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Memoirs of Museum Victoria 63(1): v-v (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

t pedal issue - New species of Australian fishes

Memoirs of Museum Victoria volume 63 issue 1

Guest Editors

Martin F. Gomon

Ichthyology, Museum Victoria, GPO Box 666, Melbourne Vic 3001, Australia (mgomon@museum.vic.gov.au)

Jeffrey M. Leis

Ichthyology, Australian Museum, 6 College Street, Sydney NSW 2010, Australia (jeff.leis@austmus.gov.au)

This issue of the Memoirs of Museum Victoria presents a

selection of papers on the taxonomy of fishes intended to

improve our understanding of Australian fish biodiversity. It is

the third in a series of such publications, the first appearing in

1987 as Volume 48, Number 1 of the Memoirs of the Museum of

Victoria. Intended as a venue for new names and descriptions of

recognised, but formally undescribed, species, the volume

contained 20 papers by 15 authors that described as new two

genera and 33 species. As a continuing initiative of OzFishNet,

a consortium of collection-based Australian ichthyologists

formed to promote research, data access and other issues of

mutual interest to the Australian community (http://www.

amonline.net.au/FISHES/ozfishnet/), a second publication.

Volume 56, Number 2 of the Records of the Australian Museum

published in 2004 contained descriptions of an additional genus

and 33 species. In that volume, the 17 papers authored by 21

scientists also clarified inadequate perceptions of related

Australian forms.

The current issue was conceived prior to the publication of

the 2004 volume of the Records to pick up the overflow of

papers expected for that publication, knowing that its accelerated

publication time line would be beyond the capability of some

intended contributors. Like its predecessors it is also intended

to continue the publication of names and descriptions so that

scientific colleagues have the means to better understand the

natural environment with which they are working. The present

contribution contains 12 papers by 19 authors and describes a

new genus and 15 species, as well as clarifying our understanding

of closely related forms and laying out a worldwide taxonomic

framework for a family having more than 70 nominal species.

Although these publications provide important steps toward

a full documentation of Australia’s fish fauna, the job is far from

complete. Despite a decline in number of taxonomic ichthyologists

in Australia and a reduction in number of positions for members

of the profession in Australian institutions, new fishes continue

to be discovered at about the same rate as they have at any time

in the last 25 years. The rate of their discovery approximates the

rate at which new species are being described. A dedicated

group of scientists continue to work away, behind the scene, to

further this tradition.

Contents

Volume 63 Number 1 2006

1 > A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean

Thomas H. Fraser and Gerald R. Allen

7 > Description of two new species of Nesogoblus (Pisces: Gobioidei: Gobiidae) from

southern Australia

Douglass F. Floese and Flelen K. Larson

15 > Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae) from

south-eastern Australia

Douglass F. Floese and Sally Reader

21 > Description of a new species of Fleterocllnus (Blennoidei: Clinidae) from southern Australia

By Douglass. F Floese and Denise S. Rennis

25 > Description of two new species of shore-eels (Gobiesocidae: Cheilobranchinae: Alabes) from

south-eastern Australia and Norfolk Island

Barry Flutchins

29 > Revision of the genus Flapalogenys (Teleostei: Perciformes) with two new species from the

Indo-West Pacific

Yuklo IwatsukI and Barry C. Russell

47 > Two new species of Parapercis (Perciformes: Pinguipedidae) from north-eastern Australia,

and rediscovery of Parapercis colemani Randall & Francis, 1993

Jeffrey W. Johnson

57 > SImIpercIs trispinosa, a new genus and species of sandperch (Perciformes: Pinguipedidae)

from eastern Australia

Jeffrey W. Johnson and John E. Randall

65 > Notoraja hirticauda, a new species of skate (Chondrichthyes: Rajoidei) from the south-eastern

Indian Ocean

Peter R. Last and John D. Mceachran

77 > Nomenclature and distribution of the species of the porcupinefish family Diodontidae

(Pisces, Teleostei)

Jeffrey M. Lels

91 > A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East China Sea

and off Western Australia

TetsujI Nakabo, Dianne Bray and UmeyoshI Yamada

97 > Australian trevallies of the genus Pseudocaranx (Teleostei: Carangidae), with description of a

new species from Western Australia

William F. Smith-Vaniz and Floward L. Jelks

Memoirs of Museum Victoria 63(1): 1-5 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean

Thomas H. Fraser^ and Gerald R. Allen^

'Mote Marine Laboratory, 1600 Ken Thompson Parkway, Sarasota, Florida, 34236 USA (cardinalfish@comcast.net)

^Department of Aquatic Vertebrates, Western Australian Museum, Frances St., Perth, Western Australia 6000, Australia

Abstract Fraser T.H. and Allen G.R. 2006. A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean.

Memoirs of Museum Victoria 63(1): 1-5.

A new species of apogonid fish, Neamia articycla, is described from Australia, Indonesia, Philippines and Fiji,

bringing the total number of described Neamia spp to three. It is distinguished from Neamia notula in preservation by: a

circular, ocellated dark spot rather than one that is oval-shaped and flattened along its anterior margin, with an outer,

narrow darkish edge clockwise dorsally from abutting edge of preopercle around to meeting ventral abutting edge of

preopercle, afaint, narrow, horizontal line of melanophores below the eye reaching the preopercle ridge; in life no darkish

marks behind the eyes, a circular spot with pale outer band without dark edging instead of an oval spot flattened along

preopercle edge with a darkish edge; five predorsal scales instead of four; usually 22 pored, lateral-line scales instead of

23 and a longer pectoral fin. Neamia articycla can be distinguished from Neamia octospina by having 14 pectoral rays

instead of 17-21 and by having seven visible first-dorsal spines instead of eight.

Keywords Apogonidae, Neamia articycla, Neamia notula, cardinalfish, new species Apogonichthys, Foa, Fowleria.

Introduction

Ichthyologists collecting from poorly sampled habitats in depths

of 10^0 m continue to find new species of cardinalfish. We

describe here a new Neamia which is associated with Halimeda

beds growing in deeper coral reef systems of the western Pacific

Ocean. The new species represents the third known species in

this clade. There may be other material in museums identified as

Fowleria because of the similar physiognomy. Neamia spp can

be distinguished from similar Fowleria species by the following

characters: fused hypurals in the caudal-fin skeleton, completely

pored lateral-line scales in adults (versus partially pored and

then pitted, except one undescribed species of Fowleria),

cycloid scales on the head and anterior portions of the body

(weakly ctenoid scales on the nape for species of Fowleria,

elsewhere on the head, weak to strongly ctenoid scales and

ctenoid on entire body).

Methods for counts and measurements were described in

detail by Fraser and Lachner, 1985. All measurements are in

mm to the nearest 0.1. Percentages are of standard length.

Institutional acronyms follow Eschmeyer, 1998 and Leviton et

ah, 1985. Field station numbers are listed for additional

collection information, for example, VGS 69-23. Radiographs

(X-rays) have been taken from type material. A camera lucida

attachment on a Wild microscope and a needle blowing air

were used to make the diagrammatic drawings of the pores.

The description of pores refers to larger and to smaller

perforations (referred to as minipores) of the skin over the

canal systems of the head. The latter are sometimes obscured

by congealed mucous.

Neamia Smith and Radcliffe in Radcliffe, 1912

Type species. Neamia octospina Smith and Radcliffe in Radcliffe,

1912.

Diagnosis. An apogonine with 3 hypurals, 1 and 2 fused, 3 and 4

fused, 5 free, hypurals 3-1-4 fused to urostylar centrum; 1 pair of

reduced or slender uroneurals; 3 epurals; 3 supraneurals; 2

supernumerary dorsal-fin spines; smooth preopercle edges and

ridge, smooth post-temporal; no basisphenoid; a reduced, tiny

supramaxilla; no palatine teeth; scales on cheek and opercle

cycloid; lateral-line scales extending from post-temporal to base

of caudal fin, all ctenoid with simple pores; rounded caudal fin;

9 dorsal-fin spines, 8th spine visible or hidden by skin, dorsal fin

deeply notched and considered separate, the 9th spine at

beginning of 2nd dorsal fin; dorsal-fin rays 9; anal-fin rays 8;

pectoral-fin rays 13-21; pale stomach, intestine and peritoneum.

Neamia articycla sp. nov.

Figures 1-4

Holotype. AMS 1.25121-005; 1, 35.5; Australia, Great Barrier

Reef, Qld, 14°41'S, 145°30'E, east of Lizard L; 31 Jan 1982; 36-38 m,

J. Leis.

Thomas H. Fraser and Gerald R. Allen

Figure 1. Neamia articycla: A. Holotype, AMS 1.251221-005, 35.5

mm SL (preserved), Australia, Qld, east of Lizard L. B. Paratype,

USNM 370291, 23.5, mm SL, Philippines, Mindanao L, modified

from colour image of afresh specimen by J. T. Williams. C. Paratype,

USNM 209665, 32.4 mm SL (preserved), Indonesia, Moluccas,

Ceram 1. D. Paratype CAS 223500, 32.9 mm SL (preserved), Fiji,

Viti Levu, Nananui-i-cake 1.

Figure 3. Post-ocular melanophore patterns on the cheek (faint line)

and circular spot with pale interval and outer edging of melanophores

starting along posterior edge of preopercle (POP). USNM 209665

paratype, 32.4 mm SL (preserved).

Figure 2. Cephalic lateralis system of pores on the head of the

holotype, Neamia articycla. A. Dorsal view. B. Lateral view

slightly tilted. C. Ventral view of dentary-articular, d. ventral

mandibular pore. e. posterior articular pore. f. post-temporal pore.

A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean

Paratypes. Pacific Ocean: Indonesia: USNM 209665; 8, 14.2-32.3;

Ceram, Pirn Bay; VGS 73-6; 10 Jan 1973; 8 m. USNM 209929; 5, 22.5-

27.7; Saparua; VGS 73-14; 18 Jan 1973; 9 m; X-ray. Fiji: CAS 223500;

1, 32.9; 17°19.462'S, 178°14.250'E, Nananui-i-cake L, Viti Levu; G02-

76; 15 Mar 2002; 1-3 m, colour transparency. Australia, Qld: AMS

1.40665-004; 1, 34.3; & AMS 1.40665-005; 1, 17.0; 14°38.20'S,

145°32.30'E, between Lizard I. & Yonge Reef, Great Barrier Reef; 26

Jan 2001; 27 m.AMS 1.40666-012; 1,31.1 &AMS 1.40666-013; 1, 19.0;

14°38.16'S, 145°32.16'E, between Lizard I. & outer barrier reef. Great

Barrier Reef; 29 Jan 2001; 25 m, colour digital photos. Philippines:

USNM 370291; 1, 23.5; 13°10.04'N, 120°35'12'E, Mindanao L; MinOO-

62; 3 Jun 2000; 30 m; X-ray, colour digital photos.

Non-type material. Indonesia: USNM 209847; 3, 10.0-20.4; Ceram,

Pirn Bay; VGS 73-5; 9 Jan 1973; 5-7 m, WAM P.32771-002; 23.4;

01°36.252'S, 135°24.553'E, Tanjung Woka, Yapen L, Papua, Indonesia;

13 Feb 2006; 34 m. Philippines: USNM 369983; 2, 18.0-27.4; 13°45'N,

120°55'E, Anilao, Batangas; 26 Apr 1980; 22 m. USNM 268318; 1,

32.6; 9°04'38'N, 123°16'44'E, Apo L; SP78-36; 7 Jun 1978; 39 m.

Other material. Fowleria sp. AMS 1.23585-001; 1, 45.0; 14°34'S,

145°34' E, south-west of Carter Reef, Great Barrier Reef, Australia; 31

Jan 1982; 30 m, J. Leis. Apogonichthys nafae Holotype, USNM 62947;

22.1; Japan, Okinawa, Naha.

Diagnosis. A species of Neamia with 7 visible first-dorsal

spines, 8th spine hidden by skin, darkish circular spot on the

opercle below opercular spine within a pale area; in life, body

uniformly reddish to brownish, all fins pale, 13-14 pectoral

rays, 5 predorsal scales, eye diameter 10-12%, pectoral fin

25-31% and body depth 37-40% of standard length.

Description. For general body shape see fig. la-d. Holotype

proportions and characteristics first with range of values in

parentheses for paratypes and other material. Proportions as %

of standard length: greatest body depth 37.4 (34-40); head

length 44.6 (41-46); eye diameter 10.0 (11-12); snout length

8.3 (8-10); bony interorbital width 3.7 (4-6); upper jaw length

20.3 (20-22); caudal peduncle depth 15.7 (15-17); caudal

peduncle length 20.9 (19-22); 1st dorsal-fin spine length 1.7

(2-4); 2nd dorsal-fin spine length 6.6 (6-10); 3rd dorsal-fin

spine length 16.9 (16-21); 4th dorsal-fin spine length 16.0 (16-

19); spine in 2nd dorsal fin 9.7 (9-12); 1st anal-fin spine length

2.0 (2-5); 2nd anal-fin spine length 10.0 (8-12); pectoral fin

length 28.9 (25-31); pelvic fin length 25.7 (24-30).

Dorsal fin VII(I)-I,9, 8th spine hidden under skin, 3rd and

4th spines about the same thickness; anal fin 11,8; pectoral fin

14; pelvic fin 1,5; principal caudal rays 9-1-8; caudal fin rounded;

pored lateral-line scales 22 (11-12 for specimens 17-20 mm

SL, 18 for specimens 20-24 mm SL, 22-23 in specimens

greater than 26 mm SL); transverse scale rows above lateral

line 2; transverse scale rows below lateral line 6; median

predorsal scales 5; circumpeduncular scale rows 12 (5-I-2-I-5).

Total rudiments and gill rakers 15 (13-15), 2-I-1-5-I-7 (1-2 -1-1-5-

6-I-5-7), well developed gill rakers 6 (6-7), upper arch 1(1),

lower arch 5 (5-6). Rudiments on lower arch as flattened tooth

patches. 2nd arch with 1 short, nob-like raker in angle, upper

arch with 1-2 flattened tooth patches, lower arch with 5-6

nob-like rakers followed by flattened tooth patches.

Villiform teeth in a band of about 5 rows becoming 2-3

rows on side of premaxilla; villiform band of about 5 rows

becoming 2-3 rows on side of dentary; 3-4 rows of villiform

teeth on vomer; none on palatine, ectopterygoid, endopterygoid

or basihyal.

Vertebrae 10-1-14. 3 hypurals 1-1-2 fused, 3-1-4 fused to

urostylar centrum, 1 pair uroneurals, 2 large epurals (a 3rd

epural could not be identified from the radiograph), a free

parhypural. 3 supraneurals, 2 supernumerary spines on 1st

dorsal pterygiophore, 8th dorsal-fin spine hidden as a nubbin.

Basisphenoid status unknown, lower portion apparently absent,

but unclear on X-ray. Status of suspensory pharyngeal

unknown. Supramaxilla present. Posttemporal smooth on

posterior margin. Preopercle smooth on vertical and horizontal

margins. Infraorbital edges smooth.

Cephalic pored sensory system (fig. 2) - all larger pores

bilateral. No variation for the following pores: 2 nasal pores at

anterior end of supraorbital canal system, 1 medial to tubular

anterior nare, 1 slightly anterior and medial to posterior nare,

1 near edge of orbit on interorbital; 4 infraorbital pores at

anterior part of infraorbital canal, 1 between ventral edge of

lachrymal and flat posterior nare, 2 on ventral edge of

lachrymal, 1 on ventral edge of 2nd infraorbital; dentary with

3 pores at anterior end of mandibular canal, 1 on tip of lip, 1

behind lip, and 1 lateral; 7 articulo-preopercular pores near

posterior edge along the preopercular canal. Variation in

number and location of minipores: supraorbital with none or 1

pore near edge of orbit on interorbital area; paired pores along

upper and lower portions of infraorbitals below orbit; dentary

and articular with 4 pairs (inner and outer) of posterior pores,

with several unpaired pores on dentary and articular;

preopercular minipores along narrow upper portion of

preopercle. Minipores variably present posterior to eye on

upper portion of infraorbitals, on nape and above preopercle

ending on the post-temporal.

All scales cycloid on head, nape and breast. Scales

becoming ctenoid above breast and behind pelvic fin. Scales

above lateral line ctenoid. All lateral-line scales ctenoid,

except first. Pored lateral-line scales simple (with 1 opening

above and below main pore), extending from post-temporal to

base of caudal fin.

Life colours. Colours based on photographs are from fresh

dead specimens shortly after collection. From B. M. Carson-

Ewart photograph (AMS 1.40666-012) - body, pelvic, caudal

and dorsal fins orangish red, flank grading to white on abdomen;

head without marks extending from orbit edge, lips whitish;

circular spot on opercle yellowish with faint outer margin of

melanophores dorsally, large inner circle of dark melanophores

about size of pupil; iris whitish with narrow inner ring of light

yellow surrounding pupil.

From B. M. Carson-Ewart photograph (AMS 1.40666-013)

- body and caudal fins a pale orangish red to more yellowish

posteriorly, with brownish marks in caudal fin, flank grading to

white on abdomen; head without marks extending from orbit

edge, lips whitish; circular spot on opercle yellowish with outer

margin ring (^-270°) of melanophores except posteriorly, large

inner circle of dark melanophores about size of pupil; iris whitish

with narrow inner ring of light yellow surrounding pupil.

From D. W. Greenfield photograph (CAS 223500) - body

and caudal fin dark brownish, head without markings extending

Thomas H. Fraser and Gerald R. Allen

from orbit edge, lower half of head and breast tannish white

extending posteriorly to opercular spot and then above

pectoral-fin base descending to near anal base; pelvic fins

whitish; spot on opercle whitish tan with inner darkish ring

smaller than eye diameter; iris tannish white with narrow

inner ring of light yellow surrounding pupil.

From J. T. Williams’ photograph (USNM 370291) - body

generally uniform brownish, with 2 rows of small spots along

caudal peduncle, about 5 pairs, 3 spots extended anteriorly in

the lower row with 2 spots below the 3 spots; head without

markings extending from orbit edge; all fins generally pale

with numerous small melanophores in caudal fin; round spot

on opercle with complete brownish outer edge of body colour

around pale area with large dark inner spot; narrow whitish

bars on lower lip; iris orangish.

Colour in ethyl alcohol. Holotype uniform light tannish in

alcohol with tiny melanophores on head, body, both dorsal fins,

denser on membranes of 2nd dorsal fin, melanophores absent

on anal, pelvic and pectoral fins; large rounded dark spot on

opercle, surrounded by pale area, melanophores at edge of pale

zone forming about a half-circle; faint series of melanophores

in a horizontal line from lower orbit to preopercular ridge (fig.

