29° 28' S, 153° 35' E, 66-73 m, 31 August

1990; AMS 1.31473-001* (106) lluka, 29° 20' S, 153° 34' E-29°

28' 28" S, 153° 36' E, 67-77 m, 7 May 1990; AMS 1.31473-003*

(6,88.5-117) lluka, 29° 20' S, 153° 34' E-29° 28' 2 S, 153° 36' E,

67-77, 7 May 1990; AMS 1.32120-002 (4, 99-105) Clarence

River, 29° 206' S, 152° 34' E-29° 25' 28" S, 153° 37' E, 67-73 m,

2 May 1990; AMS 1.33510-005* (115) Clarence River, 29° 26' S,

153° 34' E-29° 25' 28" S, 153° 35' E, 6^68 m, 2 April 1990;

AMS 1.44773.030+ (1) Ballina, 28° 33' 12" S, 153° 44' 30" E-28°

35' 12" S, 153° 44' 24" E, 69-71 m; AMS 1.44773-013* (6,86.5-

127) Ballina, 28° 33' 12" S, 153° 44' 30" E-28° 35' 12" S, 153°

44' 24" E, 69-71 m; AMS 1.44773-014+ (143) Ballina, 28°

33' 12" S, 153° 44' 30" E, 28° 35' 12" S, 153° 44' 24" E, 69-71 m;

AMS 1.44773-015+ (143) Ballina, 28° 33' 12" S, 153°

44' 30" E-28° 35' 12" S, 153° 44' 24" E, 69-71 m; CSIRO H

4773-07* (116) Yamba, 29° 24' S, 153° 35' E, 29° 23' S, 153°

35' E, 68-71 m, 17 April 1996; CSIRO H 4773-08* (116) Yamba,

29° 24' S, 153° 35' E-9° 23' S, 153° 35' E, 68-71 m, 17 April

1996; CSIRO H 4773-09* (79.8) Yamba, 29° 24' S, 153°

35' E-29° 23' S, 153° 35' E, 68-71 m, 17 April 1996; CSIRO H

4773-10* (5, 63.8-95.2) Yamba, 29° 24' S, 153° 35' E-29° 23' S,

153° 35' E, 68-71 m, 17 April 1996; NMV A2781 (2,90-110) off

Southport, 27° 42' 00.00" S, 153° 34' 48.00" E, 57 m, 6 November

1981; NMV A2792* (2,110-115) off Southport, 27° 53'59.99" S,

153° 40' 11.99" E, 83 m, 6 November 1981; Queensland: AMS

1.12536* (136, lectotype of Choerodon frenatus) Double Island

Point, 25° 56' S, 153° 11' E; CSIRO H 6909-01 (94.3) Bowling

Green Bay, 18° 54.20' S, 147° 49.93' E-18° 54.51' S, 147°

50.40' E, 64 m, 1 December 2003; CSIRO H 7898-01 (68) N of

Abbot Bay, 19° 05.89' S, 148° 09.05' E, 13 December 2003;

NMV A31424-001 (4,43.2-84.2) N of Townsville, 18° 27.94' S,

147° 03.66' E, 64 m, 29 November 2003; NMV A 31425-001

(89.3) and NMV A 31425-002 (2, 69.5-74.9) NE of Great Palm

Island, 18° 29.15' S, 147° 00.10' E, 59 m, 29 November 2003;

NMV A31426-001 (104) NE of Whitsunday Group, 19° 28.36' S,

149° 29.79' E, 72 m, 7 December 2005; QMB 1.475* (105,

paralectotype of Choerodon frenatus) Double Island Point, 25°

56' S, 153° 15' E; QMB 1.19601+ (?) Noggin Reef, 17° 5' S, 146°

23' E, 51.2 m, 10 October 1979; QMB 1.19624+ (?) Geranium

Passage, 17° 37' S, 146° 34' E, 64 m, 13 October 1979; QMB

1.20326+ (?) Elora Passage, 17° 3' S, 146° 14' E, 37^2 m, 1981;

QMB 1.23063+ (?) Swain Reefs, 21° 17' S, 150° 47' E, 39 m, 16

September 1986; QMB 1.23075+ (?) Swain Reefs, 21° 7' S, 151°

27' E, 54 m, 13 September 1986; QMB 1.23497+ (?) Britomart

Reef, 18° 19' S, 146° 38' E, 41 m, 20 January 1985; QMB

1.26614+ (?) Eraser Island, 25° S, 153° 30' E, 30 m, 16 May 1990;

QMB 1.29540+ (?) Eeather Reef, 17° 33' S, 146° 26' E, 47.5 m, 11

October 1979; QMB 1.29707+ (?) Stradbroke Island, Point

Lookout, 27° 31' S, 153° 40' E, 80 m, 21 March 1995; QMB

l. 30199+ (?) Stradbroke Island, Point Lookout, 27° 29' S, 153°

36' E, 73 m, 7 October 1995; QMB 1.32922+ (?) Llewellyn Reef,

23° 45' S, 152° 9' E, 47 m, 3 October 2000; QMB 1.32935+ (?)

North West Island, 23° 3' S, 151° 38' E, 36 m, 10 October 2000;

QMB 1.33021+ (?) Caloundra, 26° 48' S, 153° 29' E, 78 m, 21

Eebruary 2001; QMB 1.35047+ (?) 17° 54.3' S, 146° 48.9' E, 17°

54' S, 146° 49' E, 25 September 2003; QMB 1.35367+ (?) 18°

51.9' S, 147° 35.1' E-18° 52' S, 147° 35' E, 22 September 2003;

QMB 1.35566+ (?) 18° 46.5' S, 147° 34.5' E-18° 46' S, 147°

34' E, 22 September 2003; QMB 1.35598+ (?) 18° 50.1' S, 147°

41.1' E-18° 50' S, 147° 41' E, 22 September 2003; QMB 1.35605+

(?) 18° 49.5' S, 147° 45.3' E-18° 49' S, 147° 45' E, 22 September

2003; QMB 1.35878+ (?) 19° 8.7' S, 147° 20.7' E, 27 m, 23

November 2003; QMB 1.35934+ (?) 19° 27.3' S, 148° 17.1' E, 50

m, 27 November 2003; QMB 1.36244+ (?) 21° 31.5' S, 150°

3.3' E, 28 m, 8 May 2004; QMB 1.36255+ (?) 18° 20.7' S, 146°

56.7' E, 65 m, 29 April 2004; QMB 1.36282+ (?) 18° 45.3' S, 147°

7.9' E, 52 m, 30 April 2004; QMB 1.36378+ (?) 23° 29' S, 151°

56.1' E, 40 m, 22 December 2004; QMB 1.36389+ (?) 23° 58.5' S,

152° 38.1' E, 45 m, 23 May 2004; QMB 1.36687+ (?) 15° 57.9' S,

147° 50.7' E, 58 m, 8 September 2004; QMB 1.37338 (111) 19°

20.2' S, 148° 17.8' E, 52 m, 19 March 2005.

Choerodon gomoni. (30 specimens examined, 71.5-106 mm

SL). Coral Sea: AMS 1.41000-001 (105, paratype of Choerodon

gomoni) Chesterfield Bank, 21° 05' S, 158° 57' 36" E, 73 m, 24

August 1988; BPBM 33840+ (88.5, paratype of Choerodon

gomoni) Chesterfield Bank, approximately 20° 45' S, 158°

50' 06" E, 75 m, 22 August 1988; BPBM 33850+ (101, holotype

of Choerodon gomoni) Chesterfield Bank, approximately 20°

45' S, 158° 35' 04" E, 82 m, 22 August 1988; MNHN 2001-2358+

(92.5) same collection data as AMS 1.41000-001; USNM

366799+ (97.0) same collection data as BPBM33840; MNHN

2001-2358* (2, 97.3-99.5) Chesterfield Bank, 21° 05' 0" S, 158°

57' 6" E, 72 m, 24 August 1988; MNHN 2016-0093* (93.3)

Chesterfield Bank, 21° 05' 0" S, 158° 57' 6" E, 22 August 1988.

New Caledonia: MNHN 2016-0094* (2, 71.5-88.1) lies Belep,

102

M.F. Gomon

19° 38' 7" S, 163° 50' E, 35 m, 7 July 1986. Australia,

Queensland: CSIRO H 3436-04* (92.0) Cape York, Blackwood

Channel, 11° 43.8' S, 143° 43.8' E, 24m, 26 March 1993; CSIRO

H 3441-01* (106) Cape York, Blackwood Channel, 11° 40.2' S,

143° 43.2' E, 28 m, 26 March 1993; CSIRO H 6926-07* (2,

81.1-88.8) Torres Strait, Cape York, 10° 21.84' S, 143°

40.26' E-10° 21.57' S, 143° 40.78' E, 40 m, 31 January 2004;

CSIRO H 6926-09 (86.3) Torres Strait, Cape York, 10° 21.84' S,

143° 40.26' E-10° 21.57' S, 143° 40.78' E, 40 m, 31 January

2004; CSIRO H 7027-02* (2, 83.1-80.4) NE of Townsville, 18°

26.95' S, 147° 20.01' E, 60 m, 15 December 2003; CSIRO H

7460-03* (84.7) NE of Townsville, 18° 31.23' S, 147° 35.45' E,

76 m, 14 December 2003; CSIRO H 7894-01* (96.1) Torres

Strait, Seven Reefs, 10° 19.14' S, 143° 44.99' E, 54 m, 31 January

2004; CSIRO H 7896-01* (84.6) 19.1091° S, 148.6238° E, 53.6

m, 13 December 2003; CSIRO H 6442-06* (85.2) 19.6653° S,

150.0759° E, 72.5 m, 5 December 2003; NMV A 31427-001

(104) Capricorn Group, North Reef, 23° 10.58' S, 151° 48.51' E,

49 m, 22 April 2004; NMV A 31428-001 (6, 76.3-81.0) NE of

Townsville, 18° 26.95' S, 147° 20.01' E, 60 m, 15 December

2003; NMV A 31428-002 (5, 77.8-83.0) NE of Townsville, 18°

26.95' S, 147° 20.01' E, 60 m, 15 December 2003; QMB 1.23062+

(?) Swain Reefs 21° 23' S, 152° 12' E, 46 m, 19 September 1986;

QMB 1.34054+ (?) Bunker Group, f 23° 47' S, 151° 58' E, 40 m,

10 October 2002; QMB 1.36605+ (?) 18.49.5' S, 148.12.3' E 18°

49' S, 148° 12' E, 72 m, 9 September 2004.

Choerodon graphicus. (25 specimens examined, 13.4—368 mm

SL). Australia, Queensland: AMS 1.15458-001* (134) Heron

Island, 23° 27' S, 151° 57' E, 1965; AMS 1.15620-026 (2, 240-

290) One Tree Island, 23° 30' S, 152° 05' E, 26 November 1966;

AMS 1.15683-045 (250) One Tree Island, 23° 30' S, 152° 05' E,

3-5 m, 30 November 1969; AMS 1.17445-091* (2, 94.1-98.5)

One Tree Island, 23° 30' S, 152° 05' E, 3^ m, 19 September

1968; AMS 1.20773-094 (18.2) Nymph Island, 14° 39' S, 145°

15' E, 1-2 m, 6 December 1978; AMS 1.20787-067 (2, 13.4-

17.3) Linnet Reef, 14° 47' S, 145° 21' E, 2-3 m, 9 December

1978;AMS 1.20793-015 (47) Cape York, Clack Island, 14° 03' S,

144° 16' E, 3-7 m, 24 Eebruary 1979; AMS 1.21260-015 (2,

28.3-30.8) Cape Tribulation, Noah Beach, 16° 06' S, 145° 28' E,

1 m, 12 September 1979; AMS 1.34316-014+ (6, 20-230)

Townshend Island, 22° 12' 14" S, 150° 29' 25" E, 10 m, 16

September 1993; AMS 1.34323-027* (157) Ereshwater Beach,

22° 33' 42" S, 150° 47' 41" E, 6 m, 18 September 1993; AMS

1.34324-009 (2, 39.2^.5) Dome Island, 22° 24' 47" S, 150°

44' 39" E, 3-6 m, 19 September 1993; AMS 1.34344-023 (20)

Ereshwater Beach, 22° 38' 50" S, 150° 47' 55" E, 2-A m, 26

September 1993; AMS 1.34370-014+ (4, 58-77) Townshend

Island, 22° 13' 52" S, 150° 34' 26" E-22° 13' 37" S, 150°

33' 49" E, 36-37 m, 24 October 1993; AMS 1.34386-001+ (114)

Island Head Creek, 22° 21' 59" S, 150° 38' 41" E, 7-27 m, 25

October 1993; AMS IB.3527* (296, holotype of Choerodon

transversalis) 23° 26' S, 151° 55' E, 1956; AMS IB.3908 (29)

Heron Island, 23° 26' S, 151° 55' E, December 1957; BPBM

14373 (44) One Tree Island; CSIRO B 1385 (65.5) Heron Island,

23° 27' S, 151° 55' E, 2 June 1977; QMB 1.944* (288, holotype

of Choerops graphicus) Cardwell, 18° 16' S, 146° 1' E; QMB

1.8848+ (?)Maroochydore, 26° 39' S, 153° 7' E; QMB 1.9743 (1)

Bundaberg, Bargara Reef, 24° 52' S, 152° 21' E, 2.9 m, 6 June

1969; QMB 1.10269+ (?) Bundaberg, December 1972; QMB

1.11323+ (?) Bundaberg, 6 October 1973; QMB 1.11376+ (?)

Heron Island, 23° 27' S, 151° 55' E; QMB 1.13780+ (?) Noosa

River, 26° 24' S, 153° 4' E, January 1977; QMB 1.27489+ (?)

Mooloolaba, 26° 40' S, 153° 15' E, 1 September 1991; QMB

1.30310+ (?) Southport, 27° 56' S, 153° 26' E, 6 m, 26 March

1996. New Caledonia: AMS IB .4436* (157) Noumea, 1959;

BPBM 11413 (2, 187-287) 12 August 1971; USNM 438440*

(212) Noumea, He Te Nd, 9 January 1961; USNM 438441 *

(233) 1963; USNM 206521* (368).

Choerodon gymnogenys. (20 specimens examined, 99.5-140

mm SL). Arabian Gulf: MCZ 14344* (2, 99.5-139) 1861.

Seychelles: NSMT-P116909* (127) Seychelles Bank, 4° 52.8' S,

55° 54' E, 58 m, 23 November 1968; NSMT-P 116910* (122)

Seychelles Bank, 4° 1' S, 55° 55' E, 59 m, 30 November 1968;

SAIAB 98483 (1) 4.74083° S, 56.4380° E, 9 November 2008;

SAIAB 98487 (1) Seychelles Bank, 60 m; SAIAB 98483 (1)

Seychelles Bank, 59-61 m; SAIAB 98491 (1) Seychelles Bank,

4.61683° S, 54.3643° E, 57-59 m, 12 November 2008. St

Brandon Shoal: SAIAB 83986 (125) 16.8393° S, 59.5893° E,

58-60 m, 13 October 2008; SAIAB 84010 (110) 16.8393° S,

59.5893° E, 58-60 m, 13 October 2008; SAIAB 190077 (1) 52-

53 m. Zanzibar: MCZ 4466 (110) 1862; MCZ 14343 (128)

1862; MCZ 24333 (4,117-119) 1861;RUSI2984* (3,131-135);

RUSI 2985* (119); BMNH 2016.6.6.1 (118) Playfair; BMNH

1864.11.15.28* (132, skin) Playfair; BMNH 1865.2.27.8 (118)

Playfair; BMNH 1866.1.19.17* (140, lectotype of Xiphocheilus

gymnonenys) Playfair; BMNH 1866.1.19.18* (132, paralectotype

of Xiphocheilus gymnonenys) Playfair; BMNH 1868.5.30.138

(130) Playfair. Mozambique: SAIAB 81739 (2, 145-150) Xai-

Xai, 26° 9.4' S, 32° 58.6' E, 43-45 m, 30 September 2007; SAIAB

81802 (131) off Ponta do Ouro, 26.835° S, 32.935° E, 56 m, 2

Eebruary 2007.

Choerodon jordani. (49 specimens examined, 23-136 mm SL).

Japan, Okinawa: OCE-P 20141113-1* (116) near Motobu, 6

October 2014; OCE-P 20141113-2* (136) near Motobu, 6

October 2014; URM-P 32669* (139) Naha fish market, 17

October 1994; USNM 62235* (120, holotype of Choerodon

jordani) Naha, Albatross; USNM 74549* (120, paratype of

Choerodon jordani) Naha, Albatross. Australia, Queensland:

AMS 1.15625-030* (2, 114-129) Lizard Island, 23° 30' S, 152°

05' E, 32 m, 8 December 1966; AMS 1.15637-035* (118) One

Tree Island, 23° 30' S, 152° 05' E, 23-26 m, 5 October 1967;

AMS 1.17929-005* (79.2) One Tree Island, 23° 30' S, 152° 05' E,

34 m, 3 December 1973; AMS 1.19476-012* (83.3) Yonge Reef,

14° 35' S, 145° 36' E, 20-35 m, 25 November 1975; AMS

1.20716-002 (64) One Tree Island, off Wreck Bank 23° 30' S,

152° 05' E, 13 m, 13 September 1973; AMS 1.22587-009 (24.7)

Escape Reef North, 15° 49' S, 145° 50' E, 40 m, 31 October 1981;

AMS 1.22612-011* (93.8) Escape Reef North, 15° 49' S, 145°

50' E, 20 m, 1 November 1981; BPBM 14379* (3,102-119) One

Tree Island; CSIRO H 6748-04 (101) SE of Bunker Group, 23°

58.71' S, 152° 37.79' E, 54 m, 15 April 2004; CSIRO H 6913-03

(78.1) 23.3715° S, 151.9723° E, 44 m, 22 April 2004. Western

Australia: WAM P.26068-005 (46) Houtman Abrolhos, Beacon

Island, 28° 29' S, 113° 47' E, 40 m, 9 April 1978; WAM P.27591-

002 (37) Houtman Abrolhos, Long Island, 28° 29' S, 113° 46' E,

30-33 m, 17 April 1982; WAM P.27595-011 (18, 23-82)

Houtman Abrolhos, Long Island, 28° 29' S, 113° 46' E, 30-33 m,

19 April 1982; WAM P.25311-003 (4, 40.1-59.0) Houtman

Abrolhos, Beacon Island, 28° 30' S, 113° 44' E, 0-30 m, 18 May

1975; WAM P.25313-004 (102) Houtman Abrolhos, Beacon

Review of Choerodon tuskfishes

103

Island, 28° 30' S, 113° 44' E, 1-5 m, 19 May 1975; WAM

P.27590-014 (3, 23^5) Houtman Abrolhos, Dicks Island, 28°

30' S, 113° 46' E, 30-32 m, 17 April 1982. New Caledonia:

AMS 1.17466-015* (72.2) Puetege Reef, 22° 22' S, 167° 08' E,

6-25 m, 15 June 1973; Vanuatu: AMS 1.17474-013* (82.2)

Efate Island, Undine Bay, 17° 31' S, 168° 21' E, 25 m, 24 June

1973. Fiji: BPBM 31163^ BPBM 31167^; USNM 2811W (2)

Rotuma, Northern Reef, Due North of Motusa, 12.5° S, 177.083°

E, 17 May 1986. Tonga: USNM 33320U (3) Tongatapu, W side

of Hakau Mama’o Reef, 21° S, 175.215° W, 23 October 1993;

USNM 334166^ (2) Tongatapu, reef just N of Atata Island, on N

side of reef, 21.0194° S, 125.228° W, 28 October 1993; USNM

334543^ (4) W side of Makaha’a off the N central coast of

Tongatapu Island, 21.1117° S, 175.16° W, 29 October 1993;

USNM 336905^ (1) Uoleva Island, outside fringing reef on W

side, 19.8356° S, 174.422° W, 9 November 1993. American

Samoa: BPBM 150ir (61).