3). Pale stomach, intestine and peritoneum. Paratypes and non-

type material uniform brownish to tannish in alcohol with tiny

melanophores on head, body, both dorsal fins, denser on

membranes of 2nd dorsal fin, melanophores absent on anal,

pelvic and pectoral fins; large rounded dark spot on opercle,

surrounded by pale area, melanophores at edge of pale zone

forming about a half-circle; faint, darkish, horizontal line of

melanophores from lower orbit to preopercular ridge; lips with

a few narrow pale bars or lips indistinct.

Etymology. The Greek words artios, complete or perfect and

cyclus, circle as articycla an adjective referring to the dark spot

on the opercle with its edge as a complete or perfect circle.

Distribution. The new species is apparently widespread in the

western Pacific from Sumatra to Fiji, and eastern Australia

north to the Philippines (fig. 4). Although it has been collected

at relatively few locations, future collections will no doubt

expand the known distribution. Due to their cryptic habits, the

species of Neamia are seldom observed by divers, rather they

are collected with ichthyocides, dredges and sleds. Collection

data suggest N. articycla frequents the 10-40 m depth zone, in

association with beds of a calcareous algae, Halimeda or

calcareous rubble, sometimes mixed with the pocilloporid

coral Seriatopora hystrix. The URL http://www.amonline.net.

au/fishes/about/research/halimeda.htm has more information

about other fishes collected in such Halimeda beds and a link

to colour photographs of this species.

Remarks and discussion. Neamia articycla has an ocellated

opercular spot that in life lacks an outer dark margin posteriorly,

but in preservation has a darkish edge from dorsal edge abutting

preopercle around to ventral edge abutting preopercle. It has no

darkish marks behind the eyes in life, but in preservation does

have a faint, narrow, horizontal line of melanophores below the

eye reaching the preopercle ridge. In contrast, N. notula has an

oval spot flattened along the preopercle edge with a darkish

edge outside of the pale area in life, four predorsal scales instead

of five, usually 22 pored, lateral-line scales instead of 23 and a

longer pectoral fin. Neamia articycla can be distinguished from

Neamia octospina by having 14 pectoral rays instead of 17-21

and by having seven visible first-dorsal spines instead of eight.

Fraser and Allen, 2001 discussed the relationships of

Neamia with other apogonid genera. Neamia articycla does

not have characters that alter that discussion. Neamia articycla

is more similar to N. notula based on the presence of the

opercle spot, the reduced, hidden, eighth dorsal spine, larger

eye and the lower pectoral fin-ray count than to N. octospina.

Specimens of N. articycla less than about 25 mm SL have

incompletely pored lateral-line scales (the number of pored

scales varies with size) and may therefore be confused with

Fowleria, most species of which have an incomplete lateral

line - see the figure of Fowleria nafae, Snyder, 1909 in Snyder,

1912 one of the nominal species originally described as

Apogonichthys nafae.

Some similarities exist between N. articycla and species of

Fowleria. The ocellus-like opercular spot, tubular anterior nare,

lack of palatine teeth and body shape are characteristics N.

articycla shares with most species of Fowleria. However, these

genera can be distinguished by fused caudal skeletal

characteristics in Neamia, completely pored lateral-line scales

in adult Neamia, (number of pitted scales in N. articycla

decreases posteriorly with growth, unlike Fowleria), and

cycloid scales on the head and anterior portions of the body

(weakly ctenoid scales on nape for species of Fowleria,

elsewhere on head weak to strongly ctenoid scales and ctenoid

on body). Bergmann, 2004 noted some variation in the pore

pattern on the dentaries for the four species of Fowleria

examined. None appeared to be similar to N. articycla in spatial

organization, with F. vaiulae perhaps having a few of its slightly

larger minipores spaced and located in a similar fashion.

Ida and Moyer, 1974, fig. 3 compared the cephalic lateralis

and free neuromasts of five genera of cardinalfishes (including

three subgenera), Apogon {Ostorhinchus), Apogon (Apogon),

Fowleria, Cheilodipterus,Rhabdamia(Verulux), and Pseudamia.

Bergmann, 2004 provided a more extensive survey of these

systems for these same genera and another 30 apogonid genera

and subgenera in two subfamilies. Species of Neamia differ

most obviously from illustrations of Fowleria, Foa, and

Apogonichthys in having fewer pores on the dentary. A

comparison of the descriptions of Neamia suggest that N.

octospina and the holotype of N. articycla show similarities in

the three slit-like pores along the ventral edge of the lachrymal

(first infraorbital) and second infraorbital, three suborbital pores

and the paired dentary pores. Differences were apparent: location

of the anterior-most supraorbital pore back near the anterior

nare in N. articycla, slit-like in N. octospina located along the

margin with the premaxilla; larger pores along the posterior

edge of the subopercle-preopercle in N. articycla than for N.

octospina, and no mini-pores just posterior to the preopercle

ridge in N. articycla versus numerous minipores; few paired

pores on the dentary and articular in N. articycla versus

numerous paired pores; no minipores in N. articycla versus

many minipores on snout and interorbit; and fewer minipores on

the nape in N. articycla versus widespread and numerous.

A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean

Ida and Moyer, 1974 failed to indicate the taxonomic value

of pore characteristics, although their fig. 3 was suggestive.

Bergmann, 2004 found four characteristics of the cephalic

lateralis which were shared by all cardinalfishes examined.

Neamia articycla has these four characteristics: a terminal

lachrymal pore, a pair of ventral lachrymal pores, an anterior

dentary pore in the lower lip and a mental (ventral dentary

pore) immediately behind the lower lip. Bergmann, 2004

identified eight characteristics that had variation, and proposed

that these various states had phylogenetic value within

cardinalfishes. Neamia articycla has: (1) the terminal end of

the supra-orbital canal ending in a large pore near anterior nare;

(2) the lateral margin of the supra-orbital canal near the

posterior nare with a single large pore; (3) the lateral margin of

the supra-orbital canal midway on interorbit with a large pore,

but also with an additional smaller anterior pore (variably

present) apparently intermediate to having multiple canal

projections; (4) the postorbit pores with simple canal projections

with minipores; (5) the supratemporal canal anterior margin

smooth; (6) the supratemporal canal posterior margin with few

canal projections with minipores; (7) the lateral margin of the

mandibular canal with relatively large and simple pores; and

(8) a terminal mandibular pore (here called the posterior

articular pore) near the boundary with the preopercle. The

usefulness of these eight characteristics and their character

states will be further developed through examination of many

species not yet reported in the literature. Many of the other

minipores had some variation in number and location among

individuals of N. articycla.

Acknowledgments

We thank D. G. Smith, J. T. Williams (USNM), David Catania

and David W. Greenfield (CAS), and Mark McGrouther (AMS)

who aided in curatorial processes and loan of fishes. D. G.

Smith provided X-rays. L. M. R. Bergmann allowed us the use

of illustrations and information from her dissertation. D. W.

Greenfield, J. M. Leis (taken by B. M. Carson-Ewart) and J. T.

Williams provided colour images of the new species. Useful

comments were provided by D. W. Greenfield, J. M. Leis and

an anonymous reviewer.

References

Bergman, L. M. R. 2004. The cephalic lateralis system of cardinalfishes

(Perciformes: Apogonidae) and its application to the taxonomy

and systematics of the family. Unpublished PhD Dissertation,

University of Hawaii, i-vi +1-226, 56 figs, 1 tab., 3 app.

Eschmeyer, W. N., 1998. Introduction. Pp. 16-22 in: W. N. Eschmeyer,

ed.. Catalog of fishes. Volume I. California Academy of Sciences,

958 pp.

Eraser, T. H. and G. R. Allen. 2001. A New Species of Cardinalfish in

Neamia (Apogonidae, Perciformes) with a Review of Neamia

octospina. Records of the Western Australian Museum 20(2): 159-

165, 2 figs, 1 tab.

Eraser, T. H. and E. A. Lachner. 1985. A revision of the cardinalfish

subgenera Pristiapogon and Zoramia of the Indo-Pacific region

(Teleostei: Apogonidae). Smithsonian Contributions to Zoology

412:1-47, 20 figs, 4 tabs.

Ida, H. and J. T. Moyer. 1974. Apogonid fishes of Miyake- Jima and

Ishigaki-Jima, Japan, with descriptions of anew species. Japanese

Journal of Ichthyology, 21(3): 113-128, 5 figs, 3 tabs.

Leviton, A. E., R. H. Gibbs, Jr., E. Heal and C. E. Dawson. 1985.

Standards in Herpetology and Ichthyology: Part I. Standard

symbolic codes for institutional resource collections in

Herpetology and Ichthyology. Copeia, 1985(3):802-832.

Radcliffe, L. 1912. Descriptions of fifteen new fishes of the family

Cheilodipteridae from the Philippine Islands and contiguous

waters. Proceedings of the United States National Museum,

41(1868):431-446, pls.34-38.

Snyder, J. O. 1909. Descriptions of new genera and species of fishes

from Japan and the Riu Kin Islands. Proceedings of the United

States National Museum, 16(1688):597-610.

Snyder, J. O. 1912. Japanese shore fishes collected by the United States

Bureau of Eisheries steamer ‘Albatross’ expedition of 1906.

Proceedings of the United States National Museum,42(l909):399-

450, pis. 51-61.

Memoirs of Museum Victoria 63(1): 7-13 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Description of two new species of Nesogobius (Pisces: Gobioidei: Gobiidae) from

southern Australia

Douglass F. Hoese^ and Helen K. Larson^

'Fish Section, Australian Museum, 6 College St., Sydney, NSW 2010, Australia (Doug.Hoese@austmus.gov.au)

^Museum and Art Gallery of the Northern Territory, PO Box 4646 Darwin, NT 0801 Australia (Helen.Larson@nt.gov.au)

Abstract Hoese, D.F. and Larson, H.K. 0000. Description of two new species of Nesogobius (Pisces: Gobioidei: Gobiidae) from

southern Australia. Memoirs of Museum Victoria 63(1): 7-13.

Two new species of Nesogobius are described from southern Australia and compared with described species

Nesogobius pulchellus Castelnau and Nesogobius hinsbyi (McCulloch and Ogilby). Nesogobius greeni, sp. nov. differs

from N. pulchellus in lacking a spine in the second dorsal and anal fin and in lacking head pores. Nesogobius maccullochi

sp. nov. differs from N. hinsbyi in having head pores and fewer dorsal spines and dorsal and anal rays.

Keywords Fish, Gobioidei, Gobiidae, gobies, Nesogobius, new species

Introduction

The genus Nesogobius is confined to southern Australia. The

species live over and bury in fine sand. Currently the genus

contains two previously described species, Nesogobius hinsbyi

(McCulloch and Ogilby) and Nesogobius pulchellus

(Castelnau). Several species are known (Hoese and Larson,

1994). The two species described here have been confused

with the previously described species and are named to remove

that confusion. Two species have been confused under the

name Nesogobius hinsbyi. That species is known from deep

water, but the name has been incorrectly applied to a common

shallow-water species. Similarly two species have been

confused under the name Nesogobius pulchellus, but both of

these species are known from shallow water and often occur

together. Other species will be treated in a more comprehensive

review of the genus in preparation by the authors.

Hoese (1991) suggested a possible relationship of Nesogobius

with Tasmanogobius, also restricted to southern Australia. Both

genera have a poorly developed posterior connection of the

neural arch to the centrum on all vertebrae. The genus is atypical

in having a mixture of characteristics of taxa belonging to the

Gobionellinae and the Gobiinae. For example, species of

Gobionellinae normally have two epurals (Pezold, 1993), but

Nesogobius is variable, with some species having one epural

and some two. In gobionellines, the interorbital canal is

separated anteriorly between the eye, with paired anterior

interorbital pores and posterior and anterior nasal pores. In

gobiines, when pores are present, the canals are fused between

the eyes, usually with a median anterior interorbital pore (only

paired in one species of Glossogobius or rarely paired in

aberrant specimens) and no anterior nasal pore. In Nesogobius,

when pores are present, the head canals are fused between the

eyes, with a single anterior interorbital pore, as in gobiines, but

with posterior and anterior nasal pores as in gobionellines.

Methods

Counts and measurements follow Hoese (1991). Institution

abbreviations follow Levinton et al. (1985). The longitudinal

scale count was taken from behind the pectoral base to the end

of the caudal peduncle and is a count of scale rows, rather than

a straight line count. The transverse scale count (TRB) is taken

from the anal origin upward and backward to the second dorsal

base. In descriptions an asterisk indicates count of holotype.

Comparative material examined: Nesogobius hinsbyi -

Holotype TMH D.142, 1(62), Derwent R., Tas. AMS 1.22569-

001, 5(37-50), Derwent R., Tas. Nesogobius pulchellus - AMS

1.16251-001, 7(37-42), Sydney Harbour, NSW.

Nesogobius Whitley, 1929

Nesogobius Whitley, 1929: 62 (type species Gobius hinsbyi

McCulloch and Ogilby, 1919, by original designation).

The genus is unique in Australia in having only 13 segmented

caudal rays. The genus contains 11 species all confined to

southern Australia from central New South Wales to Perth,

Western Australia. Most species are known only from Victoria,

Tasmania and South Australia.

Douglass F. Hoese and Helen K. Larson

Nesogobius greeni sp. nov.

Figures 1-4

Nesogobius sp. 2.- Last, Scott and Talbot, 1983; 450, fig. 30.127

(Tas.).

Nesogobius sp. 3. - Hoese and Larson, 1994: 797, fig. 703

(southern Australia)

Material examined. Holotype: AMS 1.25944-011, 30.5 male, Richards

Point, Port Phillip Bay, Jan 1981, R. Kuiter. Paratypes: NSW; AMS

1.20021-006, 1(27), Merimbula. Vic.: AMS 1.25944-001, 6(17-30),

taken with holotype; NMV A.29344-001, 2(30-30), Crib Point,

Western Port, 4 Sep, 1974, Melbourne Univ. Zool. Dept., 0415 hr;

NMV A.29345-001, 1(32), Crib Point, Western Port, 18 Oct, 1974,

Melbourne Univ. Zool. Dept., 1100-1130 hrs; NMV A.29346-001,

2(30-30), Crib Point, Western Port, 20 Aug, 1974, Melbourne Univ.

Zool. Dept., 0500 hrs; NMV A.29348-001, 6(27-29), reef at

Beaumaris, Port Phillip Bay, 9 Jun, 1967, R. Frankenburg; NTM

S.16206-001, 3(21-29), taken with holotype; AMS 1.22572-006,

2(25-26), Swan Bay, Port Phillip Bay. Tas.; AMS IA.3621, 6(16-32),

Southport, 1.5 fathoms, 9 Feb, 1928, T.T. Flynn; AMS 1.43821001,

1(24), D’Entrecasteaux Channel, P. Last; AMS 1.43824-001, 4, (26-

30) , D’Entrecasteaux Channel, P. Last, 3 Jul 1974; AMS 1.17549-001,

6(15-24), Oyster Cove, 1 Dec, 1972, D. Hoese and W. Ivantsoff; AMS

1.43825-001, 1(25), Margate, 11 Nov, 1973, T. Walker; AMS 1.43822-

001, 1(25), Margate, 12 Dec, 1973, T. Walker; AMS 1.43818001, 8(19-

31) , Margate, 27-28 Jan, 1974, T.M. Walker; AMS 1.17193-006, 2(27-

31), Wedge Bay, May, 1976, T. Garrard; AMS 1.43823-001, 1(25),

Margate, 16 Jun, 1976, T. Walker; CSIRO T.1400, 1 (37), Port Davey,

Kelly Basin, southwest Tas., 2 m. Mar 1979, P. Last; NMV A 29347-

001, (ex QVM 1972/5/425E), 10(32-37), Kelso, R. Greene, 5 Feb

1967; QVM 1972/5/2275, 3,(19-35), Greens Beach, 8 Jan, 1967; QVM

223, 5(17-37), Greens Beach, 26 May, 1972; QVM 224, 10(26-34),

Greens Beach, 5-7 Nov, 1966, R. Green; QVM 225, 6(26-34), Greens

Beach, 5 Feb, 1967, R.H. Green; QVM 226, 1(35), Greens Beach, 13

Jan, 1968, R. Green. SA: AMS 1.20178-010, 3(19-20), Pelican Lagoon,

Kangaroo L, 8 Mar, 1978, D. Hoese and Party.

Non-type material. Tas.; CSIRO (unreg.) 3(26-36),

D’Entrecasteaux Channel, 3 Jul 1974; CSIRO (unreg.) 4(26-32),

Fortesque Bay, 10 m, P. Last; CSIRO T.103, 1(28), Parsons Bay,

Nubeena, 2 Nov 1978; CSIRO T.123, 1(31), Bayview Beach, Georges

Bay, 2 Nov 1978; CSIRO T.1665, 1(26), Dm Point, 10 Apr 1980,

University of Tasmania; CSIRO T.174, 1(36), Ansons Bay, 25 Mar

1978,P.J. Miller; CSIRO T.185, 1(37), Ansons Bay, 11 Oct 1978;

CSIRO T.186, 1(29), Ansons Bay, 11 Oct 1978; CSIRO T.184, 1(38),

Boggy Creek Beach, St. Helens, Jul 1978.

Diagnosis. 1st dorsal fin VII; 2nd dorsal-fin rays 8-11, rarely

8 or 11; anal-fin rays 8-10; no spine in 2nd dorsal or anal fin;

branched caudal-fin rays 11-12; pectoral-fin rays 16-20; no

head pores; gill opening wide, reaching forward to below or

slightly before posterior preopercular margin; head with

scales reaching to above preoperculum, sometimes almost to

eye; dorsal mid-line of nape naked or rarely with a single

scale just before 1st dorsal fin, but scales often present just to

side of midline; body scales ctenoid, in 25-30 rows; midline

of belly without scales or with a few scales posteriorly;

pectoral base usually without scales or with 1 or 2 cycloid

scales ventrally; area before pelvic fin with cycloid scales;

body deep, depth at anal origin subequal to or greater than

caudal fin length; 1st dorsal fin low, with rounded or triangular-

shaped margin.