Choerodon margaritiferus. (11 specimens examined, 82.8-115

mm SL). Japan, Okinawa: OCE-P 20140825-3* (96.4) off

Sesoko Island, 26° 39' N, 127° 51' E, 30 m, 24 August 2014;

OCE-P 20140827-1* (107) off Sesoko Island, 26° 39' N, 127°

51' E, 30 m, 27 August 2014; OCE-P 20140827-2* (103) same

collection data as OCE-P 20140827-1; OCE-P 20140827-3*

(94.5) same collection data as OCE-P 20140827-1; OCE-P

20140827-4* (97.4) same collection data as OCE-P 20140827-1.

Philippines: MNHN 2016-0095* (110) Philippines; USNM

135558* (115, holotype of Choerodon margaritiferus) Jolo;

USNM 435387 (114) Cebu, Kawit, Medellen, Visayan Sea;

USNM 435389* (115) Cebu, Kawit, Medellen, Visayan Sea;

USNM 435391 (108) Cebu, Kawit, Medellen, Visayan Sea;

Indonesia: CSIRO H 7218-03* (94.9) Lombok, Tanjung Luar,

8° 48' S, 116° 29' E, 26 January 2011, LM941; WAM P.31498-

003* (82.8, paratype of Choerodon gomoni) Banggai Islands,

Bangang Island, 1° 41.12' S, 123° 27.50' E, 30 m, 4 November

1998.

Choerodon monostigma. (116 specimens examined, 55.4—225

mm SL). Australia, Queensland: AMS E.2531 (123) Bowen,

19° 55' S, 148° 18' E, 29 m, 3 August 1910; AMS E.2683 (9,

95-147) Pine Peak, 21° 25' S, 150° 08' E, 48 m, 1 August 1910;

AMS E.2777 (4, 120-133) Gloucester Island, 19° S, 148° E, 64

m, 2 August 1910; AMS E.2778 (3, 100-125) same collection

data as AMS E.2777; AMS E.2779* (2,147-158) same collection

data as AMS E.2777; AMS 1.12518* (104, syntype of Choerodon

monostigma) Pine Peak, 21° 31' S, 150° 16' E, 1910; AMS

1.15557-204 (2, 118-123) Gulf of Carpentaria, 17° 25' S, 140°

10' E, 10.1 m, 27 November 1963; AMS 1.16013-001* (120)

Townsville, 19° 15' S, 146° 50' E, 1971; AMS 1.20826-038 (99.9)

Palm Islands, 18° 44' S, 146° 31' E, 20 m, 5 Eebruary 1979; AMS

1.20827-009 (91.4) Cape York, Hannibal Island, 11° 33' S, 142°

57' E, 22-23 m, 15 Eebruary 1979; AMS 1.34372-001 (80)

Leicester Island, 22° 09' 42" S, 150° 26' 07" E, 22° 09' 56" S,

150° 25' 55" E, 28 m, 24 October 1993; AMS 1.34373-002* (4,

55.^71.4) Collins Island, Shoalwater Bay, 22° 11' 51" S, 150°

19' 18" E, 22° 11' 32" S, 150° 18' 26" E, 19 m, 24 October 1993;

AMS 1.34374-009 (3 , 62-155) Shoalwater Bay, Akens Island,

22° 19' 54" S, 150° 15' 48" E, 22° 20' 08" S, 150° 14' 57" E, 9 m,

24 October 1993; AMS 1.34398-030 (115) Port Clinton, 22°

33' 29" S, 150° 45' 19" E, 22° 33' 14" S, 150° 45' 25" E, 11 m, 25

October 1993; AMS IA.4196- (1) Port Curtis, 23° 55' S, 151°

25' E, December 1929; AMS IB.3303* (85.6) Repulse Bay, 20°

35' S, 148° 45' E, October 1954; AMS IB.3304* (77.8) Repulse

Bay, 20° 35' S, 148° 45' E, October 1954; CSIRO A 2153 (102)

Gulf of Carpentaria, 16° 34' S, 139° 56.4' E, 29.3 m, 29 August

1963; CSIRO A 2154 (96.1) Gulf of Carpentaria, 16° 34' S, 139°

56.4' E, 29.3 m, 29 August 1963; CSIRO A 2418 (70.2) Gulf of

Carpentaria, 16° 40.0' S, 139° 57.7' E, 29.3 m, 28 August 1963;

CSIRO A 2495 (85.1) Pisonia Island, 16° 29' S, 139° 51' E, 25.2

m, 2 September 1963; CSIRO A 2496 (71.4) Gulf of Carpentaria,

Pisonia Island, 16° 29' S, 139° 51' E, 25.2 m, 2 September 1963;

CSIRO A 2515 (96.3) Gulf of Carpentaria, Mornington Island,

15° S, 139° E, 45 m, 2 September 1963; CSIRO A 3255 (93.6)

Gulf of Carpentaria, 16° 44.9' S, 140° 03.2' E, 20.1 m, 28 August

1963; CSIRO C 3718 (122) Gulf of Carpentaria, Weipa, 12°

00' S, 141° 30' E, 39 m, 13 December 1968; CSIRO C 4055 (134)

Gulf of Carpentaria, Wellesley Islands, 16° 38.7' S, 140° 05.1' E,

25.2 m, 28 August 1963; CSIRO CA 2170 (146) Gulf of

Carpentaria, Port Musgrave, 11° 04' S, 139° 58' E-ll° 06' S,

139° 58' E, 56-57 m, 13 June 1981; CSIRO H 3300-02 (128)

Clerke Island, 11° 53.1' S, 143° 16.0' E, 39 m, 13 January 1993;

CSIRO H 3309-09 (126) Hannibal Island, off Shelburne Bay, 11°

37.9' S, 143° 01.7' E-ll° 39.1' S, 143° 02.3' E, 22 m, 14 January

1993; CSIRO H 3895-15 (100) Cape Grenville, 11° 37' S, 143°

28' E, 18 m, 21 October 1994; CSIRO H 3932-02 (100) Cape

Grenville, 11° 37' S, 143° 28' E, 18 m, 21 October 1994; CSIRO

H 6736-06* (78.4) Torres Strait, NE of Bramble Cay, 9° 05.14' S,

143° 56.40' E, 51 m, 27 January 2004; CSIRO H 6895-06 (88.4)

Torres Strait, Endeavour Strait, 10° 49.22' S, 142° 09.42' E-10°

49.76' S, 142° 09.55' E, 15 m, 19 January 2004; CSIRO H 6900-

02 (55.8) Torres Strait, NE of Banks Island, 9° 51.72' S, 142°

35.40' E, 13 m, 21 January 2004; CSIRO H 6904-04* (119)

Torres Strait, NE of Newcastle Bay, 10° 28.95' S, 142° 52.48' E,

19 m, 12 January 2004; CSIRO H 6927-02* (102) Torres Strait,

NE of Cape York Peninsula, 10° 09.57' S, 143° 07.04' E, 25 m, 1

Eebruary 2004; CSIRO H 6936-03* (86.8) Torres Strait, NE of

Cape York Peninsula, 10° 12.32' S, 143° 09.43' E, 29 m, 1

Eebruary 2004; CSIRO H 7018-04 (4, 83-100) Gulf of

Carpentaria, Mornington Island, 16° 19.00' S, 138° 39.53' E-16°

18.16' S, 138° 38.12' E, 24-23 m, 23 Eebruary 2009; CSIRO H

7677-02* (112) W of Hook Island, 20° 06.04' S, 148° 41.01' E, 4

December 2003; NMV A2950 (2, 90-105) Gulf of Cai-pentaria,

105 km WNW of Cullen Point, 11° 25' 11.99" S, 141°

or 48.00" E, 45 m, 8 September 1982; NMV A2961 (2, 90-130)

Gulf of Carpentaria, 130 km W of Crab Island, 10° 49' 12.00" S,

140° 54' 29.99" E, 43 m, 10 September 1982; NMV A2967 (105)

Gulf of Carpentaria, 140 km W of Prince of Wales Island, 10°

39' 00.00" S, 140° 45' 00.00" E, 46 m, 13 September 1982; NMV

A3112 (125) 12 km SW of Cape Grenville, 12° 04' 40.79" S,

143° 10' 10.80" E, 10 m, 27 Eebruary 1983; NMV A3132* (2,

107-130) Gulf of Carpentaria, 180 km W of Cape York, 10°

50' 13.80" S, 140° 55' 06.59" E, 44 m, 26 Eebruary 1983; NMV

A3136 (2,105-110) Gulf of Carpentaria, 60 km W of Weipa, 12°

36' 18.00" S, 141° 19' 55.80" E, 40 m, 25 Eebruary 1983; NMV

A3156 (127) Great Barrier Reef, near Haggerstone Island, 12°

03' 16.92" S, 143° 15' 10.08" E, 26-29 m, 27 Eebruary 1983;

NMV A13295 (121) 11 km NNE of Cape Grenville, 11°

53' 05.99" S, 143° 16' 12.00" E, 39 m, 13 January 1993; NMV

A13296(2,118-225) 315 km E of Cape Wessel, 11° 06'35.99" S,

139° 42' 24.00" E, 54 m, 5 Eebruary 1993; NMV A13297 (105)

Gulf of Cai-pentaria, 10° 54' 29.99" S, 140° 34' 18.00" E, 51 m, 5

Eebruary 1993; NMV A13298 (119) 16 km N of Cape Grenville,

11° 48' 42.00" S, 143° 12' 42.00" E, 31 m, 13 January 1993;

104

M.F. Gomon

NMVA13299* (119) Gulf of Carpentaria, 12° 35' 23.99" S, 140°

46' 12.00" E, 60 m, 30 January 1993; NMV A13300* (125) 12

km ESE of Hannibal Island, 11° 37' 54.00" S, 143° 01' 41.99" E,

22 m, 14 January 1993; NMVA13301 (154) Gulf of Carpentaria,

11° 16' 54.00" S, 140° or 41.99" E, 59-51 m, 5 Eebruary 1993;

NMV A 31433-001* (128) NE of Shoalwater Bay, 21° 45.31' S,

150° 57.76' E, 73 m, 8 December 2003; NMV A 31434-001*

(82.6) NE of Cairns, 16° 47.27' S, 145° 58.01' E, 39 m, 17

October 2004; QMB 1.1563* (131, paralectotype of Choerodon

monostigma) Pine Peak, 21° 31' S, 150° 16' E, 47.5 m, 1910;

QMB 1.9904 (1) Lindeman Island, 20° 27' S, 149° 2' E, 1936;

QMB I.1125P (?) Townsville, 19° 16' S, 146° 49' E; QMB

1.11252^ (?) Mackay, 21° 9' S, 149° 12' E; QMB I.1180P (?)

Prudhoe Island, 21° 19' S, 149° 41' E; QMB 1.15732^ (?) Palm

Islands, 18° 44' S, 146° 35' E, 20 m, 5 Eebruary 1979; QMB

1.17552+ (?) Torres Strait, P D Sand Cay, 10° 10' S, 143° 14' E, 6

December 1974; QMB 1.17635+ (?) Torres Strait, Aureed Island,

9° 57' S, 143° 17' E, 17 November 1974; QMB 1.17636+ (?)

Torres Strait, P D Sand Cay, 10° 11' S, 143° 14' E, 18 October

1974; QMB 1.17637+ (?) Torres Strait, Stephen to Bramble Cay,

9° 20' S, 143° 42' E, 13 October 1974; QMB 1.17638+ (?) Torres

Strait, Yam Island, 9° 53' S, 142° 45' E, 22 November 1974;

QMB 1.17639+ (?) Torres Strait, Caldbeck Reef, 10° 10' S, 143°

13' E, 4 December 1974; QMB 1.17640+ (?) Torres Strait, Keats

Island, 9° 41' S, 143° 27' E, 8 October 1974; QMB 1.17641+ (?)

Torres Strait, 9° 26' S, 142° 50' E, 7.3 m, 24 March 1974; QMB

1.17642+ (?) Torres Strait, 9° 24' S, 142° 54' E, 7.3-9.1 m, 23

March 1974; QMB 1.17643+ (?) Torres Strait, 9° 30' S, 142°

59' E, 7.3-8.2 m, 25 April 1974; QMB 1.17644+ (?) Torres Strait,

9° 35' S, 142° 47' E, 26 March 1974; QMB 1.17645+ (?) Torres

Strait, 9° 37' S, 142° 54' E, 8.2 m, 27 March 1974; QMB 1.17646+

(?) Torres Strait, 9° 55' S, 141° 33' E, 12.8-14.6 m, 31 May 1974;

QMB 1.17647+ (?) Torres Strait, 9° 40' S, 142° 56' E, 29 March

1974; QMB 1.17648+ (?) Torres Strait, 9° 51' S, 142° 49' E, 14.6-

26.8 m, 5 April 1974; QMB 1.18163+ (?) Cape Weymouth, 12°

29' S, 143° 20' E, 18.3 m, 24 September 1979; QMB 1.23129+ (?)

Hinchinbrook Island, 18° 27' S, 146° 25' E, 24 m, 15 April 1985;

QMB 1.26705+ (?) Torres Strait, Warrior Reefs, 9° 48' S, 142°

58' E, 30 March 1990; QMB 1.26845+ (?) Torres Strait, Warrior

Reefs, 9° 48' S, 142° 58' E, 27 March 1990; QMB 1.27405+ (?)

Gulf of Carpentaria, 12° 29' S, 139° 42' E, 58 m, 8 December

1990; QMB 1.27765+ (?) Gulf of Carpentaria, 11° S, 140° 41' E,

47 m, 4 December 1990; QMB 1.27903+ (?) Gulf of Carpentaria,

12° 29' S, 139° 42' E, 58 m, 8 December 1990; QMB 1.36056+ (?)

20° 17.1' S, 148° 53.1' E, 35 m, 29 November 2003; QMB

l. 36337+ (?) 23° 22.5' S, 151° 2.7' E, 20 m, 25 January 2004;

QMB 1.36573+ (?) 20° 51.3' S, 149° 33.3' E, 43 m, 30 September

2004; QMB 1.36588+ (?) 21° 11.1' S, 150° 19.5' E, 53 m, 29

September 2004; QMB 1.36706+ (?) 21° 11.7' S, 149° 57.3' E, 48

m, 29 September 2004; QMB 1.36780+ (?) 22° 00.3' S, 149°

32.7' E, 11 m, 5 October 2004; Northern Territory: AMS

l. 21842-029+ (9, 80-125) Arafura Sea, 10° 35' S, 133° 45' E, 10°

37' S, 133° 47' E, 60 m, 16 November 1980; AMS 1.21944-005+

(140) Arafura Sea, 11° 35' S, 135° 00' E, 11° 34' S, 134° 54' E, 35

m, 19 November 1980; CSIRO B 2082 (8, 70.5-115) Darwin,

12° 15.0' S, 130° 03.0' E-12° 13.0' S, 130° 02.0' E, 50 m, 3 July

1980; CSIRO C 3725 (152) Arafura Sea, 08° 57' S, 137° 39' E, 58

m, 4 December 1968; CSIRO CA 2548 (90) Groote Eylandt,

Tasman Point, 14° 20' S, 136° 10' E, 18 m, 27 May 1981; NMV

A1731 (7, 115-170) Arafura Sea, 10° 10' 11.99" S, 135°

58' 11.99" E, 52 m, 22 November 1980; NTM S.01080-001+ (82)

York Sound, July 1975; NTM S.11684-007+ (3,115-167) Arafura

Sea, Cape Wessel, 10° 09' S, 136° 48' E, 57 m, 17 March 1985;

NTM S.11613-019+ (136) Arafura Sea, Cape Wessel, 10°

15' 00" S, 136° 19' 59" E, 10 March 1985; NTM S. 11953-005+ (2,

76-93) Arafura Sea, Goulbum Island, 10° 34' 59" S, 134°

33' 00" E, 55 m, 25 August 1986; NTM S.10052-001+ (118) Van

Diemen Gulf, Murganella Creek, 11° 52' 01" S, 132° 34' 59" E,

17 January 1978; NTM S.10051-001+ (3, 59-125) Van Diemen

Gulf, Murganella Creek, 11° 52' 01" S, 132° 31' 01" E, 26

October 1977; NTM S. 11786-012+ (5, 109-156) Arafura Sea,

Goulbum Islands, 10° 49' 01" S, 133° 49' 59" E, 59 m, 7

December 1985; NTM S.16134-026 (63) + Bynoe Harbour, Knife

Island, 12° 42' 04" S, 130° 35' 31" E, 7 m, 20 April 2002; NTM

S. 13790-002+ (28) Beagle Gulf, Bynoe Harbour, 12° 40' 59" S,

130° 33' 00" E, 9 m, 7 October 1993; NTM S. 10069-001+ (83)

Darwin Harbour, Shoal Bay, 12° 16' 59" S, 130° 58' 01" E, 5

April 1977; NTM S.13534-002+ (3, 120-127) Arafura Sea, 10°

30' 18" S, 134° 30' 00" E, 58 m, 24 September 1992; NTM

S.16200-002+ (14) Port Patterson, South Moira Reef, 12°

3 r 01" S, 130° 31' 01" E, 8 November 2002; NTM S. 12935-004+

(140) Arafura Sea, Croker Island, 10° 43' 59" S, 132° 31' 01" E,

55 m, 21 October 1990; NTM S. 13282-022+ (95) Beagle Gulf,

Charles Point, 12° 15' 25" S, 130° 37' 52" E, 29 m, 2 September

1992; NTM S.12938-005+ (119) Arafura Sea, 10° 48' 00" S, 133°

05' 60" E, 60 m, 21 October 1990; NTM S. 12942-006+ (2, 108-

155) Arafura Sea, 11° 22' 01" S, 134° 07' 01" E, 22 October

1990; NTM S.12948-004+ (3, 111-150) Arafura Sea, 10°

45' 00" S, 133° 52' 59" E, 61 m, 24 October 1990; NTM S. 12952-

018+ (152) Arafura Sea, 10° 34' 01" S, 134° 28' 59" E, 60 m, 24

October 1990; NTM S.12961-001+ (144) Arafura Sea, 11°

22' 59" S, 136° 05' 60" E, 34 m, 26 October 1990; NTM S. 16159-

036+ (39) Port Patterson, Moira Reef, 12° 30' 50" S, 130°

30' 40" E, 8 November 2002; NTM S. 13718-012+ (29) Darwin

Harbour, Jones Creek, 12° 33' 18" S, 130° 52' 59" E, 4 m, 15 July

1993; NTM S. 13724-005+ (25) Darwin Harbour, Pearl Raft

Creek, 12° 31' 01" S, 130° 54' 00" E, 3 m, 16 July 1993; NTM

S. 13735-012+ (2, 1^15) Darwin Harbour, West Arm, 12°

33' 18" S, 130° 46' 41" E, 12 m, 9 September 1993; NTM

S.13736-015+ (15) Darwin Harbour, West Arm, 12° 34' 19" S,

130° 45' 25" E, 8 m, 9 September 1993; NTM S.13927-002+ (16)

Darwin Harbour, Eannie Bay, 12° 25' 55" S, 130° 48' 36" E, 12

m, 18 April 1994; NTM S.13333-006+ (3,129-175) Arafura Sea,

11° 19' 01" S, 134° 28' 01" E, 39 m, 23 October 1990; NTM

S. 10050-002+ (3, 62-98) Darwin Harbour, Shoal Bay, 12°

16' 01" S, 130° 54' 00" E, 4April 1977; NTM S.15134-001+ (70)

Timor Sea, Elat Top Bank, 12° 27' 18" S, 129° 13' 37" E, 60 m,

12 October 1996; NTM S. 15916-019+ (4, 23-26) Kakadu

National Park, Pield Island, 12° 03' 25" S, 132° 24' 32" E, 1 m,

19 August 2004; NTM S.11657-038+ (2, 150-170) Arafura Sea,

Cape Wessell, 10° 24' 00" S, 136° 30' 00" E, 38.5 m, 2 Eebmary

1985; NTM S.11655-013+ (113) Arafura Sea, Cape Wessell, 10°

25' 01" S, 136° 33' 00" E, 60 m, 1 Eebruary 1985; NTM S. 11660-

022+ (4,120-146) Arafura Sea, Cape Wessell, 10° 24' 00" S, 136°

35' 60" E, 62 m, 3 Eebmary 1985; NTM S. 11658-019+ (148)