Description. Based on 44 males and 55 females. 1st dorsal

6(3), 7(74*); 2nd dorsal-fin rays 8-10 (see Table 1); anal-fin

rays 8(15), 9(77*), 10(25); pectoral-fin rays 16(6), 17(41*),

18(52), 19(2); segmented caudal-fin rays 13(54*), 14(2);

branched caudal-fin rays 10(1), 11(14*), 12(27), 13(4) midline

predorsal scales 0(38), 1(2); total gill rakers 4(2), 6(5), 7(3),

8(3), 9(3); lower gill rakers on 1st arch 4(2), 5(1), 6(7), 7(2),

8(4), 9(1); lower gill rakers on 2nd arch 4(1), 5(5), 6(5);

longitudinal scale count 25(6), 26(9), 27(16*), 28(7), 29(1),

30(1); TRB 7(3), 8(24), 9(2*). Head (28-32% SL), about as

broad as deep; mouth small, oblique, forming an angle of

30-40° with body axis, rear end of jaws below front quarter

of eye; tongue tip truncate to slightly emarginate; posterior

nostril at end of short tube, almost touching eye; anterior

nostril at end of short tube, positioned midway between eye

and upper jaw, close to posterior nostril; snout convex in side

view, forming an angle of about 45° with body axis; upper lip

thick anteriorly, thin posteriorly; lower lip thin with shallow

free ventral margin separating lip from mental frenum; chin

with round mental frenum with a small sensory papilla at

each side; eye large subequal to snout; gill rakers on outer

face of 1st arch 0-1 -i- 4-9 = 4-8, rarely 4 or 5; rakers very

short on both faces of all arches; teeth in upper jaw small,

conical and wide-set, 3-4 inner rows of close-set teeth

anteriorly tapering to 2 rows laterally; teeth in lower jaw

small, conical and wide-set in outer row, 3-5 inner rows of

smaller close-set teeth, rows tapering laterally to 1 row; body

robust, body depth at anal origin 19-22% SL. 1st dorsal-fin

origin above and just behind pelvic-fin insertion; 2nd dorsal-

fin origin just behind 1st dorsal fin; anal-fin origin below and

just behind 2nd dorsal-fin origin; pelvic-fin origin below

pectoral-fin insertion; pectoral-fin margin rounded; caudal fin

small, with rounded margin.

Head and body brown, green, or dark grey; lower surface

of head often dark grey to black; an irregular diffuse dark

brown blotch from eye to middle of jaws, and a vertically

elongate blotch below eye; body with 6-8 small dark-brown to

black spots on mid-side; dorsal midline often with black

blotches above each mid-side spot; a series of white dashes

just below mid-side; end of caudal peduncle with a <-shaped

mark, extending onto caudal-fin base, sometimes broken into

2-3 separate spots at apices of triangle; females with 1-3 dark-

brown vertical bars with white interspaces below 1st dorsal

fin; body with scattered mottling, often forming irregularly

shaped longitudinal lines; 1st dorsal fin with 2 black irregular

oblique bands, with orange interspaces; 2nd dorsal fin with

irregular oblique orange to brown stripes; anal fin grey; caudal

fin clear to grey; pectorals and pelvic fins clear to white in

males; pelvic fins almost black in female; pectoral base with

an elongate brown spot dorsally.

Variation. Sex ratios were found to be almost even with 44

males and 55 females. Too few specimens were available from

localities other than Tas. for a detailed analysis of variation.

However, 2nd dorsal ray counts average slightly higher in

southern Tas. Because of the slight differences in southern Tas.

material, most of that material is excluded from the type series.

Description of two new species of Nesogobius (Pisces: Gobioidei: Gobiidae) from southern Australia

Figure 1. Holotype of Nesogobius greeni, AMS 1.25944-001, 30.5 mm SL male, drawing by H.K. Larson.

Figure 2. Paratype of Nesogobius greeni, AMS 1.25944-002, 29.5 mm SL female, drawing by H.K. Larson.

Figure 3. Head of Nesogobius greeni, showing papilla pattern,

based on several specimens; size of papillae exaggerated, drawing

D.F. Hoese.

Table 1. Second dorsal rays in various populations of Nesogobius

greeni. Asterisk indicates count of holotype.

Northern Tasmania

Southern Tasmania

Victoria

New South Wales

South Australia

Douglass F. Hoese and Helen K. Larson

Figure 4. Underwater photo of Nesogobius greeni from type locality, photo R. Kuiter.

Distribution. Nesogobius greeni is known from Merimbula,

NSW, throughout Tas., Vic., and east to Kangaroo I., SA. The

species is normally found on sand around sea grass beds and

around rocky reefs from the intertidal to depths of 8 m.

Etymology. Named for R.H. Green formerly of the Queen

Victoria Museum, Launceston, Tas., who collected much of the

material used in this study.

Remarks. Nesogobius greeni is Nesogobius pulchellus

in its coloration and deep body. Both species are often collected

together at the same station. Nesogobius greeni differs from N.

pulchellus in lacking second dorsal and anal spines (present in

N. pulchellus), low first dorsal fin subequal in height to second

dorsal fin (versus usually higher than second dorsal).

Nesogobius maccullochi sp nov.

Figures 5-7

Gobius hinsbyi - McCulloch and Ogilby, 1919: 215, pi. 33, fig. 1

(in part, including figured specimen).

Nesogobius sp. 1 - Last, Scott and Talbot, 1983: 449, fig. 30.126

(Tas.); Hoese and Larson, 1994: 795, fig. 701 (southern Australia)

Material examined. Holotype AMS 1.17575-008, 64 mm SL female,

Pinalong Bay, Tas., 6 Dec 1972, D. Hoese and W. Ivantsoff.

Paratypes: Vic.: AMS 1.16987-007, 67(28-74), Peterborough, 21

Mar, 1972, D.F. Hoese and W. Congleton; AMS 1.16990-002, 4(42-

46), Port Phillip Bay, 23 Mar, 1972, D.F. Hoese and W. Congleton;

AMS 1.22943-001, 5(45-56), Rhyll, Phillip L, B. Rigby, 31 May, 1979;

NMV A.3254, 1(48), Bruthen Creek estuary, Gippsland, 6 Aug, 1979,

J. Buemer. SA: AMS 1.17575-003, 4(50-60), taken with holotype;

AMS 1.17629-001, 1(38), Salt Creek Bay, south of Coobowie, St.

Vincent Gulf, 0-1 m, 23 Dec, 1973, D. Hoese and Party; AMS 1.20184-

005, 4(35-42), Bay of Shoals, Kangaroo L, 11 Mar, 1978, D. Hoese

and B. Russell. Tas.: AMS 1.17562-002, 52(19-78), Browns R.,

Kingston, 0-1 m, 30 Nov, 1972, D. Hoese and W. Ivantsoff; AMS

1.17575-003, 6(57-68), inlet 6 km north of Binalong Bay, 0-1 m, 6

Dec, 1972, D. Hoese and W. Ivantsoff; NMV A.3257, 10(28-58),

Greens Beach, 8 Jan, 1967, R.H. Green; NTM S.16210-001, 1(47), St.

Helens, P. Last; QVM 220, 71(25-68), Kelso, 5 Feb, 1967, R.H. Green;

QVM 221, 31(27-47), Greens Beach, 17 Oct, 1965, R.H. Green; QVM

222, 1(28), Greens Beach, 5 Feb, 1967, R.H. Green.

Non-type material: Vic.: AMS 1.23456, 13(16-43), Stoney Point,

Western Port; NMV A.3553, 1(24), Ricketts Point, Port Phillip Bay, 17

Feb; NMV A.3523, 2(33-35), near Geelong; NMV A.2157, 3(54-65),

Portland Harbour; NMV A.3513, 1(40), Rye, Port Phillip Bay; NMV

A.3522, 1(47), Rye, Port Phillip Bay; NMV A.3527, 1(48), Crib Point,

Western Port; NMV A.3533, 1(51), Hovells Creek, near Geelong, Port

Phillip Bay; NMV A.3534, 3(42-50), Crib Point, Western Port; NMV

A.3537, 2(38-52), Crib Point, Western Port; NMV A.3538, 2(54-59),

Crib Point, Western Port, 3 Sep, 1974; NMV A.3539, 1(38), Crib Point,

Western Port; NMV A.3541, 2(37-41), Crib Point, Western Port;

NMV A.3542, 3(36-42), Crib Point, Western Port, NMV A.3548,

1(49), 3 km W of Sandringham, Port Phillip Bay, 30 Mar, 1971; NMV

A.3552, 1(41), Sorrento, Port Phillip Bay, 31 Jul, 1972. SA: AMS

1.20162-027, 1(40), Stokes Bay, Kangaroo L; AMS 1.20177-013, 1(45),

American R., Kangaroo I. Tas.: AMS 1.14200, 1(69), Wedge Bay,

paratype and figured specimen of Nesogobius hinsbyi.

Diagnosis. 1st dorsal fin VI-VIII, usually VII; 2nd dorsal-fin

rays usually I, 8-9; anal-fin rays usually I, 8, branched caudal-

fin rays usually 10; pectoral fin 17-21, rarely 21; an anterior

nasal pore medial to and slightly below level of each anterior

nostril and a posterior nasal pore by each posterior nostril, a

median anterior interorbital pore between front of eyes, a

median posterior interorbital pore between end of eyes, an

Description of two new species of Nesogobius (Pisces: Gobioidei: Gobiidae) from southern Australia

Figure 5. Holotype of Nesogobius maccullochi, AMS 1.17575-008, 64 mm SL female, photo D. Hoese.

Figure 6. Paratype of Nesogobius maccullochi AMS 1.14200, 69 mm SL male. Wedge Bay, from McCulloch and Ogilby (1919), image reversed.

Douglass F. Hoese and Helen K. Larson

Figure 7. Head of Nesogobius maccullochi, showing papilla pattern,

based on several specimens; size of papillae exaggerated, drawing by

D. Hoese.

infraorbital pore behind each eye and a lateral canal pore along

dorsal part of operculum, no preopercular pores; gill opening

wide, extending to below posterior preopercular margin; top of

head scaled to behind eyes, in 11-18 rows, from 1st dorsal-fin

origin; operculum and preoperculum (sometimes absent on

preoperculum) with a small patch of scales near dorsal margin,

remainder of head without scales; body scales ctenoid, in 37-49

rows; pectoral base and area before pelvic fin scaled (about 15

rows); 1st dorsal fin with rounded or triangular margin.

Description. Based on 109 males and 155 females. 1st dorsal

5(1), 6(8), 7(159*), 8(14); 2nd dorsal rays 1,7(3), 1,8(93*), 0,9(2),

1,9(89), 0,10(2), 1,10(3), anal rays, 1,7(19), 1,8(134*), 0,9(2),

1,9(28), 0,10(2), 1,10(1); pectoral rays 17(17), 18(71*), 19(62),

20(13), 21(1); predorsal scales 12(1), 13(7), 14(27), 15(31),

16(24), 17 (15), 18(1), 19(1*); segmented caudal rays 12(2),

13(78*); branched caudal rays 9(8), 10(43), 11(1), midline

predorsal scales 11(1), 13(4), 14(26*), 15(25), 16(30), 17(9),

18(1), total gill rakers 4(2), 6(5), 7(3), 8(3), 9(3); lower gill

rakers on 1st arch 4(9), 5(22), 6(1); lower gill rakers on 2nd

arch 4(17), 5(9), 6(4), 7(2); longitudinal scale count 37(1), 38(5),

39(6), 40(7), 41(11), 42(11), 43(8), 44(14), 45(6), 46(6*), 47(4),

48(4), 49(1); TRB 11(6), 12(14), 13(28), 14(18*), 15(4). Head

(29-32% SL), broader than deep; mouth small, oblique, forming

an angle of 20-25° with body axis, rear end of jaws below front

margin of eye; tongue tip rounded; posterior nostril at end of

short tube anterior to eye; anterior nostril at end of short tube

positioned anteroventrally from posterior nostril, separated

from posterior nostril by 2-3 nostril diameters; snout with an

elevated bump before eyes, formed by distal tips of ascending

process of maxilla; upper lip thick; lower lip thin with shallow

free ventral margin anteriorly; chin with a minute round lobe,

with sensory papillae from inner preopercular mandibular

papilla line meeting sides of lobe; eye large, slightly shorter

than snout length; gill rakers on outer face of 1st arch 0-1 -i-

3-5 = 4-6; rakers short on both faces of all arches, rakers on

outer face of 1st arch not larger than rakers on other arches;

outer row of teeth in upper jaw enlarged and directed posteriorly,

followed by 2 inner rows of smaller teeth tapering laterally to 1

row; outer row of teeth in lower jaw slightly enlarged and

curved posteriorly, 1 or 2 inner rows of smaller teeth, tapering

laterally to 1 row; body slender, body depth at anal origin 11-

13% SL. Body robust anteriorly, slender posteriorly. 1st dorsal-

fin origin just behind pelvic-fin insertion, dorsal fin low,

subequal to body depth at anal-fin origin; 2nd dorsal-fin origin

separated from 1st dorsal fin by 2-3 rows of scales, height of

2nd dorsal fin subequal to 1st dorsal fin; anal-fin origin below

and just behind 2nd dorsal-fin origin, anal fin slightly lower

than dorsal fins; pelvic-fin origin behind pectoral -fin insertion;

pectoral-fin margin rounded; pelvic and pectoral fins subequal

in length, slightly shorter than head length; caudal fin short,

length slightly shorter than pelvic-fin length, caudal fin with

truncate or slightly rounded margin.

Head and body light-grey to brown, often with scattered

white and brown flecks; a black bar from eye, extending across

middle of jaws; a black vertical bar from eye to just behind rear

end of jaws; a vertical bar just behind posterior preopercular

margin; mid-sides with 4-6 horizontally elongate dark brown

spots; a round black spot at rear end of caudal peduncle, followed

by and often connected to a black C-shaped mark at base of

caudal fin; mature males with a series of 6-12 vertical dark

brown bars on body extending onto belly; bars much thinner

than intervening spaces, but variable in width and position;

lower operculum, pectoral base, and belly white; dorsal and anal

fins with black spots forming more or less horizontal lines;

pectoral and caudal fins with small black spots forming wavy

vertical bands; pelvic fins white, often with irregular mottling.

Variation. Nesogobius maccullochi shows considerable

variation. Males differ considerably in coloration from females.

Overall females outnumbered males 1.5 times. However, only

two large samples were available to compare ratios and size.

One sample from Kelso, Tas. contains 46 females, 21 males

and four immature specimens. In a sample from Peterborough,

Vic., there are 33 females and 35 males. There was no significant

difference in sizes between males and females in either sample.

The largest female in all the samples is 78 mm SL and the

largest male 70 mm SL. In most samples the largest individual

was a female. Comparisons of fin-ray counts from various

populations showed no significant differences, but large

samples were available from few localities. The second dorsal

and anal spine are absent in less than 2% of individuals

examined.

Distribution. Tas., Vic., and SA. Normally found on sandy

areas in bays and estuaries, from the intertidal to depths of a

few metres.

Etymology. The species is named for A.R. McCulloch, formerly

Curator of Fishes at the Australian Museum. The name is given

not only in recognition of his work, but indicates that this is the

species which McCulloch and Ogilby confused under the name

Gobius hinsbyi.

Description of two new species of Nesogobius (Pisces: Gobioidei: Gobiidae) from southern Australia

Remarks. This species is the most abundant species in the

genus in shallow areas in southern waters. It has typically

been misidentified as Nesogobius hinsbyi. The species

differs from Nesogobius hinsbyi in having head pores

(absent in N. hinsbyi), opercular scales dorsally only (versus

operculum completely scaled), normally with seven dorsal

spines (versus usually eight) and second dorsal-fin rays

usually 1,8-9 (versus 1,9-10). Nesogobius hinsbyi occurs in

deeper water and is generally taken by dredge and trawl. It

should be noted that both species were included in material

used for the description of Gobius hinsbyi in McCulloch and

Ogilby (1919). The name originated from a Johnston

manuscript, where it was not described and was a nomen

nudum. That paper was later published by Whitley (1929).

The species was described by McCulloch and Ogilby (1919),

based largely on one specimen (AMS 1.14200), which is

figured here as Nesogobius maccullochi. McCulloch and

Ogilby (1919) mention the Tasmanian Museum specimen as

the type and give a brief description of the specimen and

indicate that they believed it to be identical to the described

specimen. It is regarded here that the use of the wording

“the type” clearly indicates that the holotype is the specimen

in the Tasmanian Museum. Eschmeyer (1998) listed the

specimen as a lectotype, indicating that he believed the

figured specimens should have been designated the lectotype.

Whether the Tasmanian Museum specimen is a lectotype or

holotype does not affect the identity of the species because

literature references to “the type” and to the lectotype refer

to the same specimen.

Acknowledgements

We thank T. Walker, R. Kuiter, R Last and R. Green for supplying

much of the material used in this study. Fresh material and photos

of live material were kindly provided by R. Kuiter. W. Ivantsoff

and W. Congleton assisted with field work. We are grateful to D.

Bray, T. Sim, R. Green and A. Graham for loan of material.

References

Eschmeyer, W.N. 1998. Catalog of Fishes. San Francisco: California

Academy of Sciences Vols 1-3 2905 pp.

Hoese, D.F. 1991. A revision of the temperate Australian gobiid

(Gobioidei) fish genus Tasmanogobius with a comment on the

genus Kimberleyeleotris. Memoirs of the Museum of Victoria

52(2): 361-376.

Hoese, D.F. and Farson, H.K. 1994. Family Gobiidae. pp. 781-810,

figs 690-714 in: Gomon, M.F., Glover, C.J.M. & Kuiter, R.H.

(eds) The Fishes of Australia's South Coast. Adelaide: State

Printer. 992 pp. 810 figs.

Fast, P.R., Scott, E.O.G. and Talbot, F.H. 1983. Fishes of Tasmania.

Hobart: Tasmanian Fisheries Development Authority 563 pp.

Feviton, A.E., Gibbs, R.H. Heal, E. and Dawson, C.E. 1985. Standards

in Herpetology and Ichthyology: Part 1. Standard symbolic codes

for institutional resource collections in Herpetology and

Ichthyology. Copeia 1985 (3): 802-832.

McCulloch, A. R. and Ogilby, J.D. 1919. Some Australian fishes of the

family Gobiidae. Records of the Australian Museum 12(10): 193-

291.

Pezold, F. 1993. Evidence for a monophyletic Gobiinae. Copeia

1993:634-643.

Whitley, G.P 1929. R.M. Johnston’s memoranda relating to the fishes

of Tasmania. Papers and Proceedings of the Royal Society of

Tasmania 1928: 44-68.

Memoirs of Museum Victoria 63(1): 15-19 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae)

from south-eastern Australia

Douglass F. Hoese^ and Sally Reader^

Abstract

Keywords

'Fish Section, Australian Museum, 6 College St., Sydney, NSW 2010, Australia (Doug.Hoese@austmus.gov.au)

^Fish Section, Australian Museum, 6 College St., Sydney, NSW 2010, Australia (Sally.Reader@austmus.gov.au)

Hoese D.F. and Reader S. 2006. Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae)

from south-eastern Australia. Memoirs of Museum Victoria 63(1): 15-19.

The present paper describes a single species from the coastal drainages in the Coffs Harbour area of New South

Wales. The species differs from the only previously described species, Philypnodon grandiceps, in having a narrower gill

opening (ending under posterior preopercular margin versus under eye), more vertebrae (30-32 versus 29-30), smaller

size (maximum size 60 mm SL versus 90 mm SL) and in coloration, particularly lacking the thin vertical bands on sides

of the belly characteristic of Philypnodon grandiceps. The new species occurs in freshwater and estuarine localities in

south-eastern Australia, often occuring with Philypnodon grandiceps.

Fish, Gobioidei, Eleotridae, gudgeon, Philypnodon, new species

Introduction

We follow Nelson (2006) here in recognising the family

Eleotridae as distinct from the Gobiidae. Hoese and Gill (1993)

recognised two subfamilies, the Butinae and the Eleotridinae,

placing them as subfamilies of the Gobiidae. Currently 18

described genera and 40 described species of eleotrid fishes are

known from Australia. Most are found in estuarine

environments. A number of genera of eleotrid fishes occurs in

freshwaters of Australia. Some genera, such as Bunaka Herre,

Eleotris Bloch and Schneider, Giuris Sauvage and Ophiocara

Gill are found in estuaries and freshwaters, although often in

the lower reaches of rivers. Other genera are largely confined as

adults to freshwater. This group includes Gobiomorphus Gill,

some species of Eleotris, most species of Hypseleotris Gill,

Kimberleyeleotris Hoese and Allen, Milyeringa Whitley,

Mogurnda Gill, Oxyeleotris Bleeker and Philypnodon Bleeker.

Philypnodon and Hypseleotris compressa (Krefft) occur in

both freshwater and estuarine environments as adults.

Philypnodon was long thought to be monotypic, with

Philypnodon grandiceps found in temperate coastal and inland

streams from south-eastern Australia. Hoese, Larson and

Llewellyn (1980) first noted that a dwarf species of Philypnodon

was found in southern Australia. Extensive collecting efforts

in recent years has yielded considerable material. Studies of

that material are revealing a high degree of variability both

within and between samples. Because of the high variability

in the dwarf forms, it was previously uncertain how many

species existed in the genus. The present paper describes a

single species from the coastal drainages in the Coffs Harbour

area of New South Wales in order to fix a single form for future

comparison. A more detailed discussion of geographical

variation and description of the other species will be treated in

a separate revision.

Methods

Counts and measurements follow Hoese and Allen (1987).

Institution abbreviations follow Leviton et al. (1985).

The longitudinal scale count was taken from behind the

pectoral base to the end of the caudal peduncle and is a count

of scale rows, rather than a straight line count. The transverse

scale count (TRDB) is taken from the second dorsal origin

downward and backward to the anal base. In descriptions

an asterisk indicates count of holotype. Measurements

were taken on the holotype and the ten best preserved paratypes

and are expressed as percent standard length. In the description

the first length given is for the holotype followed by the

range for the paratypes in parentheses. In lists of material

examined the registration number is given, followed by

the number of specimens, then the size range of the specimens

in parentheses.

Douglass F. Hoese and Sally Reader

Philypnodon Bleeker

Philypnodon Bleeker, 1874: 301, (type species: Eleotris nudiceps

Castelnau, 1872, by original designation), Syntypes MNHN 1509.

Gymnobutis Bleeker, 1874: 304 (type species: Eleotris

gymnocephalus Steindachner, 1866, by original designation and

monotypy), Syntypes NMW 22519

Ophiorrhinus, Ogilby, 1897: 745 (type species: Eleotris grandiceps

Krefft, 1864, by original designation) Syntypes BMNH 1864.7.22.40-

44; AMS 1.2671-2672.

The genus is distinctive in the following combination of

characters: no head pores; no scales on cheek or operculum;

nape naked to fully scaled; body fully scaled, largely with

ctenoid scales; midline of belly sometimes without scales;

anterior nostril at end of short tube above and almost in contact

with middle of upper lip, posterior nostril with raised rim slightly

in front of anterior margin of eye; usually seven dorsal spines;

first dorsal-fin origin well posterior to pectoral-fin insertion;

pelvic-fin origin well posterior to pectoral-fin insertion to almost

below pectoral-fin origin; transverse papilla pattern; large

mouth, longer in males than females; bulbous cheeks in adults;

wide gill opening extending to below eye or preoperculum; 15

segmented caudal rays; vertebrae 29-32; an intemeural gap

between neural arches after the first dorsal fin and before the

second dorsal fin, without a pterygiophore; vomer and palatine

without teeth. The genus is easily distinguished from other

eleotridine Australian genera by the combination of seven dorsal

spines and transverse papilla pattern.

The genus Philypnodon was placed within the subfamily

Eleotrinae of the Gobiidae by Hoese and Gill (1993), but did

not treat relationships within the subfamily. Thacker and

Hardman (2005) suggested a close relationship of Philypnodon

with the New World genera Microphilypnus and

Leptophilypnus, based on molecular studies. Both genera have

features in common with Philypnodon including an intemeural

gap (a space between neural arches after the first dorsal fin and

before the second dorsal fin without a pterygiophore; a derived

feature within the Eleotrinae) and a transverse papilla pattern,

a combination found only in these genera and in Thalasseleotris

in the Eleotrinae (Hoese and Gill, 1993).

The genus is confined to the freshwaters and estuaries of

south-eastern Australia from the Burnett R. in Queensland to

South Australia and from scattered localities in the Murray-

Darling River system in New South Wales and South Australia.

Philypnodon macrostomus sp. nov.

Eigures 1-3

Philypnodon sp: Hoese, Larson and Llewellyn, 1980: 171, fig. 31.3

Philypnodon sp. 1: Merrick and Schmida, 1984: 306, fig. 256

Philypnodon sp: Hoese and Larson, 1994: 810, fig. 715

Philypnodon species: Allen, Midgley and Allen, 2002: 317

Material examined. Holotype AMS 1.20111-002, 36.5 mm SL male,

creek 8 km from Glenreagh, N of Coffs Harbour, NSW, 5 May 1977,

D. Hoese and J. Bell. Paratypes: AMS 1.20111-016, 31(25-41), taken

with holotype; AMS 1.20111-003, 3(28-36), cleared and stained and-

004, 4(39-40), cleared and stained, taken with holotype; AMS

1.29683-008, 1(33), Karinga Creek W of Coffs Harbour, NSW, 19

Aug 1977, D. Hoese and R. McDowall; AMS 1.33875-001, 1(38),

Dingo Creek, 27 km W of Coffs Harbour, 5 May 1977, D. Hoese and

J. Bell; NMV A.29399-001, 4(21-36), taken with holotype; NTM

S.16213-001, 3(31-39), taken with holotype; QM 1.37768, 3(30-31),

taken with holotype.

Diagnosis. Gill opening wide, extending anteroventral to

preoperculum, but not reaching to below eye, usually ending

below posterior preopercular margin, sometimes as far forward

as midway between posterior preopercular margin and eye.

Head papillae in transverse pattern (figs 2-3). Body generally

dark brown, with darker brown mottling on sides; 1st dorsal

with black stripes in adult males. Jaws reaching to behind eye

in mature males, variably developed in juvenile males and

females, usually reaching below middle of eye to near end of

eye. Tongue tip rounded to truncate.

Description. An asterisk indicates count of holotype. 1st dorsal

fin VI (in 8*), VII (23), VIII (4); 2nd dorsal-fin rays 1,8 (8), 1,9

(2H), 1,10 (5); anal-fin rays 1,8 (10), 1,9 (22*), 1,10 (3); pectoral

rays 15 (4), 16 (21*), 17 (9). 18 (1) segmented caudal rays 8/7

(12), 8/8 (1*); branched caudal rays 7/6 (12*), 7/7 (1); procurrent

caudal rays 9/8 (1), 9/9 (1), 10/8 (1), 10/10 (1), 11/10 (3);

predorsal scale count 4(1), 5(1), 7(1), 8 (3), 9 (2), 10 (1), 11 (6),

12 (2), 13 (6), 14 (6*), 15 (1); longitudinal scale count 34 (5), 35

(6*), 36 (6), 37 (10), 38 (5), 39 (1), 42 (1); transverse scale count

(TRDB) 9 (16*), 10 (12), 11 (7); gill rakers on outer face of 1st

arch 2+1+8 (7), 3+1+8 (1), 2+1+9 (22), 2+1+10 (2); lower rakers

on outer face of 2nd arch 7 (9), 8 (22), 9 (3); vertebrae 12+19

(3), 13+18 (2), 13+19 (2).

Head distinctly depressed, flat on top, length 29.5% SL in

holotype (range 29.5-33.6% SL); eyes dorso-lateral, 6.8% SL

(6.4— 7.4% SL), interorbital wide, slightly less than eye diameter

in juveniles to about 1.5 times eye diameter in adult; snout short,

about equal to eye diameter, 8.1% SL (7.8-9.2% SL); cheeks

distinctly bulbous in adult males; mouth slightly oblique forming

an angle of about 35^5° with body axis; posterior margin of jaws

below mid-eye in females, to below or just beyond posterior end

of eye in males, length of upper jaw 15.4% SL (13.7-18.7% SL)

in males and 10.2-14.8% SL in females; teeth in both jaws small

and curved, anteriorly in 2 or rarely 3 rows, tapering laterally to a

single row; posterior nostril with raised rim, approximately 1

nostril diameter in front of eye in a horizontal line between mid-

eye to upper pupil margin; anterior nostril at end of short tube,

just behind middle of upper lip, in a horizontal line between mid-

eye to lower margin of pupil; gill rakers slender, much shorter

than filament length on outer face of 1st arch, longest rakers near

angle of arch about one-quarter length filament length; rakers on

inner face of 1st arch and following arches short and denticulate.

Body slender, slightly compressed anteriorly, becoming

very slender posteriorly; depth at pelvic-fin origin 14.6% SL

(14.5-18.8% SL); depth at anal-fin origin 14.1% SL (13.1-17.7);

caudal peduncle slender and elongate, length 29.8% SL (24.1-

30% SL), least depth 9.2% SL (8.2-10.5% SL).

Head naked; predorsal partly scaled, midline scaled to above

posterior preopercular margin to midway between posterior

preopercular margin and eye (just before posterior preopercular

margin in holotype), often naked in specimens less than 15 mm

SL; scales on side of nape variable, sometimes with scales as far

forward as midline scales (in holotype), but often reduced.

Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae) from south-eastern Australia

Figure 1. Holotype of Philypnodon macrostomus, AMS 1.20111-002, 36.5 mm SL male.

sometimes naked forward of pectoral-fin base. Body scales

ctenoid; body fully covered with scales, becoming cycloid above

a line from upper gill attachment to below posterior quarter of

1st dorsal fin; belly covered with small cycloid scales, midline

naked in specimens with reduced predorsal scales, pectoral-fin

base normally covered with small scales, naked in specimens

with reduced predorsal scales, prepelvic area covered with small

cycloid scales, except for a triangular area behind gill opening to

fully naked in specimens with reduced predorsal scales.

1st dorsal fin low, with rounded margin, fin reaching to or

just short of 2nd dorsal fin when depressed; 1st dorsal spine

slightly shorter than 2nd spine, 2nd to 4th spines subequal in

length, 5th spine subequal in length to 1st spine; 6th spine

slightly shorter than 5th spine, 7th spine shorter than 6th spine;

2nd dorsal fin separated from 1st dorsal fin, fin elevated,

slightly higher than 1st dorsal fin, subequal to body depth, 1st

segmented ray usually branched, other rays always branched;

anal fin origin below and slightly behind 2nd dorsal-fin origin,

usually below 2nd segmented dorsal ray, fin subequal in height

to 2nd dorsal fin, 1st segmented rays usually branched, other

rays branched; pelvic fins completely separate, pelvic fin short,

reaching approximately half distance from pelvic-fin origin to

anal-fin origin, length 18.4% SL (14.1-18.4% SL), pelvic-fin

rays 1,5, all segmented rays branched, rarely innermost ray

unbranched; pectoral fin with broadly rounded margin,

reaching to above or slightly before anus, length 22.8% SL

(19.6-24.2% SL), pectoral rays branched, upper and lowermost

sometimes unbranched; caudal fin with rounded posterior

margin, caudal length 17.9% SL (17.0-25.1% SL).

Urogenital papilla of male flattened dorso-ventrally, with

small lobes laterally; papilla of female broad and rounded,

with an indentation posteroventrally.

Figure 2. Lateral view of head showing sensory papillae, largely

based on holotype, drawing by S. Reader.

Sensory papillae. Transverse papillate pattern; all papilla lines

with papillae at right angle to axis of line. Sides of head with 6

lines radiating from eye; preopercular mandibular series with

10-14 transverse lines from chin to middle of preoperculum.

Other papillae as shown in figs 2 and 3.

Coloration of freshly collected specimens, (holotype and male

paratype). Head and body dark brown, lighter brown to pale

orange ventrally. Lips dark brown with diffuse orange pigment

on posterior half; 2 dark-brown stripes extending postero-

ventrally from posteroventral margin of eye onto cheeks, not

reaching posterior preopercular margin; a large light-brown oval

area behind tips of dark stripes; an irregularly shaped light-

brown stripe extending from below eye posteroventrally to just

Douglass F. Hoese and Sally Reader

Figure 3. Dorsal view of head showing sensory papillae, largely

based on holotype, drawing by S. Reader.

above posterior end of jaws; operculum with light-brown almost

horizontal stripe in line with interspace between dark stripes

extending from eye; a similar light-brown stripe dorsally on

operculum; rest of operculum dark brown; body scales edged in

dusky -brown pigment, dorsally broadly edged with dark-brown

pigment, scale centres often with light thin vertical line; series of

2-3 small black spots above pectoral-fin base; pectoral-fin base

black dorsally followed by white irregular bar extending full

length of pectoral-fin base; caudal peduncle with dark-brown

vertical bar, with a short anterior extension. Dorsal fin with 2

black curved stripes following contour of fin; distal margin of

dorsal fin with broad orange band, followed by black stripe

curving to meet body, followed ventrally by a lighter orange to

brown stripe, followed ventrally by 2nd black stripe; anteriorly

white blotches below stripe; 2nd dorsal fin with black distal

margin (about one-quarter of fin), followed ventrally by 4

curved, oblique stripes, with orange to white interspaces. Anal

fin with scattered melanophores giving fin a gray appearance;

base of fin lighter, followed distally by pale black stripe followed

by lighter stripe, followed by dark gray stripe, distal tip lighter

than rest of fin. Pectoral and pelvic fins white to dusky. Caudal

fin pale orange basally, becoming dark gray posteriorly; series

of 2^ curved dark bands, extending across all, but uppermost 1

or 2 and lowermost 1 or 2 segmented caudal rays, each band

formed from 2-3 rows of small black spots on fin rays, generally

no spots on membranes between rays.

Coloration in alcohol. Head and body light brown, paler

ventrally, males generally darker than females. 2 faint brown

stripes extending posteroventrally from posteroventral margin

of eye onto cheeks, not reaching posterior preopercular margin;

distinct light-brown areas and stripes on head present in fresh

material not visible; operculum light brown; body scales edged

in light-brown pigment, dorsally broadly edged with dark-brown

pigment; large black spot above pectoral-fin base with pale

areas; black spot dorsally on pectoral-fin base, a thin brown

band extending from spot ventrally covering pectoral-fin base,

no pale band visible; large dark-brown area at posterior end of

caudal peduncle forming a vertical band, a vertical band with a

horizontal extension, or triangular mark. Dorsal fin with 2 black

curved stripes following contour of fin as in fresh material,

orange and white areas in fresh material becoming light brown;

2nd dorsal fin with black to gray distal margin (about one-quarter

of fin), followed ventrally by 4 curved, oblique dark stripes, with

light-brown interspaces. Anal fin with scattered melanophores

giving fin a uniform gray appearance. Pectoral and pelvic fins

clear, without pigment. Caudal fin pale light-brown to gray, with

series of 1-A curved dark bands, extending across all but

uppermost 1 or 2 and lowermost 1 or 2 segmented caudal rays,

each band formed by 2-3 rows of small dark brown spots on fin

ray, generally no spots on membranes between rays.

Distribution. Philypnodon macrostomus is widely distributed in

coastal rivers from Brisbane, Qld, throughout NSW, Vic. and

eastern SA in freshwater and in brackish to full strength seawater

in upper reaches of estuaries. It is also found in the Lower Murray

R. system from scattered localities. The related Philypnodon

grandiceps occurs north to the Burdekin R., Qld. Specimens of

the dwarf species from the Mary R. north of Brisbane are

currently under study, but do not appear to be conspecific and

will be reported on later in a revision of the genus. Similarly,

specimens from the Cudgegong R. in Macquarie R. drainage in

central NSW lack predorsal scales and possibly represent a

separate species to be treated in the revision.

Etymology, from the Latin macro - large and stomus - mouth,

alluding to the large mouth that develops in males.

Remarks. Philypnodon macrostomus differs from Philypnodon

grandiceps in having a narrower gill opening (below posterior

preopercular margin versus below eye in P. grandiceps)-, sides

of belly without vertical bands (versus 4-5 thin brown almost

vertical lines); tongue tip rounded to truncate (versus bilobed);

body often mottled (versus body uniformly coloured) and

having more numerous vertebrae (usually 31-32, versus usually

Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae) from south-eastern Australia

29-30). The species also reaches a much smaller size, usually

less than 50 mm SL, with the largest specimen known 60 mm

SL (versus commonly reaching a size of 70 mm SL and reaching

a maximum size of almost 90 mm SL).

While the species is widely distributed, it shows considerable

variation over the range of the species. The intensity of

coloration varies from light brown to almost black. Fin ray

counts vary considerably, but without any clear geographical

pattern. More detailed analysis of geographical variation will

be presented in a revision by the authors at a later time.

The differences in the urogenital papilla were minor in

comparison with other gobioid fishes and it was difficult to

sex the species. Poor preservation can result in distortion of

the papilla.

Acknowledgements

We thank J. Bell and R.M. McDowall for assisting with

collection of type material.

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Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei).

Molecular Phylogenetics and Evolution 37: 858-871.

Memoirs of Museum Victoria 63(1): 21-24 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Description of a new species of Heteroclinus (Blennoidei: Clinidae) from southern

Australia

By Douglass F. Hoese^ and Denise S. Rennis^

'Fish Section, Australian Museum, 6 College St., Sydney, NSW 2010, Australia (Doug.Hoese@austmus.gov.au)

^Folly Beach, South Carolina, U.S.A.

Abstract Hoese, D.F and D. S. Rennis 2006. Descriptions of a new species of Heteroclinus (Blennoidei: Clinidae) from southern

Australia. Memoirs of Museum Victoria 63(1): 21-24.

A new species of Heteroclinus is described from Victoria, South Australia and Western Australia. The species is

distinctive from other Australian clinids in having two well-developed segmented pelvic rays, first dorsal fin on head

originating just behind or at posterior end of eye, a reduced lateral line and in fin-ray counts.

Keywords Fish, Blennioidei, Clinidae, Heteroclinus, new species

Introduction

Clinid fishes represent one of the most speciose groups of fishes

from southern waters of Australia. Over 35 species are known,

with only three extending to the tropics or southern parts of the

tropics. George and Springer (1980) reviewed the species of the

tribe Ophiclinini. Hoese (1976) described one new species of

Heteroclinus and reviewed the historical studies of clinid fishes

of Australia. Rennis et al. (1994) placed the temperate species

of the tribe Clinini into the Heteroclinus and Cristiceps.

The description provided here forms part of a revision of the

remaining clinid fishes of Australia and contrasts all the species

in the Heteroclinus adelaidae complex.