Arafura Sea, Cape Wessell, 10° 24' 00" S, 136° 31' 59" E, 56.7 m,

2 Eebruary 1985; NTM S.16951-004+ (5, 18-26) Darwin

Harbour, East Point, 12° 25' 01" S, 130° 48' 18" E, 16 November

1982; NTM S. 16750-002+ (82) Blue Mud Bay, Nicol Island, 13°

27' 00" S, 136° 26' 60" E, 22 m, 27 June 1987; NTM S. 11612-

037+ (2,133-158) Arafura Sea, Cape Wessel, 10° 25' 59" S, 136°

25' 59" E, 57 m, 9 March 1985; NTM S. 17296-002+ (46) Beagle

Review of Choerodon tuskfishes

105

Gulf, Loma Shoals, 12° 12' 00" S, 129° 59' 17" E, 41 m, 20

September 2009; WAM P.14260='= (70.2) Darwin, 12° 27' S, 130°

50' E, 4 September 1965; WAM P. 14426-00 E (1) same collection

data as WAM .P14260; WAM P.14427-001 (1) same collection

data as WAM P.14260. Western Australia: AMS 1.20402-011

(124) Bonaparte Archipelago, Admiralty Gulf to Camden Sound,

14° 00' S, 126° 00' E, 15° 20' S, 124° 25' E, 12-60 m, April 1978;

AMS 1.22801-012 (2, 72-87) North West Shelf, Port Hedland,

19° 32' S, 118° 09' E, 50-52 m, 26 March 1982; AMS 1.22802-

009* (2, 121-143) North West Shelf, Port Hedland, 19° 39' S,

117° 53' E, 52-53 m, 26 March 1982; CSIRO CA 463 (119)

Lagrange Bay, 18° 34' S, 120° 21' E-18° 30' S, 120° 23' E, 85-88

m, 2 June 1978; CSIRO CA 2169 (114) Eorestier Island, 19°

45.0' S, 118° 22.0' E-19° 46.0' S, 118° 21.0' E, 38 m, 3 June

1980; CSIRO H 3240-01 (101) Port Hedland, 19° 15.7' S, 118°

27.5' E-19° 15.0' S, 118° 29.1' E, 75-76 m, 3 October 1990;

CSIRO H 4352-01 (120) Port Hedland, 19° 29.5' S, 118°

52.2' E-19° 30.0' S, 118° 52.1' E, 37 m, 24 October 1983; CSIRO

H 4353-01 (2,72-88) Port Hedland, 19° 29.6' S, 118° 52.2'E-19°

29.7' S, 118° 52.6' E, 38-37 m, 25 October 1983; NTM S.10959-

027+ (3,101-139) Port Hedland, 19° 01' 01" S, 119° 25' 01" E, 18

April 1983; NTM S.11671-015+ (145) North West Shelf, Rowley

Shoals, 19° 03' 00" S, 118° 24' 00" E, 88 m, 2 June 1985; NTM

S. 11688-009+ (2, 102-149) North West Shelf, King Sound, 16°

31' 59" S, 121° 25' 59" E, 42 m, 17 April 1985; NTM S. 14362-

013+ (3, 61-99) Holothuria Banks, West Holothuria Reef, 13°

25' 12" S, 125° 39' 18" E, 50 m, 12 June 1996; NTM S. 15060-

003 (96) Joseph Bonaparte Gulf, 13° 13' 59" S, 128° 07' 01" E,

90 m, 14 July 1997; NTM S.16591-016+(148) Holothuria Banks,

Penguin Shoal, 12° 58' 59" S, 125° 35' 60" E, 67 m, 27 June

1988; WAM P.26259-009 (86) Cape Leveque, 16° 20' S, 122°

19' E, 50-52 m, 23 June 1978; WAM P.33928-001+ (82)

Kimberley, 13.79011647° S, 127.2439084° E, 54.5-55.1 m, 10

June 2013. Indonesia: WAM P.33092-002+ (8, 40-103) West

Papua, Ogar Island, 02° 39' S, 132° 30' E, 8-25 m, 22 March

2009.

Choerodon oligacanthus. (68 specimens examined, 50.5-250

mm SL). BMNH 1870.6.14.63 (1) unknown locality. Day.

Malaysia: AMS 1.28082-027* (97.6) 1922; MCZ 30358 (73.8)

Penang; RMNH 17917 (50.5) Port Dickson. Singapore: AMS

1.28997-002* (121) Eebruary 1923; AMS IA.3275* (134) 1923;

AMS IA.3276* (138) 1923; CAS 29237* (93.1); CAS 30378

(1); CAS 34609 (1); MCZ 1133 (111); CAS 34609 (1); MCZ

1133 (111); MCZ 25258 (222); MCZ 14374 (9, 70.5-143).

Borneo: CAS 27769 (1); CAS 33637 (1) 5.989066° N,

118.166168° E, 29 June 1929. Sabah: USNM 152167 (1)

Sandakan; USNM 153485 (1) Sandakan; USNM 153980 (7)

Sandakan. Philippines: BPBM 22093* (2, 113) Cebu; CAS

20653 (1); CAS 25931 (1) Palawan, 11.896447° N, 120.018080°

E, 6 May 1931; CAS 25932 (2) Tawi-Tawi, 4.717582°

N, 119.401948° E, 9 August 1931; NSMT-P 50478* (65.3) Cebu

fish market, 13 Eebruary 1979; USNM 51988* (250) Negros

Island; USNM 152152 (1) Cebu; USNM 153484 (1) Palawan;

USNM 153981 (1) Ulugan Bay, near mouth of Baheli River;

USNM 153982 (1) Cebu; USNM 153983 (3) Cebu; USNM

153985 (2) Iloilo; USNM 153986 (1) Sirinao Island, Nakoda

Bay; USNM 200221* (212) Palawan, Puerto Princessa. Batavia:

MCZ 14371 (2, 138-151); MCZ 22798 (248). Indonesia:

CSIRO H 7986-01* (224) Bali, Kedonganan, 8° 45' S, 115°

09' E, 25 Eebruary 2009; CSIRO H 7987-01* (229) same locality

as CSIRO H 7986-01, 1 October 2009; CSIRO H 7406-07 (107)

Lombok, Tanjung Luar, 8° 48' S, 116° 29' E, 4 March 2009;

CSIRO H 7694-02* (142) same collection information as CSIRO

H 7987-01; NMV 46066 (76.4) Bleeker collection; NMV A

31430-001* (201) same locality as CSIRO H 7986-01,2 October

2009; MZB 23113* (177) same collection information as CSIRO

H 7986-01; MZB 23114 (94.8) same collection information as

CSIRO H 7406-07; RMNH 6532* (4, 77.0-88.6, 88.6 mm SL

specimen regarded as lectotype of Crenilabrus oligacanthus)

Sumatra, Riau.

Choerodon robustus. (32 specimens examined, 146-285 mm

SL). Unknown locality: BMNH 1867.3.9.20* (242, skin). Red

Sea: BPBM 20841 (265) Gulf of Aqaba, Eilat; MNHN 1966-

196* (200) Gulf of Suez; MNHN 1966-197* (176) Gulf of Suez.

Persian Gulf: BPBM 21188 (203) Bahrain. INDIAN OCEAN,

Somalia: USNM 438442* (185) 09° 41' N, 51° 03' E, 60-70 m.

Mauritius: BMNH 1840.12.12.10* (275, holotype of

Xiphocheilus robustus) Janvier; BMNH 1941.4.21.6* (236);

MCZ 5803 (224); MCZ 6101 (263); RMNH 6534* (204,

holotype of Choerops dodecacanthus). Reunion: MNHN

A.8264* (275, syntype of Cossyphus maxillosus, mount);

MNHN A.8265* (245, syntype of Cossyphus maxillosus, mount);

MNHN A.8266* (256, syntype of Cossyphus maxillosus, mount).

Seychelles: ANSP 114058* (3, 204-228) Curieuse Island, 20 m,

23 Eebruary 1964. Sri Lanka: BMNH 1891.10.29.32* (236,

skin) H. Nevill; USNM 438443 (234) Wadge Banks, 07° 52' N,

77° 09' E, 51 m; USNM 206124* (278) Wadge Banks.

WESTERN PACIEIC OCEAN, Japan, Okinawa: NSMT P

66574* (188) Nakagusuku Bay, 7 November 2000; SMBL-E

73327 (285) Naha fish market, 100 m, 6 August 1973; SMBL-E

73328 (261) same collection data as SMBL-E 73324. URM-P

18745* (199) off Yasuda, 200 m, 25 October 1986; URM-P

23316* (207) Naha fish market, 30 December 1989; URM-P

24632* (164) Naha fish market, 20 October 1990; URM-P

24825* (221) Naha fish market, 14 November 1990; URM-P

40999* (237) Awase, 7 November 1990. Indonesia: CSIRO H

7989-01* (146) West Java, Pelabuhanratu, 7° 00' S, 106° 30' E,

11 March 2009; CSIRO H 7990-01* (194) West Java,

Pelabuhanratu, 7° 00' S, 106° 30' E, 10 March 2009; CSIRO H

7361-02* (229) Bali, Kedonganan, 8° 45' S, 115° 09' E, 14 March

2010; MZB 23115* (220) West Java, Pelabuhanratu, 07° 02' S,

106° 32' E, 14 October 2008; NMV A 31431-001* (204) same

collection data as CSIRO H 7361-02.

Choerodon rubescens. (15 specimens examined, 54.4—521 mm

SL). Australia, Western Australia: AMS 1.20230-010* (111)

Cockbum Sound, Woodman Point, 32° 08' S, 115° 45' E, 1-9 m,

27 March 1978; AMS 1.44129.002+ (1) May 2007; AMS 1.44129-

001+ (390) May 2007; BMNH 1844.2.15.54* (521, mount)

Houtman Abrolhos; BMNH 1844.2.15.66* (mount) Harvey

River; BMNH 1844.2.15.68* (279, holotype of Choerops

rubescens, skin) Houtman Abrolhos; CSIRO H 4875-01* (380)

October 1998; CSIRO H 4875-02* (324) October 1998; CSIRO

H 4875-03* (380) October 1998; NMV A2495 (252) Houtman

Abrolhos, off Beacon Island in Goss Passage, 28° 28' 48.00" S,

113° 46' 11.99" E, 30 m, 19 April 1982; WAM P.4186-001* (177)

Lancelin Island, 31° 00' S, 115° 19' E; WAM P.4515-001+ (1)

Shark Bay, 25° 21' S, 113° 44' E, November 1958; WAM P.4514-

001+ (1) Peron Peninsula, 26° 03' S, 113° 37' E, 11 January 1958;

WAM P.25310-002* (2, 54.4-67.7) Houtman Abrolhos, Beacon

Island, 28° 30' S, 113° 44' E, 2-15,18 May 1975; WAM P.25749-

106

M.F. Gomon

001* (206) Rottnest Island, Kingston Reefs, 31° 59' S, 115°

33' E, 8-10 m, 3 March 1977; WAM R 25759-001* (102)

Rottnest Island, Charlotte Point, 31° 59' S, 115° 29' E, 2^ m, 10

March 1977; WAM P.25913-002+ (183) Garden Island, 32° 12' S,

115° 40' E, January 1977; WAM P27955-018 (60) Port Denison,

29° 16' S, 114° 55' E, 9-10 m, 13 April 1983; WAM P27957-

026^ (42) Port Denison, 29° 16' S, 114° 55' E, 7-8 m, 14 April

1983; WAM P32774-00P (52) Cervantes, 30° 33.290' S, 115°

05.508' E, 3.9 m, 9 March 2006.

Choerodon schoenleinii. (113 specimens examined, 21-530 mm

SL). Japan: USNM 57670* (2, 196-229). China: ANSP 76600

(1) Hong Kong; BMNH 1968.3.11.15* (280, holotype of

Cossyphus cyanostolus, skin); BMNH 1968.3.11.16* (124,

holotype of Cossyphus ommopterus, skin); CAS 27904 (1) Hong

Kong; MCZ 14377 (1) Hong Kong; USNM 153846* (218) Hong

Kong; USNM 153847* (227) Hong Kong; USNM 153849 (227)

Hong Kong; USNM 148431 (1) Shanghai. Taiwan: USNM

192861 (2) Peng-Hu Hsien; USNM 276983* (104) Yeh-liu.

Vietnam: MNHN 97-187* (410) Tonkin Gulf. Thailand:

NSMT-P 55149* (175) Rayong, Ko Samet, Ao Phao, 4 m, 29

November 1986. Philippines: CAS 25933 (150) Sulu

Archipelago, Jolo, 5.925203° N, 121.153284° E-5.925203° N,

121.153284° E, 2 August 1931; CAS 32363 (2, 159-187) Sulu

Archipelago, Jolo, 5.925203° N, 121.153284° E-5.925203° N,

121.153284° E, 30 December 1936; CAS 38730 (166)

Zamboanga, 6.907358° N, 122.069655° E-6.907358° N,

122.069655° E, 3 September 1940; USNM 88065 (2) Masbate;

USNM 150210* (150) Zamboanga; USNM 152166* (250)

Zamboanga; USNM 152252 (1) Masbate; USNM 153470* (389)

Cebu; USNM 153630 (1) Masbate, Cataingan Bay; USNM

153628 (2) Cebu; USNM 153629 (2) Cebu; USNM 153469*

(223) Cebu; USNM 135559 (1) Luzon, Ragay Rive. Singapore:

MCZ 14373 (2). Sabah: CAS 27894 (1) Sandakan, 5.989066°

N, 118.166168° E, 29 June 1929. Indonesia: ANSP 27703 (1)

Sumatra, Padang; CSIRO H 7899-03 (320) Bali, Kedonganan,

08° 45' S, 115° 09' E, 21 October 2008; CSIRO H 7362-02 (160)

Bali, Kedonganan, 8° 45' S, 115° 09' E, 21 January 2011; MNHN

A7395* (110) Sulawesi; MZB 23116 (128) Bali, Kedonganan,

8° 45' S, 115° 09' E, 1 October 2009; NMV46065* (122); USNM

153631 (1) Sulawesi, Macassar; WAM P.31305-005 (86) Riau

Islands, Bintan Island, 01° 11' N, 104° 19' E, 13 May 1997.

Timor: RMNH 2364* (140). Papua New Guinea: AMS

IB.476U (1) 06° 25' S, 147° 12' E, 1960; WAM P.28194-036

(262) Daru Island, Daru, 9° 09' S, 143° 17' E, 3^ m, 6 November

1983. Australia: AMS 1.2972^ (1) unknown locality, 1891; AMS

1.4842^ (1) unknown locality, 1901; AMS IA.69' (1) unknown

locality, 1920. New South Wales: AMS 1.1833^ (1), Richmond

River, 29° 04' S, 153° 21' E, 1888; AMS 1.1845+ (1) Clarence

River, 28° 40' S, 152° 21' E, 29° 30' S, 153° 06' E, 1888; AMS

1.1846+ (1) Clarence River, 28° 40' S, 152° 21' E, 29° 30' S, 153°

06' E, 1888; AMS 1.2342+ (1) Clarence River, 29° 30' S, 153°

06' E, 1888; AMS 1.3298+ (1) Sydney Eish Market, 1894; AMS

1.4181+ (1) Port Jackson, 33° 51' S, 151° 16' E, 1899; AMS

1.4181.001+ (1) Port Jackson, 33° 51' S, 151° 16' E, 1899; USNM

59849* (423) N coast. Queensland: AMS A.7408 (152) Torres

Strait, 09° S, 142° E-10° S, 143° E, 1879; AMS A.7409 (157)

Torres Strait, 09° S, 142° E-10° S, 143° E, 1879; AMS A.7408+

(1) Torres Strait, 09° S, 142° E, 10° S, 143° E, 1879; AMS

A.7409+ (1) Torres Strait, 09° S, 142° E, 10° S, 143° E, 1879;

AMS 1.11937+ (1) Cape York, 10° 00' S, 142° 00' E, 1907; AMS

1.12520 (134) Great Sandy Strait, 25° 35' S, 152° 58' E, 1912;

AMS 1.12633 (1) Masthead Island, 23° 32' S, 151° 44' E, August

1912; AMS 1.15557-205* (3, 95.5-181) Gulf of Carpentaria, 17°

25' S, 140° 10' E, 10.1 m, 27 November 1963; AMS 1.16360-

001* (210, holotype of Choerops notatus) Cape Grenville, 12° S,

143° 15' E, 1875; AMS 1.19474-001* (133) Linnet Reef, 14°

47' S, 145° 21' E, 25, 22 November 1975; AMS 1.20753-010*

(60.5) Lizard Island area, 2 miles NW of Nymph Island, 14°

36' S, 145° 14' E, 1^15 m, 8 Eebruary 1979; AMS 1.20754-013

(58.3) Lizard Island area, 10 miles NW of Nymph Island, 14°

33' S, 145° 06' E, 16 m, 8 Eebruary 1979; AMS 1.20787-075 (47)

Linnet Reef, 14° 47' S, 145° 21' E, 2-3 m, 9 December 1978;

AMS 1.33802-001 (1) Lizard Island, Mermaid Cove, 14° 40' S,

145° 28' E, 4 m, 17 January 1993; AMS 1.34302-003+ (237)

Shoal water Bay, Sabina Point, 22° 23' 49" S, 150° 18' 15" E, 2 m,

14 September 1993; AMS 1.34311-021 (104) Shoalwater Bay,

Collins Island, 22° 14' 47" S, 150° 19' 08" E, 3 m, 15 September

1993; AMS 1.34311-028 (138) Shoalwater Bay, Collins Island,

22° 14' 47" S, 150° 19' 08" E, 3 m, 15 September 1993; AMS

1.9499 (95.9) Dunk Island, 17° 57' S, 146° 09' E, 1908; AMS

IA.6799* (91.3) Lindeman Island, 20° 27' S, 149° 02' E, 1936;

AMS IA.6800* (75.9) Lindeman Island, 20° 27' S, 149° 02' E,

1936; AMS IB.2742 (77.1) Moreton Bay, 27° S, 153° E, 1951;

AMS IB.5872 (222) Townsville, 19° S, 146° E, 1962; BMNH

1850.7.20.71* (180, skin) Cape York, HMS Rattlesnake; CSIRO

C 211+ (280 TL) Torres Strait, Bramble Cay, 22 January 1949;

CSIRO C 4004 (148) Gulf of Carpentaria, Gulf of Carpentaria,

Bentinck Island, 17° 06.2' S, 139° 44.2' E, 12.6 m, 12 August

1963; CSIRO C 4478 (152) Gulf of Carpentaria, Bentnick Island,

16° 55' S, 139° 37' E, 12.5 m, 1 November 1972; CSIRO C 4505

(150) Gulf of Carpentaria, Bentnick Island, 16° 55' S, 139° 37' E,

12.5 m, 1 November 1972; CSIRO CA774 (87.9) Torres Strait,

Thursday or Prince of Wales Islands, 10° 40' S, 142° 15' E, April

1979; CSIRO H 6558-05 (87) Torres Strait, NW of Prince of

Wales Island, 10° 31.02' S, 141° 42.30' E, 17 January 2004;

CSIRO H 6558-06 (2, 67-69) Torres Strait, NW of Prince of

Wales Island, 10° 31.02' S, 141° 42.30' E, 17 January 2004;

CSIRO H 6898-08 (54.1) Torres Strait, Prince of Wales Island,

10° 55.24' S, 141° 49.73' E-10° 55.23' S, 141° 49.15' E, 16 m, 14

January 2004; CSIRO H 6900-03 (62) Torres Strait, NE of Banks

Island, 9° 52.30' S, 142° 35.43' E, 21 January 2004; QMB I. 82*

(~80, holotype of Torresia lineatea) Cardwell, 18° 16' S, 146°

1' E; QMB 1.5877+ (?) Whitsunday Group, Shaw Island, 20°

29' S, 149° 5' E; QMB 1.6108 (1) Townsville; QMB 1.6110 (1)

Dunk Island, 17° 57' S, 146° 9' E; QMB 1.6111 (1) Moreton Bay,

27° 15' S, 153° 15' E; QMB 1.6112 (1) Moreton Bay, 27° 15' S,

153° 15' E; QMB 1.6330 (1) Torres Strait, Yorke Islands, 9° 44' S,

143° 25' E; QMB 1.6852+ (?) Cape Cleveland, 19° 11' S, 147°

1' E; QMB 1.7494+ (?)Magnetic Island, Cockle Bay, 19° IT S,

146° 49' E; QMB 1.9909 (1) Queensland coast; QMB 1.9910+ (?)