Currently three genera of the tribe Clinini are known from

Australia: Springeratus, with a single species, Cristiceps with

three and Heteroclinus with 25 species. Further studies on the

relationships within the group may alter the generic

classification. One species {Heteroclinus flavescens found in

southern Australia and New Zealand) has been regarded as

belonging to the separate genus Cologrammus. More detailed

information on relationships will be included in our revision

of the genus Heteroclinus.

Methods

Counts and measurements largely follow those given by Hubbs

and Lagler (1958). The last anal ray and last dorsal ray are

separate, not branched and are counted as separate rays. In

lists of material examined, institution abbreviations follow

Leviton et al. (1985). The number of specimens is given

following the registration number and the size range in mm,

standard length is given in parentheses. The dorsal-fin ray

count is partitioned into three parts, anterior first dorsal fin

(separate, or connected basally to the second dorsal fin),

second dorsal spines and dorsal segmented rays (part of second

dorsal fin). Circumorbital pore count includes pores in contact

with or immediately adjacent to the eye and includes all pores

in the infraorbital series.

Heteroclinus Castelnau, 1872

Heteroclinus adelaidae complex

Members of the Heteroclinus adelaidae complex, which

includes H. adelaidae, H. macrophthalmus and the species

described below, all have only two developed segmented pelvic

rays and the anal fin broadly connected by membrane to about

the middle of the caudal peduncle.

Heteroclinus kuiteri, sp. nov.

Figures 1-3

Heteroclinus sp. 1: Rennis, Hoese & Gomon, 1994: 746, fig. 652

(southern Australia); Hutchins, 2005 (Western Australia)

Heteroclinus species 2: Hutchins, 1994: (Western Australia)

Material examined. Holotype: AMS 1.19777-009, a 44 mm SL male,

PortseaPier, Port Phillip Bay, Vic., R. Kuiter, 12 Apr, 1977. Paratypes.

Vic.: AMS 1.19921-009, 1(41), Port Phillip Bay, R. Kuiter, 30 Jul,

1977; AMS 1.19775-002, 1(42), Portsea, Port Phillip Bay, R. Kuiter,

AMS 1.19776-005, 3(46-49), Flinders Pier, R. Kuiter, 13 Apr, 1977;

Douglass F. Hoese and Denise S. Rennis

Figure 1. Holotype of Heteroclinus kuiteri AMS 1.19777-009, a 44 mm SL male. Photo is of the right side and reversed to better show the fins

(tip of dorsal fin hidden on left side) and allow comparison with other specimens, photographed several years after collection.

Figure 2. Male paratype of Heteroclinus kuiteri, AMS 1.19777-006, 39 mm SL male, photographed a few weeks after collection.

Description of a New Species of Heterodinus (Blennoidei: Clinidae) from Southern Australia

Figure 3. Head of Heterodinus kuiteri, composite based on holotype

and paratypes showing distribution of head pores. Note that only the

upper 5 pectoral rays shown.

AMS 1.19777-006, 3(34-38), collected with holotype; AMS 1.24050-

001, 2(47-60), Port Phillip Bay, R. Kuiter, 1981; AMS 1.24051-001,

1(58), Port Phillip Bay, R. Kuiter, 1981; AMS 1.24218-001, 4(39-53),

Port Phillip Bay, R. Kuiter, 6-9 Jun, 1976; NMV A.2603, 1(41),

Refuge Cove, 5 m, G. Poore & H. Lew-Ton, 9 Feb, 1982; NMV

A. 2604, 1(35), south headland. Refuge Cove, 5 m, G. Poore & H. Lew-

Ton, 10 Feb, 1983; NMV A.2606, 2(41-44), south headland. Refuge

Cove, 13 m, C. Larsen, T. Cochran and R. Wilson, 10 Feb, 1982;

NMV A. 2607, 1(42), south headland. Refuge Cove, 13 m, C. Larsen,

T. Cochran & R. Wilson, 10 Feb, 1982; NMV A.2608, 1(48), south

headland. Refuge Cove, 13 m, C. Larsen, T. Cochran & R. Wilson, 10

Feb, 1982; NMV A.3577, 1(45), Portsea, Port Phillip Bay, 5 m, B.

Ruffle, 12 Mar, 1984; NMV A.3579, 1(36), off Stony Point, Western

Port Bay, 16 m. Fisheries and Wildlife Department “Caprella”, 1 Jun,

1967; ZMUC (uncatalogued), 4(44-52), Western Port, T. Mortensen,

6 Sep, 1914. SA NMV A.498, 1(64), Port Lincoln, J. Veitsch, 16 Dec,

1968 WA: WAM P.25767-010, 3(38-47), Sandy Hook L, Recherche

Archipelago, J.B. Hutchins, 10 Apr, 1977; WAM P.26599-004, 4(34-

48), Cosy Corner (34°15'S, 115°01'E), J.B. Hutchins, 11 Apr, 1980.

Diagnosis. Dorsal fins III, XXIX-XXXI, 2-4 (usually 3); anal

II, 21-24 (usually 22-23); segmented caudal rays 10-11 (usually

11); pectoral rays 12-13 (usually 12); pelvic rays I, 2 (sometimes

with basal element of a rudimentary 3rd ray only visible on

dissection); gill rakers on outer face of 1st arch, l-2-i-5-7=6-9

(usually 7-8). Lateral line composed of an arched section

above pectoral fin curving to midline behind pectoral fin and

continuing horizontally along midline of body; posterior

straight portion of lateral line reduced, lateral line scales not

extending beyond middle of anal fin; anteriormost lateral line

scales from end of head to above middle of pectoral fin

overlapping with a single median posterior pore, posteriorly

lateral line scales separate with a median pore at each end.

Circumorbital pores uniserial (12-17 pores). Orbital tentacle

short (1.3-4% SL), with rounded margin, sometimes with

minute lateral lobes. Nasal tentacle elongate, spoon-shaped. 1st

dorsal fin elevated, higher than 2nd dorsal fin; 2nd dorsal spine

usually longest (6.1-13.9% SL, in females, 12.6-16% SL, in

males), 1st spine rarely the longest; 1st dorsal fin originating

over or just before posterior margin of eye, posterior end of fin

connected by membrane to basal one-quarter to half of 2nd

dorsal fin. Pterygiophores from 1st dorsal fin in groove on

skull. All dorsal spines followed by fleshy flaps bound in

interspinal membrane extending from tips of spines. Last dorsal

rays evenly spaced, membrane from last ray connected to upper

base of caudal fin at end of caudal peduncle. Last anal ray

broadly connected by membrane to just beyond middle of

caudal peduncle. Pelvic rays thin (a 3rd rudimentary ray

sometimes present, visible only on dissection), 2nd ray reaching

to about the anus. Gill rakers on outer face of 1st arch short and

pointed. Body slender to moderate (depth at anal origin 16.2-

20.3% SL). Often with broad dark stripe on midside.

Description. 1st dorsal III (34*); 2nd dorsal-fin spines XXIX(6),

XXX(17), XXXI(10*), segmented dorsal rays 2(4), 3(27*),

4(3); anal fin-rays II, 21(1), II, 22(13*), II, 23(18), II, 24(2);

pectoral fin-rays 12(30*), 13(4); pelvic rays 1, 2(34*); segmented

caudal rays 10(2); 11(31*) vertebrae 14+28(6), 14+29(7),

14+30(1), 15+29(1). Circumorbital pores 12(1), 13(5), 14(9*),

15(7), 16(1), 18(1); total gill rakers on outer face of 1st arch

6(4), 7(6), 8(15*), 9(2); lower gill rakers on outer face of 1st

arch 5(7), 6(18*), 7(2). Pored lateral-line scales 19-25 (arched

portion of line) + 0-15 (straight portion of line); anterior lateral

line scales 19(4), 20(3), 21(4), 22(8), 23(1)1, 24(3*), 25(4);

posterior lateral line scales 0(2), 2(2*), 3(2), 4(3), 5(1), 6(1)

7(2), 8(1), 9(1), 10(3), 11(2), 12(4), 13(1), 14(2), 15(1);

branchiostegal rays 6(14*). Vomer with single row of teeth

forming a V; palatine without teeth.

Head compressed, moderate to large (27.2-30.4% SL, 34-

37 mm, 23.2-27.7% SL, 38-61 mm); snout obtusely pointed,

snout less than eye diameter, (3. 6-6.3% SL), eye (6.9-9.7%

SL), interorbital narrow, about half to three-quarters eye

diameter; mouth short, jaws reaching to below middle of eye,

upper jaw length, 9.3-13.2% SL; anterior nostril tubular with

simple nasal tentacle; posterior nostril with elevated rim at

anterodorsal margin of eye; gill rakers very short and simple;

rakers on 2nd and following arches moderately developed;

tongue tip broadly rounded; upper jaw with outer row of

conical teeth, slightly enlarged extending to near end of

premaxilla; anteriorly 3-4 inner rows of smaller conical teeth

tapering laterally to 1 row, ending around middle of premaxilla;

lower jaw with outer row of conical teeth enlarged covering all

of dentary; 2 or 3 inner rows of smaller teeth tapering laterally

to a single row ending before middle of dentary; intromittent

organ elongate, pointed and curved forward as in Heterodinus

macrophthalmus figured by Hoese (1976).

Head pores as shown in fig. 3, circumorbital andpreopercular

pores uniserial.

Douglass F. Hoese and Denise S. Rennis

Head largely naked, body scales small and cycloid extending

forward to above operculum below end of 1st dorsal fin; scales

overlapping and forming distinct rows anteriorly, becoming

nonimbricate and irregular posteriorly.

1st dorsal fin elevated, about twice height of 2nd dorsal in

males and slightly higher than 2nd dorsal in females; 2nd spine

usually longest, with 1st and 2nd spines subequal in height and

longer than 3rd spine; membrane from 1st dorsal fin connects to

body at base of 2nd dorsal fin; 2nd dorsal origin above a point

before pectoral origin and behind pelvic insertion; 1st spine of

2nd dorsal short, spines becoming progressively longer

posteriorly, with last spine the longest; dorsal segmented rays

evenly spaced; anal origin below 11th or 12th spine of 2nd dorsal

fin, anal spines short, anal segmented rays longer and becoming

progressively longer posteriorly; caudal fin with rounded margin

(16.3-19.2% SL); pectoral fin with rounded margin, rays

unbranched, central rays longest, reaching to above or just behind

anus; pelvic fins with hidden spine, 2 developed rays and usually

a 3rd rudimentary ray visible only on dissection.

Coloration of freshly collected material. (Based on colour

photos provided by R. Kuiter from Vic. and B. Hutchins from

WA). Colour variable. Sometimes with head, tail, and dorsal

portion of trunk dark with a brown and white mottled midlateral

band extending posteriorly from eye across upper half of

preoperculum and operculum, along trunk, and breaking into

oblong patches on tail; females and some males brown with 8

dark-brown irregularly shaped vertical bars across trunk and

tail extending onto dorsal and anal fins, bars often broken

forming irregular saddle-shaped spots, those along back darkest;

some males and females with body uniform brown or lightly

mottled; 1st dorsal fin black; females with 1st spine sometimes

banded; caudal and ventral fins pale, usually with a series of red

spots forming transverse bands; snout and edges of interorbital

frequently white; lower half of head with large black spots,

large dark spot often present basally on pectoral rays 8-10;

females with belly, ventral portions of tail and midlateral

portions of trunk and tail sometimes with white spots.

Coloration in alcohol. Similar to fresh coloration, except that

pigment less intense and red becoming pale brown.

Distribution. The species is known from Vic., Port Lincoln,

SA and Cosy Corner, near Albany and Sandy Hook I. off

Esperance, WA. It is associated with rocks and algae in shallow

water from depths of 5-13 m.

Etymology. Named for Rudie Kuiter, who provided much of the

type material and valuable material of other Australian clinids.

Remarks. One sample from Western Port, Victoria (ZMUC) is

unusual in that one specimen has 13-13 pectoral rays, two have

12-13 and one has 12-12. Only one other specimen was found

with 13 pectoral rays.

Heteroclinus kuiteri is most similar to H. adelaidae

Castelnau and H. macrophthalmus Hoese, in having the last

anal ray broadly connected by a membrane to the caudal

peduncle, two slender and elongate pelvic rays and often a

rudimentary third pelvic ray, visible only upon dissection.

Heteroclinus kuiteri differs from H. macrophthalmus in

lacking free filaments extending off the first few dorsal spines

(free and branched in H. macrophthalmus), simple orbital

tentacle (versus tentacle branched with five lobes) and fewer

dorsal rays (two-four, versus usually five). Heteroclinus

kuiteri differs from H. adelaidae in having more numerous

pectoral rays 12-13 (versus 11), dorsal fin origin over or before

posterior margin of eye (versus over posterior preopercular

margin), with pterygiophores in groove in skull (versus

pterygiophores behind skull) and no black line along anterior

lateral line (versus usually present).

Acknowledgements

We thank Gregory Millen for the photograph of the paratype.

Photos and material of this species were provided by R. Kuiter

and B. Hutchins. We thank B. Hutchins and M. Gomon for

loan of material.

References

Castelnau, F.L. de 1872. Contribution to the ichthyology of Australia.

2. Note on some South Australian fishes. Proceedings of the

Zoological and Acclimatization Society of Victoria 1: 243-248.

George, A. and Springer, V.G. 1980. Revision of the clinid fish tribe

Ophiclinini, including five new species, and definition of the family

Clinidae. Smithsonian Contributions in Zoology. 307: 1-31.

Hoese, D.F., 1976. A redescription of Heteroclinus adelaidae

Castelnau (Pisces : Clinidae), with description of a related species.

Australian. Zoologist 19(1): 51-67.

Hubbs, C.L, and Lagler, K.F 1958. Fishes of the Great Lakes region.

Cranbrook Institute of Science: Bloomfield Hills Michigan 213 pp.

Hutchins, J.B., 1994. Asurvey of the nearshore reef fish fauna of Western

Australia’s west and south coasts - the Leeuwin Province. Records of

the Western Australian Museum Supplement No. 46: 1-66.

Hutchins, J.B., 2005. Checklist of marine fishes of the Recherche

Archipelago and adjacent mainland waters in: Wells, F.E., Walker,

D.I. and Kendrick, G.A. (eds). The Marine flora and fauna of

Esperance, Western Australia. Western Australian Musem: Perth.

Rennis, D., Hoese, D.F. and Gomon, M.F 1994. Family Clinidae. Pp.

741-775, figs 650-684 in: Gomon, M.F., Glover, C.J.M. & Kuiter,

R.H. (eds) The Fishes of Australia s South Coast. Adelaide : State

Printer 992 pp.

Memoirs of Museum Victoria 63(1): 25-28 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Description of two new species of shore-eels (Gobiesocidae: Cheilobranchinae:

Alabes) from south-eastern Australia and Norfolk Island

Barry Hutchins

Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986

Abstract Hutchins, B. 2006. Description of two new species of shore-eels (Gobiesocidae: Cheilobranchinae: Alabes) from south-

eastern Australia and Norfolk Island. Memoirs of Museum Victoria 63(1): 25-28.

Two new species of Alabes are reported, one from Tasmania and the other from Norfolk Island. The first is similar in

general appearance to A. parvula, but is considerably smaller in size (to 39 mm SL) and inhabits waters of much greater depth

(160-348 m). The Norfolk Island species has a uniquely enlarged buccal region and is possibly endemic to this island.

Keywords Alabes, Cheilobranchinae, Tasmania and Norfolk Island, Australia

Introduction

Following a study of the Australian subfamily Cheilobranchidae,

Hutchins and Morrison, 2004 described five new species of

Alabes, namely A. elongatus, A. gibbosus, A. obtusirostris, A.

occidentalis and A. scotti. They also recognised four previously

described species, Alabes dorsalis Richardson, A. parvula

McCulloch, A. hoesei Springer and Fraser, and A. brevis Springer

and Fraser, concurring with the findings of an earlier review

(Springer and Fraser, 1976). The latter review also reported a

single Norfolk Island specimen of doubtful identity but close to

A. parvula, and indicated that further specimens should be

collected before its status could be verified. As no new material

of the Norfolk Island form had been forthcoming in the

intervening years, Hutchins and Morrison, 2004 preferred not to

provide a description, but did include it, as Alabes sp., in a table

to the genus indicating its unique identity. Furthermore, they also

examined, but did not report on, two specimens of a form similar

to Alabes dorsalis which had been collected offshore from

Tasmania at a depth of 160 m (A. dorsalis is a common intertidal

species which occurs rarely to a maximum depth of 30 m).

However, the two nondescript specimens were small (29-39 mm

SL versus a maximum size of 120 mm for A. dorsalis), and were

suspected of being juveniles that had possibly been carried

offshore by ocean currents. Recently a third specimen of this

form was taken at an even greater depth (348 m) near King

Island, just to the north-west of Tasmania. It proved to be

conspecific with the other two Tasmanian specimens, and

although very similar to A. dorsalis, all three nevertheless

represent an undescribed species. The purposes of the present

paper, therefore, are to provide descriptions of these two rare

species from Norfolk Island and Tasmania, and to expand the key

of Hutchins and Morrison, 2004 to include them.

Methodology follows that of Hutchins and Morrison, 2004.

The following abbreviations refer to cephalic pores of the

lateral line system: ANP, anterior nasal pore; LP, lacrymal

pore; PNP, posterior nasal pore; POP, postocular pore. Standard

length is abbreviated SL and head length is abbreviated HL.

Type material is held in the Australian Museum, Sydney

(AMS), Marine Research Laboratories, CSIRO, Hobart

(CSIRO), and Museum Victoria, Melbourne (NMV).

Key to the species of Alabes

1. Pelvic fin rudiment present, located on ventral surface

immediately behind gill opening 2

- Pelvic fin rudiment absent 6

2. Pelvic fin rudiment moderate in size, fin rays present;

POP 1 3

- Pelvic fin rudiment small, pelvic fin rays (if present) not

visible; POP 2, usually widely separated 5

3. Pelvic fin rudiment width equal to or wider than length of

gill slit 4

- Pelvic fin rudiment less than length of gill slit

(Tasmania) A. bathys sp. nov.

4. Body relatively robust (depth 9.3-11.6 in SL); dorsal fin

fold relatively high, continued anteriorly to vertical through

urogenital opening (snout to origin of dorsal fin fold 2. 0-3. 5

in SL); blackish circular blotches often present on middle of

side of adult (NSW, Vic., Tas., and SA) A. dorsalis

- Body more slender (depth 12.5-14.9 in SL); dorsal-fin

fold relatively low, origin falling well short of vertical

through urogenital opening (snout to origin of dorsal fin

fold 1.3-2 .4 in SL); no dark circular blotches on side of

large adult (WA) A. elongata

Barry Hutchins

5. Head relatively large, length 61-1.6 in SL; PNP absent

(WA) A. brevis

- Head relatively small, length 9.7-11.4 in SL; PNP present

(WA) A. gibbosa

6. Gill slit small, length 5.4-22.9 in HL; POP 1 or 2; nasal

pores present or absent 7

- Gill slit moderately large, length 3. 1-4.3 in HL; no POP

or nasal pores (Tas. and WA) A. obtusirostris

7. Origin of dorsal-fin fold over vertical through urogenital

opening or behind; anal-fin fold prominent, extending

about half way from caudal fin to urogenital opening;

interorbital relatively wide, width 5.4-10.8 in HL; body

translucent posteriorly in life, sides without alternating

black and white bars 8

- Origin of dorsal-fin fold just behind head; anal-fin fold

absent or restricted to region just anterior to caudal fin;

interorbital very narrow, width 8.8-22.6 (usually 15 or

more) in HL; body orange in life (not translucent

posteriorly), sides with alternating wide black and narrow

white bars (NSW, Vic., and Tas.) A. scotti

8. Postocular pores 1; PNP absent; caudal-fin rays less than

7 or absent 9

- Postocular pores 2; PNP usually present; caudal-fin rays

7-8 (NSW, Vic., Tas., SA, WA) A. hoesei

9. Origin of dorsal-fin fold over vertical through urogenital

opening 10

- Origin of dorsal-fin fold well posterior to vertical through

urogenital opening (Norfolk I.) springeri sp. nov.

10 Lacrymal pores present; male with pattern of regular

tiger-like bars anteriorly on side of body, not merging

ventrally with longitudinal series of 4 blackish to dark-

blue blotches on side of abdomen, area of blotches

extending about half way between gill slit and urogenital

opening (NSW, Qld) A. parvula

- Lacrymal pores absent; male with irregular arrangement

of tiger-like bars anteriorly on side of body, merging

ventrally with 2-3 blackish blotches which extend about

one -third distance between gill slit and urogenital opening

(WA) A. occidentalis

Alabes bathys sp. nov.

Figures la, b; Table 1

Material examined. Holotype. NMV A14558, 39 mm SL, Tas., 30 km

NNW of Cape Sorell (42°10.9'S, 144°48.9'E), WHOI epibenthic sled

at 160 m, RS Wilson on board RV Soela (field no. S05/84 54),

20 Oct 1984.

Paratypes. NMV A28045-001, 29 mm SL, taken with holotype

(disarticulated cleared and stained skeletal material); CSIRO H.6156-

01, 38 mm SL, Tas., King I. Canyons, west of King I. (between

39°48'S, 143°08'E and 39°50'S, 143°07'E), Sherman sled at 348 m,

LRV Southern Surveyor (field no. SS0404/35), 18 Apr 2004.

Diagnosis. Alabes bathys differs from all other species of

Alabes by a combination of its moderately large gill opening

(width 3. 1-3.7 in HL), somewhat smaller pelvic fin rudiment

(width 1.4-1. 6 in gill opening) consisting of 2 apparent fin rays

in each half, 4 cephalic pores (1 POP, 1 ANP, 2 LP) in the

lateral line system, and its unique colour pattern.

Ligure la. Alabes bathys, holotype, NMV A 14558, 39 mm SL, off

Cape Sorell, Tas. (photograph courtesy of Rudie Kuiter, NMV).

l!Mllllllllll'IM'fiil f 'Ml'lMllllimil

Ligure lb. Alabes bathys, paratype, CSIRO H.6156-01, 38 mm SL, west

of King L, Tas. (photograph courtesy of Alastair Graham, CSIRO)

Table 1. Measurements (mm) and counts of the holotype and paratypes

of Alabes bathys sp. nov.

Holotype

NMV

A 14558

Paratype

NMV

A28045-001

Paratype

CSIRO

H.6156-01

Standard length

Head length

5.2

3.9

4.4

Snout length

1.3

0.8

1.4

Eye diameter

1.6

1.2

1.5

Interorbital width

0.7

0.5

0.6

Gill slit width

1.4

1.1

1.4

Head width

2.4

2.1

2.4

Body width at gill slit

1.8

1.7

1.8

Body width (max.)

1.8

1.7

1.8

Body depth (max.)

3.0

2.3

2.9

Snout to dorsal-fin fold

9.2

Snout to anal-fin fold

Snout to anus

Ventral fin width

0.9

0.8

1.0

Caudal fin rays

Postocular pores

Posterior nasal pores

Anterior nasal pores

Dorsal lacrymal pores

Ventral lacrymal pores

Sex

Two new species of shore-eels

Description. Measurements of the holotype and paratypes are

presented in Table 1 (counts and proportions in parentheses in

the following description represent those of the paratypes

where different from the holotype). Body elongate,

subcylindrical, widest anteriorly (maximum body width at

level of gill slit 21.7 [17.1-21.1] in SL), reaching a maximum

depth about middle of body (depth 13.0 [12.6-13.1] in SL),

tapering posteriorly; head small, cylindrical, length 7.5 (7.4-

8.6) in SL, slightly wider than body, width 2.1 (1. 8-1.9) in HL;

snout short, round to slightly truncate in dorsal view, length 4.0

(3. 1-4.6) in HL; nostrils small but obvious, anterior 1 tubular,

posterior 1 with low raised rim; eye moderate in size, with

prominent clear cornea, diameter of orbit 3.3 (2.9-3.3) in HL,

bony interorbital rather narrow, width 7.4 (7.3-7.8) in HL.

Skin smooth and scaleless, usually covered with mucus

layer; lateral line sensory system consists of small open pores

and minute papillae (latter very difficult to detect); 4 pores on

each side of head, consisting of 1 POP and 1 ANP, and 2 LP.

Gill opening a moderately wide slit, located on ventral surface

of head, length 3.7 (3. 1-3. 5) in HL; gills 3?; branchiostegals 3;

mouth terminal, gape not reaching vertical line through

anterior margin of eye (pigmented area), lips narrow, somewhat

fleshy; teeth in cleared and stained paratype small, conical,

caninedike, 1 row of 6-8 teeth on each side of upper and lower

jaws, largest anteriorly (premaxilla also with 3 much smaller

teeth forming a 2nd row along inner base of main row); palatine

and vomerine teeth absent. Dorsal and anal-fin folds resemble

low fins, but lack both fin rays and underlying pterygiophores;

caudal fin with 8 (7-8) fin rays, continuous with dorsal and

anal-fin folds; bases of dorsal and anal-fin folds relatively long,

insertion of dorsal-fin fold over or slightly behind vertical

through urogenital opening, insertion of anal-fin fold well

behind vertical through urogenital opening (snout to insertion

of dorsal-fin fold 2.7 [2. 6-3. 2], snout to insertion of anal fin

fold 2.0 [1.7-2. 6], snout to urogenital opening 2.8 [2.7-2. 8], all

in SL); urogenital opening with prominent papilla. Total

vertebrae not known for holotype but cleared and stained

paratype has a total of 68 vertebrae, not including hypural

plate (21 precaudal), with last epineural on 24th vertebra.

Colour in alcohol. Head and body overall pale brown, fins

more translucent.

Colour in life. (Based on colour transparency of holotype, see

fig. la): head and body pale brown, abdominal region more

pinkish ventrally; body becoming more translucent posteriorly,

vertical fins almost totally transparent; middle of side with

longitudinal series of whitish spots from head almost to caudal

fin, each spot about equal or subequal to eye diameter; dorsal

surface with longitudinal series of similarly coloured short

cross-bands, continued onto dorsal-fin fold as vertical bars; a

few pale spots on anal-fin fold. Larger paratype differs in its

ground colour, being a more pale greyish green (fig. lb).

Distribution. Alabes bathys is known only from western Tas.,

in the region of Cape Sorrell and King L.

Remarks. This species has so far only been collected by dredge

from deep coastal waters at depths between 160 and 348 m. It

shares with Alabes dorsalis a large ventral gill slit and prominent

ventral fin rudiment, but differs in the relatively narrow width of

the latter fin (maximum width 1.4— 1.6 versus 0. 8-1.0 for A.

dorsalis, all in length of gill slit). It also has a narrower body

(maximum depth 12.6-13.1 verus 9.3-11.6 in SL), lacks the

dark circular markings along the body and its small size

(maximum length 42 mm TL versus 120 mm TL in A. dorsalis).

Etymology. This species is named bathys (from the Greek

“bathys” meaning deep) with reference to its deep water habitat.

Alabes springeri sp. nov.

Figure 2 ; Table 2

Alabes parvulus {non McCulloch) Springer and Fraser, 1976; 21.

Material examined. Holotype. AMS 1.18497-001, 31 mm SL, Norfolk

L, Emily Bay, Point Hunter (29°04'S, 167°57'E), rotenone at 1 m, C.

Anderson et al., 16 Sep 1975.

Diagnosis. Alabes springeri differs from all other species of

Alabes by a combination of its small gill opening (8.8 in HL),

lack of a pelvic-fin rudiment, narrow interorbital space (8.8 in

HL), presence of only 2 cephalic pores in the lateral line system,

and the posteriorly placed origin of the dorsal-fin fold.

Description. Measurements of the holotype, the only known

specimen, are presented in Table 2. Body elongate and

subcylindrical, reaching a maximum depth at level of about

anterior 3rd of body, then tapering posteriorly, body depth 11.1

in SL, maximum body width (= width at level of gill slit) 15.5

in SL; caudal peduncle absent (caudal fin joined to dorsal and

ventral-fin folds); head small, length 7.0 in SL, a little wider

than deep (head width 1.9 in HL); snout short, rather rounded

to slightly truncate anteriorly in dorsal view, length 3.1 in HL;

nostrils small but obvious, anterior one tubular, posterior one

with low raised rim; eye moderate in size, with prominent clear

cornea, diameter of orbit 3.7 in HL, noticeably greater than

bony interorbital width (8.8 in HL).

Skin smooth and scaleless, normally covered with a thick

mucus layer; lateral line sensory system consists of minute

papillae (latter very difficult to detect); cephalic pores 2 on

each side of head, 1 POP and 1 ANP. Gill opening a narrow

slit, located on ventral surface of head, width 8.8 in HL; gills

and branchiostegals not examined; mouth subterminal, upper

jaw projecting over lower jaw, upper and lower lips rather

narrow, rear corner of mouth not reaching anterior margin of

eye (pigmented area); teeth incisorform, 1 row in upper and

lower jaws; palatine and vomerine teeth absent. Dorsal and

anal-fin folds resemble low fins, but lacking both fin rays and

underlying pterygiophores, continuous with caudal fin; caudal

fin without visible fin rays, although there is evidence that the

posterior portion of the specimen was damaged at some time

(portion of caudal and anal-fin folds lost, and some vertebral

elements also protruding through side of body); bases of dorsal

and anal-fin folds relatively short, insertion of dorsal-fin fold

well behind vertical through urogenital opening (snout to

insertion 1.8, snout to urogenital opening 2.6, both in SL),

snout to insertion of anal-fin fold 1.4 in SL; urogenital opening

with small papilla. Total vertebrae 71 (from Springer and

Fraser, 1976), with last epipleural on 20th vertebra.

Barry Hutchins

Figure 2. Alabes springeri, hototype, AMS 1.18497-001, 31 mm SL, Norfolk L, Emily Bay, Point Hunter (illustration courtesy of Victor Springer,

USNM).

Table 2. Measurements (mm) and counts of the holotype of Alabes

springeri sp. nov.

Holotype

AMS

1.18497-001

Standard length

Head length

4.4

Snout length

1.4

Eye diameter

1.2

Interorbital width

0.5

Gill slit width

0.5

Head width

2.3

Body width at gill slit

2.0

Body width (max.)

2.0

Body depth (max.)

2.8

Snout to dorsal fin fold

Snout to anal fin fold

Snout to anus

Caudal fin rays

Postocular pores

Posterior nasal pores

Anterior nasal pores

Dorsal lacrymal pores

Ventral lacrymal pores

Sex

Male?

Colour in alcohol. Head and body overall pale brown, fins

more translucent. Springer and Fraser, 1976 illustrated some

partial cross banding on the abdominal region, triangular in

shape (see fig. 2 ), but this has since faded.

Colour in life. Unknown

Distribution. Alabes springeri is known only from the type

locality (Norfolk I.).

Remarks. This species inhabits shallow coastal waters (single

specimen taken from a depth of 1 m). It is most similar to

Alabes parvula from eastern Australia, differing in the

exceptionally short base of the dorsal-fin fold, different shape

of the dark bars on the side of the abdomen (triangular-shaped

versus more squarish in A. parvula), and the expanded shape of

the buccal region (not expanded in A. parvula). Like other

members of the Alabes parvula complex (A. parvula, A.

occidentalis, and A. hoesei), it lacks a pelvic fin rudiment, has

a very small gill slit, its body probably is transparent in life (at

least posterior to urogenital opening), and its maximum size is

less than 50 mm TL (see Table 1 in Hutchins and Morrison,

2004). This species is named springeri in honour of V.G.

Springer (USNM) who, with T. Fraser in 1976, first reported

the apparent uniqueness of the Norfolk Island specimen. (NB.

registration number AMS 1.18470-002 was inadvertently

assigned to this specimen by Springer and Fraser, 1973: 21, but

that number belongs to one of the paratypes of Alabes hoesei

which was described in the same paper).

Acknowledgements

I am grateful to Alastair Graham (CSIRO), Martin Gomon

(NMV) and Mark McGrouther (AMS) who provided the material

on loan that was used in this study. Victor Springer (USNM)

kindly gave advice on the description of the new Alabes from

Norfolk Island, and offered me the use of a previously published

illustration of this species (from Springer and Fraser, 1976).

References

Hutchins, J.B. and S.M. Morrison. 2004. Five new fish species of the

genus Alabes (Gobiesocidae: Cheilobranchidae). Records of the

Australian Museum 56 (2): 147-158.

Springer, V.G. and T.H. Fraser. 1976. Synonymy of the fish families

Cheilobranchidae (=Alabetidae) and Gobiesocidae, with

descriptions of two new species of Alabes. Smithsonian

Contributions to Zoology number 234: 1-23.

Memoirs of Museum Victoria 63(1): 29-46 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Revision of the genus Hapalogenys (Teleostei: Perciformes) with two new species

from the Indo-West Pacific

Yukio Iwatsuki^ and Barry C. Russell^

'Division of Fisheries Sciences, Faculty of Agriculture, University of Miyazaki, 1-1 Gakuen-kibanadai-nishi, Miyazaki

889-2192, Japan (yuk@cc.miyazaki-u.ac.jp)

^Museum and Art Gallery of the Northern Territory, PO Box 4646, Darwin, NT 0801, Australia (barry.russell@nt.gov.au)

Abstract Iwatsuki, Y., and Russell, B.C. 2006. Revision of the genus Hapalogenys (Teleostei: Perciformes) with two new species

from the Indo-West Pacific. Memoirs of Museum Victoria 63(1): 29-46.

The Indo-West Pacific genus Hapalogenys is reviewed and two new species are described: Hapalogenys dampieriensis

sp. nov. from Australia and H. filamentosus sp. nov. from the Philippines. The genus now includes: Hapalogenys analis

Richardson, H. dampieriensis sp. nov., H. filamentosus sp. nov., H. kishinouyei Smith and Pope, H. merguiensis Iwatsuki,

Ukkrit and Amaoka, H. nigripinnis (Schlegel in Temminck and Schlegel) and H. sennin Iwatsuki and Nakabo. Hapalogenys

dampieriensis, H. filamentosus and H. kishinouyei are similar to each other in overall body appearance and are accordingly

identified as the 'Hapalogenys kishinouyei complex”, defined by having 2-5 longitudinal stripes on the body. Hapalogenys

dampieriensis has long been confused with H. kishinouyei in having similar longitudinal dark stripes, but the two species

are easily separable on meristic and morphometric values, and body colour changes with growth. Hapalogenys filamentosus

differs from H. dampieriensis in having a longer pelvic fin, with the filamentous first fin ray almost reaching to or slightly

beyond the base of first anal-fin spine when depressed (vs. slightly beyond anus but not reaching to base of first anal-fin

spine) and two faint narrow longitudinal stripes on the body (vs. four narrow longitudinal stripes in juveniles, reducing to

two with growth). A neotype is designated for H. analis. Species of Hapalogenys can be distinguished from one another

on the basis of meristic and morphometric characters, body colour pattern, maximum size and distribution. The familial

position of Hapalogenys is briefly discussed.

Keywords Taxonomy, Pisces, Perciformes, Revision, Hapalogenys dampieriensis sp. nov., Hapalogenys filamentosus sp. nov.

Introduction

In a study of the Indo-West Pacific genus Hapalogenys, Iwatsuki

et al., 2000a pointed out the taxonomic confusion among

Hapalogenys species from Japan. Subsequently, Iwatsuki and

Nakabo, 2005 redescribed H. nigripinnis and proposed a new

species, H. sennin. Ongoing investigations of all nominal

species of Hapalogenys, including those now included in the

family Dinopercidae and the genus Parapristipoma (Heemstra

and Hecht, 1986; Iwatsuki et al., 2000a, b; Heemstra and

Iwatsuki, in press; see Discussion), have resulted in the

recognition of five species of Hapalogenys from the Indo-West

Pacific, plus two new species described herein.

Hapalogenys kishinouyei Smith and Pope, 1906, described

from Japan, was long considered an endemic East Asian shelf

species. However, Gloerfelt-Tarp and Kailola, 1984 reported it

from north-western Australia, their specimens having similar

longitudinal stripes on the body, and believed it to be an

antitropical species. The East Asian H. kishinouyei though are

separable from Australian specimens on the basis of counts,

proportional measurements, different changes in colouration

with growth, and maximum body size. In this paper we

conclude that the Australian specimens represent an

undescribed species of Hapalogenys. A second new species

from the Philippines, similar to the Australian species in

overall appearance, but differing in having a long pelvic fin

with a filamentous first ray and two faint, longitudinal stripes

on the body (vs. four at the same size), is also described.

The following account reviews the genus Hapalogenys

from the Indo-West Pacific, including two new species, on the

basis of all known types and a wide range of non-type

specimens, from a wide geographic area. The familial position

of Hapalogenys is briefly discussed.

Yukio Iwatsuki and Barry C. Russell

Methods

Counts and measurements follow Iwatsuki et al., 2000a.

Terminology generally follows Johnson, 1980, 1984, although

that of the supraneural bones follows Mabee, 1988 and the

formula of Ahlstrom et ah, 1976. Institutional codes follow

Leviton et al., 1985, with the following additions: Division of

Fisheries Sciences, University of Miyazaki, Japan (MUFS);

Phuket Marine Biological Center, Thailand (PMBC);

Kanagawa Prefectual Museum, Kanagawa, Japan (KPM). The

very short dense papillae and barbels on the fleshy lower lip in

Hapalogenys analis (MUFS 7148, 12258) and H. sennin

(MUFS 2086, 12225) necessitated dissection so as to determine

the number of pores on and posterior to the chin.