Queensland coast; QMB 1.9911 (1) Queensland coast; QMB

1.9912 (1) Queensland coast; QMB 1.9913 (1) Queensland coast;

QMB 1.11824+ (?) Moreton Bay, Myora Bight, 27° 28' S, 153°

25' E; QMB 1.12212+ (?) Moreton Bay, Myora Bight, 27° 28' S,

153° 25' E; QMB 1.12554+ (?) Moreton Bay, 27° 15' S, 153°

15' E, 23 December 1952; QMB 1.15846+ (?) 2 miles NW of

Nymph Island, 14° 36' S, 145° 14' E, 15 m, 8 Eebruary 1979;

QMB 1.17553+ (?) Thursday Island Harbour, 10° 35' S, 142°

13' E, 20 March 1974; QMB 1.17554+ (?) Torres Strait, Aureed

Island, 9° 57' S, 143° 17' E, 17 October 1974; QMB 1.17555+ (?)

Torres Strait, 9° 24' S, 142° 51' E, 7.3 m, 23 March 1974; QMB

Review of Choerodon tuskfishes

107

l. 17556' (?) Torres Strait, 9° 40' S, 142° 33' E, 7.3-13.7 m, 28

March 1974; QMB 1.17557' (?) Torres Strait, 10° 21' S, 141°

44' E, 11.9 m, 3 November 1974; QMB 1.21140' (?) Weipa

Channel, 12° 37' S, 141° 52' E, 17 May 1984; QMB 1.22733' (?)

Cullen Island, SE of Sarina Beach, 21° 25' S, 149° 30' E, 9 April

1987; QMB 1.22784' (?) Sarina Inlet, 21° 24' S, 149° 19' E, 0.5

m, 10 April 1987; QMB 1.26697' (?) Torres Strait, Warrior Reefs,

9° 48' S, 142° 58' E, 28 March 1990; QMB 1.32523' (?) Pompey

Group, Renes Nook, on T line, 21° 17' S, 151° 31' E, 25 m, 13

March 2000; QMB 1.33768' (?) ~300 m S of centre of Southern

end of Sweers Island, 17° 9' S, 139° 36' E, 2-3.5 m, 15 November

2002; QMB 1.34433' (?) rocky reef ca 400 m off SW tip of

Sweers Island, 17° 8' S, 139° 36' E, 0.5 m-2 m, 16 November

2002; QMB 1.34556' (?) rocky reef ca 800 m off NE side of

Sweers Island, 17° 5' S, 139° 39' E, 0.5-3 m, 18 November 2002;

QMB 1.34628' (?) 150 m off beach at centre of E side of Sweers

Island, 17° 7' S, 139° 37' E, 0.3-0.5 m, 23 November 2002; QMB

1.34674' (?) SE of Sweers Island, ca 800 m E of Locust Rock, 17°

10' S, 139° 38' E, 0.1-2 m, 24 November 2002; QMB 1.34707'

(?) E side ofSweers Island, N of Observation Hill, 17° 8' S, 139°

37' E, 0.1-1 m, 19 November 2002; QMB 1.35843' (?) 21.43.5' S

150.13.5' E-21° 43' S, 150° 13' E, 32 m, 3 December 2003;

USNM 176733* (248) Brisbane. Northern Territory: AMS

1.24678-032* (2, 99.8-116) Darwin, East Point, 12° 25' S, 130°

49' E, 1 m, 31 August 1984; AMS 1A.1461* (181) Gulf of

Carpentaria, Pellew Group, 15° 33' S, 136° 59' E, June 1923;

AMS 1A.1462* (135) Gulf of Carpentaria, Pellew Group, 15°

33' S, 136° 59' E, June 1923; AMS 1A.4388 (78.8) Port Darwin,

12° 27' S, 130° 48' E, 1930; CSIRO CA 2378 (358) Chasm

Island, 13° 48' S, 136° 35' E, 26 July 1981; NMV A31441-001

(365) Gulf of Carpentaria, Groote Eylandt, S of North Point

Island, 13° 37' S, 136° 41' E, 3-5 m, January 1978; NTM

S.10772-003' (105) Darwin, 1978; NTM S.10432-003' (200)

Port Essington, Table Head, 11° 13' 01" S, 132° 10' 01" E, 3

July 1982; NTM S. 16134-027' (138) Bynoe Harbour, Knife

Island, 12° 42' 04" S, 130° 35' 31" E, 7 m, 20 April 2002; NTM

S.11296-006' (192) East Bremer Islet, 12° 04' 59" S, 136°

52' 01" E, 25 March 1975; NTM S.11408-005' (270) Gove, 21

August 1976; NTM S.11263-017' (2, 127-215) Cobourg

Peninsula, Coral Bay, 11° 12' 00" S, 132° 03' 00" E, 18 May

1983; NTM S.11191-002' (357) PeronIsland, 13° 10' 01" S, 130°

03' 00" E, 8 April 1983; NTM S.11260-035' (123) Cobourg

Peninsula, Coral Bay, 11° 10' 59" S, 132° 03' 00" E, 5 m, 16 May

1983; NTM S.16761-001' (218) Darwin Harbour, Nightcliff, 12°

22' 41" S, 130° 50' 13" E, 4 m, 11 January 1982; NTM S.16776-

001' (158) Darwin Harbour, Shoal Bay, 12° 10' 01" S, 130°

58' 59" E, 10 October 1977; NTM S. 10979-005' (46) Darwin

Harbour, Dudley Point Reef, 12° 12' 14" S, 130° 49' 01" E, 15

November 1982; NTM S. 16708-065' (6, 52-100) Darwin

Harbour, Vesteys Beach, 12° 26' 10" S, 130° 49' 44" E, 0.3 m, 14

November 2008; NTM S.16989-004' (180) Timor Sea, Bathurst

Island, 20° 42' 00" S, 129° 58' 59" E, 30 m, 8 July 1980; USNM

174392* (180) Yirrkala; USNM 174396* (193) Yirrkala; USNM

174402* (300) bay off Yirrkala; USNM 174407* (214) Groote

Eylandt. Western Australia: AMS 1.13270 (1) Shark Bay, 25°

20' S, 113° 35' E, 1914; CSIRO A 3388 (70.0) Exmouth Gulf,

Giralia Bay, 22° 26' S, 114° 21' E, 6 September 1966; CSIRO

CA982* (380) Nickol Bay, 20° 20.0' S, 116° 29.0' E-20° 20.0' S,

116° 26.0' E, 46 m, 31 May 1980; CSIRO H 2891-01' (205)

Monte Bello Islands, 20° 35.7' S, 115° 49.3' E-20° 35.0' S, 115°

50.3' E, 29 m, 16 September 1991; CSIRO H 4102-04' (1)

between Shark Bay & Exmouth Gulf, September 1995; SAM

E2985 (167, holotype of Choerodon rubidus) Point Samson, 23

July 1957; WAM P.21781-001 (370); WAM P.30320-005 (116)

Buccaneer Archipelago, Powerful Island, 16° 05' 00" S, 123°

27' 00" E, 0.1-1 m, 26 August 1991; WAM P.26278-002 (324)

16° 35' S, 121° 31' E, 28-34 m, 29 June 1978; WAM P.31391-

022 (21) Beagle Bay, 16° 59' S, 122° 40' E, 9-10 m, 28 August

1997; WAM P.28059-028 (93) 17° 59' S, 122° 11' E, 0-1 m, 28

March 1982; WAM P.31512-018 (106) Dampier Archipelago,

Collier Rocks, 20° 25' S, 116° 51' E, 0.1-0.4 m, 24 October 1998;

WAM P.24870-001 (3, 236-530) Monte Bello Islands, 20° 26' S,

115° 37' E, May 1973; WAM P.24668-001 (300) Dampier

Archipelago, Kendrew Island, 20° 29' S, 116° 32' E, 18 May

1974; WAM P.24774-001 (345) North West Cape, 21° 47' S,

114° 10' E, June 1974; WAM P.4597-001 (265) Houtman

Abrolhos, Wallabi Island, 28° 30' S, 113° 50' E.

Choerodon sugillatum. (114 specimens examined, 35.3-149

mm SL). Australia, Queensland: AMS 1.15557-206* (116,

paratype of Choerodon sugillatum) Gulf of Carpentaria, 17°

25' S, 140° 10' E, 10.1 m, 27 November 1963; AMS 1.19450-

037* (3, 36.2-65.7, paratypes of Choerodon sugillatum) Lizard

Island, 14° 40' S, 145° 27' E, 15 m, 10 November 1975; AMS

1.20751- 006* (8, 59.2-97.8, paratypes of Choerodon sugillatum)

Lizard Island, 14° 38' S, 145° 24' E, 25 m, 8 Eebruary 1979; AMS

1.20751- 044 (5, 77-92) Lizard Island, 14° 38' S, 145° 24' E, 25

m, 8 Eebruary 1979; AMS 1.20771-066* (2, 87.4—104, paratypes

of Choerodon sugillatum) Cape York, Captain Billy Creek, 11°

37' S, 142° 56' E, 16-18 m, 18 Eebruary 1979; AMS 1.20827-007

(81) Cape York, Hannibal Island, 11° 33' S, 142° 57' E, 22-23 m,

15 Eebruary 1979; CSIRO C 3462 (114) Gulf of Carpentaria, 16°

32' S, 140° 15' E, 12 September 1963; CSIRO CA 2165 (64.7)

Gulf of Carpentaria, Port Musgrave, 11° 18' S, 141° 41' E-ll°

18' S, 141° 40' E, 18 m, 8 June 1981; CSIRO H 3303-01 (125) off

Shelburne Bay, near Bird Island, 11° 48.7' S, 143° 12.7' E-ll°

47.3' S, 143° 11.1' E, 31 m, January 1993; CSIRO H 3442-06

(59.2) E of Cape York Peninsula, 11° 34.2' S, 143° 30.6' E, 40 m,

30 May 1993; CSIRO H 3895-10 (95) Cape Grenville, 11° 37' S,

143° 28' E, 18 m, 21 October 1994; CSIRO H 3897-04 (135)

Great Barrier Reef, Pollard Channel, 11° 51.51' S, 143° 27.83' E,

50 m, 17 October 1994; CSIRO H 3897-05 (135) Great Barrier

Reef, Pollard Channel, 11° 51.51' S, 143° 27.83' E, 50 m, 17

October 1994; CSIRO H 6151-03 (72.2) Torres Strait, Cape

York, 10° 36.82' S, 143° 08.54' E-10° 36.53' S, 143° 09.04' E, 26

m, 9 January 2004; CSIRO H 6519-03 (6,52.2-96.1) Cooktown,

15° 01.91' S, 145° 29.40' E-15° 02.24' S, 145° 28.91' E, 39 m, 20

November 2003; CSIRO H 7017-02^ (96) Gulf of Carpentaria,

Momington Island, 16° 12.92' S, 138° 58.95' E-16° 13.05' S,

139° 00.63' E, 27-28 m, 22 Eebruary 2009; NMV A13303 (127)

13 km E of Hannibal Island, 11° 38' 30.00" S, 143° 02' 23.99" E,

22 m, 15 January 1993; NMV A13304 (116) 12 km ESE of

Hannibal Island, 11° 37' 54.00" S, 143° 01' 41.99" E, 22 m, 14

January 1993; NMV A13305 (107) 11 km NNE of Cape

Grenville, 11° 53' 05.99" S, 143° 16' 12.00" E, 39 m, 13 January

1993; NMV A13306 (2, 115-129) Gulf of Carpentaria, 114 km

W of Prince of Wales Island, 10° 50' 35.99" S, 141° 12' 42.00" E,

27 m, 3 Eebruary 1993; NMV A13307 (98.8) 18 km W of

Saunders Island, 11° 41' 42.00" S, 143° 00' 53.99" E, 18-17 m,

14 January 1993; NMV A2952 (120) Gulf of Carpentaria, 90 km

NW of Cullen Point, 11° 19' 12.00" S, 141° 28' 11.99" E,30m,9

September 1982; NMV A2954 (95) Gulf of Carpentaria, 95 km

108

M.F. Gomon

NW of Prince of Wales Island, 10° IT S, 141° 15' E, 23 m, 11

September 1982; NMV A2960 (104) Gulf of Carpentaria, 120

km W of Crab Island, 10° 57' S, 141° 02' 41.99" E, 43-44 m, 10

September 1982; NMV A2968 (119) Gulf of Carpentaria, 140

km W of Prince of Wales Island, 10° 39' S, 140° 45' E, 46 m, 13

September 1982; NMV A3126* (104, holotype of Choerodon

sugillatum) Cape Bedford, 15° 13.59' S, 145° 23.36' E-15° 00' S,

145° 27.87' E, 30 m, 1 Mareh 1983; NMVA3130* (3,81.2-99.2,

paratypes of Choerodon sugUlatum) Gulf of Carpentaria, Cape

York, 10° 50.23' S, 140° 55.11' E-10° 48.01' S, 140° 55.68' E, 44

m, 26 Eebruary 1983; NMV A3137* (97.2, paratype of

Choerodon sugUlatum) Cape Tribulation, 16° 09.46' S, 145°

33.23' E-16° 12.17' S, 145° 36.06' E, 30 m, 1 March 1983; NMV

A3173 (4,100-115) Great Barrier Reef, near Haggerstone Island,

12° 03' 16.92" S, 143° 15' 10.08" E, 26-29 m, 27 Eebruary 1983;

NMV A3178 (120) Gulf of Carpentaria, 155 km W of Cape York,

10° 49' 43.20" S, 141° 08' 32.39" E, 32 m, 26 Eebruary 1983;

NTM S.13273-016^ (4, 49-76) Gulf of Carpentaria, Booby

Island, 10° 43' 59" S, 141° 52' 59" E, 10 m, 29 November 1991;

NTM S.13277-008+ (3, 88-111) Cape York, 11° 21' 00" S, 142°

58' 01" E, 22 m, 1 Deeember 1991; NTM S. 13278-011+ (2,100-

123) Cape York, 11° 22' 01" S, 142° 55' 01" E, 21 m, 1 Deeember

1991; QMB 1.10740+ (?) North Palm Island, 18° 33' S, 146°

29' E; QMB 1.12398+ (?) Cairns, 16° 50' S, 146° E, 27 November

1957; QMB 1.15602+ (?) Eel Reef, 12° 31' S, 143° 27' E, 30-32

m, 20 Eebruary 1979; QMB 1.15621+ (?) Eel Reef, 12° 22' S,

143° 20' E, 22 m, 20 Eebruary 1979; QMB 1.15626+ (?) Nymph

Island, 14° 36' S, 145° 14' E, 15 m, 8 Eebruary 1979; QMB

l. 15674+ (?) Lizard Island, 15° 30' S, 145° 22' E, 20 m, 8 Eebruary

1979; QMB 1.15698+ (?) Nymph Island, 14° 33' S, 145° 6' E, 16

m, 8 Eebruary 1979; QMB 1.16164+ (?) Hannibal Island, 11°

33' S, 142° 57' E, 21.9 m, 15 Eebruary 1979; QMB 1.16214+ (?)

Lizard Island, 14° 38' S, 145° 24' E, 25.6 m, 8 Eebruary 1979;

QMB 1.17566+ (?) Torres Strait, Aureed Island, 9° 57' S, 143°

17' E, 27 November 1974; QMB 1.17567^ (?) Torres Strait, P D

Sand Cay, 10° 11' S, 143° 14' E, 18 Oetober 1974; QMB 1.17568+

(?)Torres Strait, Aeroplane Sandbank, 10° 20' S, 143° 15' E,

18.3-25.6 m, 19 Oetober 1974; QMB 1.17569+ (?) Torres Strait,

Caldbeek Reef, 10° 10' S, 143° 13' E, 5 Deeember 1974; QMB

l. 17570+ (?) Torres Strait, 9° 44' S, 142° 51' E, 31 Mareh 1974;

QMB 1.17571+ (?) Torres Strait, 9° 49' S, 142° 48' E, 12.8-17.4

m, 4 April 1974; QMB 1.17572+ (?) Torres Strait, 9° 53' S, 142°

50' E, 6 April 1974; QMB 1.17573+ (?) Torres Strait, 9° 58' S,

142° 41' E, 14.6-16.5 m, 8 April 1974; QMB 1.17574+ (?) Torres

Strait, 10° 4' S, 142° 43' E, 25 April 1974; QMB 1.18098+ (?)

Prineess Charlotte Bay, 14° 5' S, 144° 4' E, 25.6 m, 19 September

1979; QMB 1.18212+ (?) Lizard Island, 14° 20' S, 145° 51' E,

23.8 m, 18 September 1979; QMB 1.18502+ (?) Green Island, 16°

46' S, 145° 58' E, 40 m, 9 Oetober 1980; QMB 1.19608+ (?) Elora

Passage, 17° 1' S, 146° 20' E, 49.4 m, 9 Oetober 1979; QMB

1.19655+ (?) Elora Passage, 17° 8' S, 146° 15' E, 43.9 m, 16

Oetober 1979; QMB 1.19690+ (?) Palm Islands, 18° 35' S, 146°

49' E, 45.7 m, 16 Oetober 1979; QMB 1.20322+ (?) Sudbury

Reef, 17° 3' S, 146° 7' E, 33-36 m, 3 June 1905; QMB 1.20474+

(?) Cairns, 17° S, 146° 3' E, 30 m, 25 April 1982; QMB 1.20494+

(?) Cairns, 17° S, 146° E, 25 April 1982; QMB 1.23061+ (?)

Swain Reefs, 21° S, 151° E, 62 m, 17 September 1986; QMB

1.23517+ (?) Cape Bowling Green, 19° S, 147° 25' E, 41 m, 8

May 1985; QMB 1.26151+ (?) Caprieom Group, 23° 30' S, 152°

E, 36.5-54.8 m, 2 August 1957; QMB 1.26226+ (?) Cairns, 16°

50' S, 146° E, 27 November 1957; QMB 1.27727+ (?) Gulf of

Carpentaria, 10° 30' S, 141° 9' E, 23 m, 3 December 1990; QMB

l. 27964+ (?) Gulf of Carpentaria, 15° 57' S, 138° 41' E, 25 m, 11

Deeember 1990; QMB 1.30052+ (?) Gulf of Carpentaria, 15°

32' S, 139° 42' E, 45 m, 7 Deeember 1990; QMB 1.34382+ (?)

MeCulloeh Reef, 17° 16' S, 146° 25' E, 28 September 2003;

QMB 1.34854+ (?) Gulf of Carpentaria, 12° 56' S, 141° 9' E, 36

m, 26 November 1991; QMB 1.34860+ (?) Gulf of Carpentaria,

11° 19' S, 140° 56' E, 41 m, 27 November 1991; QMB 1.35033+

(?) 17° 57.3' S, 146° 25.5' E, 27 September 2003; QMB 1.35146+

(?) 14° 42.3' S, 145° 26.1' E, 5 Oetober 2003; QMB 1.35332+ (?)

19° 5.1' S, 147° 23.7' E, 21 September 2003; QMB 1.35375+ (?)

19° 2.7' S, 147° 23.7' E, 21 September 2003; QMB 1.35544+ (?)

18° 38.1' S, 147° 16.5' E, 18 September 2004; QMB 1.35611+ (?)

18° 58.5' S, 146° 56.1' E, 18 September 2003; QMB 1.36216+ (?)