Systematics

Hapalogenys Richardson, 1844

Hapalogenys Richardson, 1844a: 462 (type species not

designated). -Bleeker, 1876: 271 (Hapalogenys nitens subsequently

designated as type species by Richardson, 1844b [=//. nigripinnis

Schlegel in Temminck and Schlegel, 1843], see Iwatsuki and Nakabo,

2005). -Johnson, 1984: 465 (placed as incertae sedis in

Percoidei). -Springer and Raasch, 1995: 93, 104 (established family

name Hapalogenidae [sic. Haplogeniidae] for this genus). -Iwatsuki et

al., 2000a: 129.-Iwatsuki and Nakabo, 2005: 854.

Definition of the genns Hapalogenys

Body compressed, elevated; mouth moderate, horizontal; upper

jaw protractile; 10 pores on and behind chin, including a pair of

very small pores near symphysis (often hidden by cluster of short

dense barbels or papillae, especially in Hapalogenys analis and

H. sennin), plus 2 pores anteroventrally on dentary (often hidden

by cluster of short dense barbels or papillae, especially in H.

analis and H. nigripinnis), a single pore ventrally, midway along

each dentary and a single pore ventrally at articulation between

dentary and angular (sometimes slit-like in H. dampieriensis sp.

nov., H. sennin and H. kishinouyei, a pit partially or entirely

covered posteriorly by membrane in H. nigripinnis and H. sennin,

especially in larger adults); a cluster of short or long, crowded

papillae and barbels on and behind chin, generally developed

with age; snout tip naked or with small papillae; teeth uniformly

small, in bands on jaws, vomer, and palatines; preopercle serrate;

operclewith 1 or2shortspines;7branchiostegals;pseudobranchiae

present; air bladder simple; pyloric appendages few; pored

lateral-line scales 41^8; soft vertical fins scaled basally; dorsal

surface of head, including snout, jaws and opercular elements

scaly; dorsal-fin spines 11 with antrorse spine anteriorly (the

antrorse spine is, in fact, an anterior projection of the 1st

pterygiophore, not a fin spine), rays 13-15; anal fin with 3 spines

(2 supernumerary spines on 1st anal pterygiophore; see Johnson,

1980), usually 9 rays (rarely 8); caudal fin generally rounded;

pectoral fin pointed; dorsal- and anal-fin pterygiophores with

separate proximal, middle and distal radials; supraneural formula

0/0-I-0/2/1-I-1/; principal caudal-fin rays 9-1-8; procurrent rays 5 or

6-1-5 or 6 (upper -i- lower); caudal skeleton with 5 hypurals, 3

epurals, 2 uroneurals and 2 autogenous haemal spines; hypural

fusions absent; procurrent spur absent; vertebrae 10-1-14.

Relationships

The genus Hapalogenys has been traditionally placed in the

Haemulidae (Akazaki, 1984; McKay, 2001, Nelson 2006),

although Johnson, 1984 included it as 'dncertae sedis'' in the

Percoidei because of its uncertain affinities. Springer and Raasch,

1995 established a new family name, Hapalogenidae (sic.

Haplogeniidae), for the genus, but without any strong supporting

evidence. McKay, 2001 also recognised Hapalogenys as removed

from the Haemulidae, although he retained it in that family for

convenience. He reported Hapalogenys is very close to the two

species of the family Dinopercidae, but lacks intrinsic muscles

on the posterior part of the swimbladder. Based on similarities of

larval morphology with Lobotes and Datnioides (= Coins), Leis

and Carson-Ewart, 2000, 2004 placed Hapalogenys in a group

they informally called Hobotes-Xike' , and suggested a possible

relationship of Hapalogenys to lobotids. Clearly, further study is

needed to clarify the familial position of Hapalogenys, and until

its relationships to other genera can be resolved we provisionally

retain Hapalogenys in the Haemulidae.

Key to species

1. Scales on maxilla (figs 3A and C) 2

— No scales on maxilla (figs 3C, 3E, 3G and 4A, E) 3

2. Head and body with 2 oblique dark bands (rarely indistinct),

1st descending from nape to behind pectoral fin and running

to posterior part of soft anal-fin rays, 2nd descending from

base of anterior 3rd or 4th dorsal-fin spines and soft dorsal-

fin base, curving backwards through lateral line to upper

part of caudal peduncle (fig. IG); posterior margin of soft

dorsal, anal and caudal fins not dense black (fig. IG);

spinous dorsal-fin membranes mostly yellowish brown, not

dense black (fig. IG) Hapalogenys nigripinnis (fig. IG)

— Head and body with 5-7 alternating whitish and dark-

brown bands, 1st (often indistinct) from just before eye to

posterior of lower jaw, 2nd somewhat oblique, extending

from nape across opercle to pelvic-fin base (becoming

wider posteroventrally), 3rd from base of 2nd and 3rd

dorsal-fin spines to just behind pelvic-fin base; posterior

margin of soft dorsal, anal and caudal fins dense black (fig.

lA); spinous dorsal-fin membrane dense black (fig. lA)

Hapalogenys analis (fig. lA)

3. Body with 2 oblique dark bands or sometimes no bands

— Body with 2-5 longitudinal dark stripes (sometimes

indistinct or faint but visible), 1st from front of 1st dorsal-

fin spine along dorsal midline, 2nd from nape to base of

mid dorsal-fin soft rays, 3rd from eye to last dorsal-fin ray

base, 4th from preopercular flange, through base of

pectoral fin, to lower caudal peduncle, last from isthmus

to base of anal spinous fin (fig. 2E) 5

4. 1st dark band on body descending from nape to behind

pectoral fin, and 2nd from base of anterior 2nd or 3rd

dorsal-fin spines and soft dorsal-fin base, curving backwards

through lateral line to soft anal-fin and caudal peduncle

(fig. IF), but bands often lost in preserved specimens; orbit

diameter large (3. 5-3. 8 in head length); papillae on fleshy

lower lip well-developed but very short on chin (figs

4A-B) Hapalogenys merguiensis (fig. IF)

Revision of the genus Hapalogenys

— 1st dark band descending from nape to behind pectoral fin,

and 2nd descending from base of 7-lOth dorsal-fin spines,

curving downwards above or through lateral line, but bands

often lost in preserved specimens (fig. IH): orbit diameter

small (7.4-12.7 in head length); barbels on fleshy lower lip

extremely well -developed and in a dense cluster on chin

(figs 4E-F) Hapalogenys sennin (fig. IH)

5. Filamentous tip of 1st pelvic-fin ray almost reaching to or

slightly beyond base of 1st anal-fin spine when depressed

(figs IC-D); 2nd and 3rd longitudinal dark stripes on body

faint (fig. 1C); posteriormost angle of jaw reaching to a

vertical through centre of eye in specimens between

130-160 mm SL (figs IC-D)

Hapalogenys filamentosus sp. nov. (fig. 1C)

— Filamentous tip of 1st pelvic-fin ray extending slightly

beyond anus but clearly not reaching to base of 1st anal-

fin spine when depressed (figs lA-B, lE-H); 4th and 5th

longitudinal dark stripes on body indistinct or absent (figs

lA, IE, 2A-F); jaw reaching to slightly behind a vertical

through anteriormost eye membrane in specimens

between 130-160 mm SL (figs 1C, 2B-C, 2E-F) 6

6. 3rd body stripe narrow, its width below base of 5th and 6th

dorsal-fin spines less than pupil diameter in specimens

smaller than about 170 mm SL, 3rd, 4th and 5th stripes lost

in specimens larger than 200 mm SL (figs IB, 2A-C)

Hapalogenys dampieriensis sp. nov. (figs IB, 2A-C)

— 3rd stripe broad, its width below base of 5th and 6th

dorsal-fin spines greater than pupil diameter at all sizes;

1st, 4th and 5th stripes lost in specimens larger than about

250 mm SL (figs IE, 2D-F)

Hapalogenys kishinouyei (figs IE, 2D-F)

Hapalogenys analis Richardson, 1845

English Name; Broadbanded Velvetchin

Japanese Name: Setodai

Figures lA, 3A-B

Hapalogenys analis Richardson, 1845: 85, pi. 43, fig. 1 (type

locality: Canton, China). -Whitehead, 1970; 215 (Canton,

China). -Bauchot et ah, 1983: 32 (Canton, China).

Pristipoma mucronata Eydoux and Souleyet, 1850 (as 1841 but

erroneous, see Bauchot et ah, 1982): 161, pi. 2, fig. 1 (type locality;

Macao, China).

Hapalogenys mucronatus Gunther, 1859: 318 (China). -Bleeker,

1865-1869: 56, 58 (Amoy, China). -Ishikawa and Matsuura, 1897; 54

(Tokyo). -Jordan and Thompson, 1912: 553 (Kobe, Hyogo and

Onomi chi , Hiroshima) . -Izukaand Matsuura, 1 920 : 1 50 (Osaka) . -Tanaka,

1925: 888, pi. 181, fig. 495 (Tadanoumi).-Eowler, 1930: 605

(Japan).-Shen, 1993: 360, pi. 101 (Kaoshung, Taiwan). -Cheng et ah,

1997: 257 (Shandong Province, China). -Sadovy and Cornish, 2000:

229 (Hong Kong, China).-Wang et al., 2001: 223 (Heibei, China).

Hapalogenis mucronatus Steindachner and Ddderlein, 1883: 11

(Osaka).

Type material. Neotype: MUES 12258 (fig. lA), 185 mm SL, East

China Sea, bottom trawl, 50-100 m depth, 12 June 1996.

Non-type material. 72-201 mm SL, n=49. EAKU 12098, 88 mm

SL, East China Sea; EAKU 12098, 72 mm SL, East China Sea; EAKU

29281, 137 mm SL, off Yamaguchi, Sea of Japan; EAKU 34879,153

mm SL, off Ehime, Bungo Channel; EAKU 10031^100316, 3: 108-

137 mm SL, Gulf of Tong-king, northern Vietnam; ERLM 8115, 8145,

2: 87-114 mm SL, East China Sea (31°9'N, 125°00'E, 31°20'N, 125°10'

E), trawl, less than 100 m depth; ERLM 8170-8172 and 8189, 4: 102-

133 mm SL, East China Sea (31°23'N, 125°05'E, 31°10'N, 125°10'E),

trawl, less than 100 m depth; HUMZ 49427^9430, 4: 87-131 mm SL,

Yahatahama Market, Ehime, Japan; HUMZ 71708, 115 mm SL,

Kaoshung Eish Market, Taiwan; HUMZ 106037, 166 mm SL, Saeki

Eish Market, Oita, Japan; HUMZ 108351, 108415, 108650, 74-95 mm

SL, East China Sea, about 81-86 m depth; lOCAS 51-118, 117 mm SL,

off Chingtao, China; lOCAS 57-1549, 2001, Ryonin, China; lOCAS

73-211, off Amoy China; MNHN 7702 (holotype of Pristipoma

mucronata), 154 mm SL, near Macao; MUES 2308, 87 mm SL, Kiryu,

Taiwan; MUES 7148, 129 mm SL, Akamizu, Nobeoka, Miyazaki,

Japan, set net, less than 30 m depth; MUES 8652, 143 mm SL, bottom

trawl, 50-100 m depth, off Nobeoka, Miyazaki, Japan; NSMT-P 2116,

Kasaoka, Seto Inland Sea; SEU 58-2573, 165 mm SL, off Shanghai,

China; SEU 3862-3863, 2: 81-181 mm SL, Shanghai, China; URM-P

751, 102 mm SL, East China Sea; URM-P 10446-10448, 3: 91-95 mm

SL, Akashi, Hyogo, Seto Inland Sea; URM-P 19090, 19692-19694,

19737, 5: 90-147 mm SL, East China Sea; URM-P 23097, 112 mm SL,

Tainan, Taiwan; ZRC 38018, 105 mm SL, W coast of Singapore; ZRC

41178, 105 mm SL, Razaras L, Singapore.

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster

of short papillae and barbels anteriorly, 10 pores on and behind

chin, 4 anterioriormost pairs hidden by papillae (figs 3A-B),

slightly scaly posteriorly (on posterior abdominal part of

angular of lower jaws, fig. 3B); small scales on maxilla (fig.

3A); 5-7 alternating whitish and dark-brown bands on head and

body, 1st (often indistinct) from just before eye to posterior of

lower jaw, 2nd somewhat oblique, becoming wider posteriorly,

from nape to between eye and pelvic-fin base, 3rd from base of

2nd and 3rd dorsal-fin spines to just behind pelvic-fin base; 4th

from 7th and 8th dorsal-fin spine base to just before 1st anal-fin

spine, 5th from 2nd and 3rd dorsal-fin soft ray base to anterior

of anal-fin soft ray base, 6th on caudal peduncle, 7th (often

indistinct) on caudal-fin base; soft rayed portions of dorsal and

anal fins somewhat rounded posteriorly and slightly angulated

posteriorly, respectively (fig. lA), spinous dorsal-fin and anal-

fin (until 1st anal-fin ray) membranes dense black; membrane

of soft dorsal- anal- (especially in juveniles and young) and

caudal-fins somewhat yellowish in fresh specimens, a dense

black margin posteriorly, about same width as pupil diameter

(juveniles) or half of this width (adults) (fig. lA); pelvic-fin tip

not reaching to base of 1st anal-fin spine when depressed (fig.

lA); 44-47 (but rarely 43 or 48) pored lateral-line scales;

procumbent spine-like process (exposed tip of 1 st ptery giophore)

at origin of dorsal fin usually covered by predorsal scales.

Description. Counts and proportional measurements as

percentage of SL for the neotype and other specimens are given

in Table 1. Data for the neotype are presented first, followed by

non-type material (if different) in parentheses. Characters

given in the diagnosis are not repeated.

Body deep, 49% of SL (47-57% of SL), relatively

compressed, covered with ctenoid scales, ctenii free on posterior

margin of exposed area; lateral line continuous until on hypural

plate; orbit diameter distinctly larger than interorbital space;

suborbital depth distinctly less than eye diameter; head covered

Yukio Iwatsuki and Barry C. Russell

Table 1. Counts and proportional measurements, and characters as percentage of standard length for the neotype and non-type specimens of

Hapalogenys analis Richardson, 1845

Hapalogenys analis

Neotype

Non-type specimens

MUFS 12258, 185 mm SL

72-201 (mean 116) mm SL

n=49

Dorsal-fin rays

XI, 15

XI, 15-16

Anal-fin rays

III, 9

III, 9 (rarely 10)

Pectoral-fin rays

19 (i + 18)

18-19 (rarely 20)

Pelvic-fin rays

1,5

Pored lateral-line scales

45 + 4

44-47 but rarely 43 or 48

Scale rows above and below lateral line

11/24

10-14 (rarely 9) / 22-24 (rarely 21)

Gill rakers including all rudiments

6 + 14

6-7 + 13-

Body depth

47-57

(52)

Body depth at anal-fin origin

42-65

(46)

Head length

36-57

(39)

Body width

17-30

(20)

Snout length

12-20

(14)

Dermal eye opening

8-14

(10)

Orbit diameter

10-16

(12)

Suborbital depth

6-10

(6)

Interorbital width

7-12

(8)

Upper jaw length

14-22

(15)

Caudal-peduncle depth

11-18

(13)

Caudal-peduncle length

13-23

(16)

Predorsal length

42-68

(48)

Preanal length

65-100

(69)

Prepelvic length

39-44

(62)

Dorsal-fin base

56-88

(62)

Anal-fin base

17-28

(20)

Caudal-fin length

21-35

(27)

Pelvic-fin spine

16-27

(19)

First pelvic-fin ray

25-39

(30)

Longest pectoral-fin ray

23-37

(27)

First dorsal-fin spine

6-10

(7)

Second dorsal-fin spine

10-19

(14)

Third dorsal-fin spine

27-42

(31)

Fourth dorsal-fin spine

21-35

(25)

Fifth dorsal-fin spine

19-32

(23)

Last dorsal-fin spine

8-14

(11)

First dorsal-fin ray

17-27

(20)

Second dorsal-fin ray (longest)

18-31

(22)

First anal-fin spine

22-24

(23)

Second anal-fin spine

8-12

(10)

Third anal-fin spine

19-28

(23)

First anal-fin ray

11-20

(14)

Second anal-fin ray

18-31

(21)

Longest anal-fin ray (third or fourth)

18-33

(22)

Revision of the genus Hapalogenys

Figure 1. The seven species of Hapalogenys. A), H. analis, MUFS 12258, 185 mm SL, East China Sea; B) H. dampieriensis sp. nov., CSIRO

C4036-02, Holotype, 224 mm SL, N of Cape Lambert, north-western Australia; C), H. filamentosus sp. nov., MUFS 7666, holotype, off Iloilo,

Panay Is., Philippines; D), Right pelvic-fin (reversed) of H. filamentosus sp. nov., MUFS 7666, holotype; E), H. kishinouyei, MUFS 12316, 266

mm SL, Miyazaki, Kyushu, Japan; F), H. merguiensis, PMBC 10985, paratype, 199 mm SL, off Satun Province near Phuket, Andaman Sea; G),

H. nigripinnis, Kanagawa Prefectual Museum’s photo database (KPM-NR0001009, photographed by T. Suzuki), about 180 mm SL, Moroyose,

Hyogo, Sea of Japan; H), H. sennin, MUFS11649, holotype, 232 mm SL, Meitsu, Miyazaki, Japan. Arrow shows tip of pelvic-fin ray.

Yukio Iwatsuki and Barry C. Russell

Figure 2. Three stages of 2 species, Hapalogenys dampieriensis sp. nov. (A-C) and H. kishinouyei (D-F). A), AMS L22805-035, paratype, 65

mm SL, Northwest Shelf, 170 km off Port Headland, Australia, B), NTM S. 13569-010, paratype, 134 mm SL, Arafura Sea, Australia, C), AMS

I 22805-035, paratype, 158 mm SL, Northwest Shelf, 170 km off Port Headland, Australia, D) USNM 55610, holotype, 81 mm SL, Tokyo, Japan,

E), MUFS 20896, 131 mm SL, Meitsu, Nango, Miyazaki, Japan, F), MUFS 14286, 168 mm SL, Meitsu, Miyazaki. Japan.

Revision of the genus Hapalogenys

Figure 3. Head (left) and lower jaw ventral view (right) of 4 species, Hapalogenys analis (A-B), H. dampieriensis sp. nov. (C-D), H.filamentosus

sp. nov. (E-F) and H. kishinouyei (G-H). A-B), HUMZ 106037, 113 mm SL, Saeki, Ooita, Japan; C-D), NTM S13510-010, 134 mm SL, Arafura

Sea, Northern Territory, Australia; E-F), MUFS 7654, 134 mm SL, off Iloilo, Panay Is., Philippines; G-H), MUFS 20986, 131 mm SL, Meitsu,

Miyazaki, Japan. Arrow indicates squamation on maxilla.