17° 29.7' S, 146° 15.9' E, 29 m, 28 April 2004; QMB 1.36287+ (?)

20° 20.7' S, 150° 23.1' E, 20 m, 11 May 2004; QMB 1.36943+ (?)

Cape York, E of Eurze Point, 11° 4' S, 142° 52' E, 21 m, 30

November 1991; QMB 1.37421+ (?) 13° 43.5' S, 144° 7.5' E, 35

m, 18 January 2005; USNM 280628* (4, 64.2-84.0) same

eolleetion data as AMS 1.20751-006; Northern Territory: NTM

S.12942-007+ (136) Arafura Sea, 11° 22' 01" S, 134° 07' 01" E,

22 Oetober 1990; NTM S. 12933-002+ (2, 76-83) Arafura Sea,

10° 24' 00" S, 129° 34' 01" E, 91 m, 17 November 1990; Western

Australia: AMS 1.22802-008 (4, 10^123) North West Shelf,

Port Hedland, 19° 39' S, 117° 53' E, 52-53 m, 26 March 1982;

CSIRO B 4020 (45.2) North West Shelf, Port Hedland, 19°

54.9' S, 118° 00.8' E-19° 54.7' S, 118° 00.5' E, 37-38 m, 5

Deeember 1982; CSIRO CA 2164 (85.8) Niekol Bay, 19° 50.0' S,

116° 28.0' E-19° 52.0' S, 116° 27.0' E, 68-70 m, 4 Deeember

1979; CSIRO CA2166 (118) Joseph Bonaparte Gulf, 11° 01.0' S,

128° 37.0' E-ll° 01.0' S, 128° 35.0' E, 30 m, 10 July 1980;

CSIRO CA 2167 (99.5) Monte Bello Islands, 20° 04.0' S, 116°

18.0' E-20° 02.0' S, 116° 20.0' E, 56-58 m, 31 May 1980; CSIRO

CA2168 (90.3) Ashmore Reef, NW of Admiralty Gulf, 12° 27' S,

124° 27' E-12° 27 S, 124° 25' E, 76-78 m, 16 July 1980; CSIRO

CA 2323 (67.6) Admiralty Gulf, 13° 23' S, 124° 48' E-13° 25' S,

124° 51' E, 120 m, 29 Mareh 1981; CSIRO CA 2324 (86.7)

Admiralty Gulf, 13° 23' S, 124° 48' E-13° 25' S, 124° 51' E, 120

m, 29 Mareh 1981; CSIRO CA2325 (92.9) Admiralty Gulf, 13°

23' S, 124° 48' E-13° 25' S, 124° 51' E, 120 m, 29 Mareh 1981;

CSIRO CA 3058 (126) Port Hedland, 20° 02.6' S, 117°

53.6' E-20° 01.2' S, 117° 53.6' E, 34-36 m, 17 Oetober 1982;

CSIRO H 648-01 (96.1) Monte Bello Islands, 20° 00.0' S, 115°

58.0' E-20° 03.0' S, 115° 57.0' E, 78-80 m, 2 Deeember 1979;

CSIRO H 656-01* (141, paratype of Choerodon sugillatum)

Dampier Arehipelago, Cape Preston, 20° 48.0' S, 116°

00.0' E-20° 46.0' S, 115° 59.0' E, 19-22 m, 1 Deeember 1979;

CSIRO H 657-01* (35.3, paratype of Choerodon sugillatum)

Monte Bello Islands, 20° 00.0' S, 115° 58.0' E-20° 03.0' S, 115°

57.0' E, 78-80 m, 2 Deeember 1979; CSIRO H 657-02* (50.6,

paratype of Choerodon sugillatum) Monte Bello Islands, 20°

00.0' S, 115° 58.0' E-20° 03.0' S, 115° 57.0' E, 78-80 m, 2

Deeember 1979; CSIRO H 658-01* (96.1, paratype of Choerodon

sugillatum) Dampier Arehipelago, 19° 47' S, 116° 33' E-19°

45' S, 116° 35' E, 6^60 m, 4 December 1979; CSIRO H 2188-02

(9, 37-113) North West Shelf, Port Hedland, 19° 29.7' S, 118°

52.0' E-19° 29.4' S, 118° 52.5' E, 39 m, 25 Oetober 1983; CSIRO

H 4351-01 (2, 75-110) Port Hedland, 19° 58.5' S, 117°

49.0' E-19° 58.1' S, 117° 49.5' E, 42 m, 26 June 1983; NMV

A1876* (3, 12^133) North West Shelf, NW of Dampier, 20°

10' 12.00" S, 116° 04' 11.99" E, 60 m, 9 Mareh 1981; NMV

Review of Choerodon tuskfishes

109

A1926 (14, 75-125) North West Shelf, NW of Dampier, 20°

07' 12.00" S, 116° 12' E, 60 m, 9 March 1981; NMVA1938 (140)

North West Shelf, NW of Dampier, 20° 10' 47.99" S, 116° 09' E,

57-58 m, 9 March 1981; NMV A1939 (3, 100-110) North West

Shelf, NNE of Cape Lambert, 19° 31' 11.99" S, 117° 17' 59.99" E,

73 m, 10 March 1981; NTM S.13974-010' (130) Dampier

Archipelago, 20° 13' 01" S, 116° 17' 60" E, 11 May 1983; NTM

S.14363-014^ (2,60-69) Holothuria Banks, Bassett Smith Shoal,

13° 13' 08" S, 125° 14' 53" E, 63 m, 12 June 1996; NTM S. 11672-

003' (112) North West Shelf, Rowley Shoals, 19° 01' 59" S, 118°

30' 00" E, 86 m, 2 June 1985; NTM S.11580-010^ (103) Dampier

Archipelago, 20° 03' 00" S, 115° 47' 60" E, 53 m, 8 May 1983;

NTM S.11554-019' (109) Dampier Archipelago, 20° 03' 00" S,

115° 47' 60" E, 7 May 1982; NTM S.11673-00E (84) North West

Shelf, Rowley Shoals, 19° 12' 00" S, 118° 40' 59" E, 80 m, 1 June

1985; NTM S. 11690-007+ (2, 76-116) North West Shelf, King

Sound, 16° 33' 00" S, 121° 28' 59" E, 46 m, 17 April 1985; NTM

S. 11582-008+ (99) 19° 48' 00" S, 116° 34' 59" E, 73 m, 19 May

1983; NTM S.16718-005+ (126) North West Shelf, Bedout

Island, 19° 04' 01" S, 119° 27' 00" E, 75 m, 25 May 1983; WAM

R25354-023* (6, 133-145, paratypes of Choerodon sugillatum)

Monte Bello Islands, 20° 25' S, 115° 30' E, April 1975.

Choerodon typus. (45 specimens examined, 38.6-123 mm SL).

INDIAN OCEAN, India: BPBM 20506 (3, 75-100) Madras;

BPBM 20654 (77.1) Madras-Tuticorin. Andaman Islands:

USNM 218485* (2, 78.2-79.5) 12° 03' N, 92° 57' E, 37 m.

Thailand: CAS 204373 (1) Gulf of Thailand, SE of Goh Chuang,

12° IT N, 100° 35' E, 25 m, 1^19 December 1960; CAS 204380

(2) Gulf of Thailand, SSE of Goh Chuang, 12° 06' N, 101° IT E,

12 m, 28 December-2 January 1961; NSMT-P 55232* (93.9)

Gulf of Thailand, Songkhla, 18 November 1986; USNM

218491* (11, 65.8-94.2) 09° 13' N, 97° 51' E, 58 m. PACIFIC

OCEAN, China: USNM 219452* (38.6) South China Sea,

Macclesfield Bank, 15° 49.4' N, 114° 31.8' E-15° 58.8' N, 114°

33.8' E. Taiwan: THUP 03367 (73.0) Koahsiung. Palau: CAS

204374 (1) Tapraki Reef, 7° 16' 7" N, 134° 27' 31" E, 0-3 m, 15

April 1959. Vietnam: MNHN 1963-604* (82.4); MNHN 1964-

632* (2, 104—110) South Vietnam. Singapore: BMNH

1884.8.14.10 (97.3) Dobson; CAS 29237 (1) 1.358430°

N, 103.803433° E. Philippines: USNM 153279* (51.6) Palawan,

Cuvayan Island; USNM 153280* (61.7) Corregidor Light;

USNM 153281* (59.6) Palawan, Observatory Island; USNM

153282* (62.8) Luzon, Sueste Point; USNM 347101 (115) Iloilo,

Panay Island; USNM 260873* (2, 95.4-96.5) Visayan Sea

between northern Negros and Masbate, 11° 35' 45" N, 123°

55' 32" E, 78.7 m, 6 June 1978; USNM 260875* (123) Visayan

Sea between northern Negros and Masbate, 11° 38' 20" N, 123°

58' 38" E, 80.5 m, 5 June 1978; USNM 347101* (2, 123-123)

Iloilo Strait between Penay and Guimaris Islands; USNM

408907* (104) Luzon, Sorsogon, Bulan. Indonesia: BMNH

1864.5.15.35* (102, holotype of Xiphocheilos typus) Nias;

BMNH 1879.5.14.34* (87.4, holotype of Xiphochilus

quadrimaculatus) Arafura StSL', USNM 153979* (59.0) Sulawesi,

Makasser Island. Australia, Queensland: CSIRO H 1319-ft2

(95.5) 16.5347° S, 146.0336° E, 54.2 m, 22 November 2003;

CSIRO H 6647-03 (93.3) Northern Great Barrier Reef, N of

Princess Charlotte Bay, 13.57° S, 144.03° E, 40 m, 5 October

2004; NMV A2949 (110) Gulf of Carpentaria, 105 km WNW of

Cullen Point, 11° 25' 11.99" S, 141° 01' 48.00" E, 45 m, 8

September 1982; NMV A13302 (105) Gulf of Carpentaria, 11°

55' 05.99" S, 139° 56' 41.99" E, 59 m, 6 Eebruary 1993.

Choerodon venustus. (56 specimens examined, 48-384 mm

SL). Australia: ANSP 82300 (1); USNM 176703* (304) Great

Barrier Reef area near Brisbane, 8-29 May 1952. New South

Wales: AMS 1.25663-008 (89) Yamba, 29° 31' S, 153° 28' E-29°

34' S, 153° 25' E, 49-53 m, 24 March 1985; AMS 1.25663-019*

(52) same collection data as AMS 1.25663-008; AMS 1.25665-

010* (2, 64.2-73.4) Yamba, 29° 21' S, 153° 29' E, 49-51 m, 21

March 1985; AMS 1.25665-034 (2, 63-71) same collection data

as AMS 1.25665-010; AMS 1.26022-002* (85) off Woody Head,

29° 21' S, 153° 27' E-29° 21' S, 153° 31' E, 4^53, 23 March

1985; AMS 1.26534-010 (2, 48-57) Iluka, 29° 22' S, 153°

27' E-29° 14' S, 153° 32' E, 46-57 m, 23 May 1986; AMS

1.27659-003* (78.4) Yamba, 29° 16' S, 153° 29' E-29° 16' S,

153° 32' E, 43-53 m, 23 March 1985; AMS IB.2616 (1)

Newcastle, 32° 56' S, 151° 46' E, 1951; NMV A22004 (30.7)

Port Jackson, 33° 51' 00.00" S, 151° 16' 12.00" E, 20 m, 8 May

1982. Queensland: AMS E.1472 (1) Double Island Point, 26°

25' S, 153° 30' E, 60 m, 28 June 1910; AMS E.1473 (144, syntype

of Choerodon ambiguus) same collection data as AMS E.1472;

AMS E.1474 (1) same collection data as AMS E.1472; AMS

E.1475 (1) same collection data as AMS E.1472; AMS E.1476

(198) same collection data as AMS E.1472; AMS E.1528+ (133)

Wide Bay, 25° 52' S, 153° 07' E, 1909; AMS E.1529* (154) same

collection data as AMS E.1528; AMS E.2873^* (10, 95.8-133)

Double Island Point, 26° 08' S, 153° 25' E, 60 m, 2 December

1910; AMS 1.10935+ (172) Double Island Point, 26° 00' S, 153°

00' E, 28 June 1910; AMS 1.10936 (225) same collection data as

AMS 1.10935; AMS 1.10991 (155) Wide Bay, 25° 52' S, 153°

07' E, 1910; AMS 1.11007 (384) Double Island, 26° 00' S, 153°

00' E, 53 m, 29 June 1910; AMS 1.12535* (134, syntype of

Choerodon ambiguus) Double Island Point, 25° 56' S, 153° 11' E,

1909; AMS 1.15625-035 (190) One Tree Island, 23° 30' S, 152°

05' E, 32 m, 8 December 1966; AMS 1.15637-040 (222) One

Tree Island, 23° 30' S, 152° 05' E, 23-26 m, 5 October 1967;

AMS 1.15684-038* (213) One Tree Island, 23° 30' S, 152° 05' E,

32-29 m, 1 December 1969; AMS 1.22002-002 (240) One Tree

Island, 23° 30' S, 152° 05' E, 47-67 m, 8 December 1966; AMS

1.24089-002* (219) Heron Island, 23° 27' S, 151° 55' E, 3

December 1977; AMS IA.2072 (335) Great Barrier Reef, Great

Palm Island, 18° 45' S, 146° 37' E, July 1924; BPBM 14507 (70)

One Tree Island, 19 January 1973; BPBM 15972 (241) Heron

Island, 30 June 1973; BPBM 14507* (66.3) Heron Island; BPBM

15972 (2, 197-241) Heron Island; CSIRO H 4307-13* (355) 31

July 1996; CSIRO H 6913-01+ (240) Gladstone, 23° 22.29' S,

151° 58.34' E, 23° 22.51' S, 151° 58.86' E, 44 m, 22 April 2004;

CSIRO H 6697-03* (138) 19.2638° S, 148.141° E, 59.3 m, 12

December 2003; CSIRO H 6748-05* (132) SE of Bunker Group,

23° 58.71' S, 152° 37.79' E, 54 m, 15 April 2004; CSIROH 6152-

03* (4, 53.3-192) 21.5096° S, 151.4815° E, 60 m, 7 December

2003; MCZ 36810 (245) Gladstone; QMB 1.4735* (234,

holotype of Choerops venustus) Moreton Bay; 1.4736 (1)

Moreton Bay; QMB 1.9457 (1) Heron Island; QMB 1.11411 (1)

Heron Island; QMB 1.535+ (?) Elat Top; QMB 1.673+ (?) Moreton

Bay, 27° 15' S, 153° 15' E; QMB 1.703+ (?) Moreton Bay, 27°

15' S, 153° 15' E; QMB 1.718+ (?) Cape Moreton, 27° 2' S, 153°

28' E; QMB 1.750+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB

1.751+ (?) Masthead Island, Capricorn Group, 23° 32' S, 151°

44' E; QMB 1.1543+ (?) Double Island Point, 25° 56' S, 153°

ir E, 60.3 m; QMB 1.2114+ (?) Moreton Bay, 27° 15' S, 153°

110

M.F. Gomon

15' E; QMB 1.2984^ (?) Moreton Bay, 27° 15' S, 153° 15' E;

QMB 1.3068+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB

1.3069+ (?) Moreton Bay, 27° 15' S, 153° 15' E; QMB 1.3201+ (?)

Moreton Bay, Myora, 27° 28' S, 153° 25' E; QMB 1.4735+ (?)

Moreton Bay, 27° 15' S, 153° 15' E; QMB 1.4736+ (?)Moreton

Bay, Snapper Banks; QMB 1.5404+ (?) Kelso Reef, 18° 24' S,

147° E; QMB 1.5415+ (?) Townsville, Lodestone Reef, 18° 41' S,

147° 5' E; QMB 1.5529+ (?) Heron Island, 23° 27' S, 151° 55' E;

QMB 1.5536+ (?) Heron Island, 23° 27' S, 151° 55' E; QMB

1.5537+ (?); QMB 1.5544+ (?) off Townsville, Lodestone Reef,

18° 41' S, 147° 5' E; QMB 1.6096+ (?) Palm Island, 18° 40' S,

146° 33' E; QMB 1.6607+ (?)Eraser Island, N Gardiner Bank, 25°

S, 153° 25' E; QMB 1.6992+ (?) Keeper Reef, 18° 45' S, 147°

16' E; QMB 1.9457+ (?) Heron Island, 23° 27' S, 151° 55' E; QMB

1.11411+ (?) Heron Island, 23° 27' S, 151° 55' E; QMB 1.13771+

(?) Noosa Heads, 26° 23' S, 153° 7' E, 1 January 1977; QMB

1.16612+ (?) Mooloolaba, 26° 41' S, 153° 16' E, 21.4^36.6 m, 2

July 1979; QMB 1.16613+ (?) Mooloolaba, 26° 41' S, 153° 16' E,

21.4r-36.6 m, 2 July 1979; QMB 1.19312+ (?) Brisbane, 5

Eebruary 1982; QMB 1.21311+ (?) Caprieom Group, 23° 30' S,

152° E, 29 July 1980; QMB 1.21331+ (?) Masthead Island, 23°

30' S, 151° 44' E, 2 Mareh 1982; QMB 1.21385+ (?) Brisbane, 1

November 1984; QMB 1.23045+ (?) Swain Reefs, 21° 16' S, 151°

10' E, 33 m, 14 September 1986; QMB 1.23046+ (?) Swain Reefs,

21° 13' S, 150° 43' E, 47 m, 16 September 1986; QMB 1.25362+

(?) Brisbane, 20 July 1988; QMB 1.26511+ (?) Brisbane, 1

January 1990; QMB 1.32731+ (?) Llewellyn Reef, 23° 45' S, 152°

9' E, 47 m, 3 Oetober 2000; QMB 1.34083+ (?) Caprieom Group,

23° 2' S, 151° 26' E, 40 m, 3 Oetober 2000; QMB 1.36123+ (?)

21° 30.9' S, 151° 28.5' E, 60 m, 25 May 2004; QMB 1.37869+ (?)

19° 51.9' S, 149° 10.5' E, 60 m, 27 November 2005; QMB

1.40027+ (?) 23° 2.7' S, 151° 5.1' E, 30 m, 3 November 2005.

Choerodon vitta. (40 speeimens examined, 29.5-174 mm SL).

Indonesia: AMS 1.12534* (141, paraleetotype of Choerodon

vitta) Am Islands, Dobo, 1912; QMB 1.1555* (148, leetotype of

Choerodon vitta) Am Islands, Dobo. Australia, Queensland:

AMS 1.15557-201* (3, 101-144) Gulf of Carpentaria, 17° 25' S,

140° 10' E, 10.1 m, 27 November 1963; AMS 1.15557-202*

(82.1) Gulf of Carpentaria, 17° 25' S, 140° 10' E, 10.1 m, 27

November 1963; AMS 1.18767-009 (4, 29.5-47.1) Great Barrier

Reef, Linnet Reef, 14° 47' S, 145° 20' E, 6-15 m, 22 November

1975; AMS 1.19461-034* (2, 63.9-68.7) Decapolis Reef, 14°

51' S, 145° 17'E, 2^m, 14 November 1975; AMS 1.20825-002*

(174) Deeapolis Reef, 14° 51' S, 145° 16' E, 3 Deeember 1978;

CSIRO C 4010 (147) Gulf of Carpentaria, 17° 06' S, 139° 44' E,

12 August 1963; CSIRO C 4481 (127) Gulf of Carpentaria,

Bentnick Island, 16° 55' S, 139° 37' E, 12.5 m, 1 November

1972; CSIRO H 6558-04 (107) Torres Strait, NW of Prinee of

Wales Island, 10° 31.02' S, 141° 42.30' E, 17 January 2004;

CSIRO H 6904-03* (120) Torres Strait, Neweastle Bay, 10°

28.80' S, 142° 52.44' E-10° 28.95' S, 142° 52.48' E, 19 m, 12

January 2004; CSIRO H 6904-05* (109) same eolleetion data as

CSIRO H 6904-03; CSIRO H 7996-01* (2, 97.1-127) Torres

Strait, Endeavour Strait, 10° 46.58' S, 142° 15.24' E, 16 m, 19

January 2004; CSIRO H 7897-01* (78.9) 10.1067° S, 142.9177°

E, 19.7 m, 12 January 2004; NMV A3181 (145) Gulf of

Carpentaria, 155 km W of Cape York, 10° 49' 43.20" S, 141°

08' 32.39" E, 32 m, 26 Eebruary 1983; NMV A 31435-001*

(103) N of Margaret Bay, 11° 54.34' S, 143° 16.54' E, 49 m, 30

September 2004; QMB 1.1331 (117) Southport, 27° 58' S, 153°

25' E; QMB I. 11331+ (?) N of Townsville, Paluma Shoals, 19°

6' S 146° 33' E; QMB I. 11375+ (?) Palm Island, 18° 40' S 146°

33' E; QMB I. 15241+ (?) Torres Strait, 9° 40' S 142° 33' E, 7.3-

8.2 m, 28 Mareh 1974; QMB I. 17558+ (?) Torres Strait, 10° 2' S

142° 37' E, 17 m, 21 April 1974; QMB I. 17559+ (?) Torres Strait,

10° 2' S 142° 44' E, 8.5 m, 20 April 1974; QMB I. 17560+ (?)

Torres Strait, 9° 52' S 142° 32' E, 13 m, 22 April 1974; QMB I.

17561+(?) Torres Strait, 10° 4'S 142° 31'E, 14 m, 25 April 1974;

QMB I. 17562+ (?) Torres Strait, 10° 7' S 142° 41' E, 26 April

1974; QMB I. 17563+ (?) Torres Strait, 10° 12' S 142° 39' E, 18

m, 2 May 1974; QMB I. 17564+ (?) Torres Strait, 9° 52' S 141°

33' E, 12.8-14.6 m, 30 May 1974; QMB I. 17565+ (?) Torres

Strait, 9° 56' S 141° 41' E, 11.9 m, 1 June 1974; QMB I. 18168+

(?) Eel Reef, 12° 26' S 143° 19' E, 18.3 m, 24 September 1979;

QMB I. 20102+ (?) Erankland Islands, 17° 11' S 146° 4' E, 6-10

m, 1 November 1982; QMB I. 23538+ (?) Cape Bowling Green,

19° 17' S 147° 21' E, 18 m, 19 Eebmary 1985; QMB 1.34896+ (?)

Gulf of Carpentaria, 10° 38' S 141° 27' E, 15 m, 29 November

1991; QMB I. 36202+ (?) 22° 3.3' S 150° 21.3' E, 28 m, 10 May

2004; QMB I. 38944+ (?) Gloueester Passage, 20° 3' 11" S 148°

25' 38" E, 9 m, 20 September 2011; Northern Territory: NTM

S. 11283-004+ (41) Darwin Harbour, Dudley Point, 12° 25' 01" S,

130° 48' 00" E, 23 May 1984; NTM S. 11274-009+ (79) English

Company Islands, Cotton Island, 11° 48' 00" S, 136° 30' 00" E,

11 m, 21 Eebmary 1984; NTM S.10696-010+ (5,58-150) Darwin

Harbour, East Point, 12° 24' 00" S, 130° 49' 01" E, 13 September

1982; NTM S.13257-003+ (157) Gulf of Carpentaria, Cape Grey,

13° 03' S, 136° 45' E, 22 m, 23 November 1991; NTM S.12961-

002+ (136) Arafura Sea, 11° 22' 59" S, 136° 05' 60" E, 34 m, 26

Oetober 1990; NTM S. 13311-005+ (3, 104-141) Timor Sea,

North Anson Bay, 12° 58' 01" S, 129° 54' 00" E, 18 m, 21

November 1990; NTM S.11252-001+ (158) Cobourg Peninsula,

Table Head, 11° 15' 00" S, 132° 10' 01" E, 7 m, 12 May 1983;

NTM S. 13236-020+ (20) Marehinbar Island, Sphinx Head Bay,

11° 16' 01" S, 136° 40' 59" E, 16 November 1990; NTM S.17174-

001+ (3, 68-87) Darwin Harbour, East Point Reef, 12° 24' 47" S,

130° 48' 36" E, 22 November 1982; NTM S.13436-002+ (15)

Elat Top Bank, 12° 16' 48" S, 129° 13' 59" E, 17 May 1992;

Western Australia: AMS 1.21630-002* (146) Timor Sea, 11°

02' S, 128° 48' E, 30 m, 26 June 1979; AMS 1.22802-007+ (142)

North West Shelf, Port Hedland, 19° 39' S, 117° 53' E, 52-53 m,

26 Mareh 1982; AMS 1.22806-010 (2, 132-135) North West

Shelf, Port Hedland, 19° 29' S, 118° 22' E, 54-60 m, 29 Mareh

1982; AMS 1.22831-028* (8, 95.2-172) North West Shelf, Port

Hedland, 20° 00' S, 117° 16' E, 50 m, 16 April 1982; CSIRO CA

233 (134) Dampier Arehipelago, 20° 35' S, 116° 14' E-20° 35' S,

116° 17' E, 32-34 m, 14 May 1978; CSIRO CA 1492 (124)

Roebuek Bay, 17° 50.0' S, 121° 46.0' E-17° 51.0' S, 121° 42.0' E,

52-58 m, 8 November 1980; CSIRO CA 2162 (152) Eorestier

Island, 19° 53.0' S, 118° 13.0' E-19° 52.0' S, 118° 12.0' E, 37-36

m, 3 June 1980; CSIRO CA 2163 (147) Eighty Mile Beaeh, 19°

27' S, 120° 21' E-19° 29' S, 120° 23' E, 35-31 m, 7 June 1980;

NMV A1874 (167) North West Shelf, NW of Dampier, 20°

10' 12.00" S, 116° 04' 11.99" E, 60 m, 9 Mareh 1981; NTM

S.10726-001+ (170) Cootamundra Shoals, 10° 49' 59" S, 129°

13' 01" E, 14 May 1982; NTM S.11688-007+ (144) North West

Shelf, King Sound, 16° 31' 59" S, 121° 25' 59" E, 42 m, 17 April

1985; NTM S. 11690-012+ (2, 12^159) North West Shelf, King

Sound, 16° 33' 00" S, 121° 28' 59" E, 46 m, 17 April 1985; NTM

S. 14362-014+ (80) Holothuria Banks, West Holothuria Reef, 13°

25' 12" S, 125° 39' 18" E, 50 m, 12 June 1996; NTM S.16718-

Review of Choerodon tuskfishes

111

008^ (172) North West Shelf, Bedout Island, 19° 04' 01" S, 119°

27' 00" E, 75 m, 25 May 1983; WAM P.31391-012^ (63) Beagle

Bay, 16° 59' S, 122° 40' E, 9-10 m, 28 September 1997; WAM

P.25397-019+ (6, 97-139) Rowley Shoals, 17° 29' S, 121° 52' E,

0^2 m, 21 December 1969; WAM P31894-00P (55) Dampier

Archipelago, Rosemary Island, Tish Point, 20° 29.94' S, 116°

38.11' E, 12-15 m, 26 My 1999; WAM P.31884-002+ (39)

Dampier Archipelago, Enderby Island, 20° 40.93' S, 116°

33.21' E, 9-9.2 m, 23 My 1999; WAM P.32707-00P (2,76-82)

Exmouth Gulf, Bundegi Reef, 21° 51.281' S, 114° 12.056' 0" E,

21.^21.2 m, 8 My 2004; WAM P.32667-003^ (120) Exmouth

Gulf, Bundegi Reef, 21° 51.815' S, 114° 11.902' 0" E, 22.4-22.6,

16 March 2004; WAM P.4550-00P (116) Exmouth Gulf, 22°

05' S, 114° 15' E, 18 September 1958; WAM P.4553-00P (114)

Exmouth Gulf, 22° 05' S, 114° 15' E, 5 October 1958; WAM

P.4583-00P (?) Exmouth Gulf, 22° 05' S, 114° 15' E, 5 October

1958; WAM P.5694-00P (?) Exmouth Gulf, 22° 05' S, 114°

15' E; WAM P.25095-050^ (126) Exmouth Gulf, 22° 05' S, 114°

15' E, 0-12 m, October 1974.

Choerodon zamboangae. (36 specimens examined, 95.6-249

mm SL). Japan, Okinawa: BPBM 19167 (222) Naha; SMBL-E

73324 (216) Naha, 100 m, 6 August 1973; SMBL-E 73325 (231)

same collection data as SMBL-E 73324. Taiwan: THUP 00956*

(213); THUP 02069* (164); THUP 02492* (210). Philippines:

AMS 1.31430.040^ (1) Dumaguete, 09° 20' N, 123° 18' E, 16

April 1986; AMS 1.31430-010' (220) Dumaguete, 09° 20' N,

123° 18' E, 16 April 1986; AMS I.40105-01U (1) Lighthouse

Point, North Mindoro Island on Escarceo Point, 13° 31' 05" N,

120° 59' 33" E, 5-20 m, 15 May 2000; BMNH 1933.3.11.497

(163) Sulu, Mo, Herre; CAS 32364 (3, 130-162) Sulu

Archipelago, Mo, 5.925203° N, 121.153284° E-5.925203° N,

121.153284° E, 30 December 1936; CAS 38731 (188) 6 miles

NE of Zamboanga, 5 September 1940; USNM 557*8 (95.6,

paratype of Choerodon melanostigma) Jolo, 11 Eebruary 1908;

USNM 5579* (164, paratype of Choerodon melanostigma) same

collection data as USNM 5578; USNM 57846* (226, holotype of

Choerodon zamboangae) Zamboanga, Mindinao, August 1906;

USNM 61154* (203, paratype of Choerodon zamboangae)

Zamboanga, Mindinao, August 1906; USNM 89967* (171,

holotype of Choerodon melanostigma) Jolo; USNM 152164*

(217) Zamboanga; USNM 152165* (2, 190-196) Zamboanga;

USNM 153475* (204) Zamboanga; USNM 153476* (203)

Zamboanga. Indonesia: BPBM 2994 (1) Lombok; BPBM

29944* (2, 131-157) Lombok, 11 Eebruary 1984; CSIRO H

7988-01* (133) Kedonganan, 8° 45' S, 115° 10' E, 24 Eebruary

2009; CSIRO H 7219-06* (130) Lombok, Tanjung Luar, 8°

48' S, 116° 29' E, 4 November 2010; CSIRO H 7219-07* (166)

same collection data as CSIRO H 7219-06; MZB 23117* (137)

Lombok, Tanjung Luar, 8° 48' S, 116° 29' E, 4 August 2010;

NMV A 31432-001* (157) Lombok, Tanjung Luar, 8° 48' S, 116°

2' E, 26 Eebruary 2011; WAM P.32927-002 (109) Raja Ampat

Islands, Misool Island, 02° 13' S, 130° 33' E, 65-75 m, 13

November 2007; NTM S.10752-00U (165) South Lombok, 09°

or 01" S, 116° 17' 60" E, September 1981; NTM S.10753-00r

(183) 1981; NTM S.10752-014+ (189) South Lombok, 09°

or 01" S, 116° 17' 60" E, September 1981 Timor-Leste: AMS

1.46112.050+ (1) Dili, 08° 32' 53" S, 125° 35' 15" E, 18 September

2012; AMS 1.46112-049+ (190) Dili, 08° 32' 53" S, 125°

35' 15" E, 18 September 2012. Australia, Northern Territory:

AMS I. 37179-001* (136) Timor Sea, 10° 07' 49" S, 129°

19' 55" E, 86 m, 2 October 1995; AMS 1.37184-001+ (210)

Arafura Sea, 09° 58' 44" S, 129° 18' 15" E, 82.3 m, 1 October

1995; AMS 1.37184-004+ (230) Arafura Sea, 09° 58' 44" S, 129°

18' 15" E, 82.3 m, 1 October 1995; NTM S.14000-001 (215)

Arafura Sea, “Tassie Shoal”, 10° 16' 48" S, 129° 38' 06" E, 88 m,

15 March 1995; NTM S.15129-002+ (236) Arafura Sea, Evans

Shoal, 10° 02' 49" S, 129° 29' 13" E, 107 m, 15 January 2000;

NTM S.15224-001+ (2, 222-243) Arafura Sea, Evans Shoal, 10°

09' 47" S, 129° 43' 55" E, 88 m, 29 March 1995; NTM S. 13728-

003+ (240) Arafura Sea, Lynedoch Bank, 10° 10' 59" S, 130°

40' 01" E, 102 m, 28 August 1993. Western Australia: AMS

l. 22804-017* (244) North West Shelf, Port Hedland, 18° 57' S,

118° 12' E, 100-104 m, 27 March 1982; CSIRO CA 2161 (194)

Joseph Bonaparte Gulf, 9° 52.0' S, 129° 12.0' E-9° 52.0' S, 129°

14.0' E, 138-158 m, 9 July 1980; NMV A 29706-001 (244) north

Western Australia, 15° 06' 31" S-15° 04' 50" S, 121°

46' 11" E-121° 48' 23" E, 90-96 m, 28 June 2007; WAM P.26251-

004* (153) Cape Villaret, 16° 03' S, 120° 58' E, 98-100 m, 25

June 1978; WAM P.26277-003 (143) 18° 53' S, 118° 35' E, 105

m, 29 May 1978; WAM P.25927-007 (249) Rankin Bank, 19°

40' S, 115° 45' E, 128 m, 25 August 1977; WAM P.31532-001

(235) Heywood Shoal, 13° 25.53' S, 124° 05.81' E, 94 m, 31 July

1998; WAM P.31605-001 (245) Browse Island, 14° 06' S, 123°

33' E, 7 December 1997.

Choerodon zosterophorus. (9 specimens examined, 44.7-153

mm SL). Philippines: BPBM 23470* (148) Luzon, Batangas,

Caban Island, 24 m, 27 July 1970; USNM 378714 (124) S tip of

Buyallao Island off SE Mindoro, 12° 21' 42" N, 121° 27' 34" E,

0-20 m, 30 May 2000; USNM 436349 (43) Oriental Mindoro,

Puerto Galera, off point immediately to W of Balatero Cove, 13°

30.813' N, 120° 55.786' E, 18-26 m, 2015. Indonesia: BPBM

29868* (103) Lombok, 6 Eebruary 1984; BPBM 32105* (109)

Elores, 8° 34' 40" S, 122° 11' 55" E, 20 m, 17 September 1987;

BMNH 1870.8.31.27* (153, lectotype of Choerops brenchleyi)

Raja Ampat Islands, Misol Island; BMNH 1870.^31.36* (147,

paralectotype of Choerops brenchleyi) Raja Ampat Islands,

Misol Island; BMNH 1870.8.31.85* (141, paralectotype of

Choerops brenchleyi) RajaAmpat Islands, Misol Island); RMNH

6537* (131, presumed holotype of Choerops zosterophorus)

Great Kei Island. New Guinea: MNHN A.8860* (117); BPBM

32434* (44.7) Laloata Island, 25 m, 27 October 1987.

Memoirs of Museum Victoria 76:113-120 (2017) Pubiished 2017

1447-2554 (On-iine)

http://museumvictoria.com.au/about/books-and-journais/journais/memoirs-of-museum-victoria/

DOi: http://doi.Org/10.24199/j.mmv.2017.76.02

Notes on Victorian lulomorphidae (Diplopoda: Spirostreptida)

Robert MeSIBOV (http://zoobank.org/urn:lsid:zoobank.org:author:24BA85AE-1266-494F-9DE5-EEF3C9815269)

West Ulverstone, Tasmania 7315, Australia (email: robert.mesibov@gmail.com)

Abstract Mesibov, R. 2017. Notes on Victorian lulomorphidae (Diplopoda: Spirostreptida). Memoirs of Museum Victoria 76: 113-120.

New locality records and illustrations are given for Atelomastix solitaria Jeekel, 2009, Victoriocambala bidentata

Jeekel, 2009 and V. buffalensis Verhoeff, 1944. Diagnostic differences are reviewed for Victoriocambala Verhoeff, 1944

versus Amastigogonus Brdlemann, 1913 and Equestrigonus Mesibov, 2017.

Keywords Diplopoda, Spirostreptida, lulomorphidae, Victoria, Australia

Introduction

During a recent visit to Museum Victoria, I sorted the Museum’s

collection of spirostreptidan millipedes to morphospecies.

Among the samples were specimens of the three previously

described Victorian lulomorphidae: Atelomastix solitaria

Jeekel, 2009, Victoriocambala bidentata Jeekel, 2009 and V.

buffalensis Verhoeff, 1944. Here I present new distribution

records for the three species and add details to their published

descriptions. Specimen locality records are available in table

form and as keyhole markup language (KML) files on the

Millipedes of Australia website, https://www.polydesmida.info/

millipedesofaustralia/localities.html.

I was unable to examine any of the type or other published

specimens of C.A.W. Jeekel for the three species. Following

Jeekel’s death in 2010, specimens in his possession and those

he had deposited in the Zoological Museum in Amsterdam

were transferred to the Naturalis Biodiversity Center in

Leiden, the Netherlands. In 2014, a search in the Naturalis

Biodiversity Center failed to yield the types of V. bidentata or

Jeekel’s published vouchers of V. buffalensis (K. van Dorp, in

litt., 17 Sep 2014). The types of A. solitaria, described from

specimens in the South Australian Museum collection, were

apparently not returned by Jeekel because they are not

currently in the museum (P. Hudson, in litt., 17 Mar 2017).

Abbreviations: NBC = Naturalis Biodiversity Center,

Leiden, the Netherlands; NMV = Museum Victoria,

Melbourne, Victoria, Australia; QVM = Queen Victoria

Museum and Art Gallery, Launceston, Tasmania, Australia;

SAM = South Australian Museum, Adelaide, South Australia,

Australia; ZSM = Zoologische Staatssammlung Munchen,

Miinchen, Germany.

Atelomastix solitaria Jeekel, 2009

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

F603EFD5-9818-4B08-9348-F5C15750BD98

Figure 1

Atelomastix solitaria Jeekel, 2009: 31-34, figs 1-4; Edward &

Harvey, 2010: 35; Korsos & Read, 2012: 45; Mesibov, 2017: 26.

Atelomastix sp. Jeekel, 1985: fig. 4.

Previous record. Male holotype and 2 male, 2 female and

4 juvenile paratypes, near Silverband Falls, Grampians, Victoria

[-37.1960 142.5250 +2 kml, J.J.H. Szent-Ivany & M.L. Szent-Ivany,

6 May 1978, under bark of Eucalyptus sp. in forest, SAM (Jeekel

2009, p. 31).

Material examined. 1 male, 3 females. Broken Falls, Grampians,

Victoria [-37.1110 142.4085 +500 m], A. Burns, 30 Oct 1949, NMV

K-13628; 1 male, 1 female, Grampians, Victoria [-37.2170 142.4350

+20 kml, collector unknown, 10 Nov 1949, NMV K-13655.

Description. Jeekel (2009) provides a detailed description of

the types, but his notes on the anterior gonopod are brief. The

Broken River male (fig. lA) has sclerite a nearly erect, curving

slightly posteriorly at apex, spatulate, concave posteriorly;

small field of ca 12 short setae on anterolateral surface; medial

margin slightly thickened subapically. Pseudoflagellum arising

at ca 1/2 gonopod height on medial surface of sclerite a, thin,

cylindrical, directed anterolaterally, tapering gradually to

rounded tip basal to sclerite a tip; posterior surface of

pseudoflagellum finely rugose. Sclerite b arising at ca 1/2

gonopod height, erect, reaching ca 5/6 gonopod height, directed

slightly posteriorly, not expanded distally; distal margin

oblique (lower posteriorly), slightly emarginate with sparse

comb of short setae; posterior margin slightly sinuous. Sclerite

c arising at ca 1/3-1/2 gonopod height, somewhat flattened

mediolaterally and tapering to rounded tip, directed slightly

laterally, strongly curving medially, reaching ca 2/3 gonopod

height with 4-5 long well-spaced setae on medial surface at ca

114

R. Mesibov

Figure 1. Atelomastix solitaria Jeekel, 2009, left anterior gonopod. A, medial and slightly posterior view of male from NMV K-13628; B,

posterior and slightly ventral (i.e. distal) view of male holotype, de-speckled scan of fig. 4 in Jeekel (2009). Pseudoflagellum (pf) and sclerites a,

b and c labelled; scale bar for A = 1.0 mm.

Victorian lulomorphidae

115

1/3-1/2 sclerite c height.

Distribution. So far known only from the Grampians National

Park in western Victoria (fig. 2).

Remarks. Jeekel (2009, p. 34) wrote that he had previously

illustrated the anterior gonopod of A. solitaria in Jeekel (1985),

where the species is referred to as 'Atelomastix spec, from

South Australia [sic]”. Fig. 5 in Jeekel (1985; p. 108) shows a

posterior view of the right anterior gonopod and is close to a

mirror image of the posterior view of the left anterior gonopod

in fig. 4 of the 2009 paper (reproduced here in modified form as

fig. IB). Neither of the two anterior gonopod illustrations in

Jeekel (2009) is in side view, leaving the shapes of the gonopod

sclerites a little unclear. The medial view in fig. lA will allow

easier comparison with Atelomastix anterior gonopods, as

illustrated in side view by Attems (1911), Edward and Harvey

(2010) and Mesibov (2017).

Victoriocambala Verhoeff, 1944

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act;

330CE9AA-124D-47EF-BCCC-6B60BE370A96

Victoriocambala Verhoeff, 1944: 35, 41; Jeekel, 1971: 115;

Hoffman, 1980: 91; Jeekel, 1981: 40; Jeekel, 1985; 106, fig 4; Mauries,

1987: 196, 198; Korsos & Johns, 2009: 3; Jeekel, 2009: 35; Edward &

Harvey, 2010: 5; Korsos & Read, 2012; 46.

Type species. Victoriocambala buffalensis Verhoeff, 1944,

by monotypy.

Remarks. Verhoeff (1944, p. 41) noted that his new, monotypic

Victoriocambala was most closely related to Amastigogonus

Brdlemann, 1913, a then-monotypic Tasmanian genus to which

he added two new Tasmanian species. By the time Jeekel

(2009) described a second Victorian Victoriocambala species,

the number of Tasmanian species had increased

to four (Hoffman 1972; Mauries et al.. 2001), and I have since

added another six Tasmanian species (Mesibov, 2017).

Consistent and taxonomically useful differences between

species in the two genera are listed in Table 1.

The two Victoriocambala species share several characters

with the Tasmanian endemic Equestrigonus tasmaniensis

Mesibov, 2017. In all three cases, there is an apical fringe of

long setae on the anterior gonopod telopodite, the leg 7 coxa is

not elongated, there is a prominent tab anteriorly on the leg 1

prefemur and the leg 2 coxa, and prefemoral tabs begin on ring

5. E. tasmaniensis differs from the Victoriocambala species

in diplosegment sculpture (suture weakly defined, longitudinal

metazonite striae curving upwards towards suture), having a

prominent setal crown on the posterior gonopod and the coxite

process on the anterior gonopod directed posterodistally

rather than parallelling the telopodite and protecting the

pseudoflagellum (Mesibov, 2017). It seems likely that in a

three-taxon molecular phylogeny, Equestrigonus and

Victoriocambala would be more closely related than either is

to Amastigogonus.

Victoriocambala bidentata Jeekel, 2009

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

CE72AC10-E464-42E6-BB72-1BE85208D37E

Figures 4C, 4D, 5A

Victoriocambala bidentata Jeekel, 2009: 35-38, figs 6-9.—Korsos

& Read, 2012: 46.

Previous record. Male holotype and 4 male, 2 female and 5

juvenile paratypes. Drummer State Forest, 15 km E of Cann River,

Victoria [-37.5765 149.3105 +1 km], C.A.W. Jeekel & A.M. Jeekel-

Table 1. Character state differences between species of Amastigogonus Brdlemann, 1913 and Victoriocambala Verhoeff, 1944.

Victoriocambala

Amastigogonus

Diplosegment suture

distinct around entire ring

(fig.3A,s)

indistinct, ventrally undetectable

Metazonite striae

longitudinal, stop at suture

(fig. 3A, ms)

curve upwards anteriorly, extend onto prozonite

(fig. 3B, ms)

Prozonite striae

semi-annular (below ozopore)

(fig- 3, ps)

absent

Male leg 1 coxosternite

in two pieces (fig. 4A)

in one piece (fig. 4B)

Male legl prefemur

nearly fused with coxa, not distinguishable

anteriorly

clearly demarcated from coxa (fig. 4B)

Male leg 1 tarsus

fused with tibia (fig. 4A)

clearly demarcated from tibia (fig. 4B)

Male prefemoral pads

begin on leg 4 or 5 (ring 5)

(fig. 3C, ptl)

begin on leg 10 (ring 8)

(fig. 3D, ptl)

Male leg 7 coxa

same as other anterior coxae

(fig. 3C,7c)

elongated and distally swollen

(fig. 3D, 7c; see also figs 2A, B in Mesibov, 2017)

Anterior gonopod telopodite

with apical fringe of setae

without apical fringe of setae

Posterior gonopod

with posterolateral bump or process bearing

minute setae (“telopodite remnant”; fig. 4C)

without posterolateral bump or process

116

R. Mesibov

Figure 2. Map of Victoria showing known localities as of 1 March 2017 for Atelomastix solitaria Jeekel, 2009 (circles), Victoriocambala

bidentata Jeekel, 2009 (triangles) and V. buffalensis Verhoeff, 1944 (squares). Mercator projection.

Rijvers, 13 Nov 1980, Australia Expedition station 84, Eucalyptus

forest along Princes Highway, under logs, Naturalis Biodiversity

Center (Jeekel 2009, p. 35).

Material examined. 1 male, 1 female, McKenzie River on Princes

Highway, 20 km NE of Cabbage Tree Creek (“196357"), Victoria

[-37.6294 148.8845 ±500 m], Australian Biological Resources Study

snail survey, 13 Apr 1975, NMV K-13634; 1 male, Marshmead Bush

Blitz, end of Lakeview Road, Victoria, 37° 31.565' S 149° 47.646' E

[37.5261 149.7941 +100 m], S. Hinkley & P. Lillywhite, 3 Dec 2016,

beating and direct search, NMV K-13651.

Distribution. Known from three localities in far East Gippsland,

Victoria (fig. 2).

Remarks. This species is easily recognised by the tooth on the

anterodistal margin of the anterior gonopod telopodite

(fig. 5A, t) and by the thinness of the tip of the coxite process

(fig- 4D).

Victoriocambala buffalensis Verhoeff, 1944

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

0BA89A30-C7D6-4437-90B7-A427AD8E2CF3

Figures 4A, 4E, 4F, 5B, 5C

Victoriocambala buffalensis Verhoeff, 1944: 42-43, figs 9-12;

Jeekel, 1971: 115; Jeekel' 1981: 40; Jeekel, 2009: 38-41, figs 10-13;

Korsos & Read, 2012: 46.

Previous records. Male holotype (Verhoeff described only one

male specimen), Mt Buffalo, Victoria [-36.7760 146.7670 +10 km],

date unknown, possibly collected by G.E. Nicholls, ZSM 20060631;

[the following five records are from Jeekel, 2009, p. 38] 1 female,

Gunyah Gunyah, 32 km SSW of Morwell, Victoria [-38.5306 146.3286

+100 m], station 92, C.A.W. Jeekel & A.M. Jeekel-Rijvers, 17 Nov

1980, timber track along Grand Ridge Road, temperate rainforest with

tree ferns, under logs and litter and in rotting trees; 1 juvenile, Mt

Taylor, 11 km NNW of Bairnsdale, Victoria [-37.7578 147.5986 +1 km]

station 87, same collectors, 14 Nov 1980, fragment of Eucalyptus

Victorian lulomorphidae

117

Figure 3. Victoriocambala buffalensis Verhoeff, 1944, NMV K-13624 (A, C) and Amastigogonus tasmanianus Brolemann, 1913, QVM 23:54414

(B, D). A, B, right lateral views (anterior to right) of midbody rings showing suture (s), metazonite striae (ms) and prozonite striae (ps). C, D,

ventral views of anterior portion of body (anterior to right) showing first prefemoral tab (ptl) and leg 7 coxa (7c). Scale bars = 1.0 mm.

118

R. Mesibov

Figure 4. Victoriocambala buffalensis Verhoeff, 1944 (A, F, NMV K-13624; E, NMV K-12000), Amastigogonus tasmanianus Brolemann, 1913

(B, QVM 23:54414) and V. bidentata Jeekel, 2009 (C, D, NMV K-13651). A, B, anterior views of left leg 1. C posterior view of right posterior

gonopod. D-F, lateral outlines of distal portion of coxite process on left anterior gonopod (anterior to left), at = anterior tab on prefemur, fe =

femur. Ip = lateral process (telopodite remnant), msd = median sternite division, po = postfemur, pr = prefemur, ta = tarsus, ti = tibia, tt = fused

tibiotarsus. Scale bars for A-C = 0.25 mm; D-F not to scale.

Victorian lulomorphidae

119

Figure 5. Victoriocambala bidentata Jeekel, 2009 (A, NMV K-13634) and F. buffalensis Verhoeff, 1944 (B, NMV K-13646; C, NMV K-13625).

Anterior views of anterior gonopods. c = coxite process, t = tooth on telopodite margin. Scale bars = 0.5 mm.

forest, along roadside between grassland, under logs; 1 male, Tarra

National Park, Victoria [-38.4289 146.5669 +1 km], station 91, same

collectors, along nature track in temperate rainforest with tree ferns,

under logs and in soil; 2 males, 2 females, Ferntree Gully National

Park, 18 km NNE of Dandenong, Victoria [-37.8800 145.3185 +500

m], station 93, same collectors, 18 Nov 1980, along nature track in

temperate rainforest with tree ferns, under logs and litter and in rotting

trees; 5 males, 13 females, 4 juveniles, Glenaladale National Park, 28

km WNW of Bairnsdale, Victoria [-37.6500 147.2667 +5 km], station

88, same collectors, 15 Nov 1980, dry rainforest along creek, along

nature track, under logs and stones, and in litter.

Material examined. 1 male, Glenrowan, Victoria [-36.4620 146.2240

+5 km], W. Kershaw, date unknown, NMV K-13633; 1 male, 2 females,

Thorpdale, Victoria [-38.2890 146.1760 +5 km], collector unknown, 1

Apr 1899, O.A. Sayce purchase 25 Jul 1911, NMV K-13625; 1 male, 2

females, Gembrook, Victoria [-37.9530 145.5540 +5 km], J. Kershaw,

Nov 1901, NMV K-13646; 1 male, 2 females. Emerald district,

Victoria [-37.9320 145.4400 +10 km], E.N. Jarvis, Aug 1904, NMV

K-13629; 2 males, 3 females, 1 juvenile (fragments), either Neerim

North Post Office or Noojee (unclear if site or collector address),

Victoria [-37.9030 145.9980 +5 km], T.G. Robinson, 8 Jul 1908, NMV

K-13623; 3 males, 1 female, Toolangi near Healesville, Victoria

[-37.5360 145.4710 +5 km], J.A. Kershaw, 16 Nov 1909, NMV

K-13624; 3 males, Gippsland, Victoria, J.E. Dixon, 1 Jul 1914, NMV

K-13640; 1 male, Bruthen, Victoria [-37.7070 147.8320 +5 km]. Dr

J.A. Leach, 12 Mar 1917, NMV K-13644; 1 male, 2 females, Bruthen,

Victoria [-37.7070 147.8320 +5 km], J. Barling, 3 Jan 1918, NMV

K-13645; 1 male (fragments), Marysville, Victoria [-37.5100 145.7480

+2 km], G.E. Hill, 19 Aug 1923, NMV K-13654; 1 male, 1 female,

Eerntree Gully, Victoria [-37.8840 145.2950 +2 km], G.E. Hill, 22 Eeb

1924, NMV K-13635; 1 male, Sherbrooke Eorest, Eerntree Gully,

Victoria [-37.8938 145.3635 +2 km], A. McEvey, 25 Sep 1969, “a bird

used this sp of millipedes for anting”, NMV K-13650; 1 male, near

Shady Creek, Dartmouth Survey locality LN, Victoria [-36.5500

147.6167 +2 km], Eield Survey Group, Eield Naturalists Club of

Victoria, 3 Mar 1973, NMV K-13643; 1 female, Tarra-Bulga National

Park, 0.5 km NNE of Tarra Valley Picnic Area, Victoria, 38° 26' 40"

S 146° 32' 30" E [-38.4444 146.5417 +1 km], G. Milledge, 14 Nov

1995, direct search, NOH 2239, NMV K-13637; 1 male. The Big

Culvert, 2.5 km ENE of Mt Observation, Victoria, 37° 33' 36" S 145°

52' 15" E [-37.5600 145.8708 +100 m] G. Milledge, 19 Eeb 1996,

direct search, NOH 1892, NMV K-13631; 1 male, Tarra-Bulga

National Park, 0.2 km W of Tarra Valley Picnic Area, Victoria, 38° 27'

S 146° 32' E [-38.4500 146.5333 +2 km], G. Milledge, 6 Mar 1996,

direct search, NOH 2238, NMV K-13639; 1 male, 2 juveniles,

Dandenong Ranges National Park, ca 0.22 km WNW of Kallista

roundabout, Kallista Project Site OSIA, Victoria, 37° 53.066' S 145°

22.006' E [-37.8844 145.3668 +25 m], 420 m a.s.l., N. Porch & A.

Lodewyke, 18 Apr 2013, open ground under tree ferns and dead holly

on E slope in mountain ash {Eucalyptus regnans) forest, Berlesate of

all leaf litter on 2 m by 2 m grid, NMV K-11998; 1 male, Dandenong

Ranges National Park, ca 0.21 km SW of Kallista roundabout, Kallista

Project Site SP2A, Victoria, 37° 53.112' S 145° 22.065' E [-37.8863

145.3678 +25 m], 395 m a.s.l., N. Porch & A. Lodewyke, 20 Apr 2013,

shield fern {Polysticlium proliferum) dominated ground under open

canopy, mountain ash (Eucalyptus regnans) forest, E slope, Berlesate

of all leaf litter on 2 m by 2 m grid; NMV K-12000

Distribution. As circumscribed here (see Remarks);

V. buffalensis is widespread in eastern Victoria (fig. 2).

Remarks. More than one species may be included here as

V. buffalensis. As noted by Jeekel (2009, p. 41), “This species

seems rather unstable in various eharacters”, among them the

shape of the coxite process. The process is fairly straight in

specimens from parts of Gippsland and the mountains north¬

east of Melbourne (figs 4E, 5B and fig. 10 in Jeekel, 2009), but

elsewhere in eastern Victoria, the tip is bent anterolaterally (figs

4F, 5C and fig. 12 in Verhoeff, 1944) the posterior surface is

depressed, and the anterior and lateral surfaces bowed outwards

around the depression (figs 4F, 5C). I have not noticed any other

differences between specimens with the two coxite process

forms, and I am reluctant to erect new Victoriocambala species

without more intensive sampling across the V. buffalensis range.

120

R. Mesibov

Acknowledgements

I am grateful to Peter Lillywhite and Catriona McPhee (NMV)

for assistance with the registration and loan of specimens

during my visit to Museum Victoria in February 2017, to

Karen van Dorp (NBC) and Peter Hudson (SAM) for advice

on specimen holdings, and to reviewers Nesrine Akkari and

Sergei Golovatch for helpful comments on the draft manuscript.

This study was funded by the author.

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di git alCollections/vi ewer popup.aspx?seid=BMAZO_

S004_1987_T009_N001

Mauries, J.-P, Golovatch, S.L, and Hoffman, R.L. 2001. On type material

and the identity of several lulus species in the collection of the

Museum national d’Histoire naturelle in Paris (Diplopoda,

Spirostreptida, Spirobolida). Zoosystema 23(3): 579-589. http://

sciencepress. mnhn.fr/sites/default/files/articles/pdf/z2001n3al2.pdf

Mesibov, R. 2017. lulomorphid millipedes (Diplopoda, Spirostreptida,

lulomorphidae) of Tasmania, Australia. ZooKeys 652: 1-36.

https://doi.org/10.3897/zookeys.652.12035

Verhoeff, K.W 1944. Zur Kenntnis der Cambaliden und fiber einige neue

australische Eormen derselben. Zoologischer Anzeiger 145: 27-45.

Memoirs of Museum Victoria 76:121-132 (2017) Pubiished 2017

1447-2554 (On-iine)

https://museumvictoria.com.au/about/books-and-journais/journais/memoirs-of-museum-victoria/

DOi: https://doi.Org/10.24199/j.mmv.2017.76.03

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

(http://zoobank.org/urn:lsid:zoobank.org:pub: 488E9F74-5E69-42B1-A12A-4E5239D9998D)

P. Mark O’Loughlin^*, Guadalupe Bribiesca-Contreras^

' Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia

^ School of Biosciences, Faculty of Sciences, The University of Melbourne, Melbourne, Victoria 3010, Australia

(lbribiesca@museum.vic.gov.au)

*To whom correspondence should be addressed. E-mail: pmoloughlin@edmundrice.org

Abstract O’Loughlin, P.M. and Bribiesca-Contreras, G. 2017. New asterinid seastars from the Pacific Ocean (Echinodermata:

Asteroidea). Memoirs of Museum Victoria 76: 121-132.

Three new Aquilonastra O’Loughlin (in O’Loughlin and Waters, 2004) species are described: Aquilonastra donia

sp. nov. for New Caledonia, lodged in the Museum national d’Histoire naturelle in Paris; Aquilonastra korora sp. nov. and

Aquilonastra starmeri sp. nov., both for Palau and lodged in the Florida Museum of Natural History.

Keywords New Caledonia, Palau, Aquilonastra, new species.

Introduction

O’Loughlin and Waters (2004) provided a comprehensive

revision of genera of Asterinidae based on morphological and

limited molecular genetic data. Within this work they

established the new genus Aquilonastra O’Loughlin and

provided a key to Asterinidae genera. O’Loughlin and Rowe

(2006) revised genus Aquilonastra based on morphological

observations. Thirteen new species were described and a key

to Aquilonastra species was provided. O’Loughlin (2009),

O’Loughlin and Mackenzie (2013), and O’Loughlin and

Bribiesca-Contreras (2015) added new species of Aquilonastra.

O’Loughlin and Bribiesca-Contreras (2015) provided a revised

key to Aquilonastra species. We recognise the need for further

revision of Asterinidae based on comprehensive genetic data.

We assign the current new species from the Pacific Ocean to

Aquilonastra based on morphological observations only, and

acknowledge some uncertainty with these assignments.

Methods

For photography purposes, the alcohol-preserved specimen

was allowed to partly air-dry. Photographs were taken using a

Cannon 5D Mark II camera with a Cannon 65 mm macro lens

and a 100 mm lens. Series of photographs were taken and

stacked using the Zerene Stacker software.

After assembling the whole-specimen montage

photographs, a ray from each of the three type specimens was

cut off for the purpose of observing external and internal

skeletal structure. The distal end of each of these three cut-off

arms was cleared briefly in bleach and then washed in water.

Photographs were taken to show internal skeletal structures.

Definitions and illustrations of terms

For definitions and illustrations of terms used, such as

superactinal plates, superambulacral plates, sacciform

spinelets and splay-pointed spinelets, see O’Loughlin and

Waters (2004).

Abbreviations

MNHN Museum national d’Histoire naturelle.

UF Florida Museum of Natural History, University of

Florida.

Asterinidae Gray, 1840

Synonymy. See Clark and Downey, 1992.

Diagnosis. See Clark and Downey, 1992.

Remarks. For a recent revision of Asterinidae, see O’Loughlin

and Waters (2004). For the addition of genus Ailsastra, see

O’Loughlin and Rowe (2005).

Aquilonastra O’Loughlin, 2004 (in O’Loughlin and Waters,

2004)

Aquilonastra O’Loughlin, in O’Loughlin and Waters, 2004: 5

(key to genus), 13-15, tables 1, 2.—O’Loughlin and Rowe, 2005:

18L-Saba and Fujita, 2006: 270.-Byrne, 2006: 244, 245, 248, 250,

251.—O’Loughlin and Rowe, 2006: 257-287 (key to

species).—O’Loughlin, 2009: 204, fig. L—O’Loughlin and

Mackenzie, 2013: 177-180, figs 1, 2.—O’Loughlin and Bribiesca-

Contreras, 2015: 33-46, figs 1-8 (revised key to species).

Diagnosis (from O’Loughlin and Bribiesca-Contreras, 2015).

Rays 5, or 5-8 in fissiparous species; inter-radial margin deeply

122

P.M. O’Loughlin & G. Bribiesca-Contreras

incurved, form stellate; rays discrete, broad at base, tapering,

rounded distally; flat actinally, convex abactinally; abactinal

plates in longitudinal series, not perpendicular to margin;

papulate areas extensive; papulae predominantly single, large, in

longitudinal series along sides of rays; abactinal plates with

glassy convexities; abactinal spinelets and actinal spines

predominantly fine, glassy, conical or sacciform or splay-pointed

sacciform, in bands or tufts, numerous (10-40 per plate); actinal

plates in longitudinal, sometimes oblique, series; superambulacral

plates present for all of ray, sometimes for part of ray or absent in

paedomorphic species; superactinal plates present.

Remarks. We have not further revised the morphological

diagnosis of genus Aquilonastra to accord with variations

observed here, such as a shallow incurved junction of rays in

two of the species below and the absence of superambulacral

plates in one of the species below. We anticipate a necessary

extensive revision when adequate genetic data become available.

Aquilonastra donia sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

C41D2993-A090-4D51-A8B9-C15A29B271B3

Figures 1, 2, 3a-d.

Material examined. Holotype. New Caledonia, Yate reef flat, coll.

Gaillande, July 1970, MNHN-IE-2014-641 (original registration

MNHN EcAsll855) (dry; many spinelets and pedicellariae lost with

handling over time).

Description. Asterinid seastar, five sub-equal rays, rays wide

basally, bluntly rounded distally, R = 22 mm, r = 15 mm, rays

merge at bases, inter-radial junction of rays shallow in-curved,

central rays elevated and rounded abactinally, distinct broad

inter-radial low non-papulate apron marginally, rays flat actinally,

margin acute. No abactinal or actinal gonopores detected. Single

madreporite. Notfissiparous. Superomarginal andinferomarginal

plates sub-equal, superomarginal plates in discrete series,

slightly larger than adjacent distal abactinal plates, inferomarginal

plates projecting outwards. Broad low marginal apron up to 7

plates across longitudinally, supported by internal superactinal

plates; superambulacral plates present, small. Glassy convexities

on cleared abactinal and actinal plates.

Abactinal: disc discrete, small, demarcated irregularly by

5 wide doubly papulate radial plates and 5 smaller inter-radial

plates; single conspicuous madreporite, above junction of

bases of two rays; pedicellariae present inter-radially,

proximally and distally, conspicuous, frequently longer than

adjacent spinelets, each comprising two stout, in-curved

pointed conical teeth, present on proximal edges of inter-radial

plates over concave indentation for papula; abactinal plates

imbricate, indented proximally for single papula; no regular

carinal series of plates, but a few doubly papulate carinal

plates along some proximal upper rays; few small secondary

plates on disc, rays and marginal apron; abactinal papular

plates in about 7 longitudinal series down each side of each

ray, plates with elevated rounded proximal edge, about 20

papular plates in uppermost series, 1-2 in lowest series; distal

inter-radial plates on apron in longitudinal series parallel to

the rays and oblique curved series from the ray to the margin.

Abactinal spinelets readily lost: up to at least 20 digitiform

spinelets over crown of proximal plates, not in tufts, distal

spinelets slender conical to subsacciform; up to about 5 slender

conical spinelets per superomarginal plate.

Actinal: inter-radial plates in longitudinal series parallel to

the ambulacrum and oblique transverse series from the

ambulacrum to the margin. Actinal spines per plate: oral 6,

long digitiform to conical proximally, to short distally; sub¬

oral about 5, long conical proximally to short distally; furrow

4-5 long conical; subambulacral 4-5 long conical; actinal 2-3,

mostly 2, long conical; inferomarginal up to about 16, thick

digitiform actinally to thin conical subsacciform abactinally.

Distribution. New Caledonia, reef flat.

Etymology. Original collection record is New Hebrides, now

New Caledonia, and named for the donia ending to Caledonia.

Remarks. The long abactinal pedicellariae, with 2 conical

pointed and curved valves that are frequently longer than

adjacent spinelets, and actinal inter-radial plates with

predominantly 2 spines, are distinguishing feature of

Aquilonastra donia sp. nov. Other Aquilonastra species with

prominent pedicellariae with differentiated valves (teeth) are

distinguished by: Aquilonastra anomala (H.L. Clark, 1921) is

fissiparous with multiple madreporites; Aquilonastra batheri

(Goto, 1914) has tufts of splayed abactinal spinelets, and up to

12 actinal inter-radial spines; Aquilonastra corallicola (Marsh,

1977) is fissiparous and has two different forms of spinelets on

the paxilliform upper ray plates; Aquilonastra coronata

(Martens, 1866) has irregularly distributed high paxilliform

abactinal plates with two different forms of spinelets;

Aquilonastra iranica (Mortensen, 1940) has up to 5 doubly

papulate proximal carinal plates and the spinelets on proximal

abactinal plates are in small groups; Aquilonastra richmondi

O’Loughlin and Rowe, 2006 has spinelets of two forms on

abactinal plates; Aquilonastra rowleyi O’Loughlin and Rowe,

2006 has a disc that is delineated by a dense band of spinelets

on wide radial plates, each plate with about 100 long thin

pencil-like glassy spinelets; Aquilonastra shirleyae

O’Loughlin, 2009 has spinelets in splayed clusters and up to 10

actinal inter-radial spines per plate; Aquilonastra watersi

O’Loughlin and Rowe, 2006 has abactinal spinelets in clusters.

Aquilonastra korora sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

765787CA-CE22-4EE5-8637-497D9E9EA85D

Figures 4, 5, 6a-d.

Material examined. Holotype. Palau, Koror, Mutremdiu reef, rock, 30 m,

coll. J. Starmer, 19 Aug 1995, code BEL-276, UE 2437 (fixed in formalin,

dry; many spinelets and pedicellariae lost with handling over time).

Description. Asterinid seastar, five sub-equal rays nearly

digitiform, rays wide basally, narrowly rounded to pointed

distally, R = 25 mm, r = 10 mm, rays merge at bases, inter-

radial junction of rays deeply in-curved, central rays elevated

and rounded abactinally, weakly evident low sloping distal

abactinal margin, no distinct apron; rays flat actinally, margin

acute. Abactinal gonopores detected. Single madreporite. Not

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

123

Figure 1. Montage photograph of the holotype of Aquilonastra donia sp. nov. (MNHN-IE-2014-641). Abactinal view showing five sub-equal

rays, distinct low marginal apron, absence of carinal series of plates, presence of rare doubly papulate carinal plates, and single large madreporite.

The abactinal plates have become denuded of most spinelets over time.

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P.M. O’Loughlin & G. Bribiesca-Contreras

Figure 2. Montage photograph of the holotype of Aquilonastra donia sp. nov. (MNHN-IE-2014-641). Actinal view showing five sub-equal rays,

spination, longitudinal and oblique-transverse series of actinal plates.

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

125

Figure 3. Montage photographs of the holotype of Aquilonastra donia sp. nov. (MNHN-IE-2014-641): a, abactinal view of disc, single large

madreporite, some remaining digitiform spinelets, and pedicellariae (highlighted); b, abactinal details of distal interradius showing pedicellariae

(highlighted), and conical to sub-sacciform spinelets on apron; c, actinal view showing oral, suboral, furrow and inter-radial spines; d, transverse

section of a ray with superactinal (highlighted left) and superambulacral plates (highlighted right).

fissiparous. Fairly irregular series of superomarginal plates,

subequal with adjacent abactinal and inferomarginal plates,

inferomarginal plates project narrowly outwards. Narrow

sloping margin up to about 5 plates across longitudinally,

supported by some internal superactinal plates, distal marginal

abactinal and actinal plates contiguous internally;

superambulacral plates absent with tissue strengthening only of

junction internally between ambulacral and actinal plates.

Glassy convexities on cleared abactinal and actinal plates.

Abactinal: disc discrete, small, demarcated irregularly by 5

bean-shaped non-papulate radial plates and 5 slightly smaller

inter-radial plates, disc plates papulate, distal abactinal plates

on rays not papulate; single conspicuous madreporite, above

junction of bases of two rays; pedicellariae present inter-

radially, proximally, low, not conspicuous, each comprising

two short thick blunt teeth, present on proximal edges of inter-

radial plates over concave indentations for papulae; abactinal

plates imbricate, indented proximally for predominantly 1

rarely 2 papulae; no regular carinal series of plates, rare doubly

papulate carinal plates present; rare small secondary plates on

disc and proximal upper rays; abactinal plates in about 3

longitudinal series down each side of each ray, plates with

elevated rounded proximal edge, up to about 20 papular plates

in uppermost series, about 10 in mid-series, about 4-7 in lowest

series; distal inter-radial plates in longitudinal and irregularly

distributed oblique series. Abactinal spinelets readily lost:

abactinal spinelets finely sacciform with needle-like long taper,

up to at about 16 spinelets across proximal raised edge of

proximal plates, not in tufts, distal spinelets slender conical to

subsacciform; up to about 2-3 small fine sacciform acicular

spinelets on crowns of superomarginal plates.

Actinal: inter-radial plates in longitudinal series parallel to

the ambulacral furrow, some oblique transverse series from

the ambulacrum, distal inter-radials irregular in distribution.

Actinal spines per plate: oral up to 9, long conical proximally,

to short distally; suboral up to 6, 3 long conical proximally,

2-3 shorter on each side of plate distally; furrow 5-6 conical

subsacciform, 2 long apically, 1-2 short laterally on each side

of plate; subambulacral up to 8, 2 long subsacciform conical

apically, 3 short conical on each side of plate; actinal

predominantly 2, subsacciform conical; inferomarginal

predominantly about 5 sacciform.

Distribution. Palau, rocky reef, 30 m.

Etymology. Named korora for Koror, the state in Palau where

the type was collected.

Remarks. A diagnostic character of genus Aquilonastra is the

presence of superambulacral plates. Aquilonastra korora sp.

126

P.M. O’Loughlin & G. Bribiesca-Contreras

Figure 4. Montage photograph of the holotype of Aquilonastra korora sp. nov. (UF 2437). Abactinal view showing residual colouration, five

sub-equal digitiform rays, absence of carinal series of plates, single large madreporite (lower arrow) and at least one conspicuous abactinal

gonopore (upper arrow). The abactinal plates have become denuded of most spinelets over time.

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

127

Figure 5. Montage photograph of the holotype of Aquilonastra korora sp. nov. (UF 2437). Actinal view showing residual actinal colouration, five

sub-equal rays, spination, longitudinal and oblique-transverse series of actinal plates with irregular distal inter-radials.

128

P.M. O’Loughlin & G. Bribiesca-Contreras

Figure 6. Montage photographs of the holotype of Aquilonastra korora sp. nov. (UF 2437): a, abactinal view of disc, showing disc plate

arrangement, single large madreporite (top right), sacciform spinelets; b, abactinal details of distal interradius showing short blunt pedicellariae

(highlighted) and residual slender conical, sub-sacciform and sacciform spinelets; c, actinal view showing oral, suboral, furrow and actinal

interradial spines; d, transverse section of a ray with a superactinal plate highlighted.

nov. is thus unique among Aquilonastra species in lacking such

plates, but the species conforms well in most ways with the

diagnosis of the genus and is assigned to this genus, with

reservations, as the most appropriate existing genus. The

presence of low pedicellariae with two short thick blunt teeth

and predominantly 2 actinal inter-radial spines per plate are

distinguishing characters. There are additional characters that

distinguish Aquilonastra korora sp. nov. from other

Aquilonastra species that have distinctive pedicellariae:

Aquilonastra anomala (Clark, 1921) and Aquilonastra

corallicola (Marsh, 1977) have up to 8 rays and are fissiparous

with multiple madreporites; Aquilonastra batheri (Goto, 1914)

has tufts of splayed abactinal spinelets and up to 12 actinal

inter-radial spines; Aquilonastra coronata (Martens, 1866) has

irregularly distributed high paxilliform abactinal plates with

two different forms of spinelets; Aquilonastra iranica

(Mortensen, 1940) has up to 5 doubly papulate proximal carinal

plates and the spinelets on proximal abactinal plates are in

clusters; Aquilonastra rowleyi O’Loughlin and Rowe, 2006 has

a disc that is delineated by a dense band of spinelets on wide

radial plates, each plate with about 100 long thin pencil-like

glassy spinelets; Aquilonastra shirleyae O’Loughlin, 2009 has

spinelets in splayed clusters and up to 10 actinal inter-radial

spines per plate; Aquilonastra watersi O’Loughlin and Rowe,

2006 has pedicellariae with differentiated valves but has

doubly papulate proximal carinal plates and abactinal spinelets

in clusters. All of these species have superambulacral plates.

We note that Aquilonastra corallicola was described for Palau.

Aquilonastra starmeri sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

8B1FF925-A45F-4268-BDFE-C5F262C420D2

Figures 7, 8, 9a-d.

Material examined. Holotype. Palau, off NE Ngeryktabe Island,

rocky islets, 7° 18.26’ N, 134° 27.25’ E, 10 m, coll. J. Starmer, 5 Mar

2003, code BPAL-081, photo GP 954, UP 1612 (in 75% ethanol; many

spinelets lost with handling over time).

Description. Asterinid seastar, five sub-equal rays, rays wide

basally, bluntly rounded distally, R = 20 mm, r = 12 mm, rays

merge at bases, inter-radial junction of rays with shallow in¬

curve, central rays with high elevation and rounded abactinally,

distinct broad inter-radial low non-papulate apron marginally,

up to about 7 plates wide longitudinally, rays flat actinally,

margin acute. Not fissiparous. No abactinal or actinal gonopores

detected. No pedicellariae detected. Superomarginal plates in

irregular series, slightly smaller than projecting inferomarginal

plates, subequal in size with adjacent irregular apron plates.

Broad low proximal marginal apron supported by internal

superactinal plates, outer marginal apron supported by

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

129

Figure 7. Montage photograph of the holotype of Aquilonastra starmeri sp. nov. (UF 1612). Abactinal view showing five sub-equal rays, distinct

marginal apron, lacking carinal series of plates, single large madreporite. The abactinal plates have become denuded of most spinelets over time.

130

P.M. O’Loughlin & G. Bribiesca-Contreras

Figure 8. Montage photograph of the holotype of Aquilonastra starmeri sp. nov. (UF 1612). Actinal view showing five sub-equal rays, spination,

longitudinal and oblique-transverse series of actinal plates.

New asterinid seastars from the western Pacific Ocean (Echinodermata: Asteroidea)

131

Figure 9. Montage photographs of the holotype of Aquilonastra starmeri sp. nov. (UF 1612): a, abactinal view of disc (left) and proximal ray

plates with short conical to digitiform spinelets; b, abactinal details of distal interradius showing acicular conical to subsacciform spinelets on

apron, and supero- and inferomarginal plates; c, actinal view showing oral, suboral, furrow and actinal interradial spines; d, transverse section

of a ray with superambulacral plate (left) and superactinal plate (right) highlighted.

contiguous abactinal and actinal plates; some small

superambulacral plates present. Glassy convexities on cleared

abactinal and actinal plates.

Abactinal: disc discrete, small, demarcated irregularly by 5

wide doubly or singly papulate radial plates and 5 smaller inter¬

radial plates; single conspicuous madreporite, above junction

of bases of two rays. Abactinal plates imbricate, indented

proximally for single papula, rarely 2; no carinal series of

plates, only one doubly papulate carinal plate; small secondary

plates on disc and few abactinally; abactinal plates in about 6

longitudinal series down each side of each ray, plates with

elevated rounded proximal edge, about 20 papular plates in

uppermost series, 1-3 in lowest series. Surface of abactinal

plates covered with up to about 40 short conical to digitiform

spinelets, not sacciform, not clustered, spinelets readily lost;

spinelets on abactinal apron acicular, conical, subsacciform,

about 6 per plate; superomarginal plates with fewer spinelets.

Actinal: inter-radial plates in longitudinal and oblique

transverse series from the ambulacrum. Actinal spines per

plate: oral 11, long conical, fused basally, shorter distally; sub¬

oral about 8, long conical proximally to short distally; furrow

5, conical, fused basally; subambulacral 4, conical, fused

basally; actinal 3-5, conical, fused basally; inferomarginal up

to about 12, digitiform, short actinally.

Distribution. Palau, off NE Ngeryktabe Island, rocky islets, 10 m.

Etymology. Named for John Starmer who collected and

documented the type specimen.

Remarks. Aquilonastra starmeri sp. nov. is distinguished from

many Aquilonstra species by lacking pedicellariae, lacking

carinal series of plates and having a non-fissiparous habit.

Aquilonastra species that lack pedicellariae but are fissiparous

are: Aquilonastra burtoni (Gray, 1840); Aquilonastra cassini

O’Loughlin and Bribiesca-Contreras, 2015; Aquilonastra

chantalae O’Loughlin and Mackenzie, 2013; Aquilonastra

colemani O’Loughlin and Rowe, 2006; Aquilonastra conandae

O’Loughlin and Rowe, 2006; Aquilonastra doranae O’Loughlin

and Rowe, 2006; Aquilonastra moosleitneri O’Loughlin and

132

P.M. O’Loughlin & G. Bribiesca-Contreras

Rowe, 2006; Aquilonastra yairi O’Loughlin and Rowe, 2006.

Aquilonastra starmeri sp. nov has 3-5 actinal inter-radial spines

per plate and is thus distinguished from those with up to eight

and more: Aquilonastra byrneae O’Loughlin and Rowe, 2006;

Aquilonastra cepheus (Muller and Troschel, 1834); Aquilonastra

limboonkengi (Smith, 1927); Aquilonastra samyni O’Loughlin

and Rowe, 2006. Aquilonastra starmeri sp. nov. has uniformly

rounded rays abactinally and is distinguished from those with

paxilliform rays: Aquilonastra lorioli (Koehler, 1910);

Aquilonastra minor (Hayashi, 1974); Aquilonastra rosea (H.L.

Clark, 1938). Aquilonastra starmeri sp. nov. is distinguished

from the remaining species assigned to Aquilonastra that are not

fissiparous and lack prominent pedicellariae as follows:

Aquilonastra alisonae O’Loughlin and Bribiesca-Contreras,

2015 has splay-pointed abactinal spinelets; Aquilonastra

halseyae O’Loughlin and Rowe, 2006 has sacciform spinelets

and the primary superomarginal plates are frequently separated

by smaller plates; Aquilonastra marshae O’Loughlin and Rowe,

2006 has some doubly papulate carinal series of plates;

Aquilonastra oharai O’Loughlin and Rowe, 2006 has spinelets

in double splayed series across the proximal edge of abactinal

plates; Aquilonastra scobinata (Livingstone, 1933) has carinal

series of singly papulate plates and long thin pencil-like sacciform

splay-pointed spinelets. We note again that Aquilonastra

corallicola was described for Palau and is fissiparous.

Acknowledgements

We are grateful to the Museum national d’Histoire naturelle in

Paris, and to Gustav Paulay and the Florida Museum of

Natural History in the University of Florida for the opportunity

to borrow and study the specimens. We are grateful for the

critical reviews of our manuscript provided by Chris Mah

(Smithsonian Institution) and Melanie Mackenzie (Museum

Victoria), and to David Paul (Museum Victoria) for assistance

with photography.

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