Yukio Iwatsuki and Barry C. Russell

with scales, extending almost to anterior nostrils; lips, chin and

ventral part of urohyal naked; cheek and opercle fully scaled;

jaws with bands of pointed conical teeth, outermost teeth

generally enlarged, but not forming canines; teeth absent on

vomer and palatines; preopercle serrate; upper opercle with 2

short spines posteriorly; single notched dorsal fin; 3rd dorsal-

fin spine and 2nd dorsal-fin ray longest; 1st dorsal- and anal-fin

soft rays not strongly bifurcated; 1st anal-fin spine short, 2nd

longest, clearly longer than 3rd; dorsal and anal fins with high

scaly sheath; caudal fin generally rounded.

Live colouration. Based on photograph of MUFS 12258 (fig.

lA), neotype, 185 mm SL and MUFS 8422, 113 mm SL: head

and body generally dark brownish, with 7 alternating whitish

and dark brown bands (see Diagnosis); iris black; lips, lower

part of cheek, preopercle and interopercle pale brownish.

Preserved colouration. Based on the neotype and non-type

specimens: head and body generally similar to live colouration.

Distribution. Hapalogenys analis is currently known from the

East Asian Shelf to the Sea of Japan and Pacific coast of southern

Japan (except Ryukyu and Ogasawara Is.), including the western

part of Taiwan and South China Sea (fig. 5). Lim, 1994

questionably reported H. analis from Singapore (3 collected

specimens in total) (K. K. P. Lim, ZRC, pers. comm.).

Ecological note. Most specimens of Hapalogenys analis have

been collected by bottom trawl in depths less than 100 m.

Smaller specimens (< about 100 mm SL) have been collected

by set nets in depths of 20-50 m.

Remarks. The date of publication of Hapalogenys mucronatus

Eydoux and Souleyet (as 1841 on cover, but presumably 1850)

and status of that species as a junior synonym of H. analis

Richardson, 1845 as given by Bauchot et ah, 1982, 1983, Lim,

1994, and Iwatsuki et ah, 2000a is followed here. The type

specimens of H. analis Richardson, 1845 are lost (J. Maclaine,

BMNH, pers. comm.) and a neotype (MUFS 12258, 185 mm SL)

of the species is herein designated in order to avoid confusion.

Hapalogenys dampieriensis sp. nov.

New English name: Australian Striped Velvetchin

Figures IB, 2C-D

Hapalogenys kishinouyei.—non Smith and Pope, 1906; Gloerfelt-

Tarp and Kailola, 1984: 197, upper 2nd left picture of p. 196 (north-

western Australia). -Sainsbury et al., 1985: 214, lowest picture on p.

215 (north-western Australia). -Allen and Swainson, 1988; 82 (north-

western Australia). -Allen, 1997: 128 (north-western Australia and

south-eastern Asia). -McKay, 2001: 2969 (in part, north-western

Australia). -Hutchins, 2001; 34 (Western Australia).

Type material. Holotype: CSIRO C4036-02, 224 mm SL, north of

Cape Lambert, WA, Australia (19°09'S, 117°26-117°28'E), 121-123

m depth, 1 Sep 1995, coll. A. Graham and G. Yearsley. Paratypes: (all

from Australia, n =8): AMS 1.22805-035, 3: 66-158 mm SL, North-

west Shelf, 170 km N of Port Hedland, WA (18°28'S, 118°15'E), 150-

156 m depth, 28 Mar 1982, coll. J. Paxton and M. McGrouther; NTM

S. 13569-010, 3: 127-134 mm SL, 104-108 m depth, Arafura Sea, NT,

18 Oct 1992, coll. R. Williams; CSIRO CA244, 290 mm SL, north-

west of Montebello L, WA (19°58'-19°57'S, 115°12'-115°14'E), 10

May 1978, coll. CSIRO; CSIRO CA1552 (voucher specimen based on

Sainsbury et al., 1985), 231 mm SL, north-east of Monte Bello L, WA

(19°34'-19°36'S, 116°09'-116°12'E), 7 Jun 1980, coll. CSIRO.

Non-type material. 66-300 mm SL, all from Australia, n=17. CSIRO

H4036-03, 194 mm SL, northern Cape Lambert, WA (19°09'S,

117°26'-117°28'E), 1 Sep 2002, coll. A. Graham and G. Yearsley;

CSIRO H4069-03, 81 mm SL, north-west of Port Hedland, WA

(18°38'-18°39'S, 118°7'-118°8' E), 8 Sep 1995, coll. A. Graham and G.

Yearsley; CSIRO H4069-04, 2: 74-75 mm SL, north-west of Port

Headland, WA (18°38'-18°39'S, 118°7'-118°8'E), 8 Sep 1995, coll. A.

Graham and G. Yearsley; CSIRO CA1106-CA1107, 2: both 192 mm

SL, off western Admiralty Bay, WA, 9 Nov, 2002; NTM S12819-002,

300 mm SL, Evans Shoal, Timor Sea, NT, 22 Apr 1990, coll. J.Lloyd;

NTM S13373-010, 172 mm SL, 87 m depth, Arafura Sea, NT, 30 Oct

1990, coll. NT Lisheries; NTM S. 13569-010, 2; 127-129 mm SL,

Arafura Sea, NT, 18 Oct 1992, coll. R. Williams; NTM S 13523-008, 169

mm SL, 97-103 m depth, Arafura Sea, NT, 18 Sep 1992, coll. R.

Williams, NTM S. 13547-002, 65 mm SL, Arafura Sea, NT, 31 Oct,

1992, coll. R. Williams; WAM P25836-002, 269 mm SL, Bernier L, WA

(24°40'S, 112°27'E), 128-161 m depth; WAM P26 194-0 18, 166mmSL,

N of Monte Bello L, off Dampier, WA (19°26'S, 116°31'E), 120-128 m

depth; WAM P26 195-0 16, 97 mm SL, north of Monte Bello L, off

Dampier, WA (19°10'S, 116°46'E), 175-178 m depth; WAM P27244-

002, 215 mm SL, Rowley Shoals, WA(18°37'S, 119°33'E), 114mdepth;

WAM P30426-001, 248 mm SL, West of Barrow L, off Onslow, WA

(20°40'S, 113°43'E), 225-230 m depth; WAM P30658-002, 221 mm SL,

6 miles, NE of Monte Bello L, WA (20°26'S, 115°32'E); WAM P30666-

001, 224 mm SL, Broome, WA (17°58'S, 122°14'E).

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster

of very short papillae anteriorly (figs 3C-D), scaly posteriorly

(on posterior abdominal part of angular of lower jaws, [fig. 3D];

10 unobstructed pores on and behind chin (posteriormost 2

sometimes slit-like) (see figs 3C-D; Gloerfelt-Tarp and Kailola,

1984: 197, fig. 2); no scales on maxilla (figs 3C-D); 4 narrow

longitudinal dark stripes (2nd and 3rd stripes most distinct, 2nd

from nape to base of mid dorsal-fin soft rays, 3rd from eye to

last dorsal-fin ray base) in specimens less than about 100 mm

SL, thereafter 2nd and 3rd stripes visible only, remainder and

3rd stripes not present in specimens less than about 200 mm SL

(figs IB, 2A-C), their width below base of 5th and 6th dorsal-

fin spines clearly narrower than pupil diameter (see figs 2A-C

in 65-158 mm SL and fig. IB); 41-45 pored lateral -line scales;

soft rayed portions of dorsal and anal fins somewhat truncated

posteriorly and slightly angulated posteriorly, respectively (figs

IB, 2A-C); pelvic-fin tip extending slightly beyond anus but

clearly not reaching to base of 1st anal-fin spine when depressed

(figs IB, 2A-C); procumbent spine-like process (tip of 1st

pterygiophore) apparent at origin of dorsal-fin but covered by

predorsal scales.

Description. For the holotype and 5 paratypes, counts and

proportional measurements as percentage of SL are given in

Table 2. Data for the holotype are presented first, followed by

non-type material (if different) in parentheses. Characters

given in the diagnosis are not repeated.

Body deep, 45% of SL (42-55% of SL), relatively compressed,

covered with ctenoid scales, ctenii free on posterior margin of

exposed area; lateral line continuous until on hypural plate; orbit

diameter slightly larger than interorbital space; suborbital depth

Revision of the genus Hapalogenys

Table 2. Counts and proportional measurements, and characters as percentage of standard length of Hapalogenys dampieriensis sp. nov.,

H.filamentosus sp. nov., and H. kishinouyei (Smith and Pope, 1908)

Hapalogenys dampieriensis sp. nov.

Hapalogenys

filamentosus sp. nov.

Hapalogenys

kishinouyei

Holotype

Paratypes*

Non-type

Holotype

Paratypes*

Holotype

Non-type

CSIRO

specimens

MUFS

USNM

specimens

C4036-02

n=8

n=17

7666

n=3

55610

n=39

Standard length (mm)

224

66-290

74-300

147

129-147

45-393

Dorsal-fin rays

XI, 14

XI, 13-

XI, 13-14

XI, 14

XI, 14 (rarely 15)

Anal-fin rays

III, 9

III, 8-9

III, 9

Pectoral-fin rays

17-18

Pelvic-fin rays

1,5

Pored lateral-line scales

41-45

41 or 42

44-47

Scale rows above and below lateral line

12/20

10-12/20-23

11/22

10-11/20

-22

11/23

11-13/23-25

Gill rakers including all rudiments

6+12

5-6 + 11-12

5-6 + 11-

-12

5+11

5 + 11-12

7+11

4-7+11

Scales on maxilla

Scales on posterior and ventral aspects

absent

of angular

present

Body depth

42-55

(48)

74-300 (173)

147

115-149 (136)

44-53

(49)

Body depth at anal-fin origin

35-47

(41)

42-55

(48)

48-50

(49)

38-46

(42)

Head length

34-46

(41)

35-47

(42)

42-43

(42)

36-42

(38)

Body width

18-23

(20)

18-25

(20)

42-44

(43)

17-21

(19)

Snout length

14-16

(15)

13-16

(14)

16-20

(18)

12-17

(14)

Dermal eye opening

8-12

(10)

7-12

(10)

13-16

(14)

7-15

(8)

Orbit diameter

8-16

(12)

8-16

(11)

10-11

(11)

8-17

(10)

Suborbital depth

5-8

(7)

5-8

(7)

(14)

5-8

(7)

Interorbital width

(8)

8-9

(9)

7-8

(7)

8-10

(9)

Upper jaw length

15-16

(16)

15-16

(16)

8-10

(9)

14-16

(15)

Caudal-peduncle depth

11-13

(12)

11-13

(12)

15-16

(16)

11-13

(12)

Caudal-peduncle length

12-17

(16)

9-17

(13)

12-13

(13)

16-18

(17)

Predorsal length

38-49

(46)

38-49

(44)

17-18

(17)

40-49

(44)

Preanal length

71-77

(73)

69-77

(72)

45-49

(48)

67-74

(70)

Prepelvic length

40-52

(47)

39-52

(45)

69-74

(71)

39-48

(41)

Dorsal-fin base

54-59

(57)

54-62

(58)

41-48

(43)

55-62

(59)

Anal-fin base

13-17

(15)

13-20

(16)

57-61

(59)

15-20

(17)

Caudal-fin length

18-29

(25)

18-29

(24)

16-18

(17)

21-31

(25)

Pelvic-fin spine

12-19

(16)

12-19

(15)

27-28

(27)

13-18

(16)

First pelvic-fin ray

24-31

(29)

21-31

(27)

18-21

(20)

22-31

(25)

Longest pectoral-fin ray

19-32

(28)

19-32

(26)

30-35

(33)

23-31

(26)

First dorsal-fin spine

4-9

(7)

4-9

(6)

30-32

(31)

4-9

(6)

Second dorsal-fin spine

7-16

(12)

7-16

(11)

7-10

(8)

6-16

(12)

Third dorsal-fin spine

14-24

(19)

13-24

(18)

13-18

(15)

15-24

(20)

Fourth dorsal-fin spine

14-27

(22)

14-27

(20)

22-26

(24)

17-26

(22)

Fifth dorsal-fin spine

13-27

(21)

13-27

(19)

25-28

(26)

17-25

(21)

Last dorsal-fin spine

3-11

(8)

3-11

(6)

23-27

(24)

6-11

(8)

First dorsal-fin ray

13-24

(14)

6-20

(13)

10-11

(10)

14-19

(17)

Second dorsal-fin ray (longest)

13-22

(18)

13-22

(16)

17-20

(19)

16-21

(19)

First anal-fin spine

5-11

(9)

13-24

(17)

20-23

(21)

(24)

Second anal-fin spine

13-24

(20)

4-11

(7)

(23)

6-11

(8)

Third anal-fin spine

10-15

(13)

13-24

(18)

9-10

(10)

14-20

(18)

First anal-fin ray

13-23

(19)

12-23

(16)

22-24

(23)

9-14

(12)

Second anal-fin ray

11-20

(18)

11-21

(16)

14-16

(15)

16-24

(19)

*Paratypes, AMS 1.22805-035 (2 specimens) and NTM 13547-002, 13523-008, S. 13551-003 and 13569-010 (3 specimens); **Paratypes, MUFS

7654, 7667-7668, 3 specimens; ***Upper and lower gill raker counts decrease with growth from 6 or 7 to 4 or 5 and from 12 or 13 to 11 or 12,

respectively.

Yukio Iwatsuki and Barry C. Russell

Figure 5. Distribution of 4 Hapalogenys species: Hapalogenys Figure 6. Distribution of 3 Hapalogenys species: Hapalogenys

analis, Hapalogenys dampieriensis sp. nov., Hapalogenys merguiensis, H. nigripinnis and H. sennin.v

filamentosus sp. nov. and Hapalogenys kishinouyei.

distinctly less than eye diameter; head covered with scales,

extending almost to snout tip; lips, chin and ventral part of urohyal

naked; cheek and opercle fully scaled; jaws with bands of pointed

conical teeth, outermost teeth generally enlarged, but not forming

canines; teeth absent on vomer and palatines; preopercle serrate;

upper opercle with 2 short spines posteriorly; single notched

dorsal-fin; 4th dorsal -fin spine and 2nd dorsal-fin ray longest; 1st

dorsal- and anal-fin soft rays not strongly bifurcated; 1st anal-fin

spine short, 2nd very robust, longest, clearly longer than 3rd;

dorsal and anal fins with high scaly sheath; caudal fin rounded in

juveniles, becoming truncate with growth.

Live colouration. Based on a colour photograph in Gloerfert-

Tarp and Kailola, 1984: 2nd upper left figure on p. 196,

registered as CSIRO CA1552, 270 mm SL, from north-western

Australia - specimen not seen by us and CSIRO C4036-02,

holotype, 234 mm SL: head and body generally pale silvery

bronze, usually lighter than in H. kishinouyei (figs IB, IE, and

2A-F); dorsal and anal fins translucent or pale brownish; soft

dorsal fin pale; pelvic-fin rays and membranes dusky; pelvic-fin

spine dark-brownish; iris golden-brown; lips, lower parts of

cheek, preopercle, and interopercle pale-brownish.

Preserved colouration. Based on all type specimens: head and

body generally dark-brownish; 4 narrow longitudinal brownish

pale-brown dark stripes (2nd and 3rd stripes most distinct) in

specimens smaller than about 100 mm SL, 2nd and 3rd stripes

only present in larger specimens, their width below base of 5th

and 6th dorsal-fin spines narrower than pupil diameter (see figs

2A-C in 65-158 mm SL and fig. IB); pelvic-fin rays, including

membrane, dusky; pelvic-fin spine pale-brown; iris, lips,

ventral portion of cheek, preopercle, and interopercle brown.

Distribution. Hapalogenys dampieriensis sp. nov. is currently

known only from north-western Australia (see fig. 5) in depths

of 87-230 m. The species is likely to have a continuous

distribution in similar depths between known localities.

Ecological note. The specific habitat at the collection sites of

the type specimens is uncertain, but likely to be dominated by

a muddy rocky bottom, similar to that of H. kishinouyei.

Etymology. The name ''dampieriensis ” refers to the Dampierian

Province (named after the explorer William Dampier), a

biogeographic region extending from approximately Geraldton

in Western Australia across northern Australia to Cape York,

approximating the distribution of the new species in Australia.

Remarks. The largest recorded size of H. dampieriensis sp.

nov. is 300 mm SL (NTM S12819-002), compared with more

than 500 mm SL for H. kishinouyei.

Revision of the genus Hapalogenys

Hapalogenys filamentosus sp. nov.

New English name: Philippines Dark Velvetchin

Figures IC-D, 3E-F

Type material. Holotype: MUFS 7666, 147 mm SL, off Iloilo, Panay

L, Philippines, 10 Mar 1981, bottom trawl, 30-80 m, coll. M. Akazaki.

Paratypes: MUFS 7654, 149 mm SF, off Iloilo, Panay L, Philippines,

bottom trawl, about 40 m depth, 11 Mar 1981, coll. M. Akazaki;

MUFS 7667-7668, 2: 129-143 mm SF, same data as holotype.

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster

of very short papillae (figs 3E-F), scaly posteriorly (on posterior

abdominal part of angular of lower jaws, (fig. 3F); 10

unobstructed pores on and behind chin (fig. 3F; see Gloerfelt-

Tarp and Kailola, 1984:197, fig. 2); no scales on maxilla (fig.

3E); 2 faint longitudinal dark stripes on body, 1st from nape to

base of mid dorsal-fin soft rays, 2nd from eye to base of last

dorsal-fin soft ray, their width below base of 5th and 6th dorsal-

fin spines clearly narrower than pupil diameter (see figs 2A-C

in 65-158 mm SL and fig. IB); soft rayed portions of dorsal and

anal fins somewhat rounded posteriorly and slightly angulated

posteriorly, respectively (fig. 1C); pelvic-fin tip almost reaching

to or slightly beyond base of 1st anal-fin spine when depressed

(figs IC-D); 41 or 42 pored lateral-line scales; a procumbent

spine-like process (exposed tip of 1st pterygiophore) apparent

at origin of dorsal fin but covered by predorsal scales.

Description. Counts and proportional measurements as

percentage of SL of the holotype and 5 paratypes are given in

Table 2. Data for the holotype are presented first, followed by

paratype material (if different) in parentheses. Characters

given in the diagnosis are not repeated.

Body deep, 50% of SL (45-55% of SL), relatively

compressed, covered with ctenoid scales, ctenii free on

posterior margin of exposed area; lateral line continuous until

on hypural plate; orbit diameter slightly larger than interorbital

space; suborbital depth clearly less than eye diameter; head

covered with scales, extending almost to snout tip; lips, chin

and ventral part of urohyal naked; cheek and opercular bones

fully scaled; jaws with bands of pointed conical teeth,

outermost teeth generally much enlarged but no distinct

canines; teeth absent on vomer and palatines; preopercle

serrate; upper opercle with 2 short spines posteriorly; single

notched dorsal fin; 4th dorsal-fin spine and 2nd dorsal-fin ray

longest; 1st dorsal- and anal-fin soft rays not strongly

bifurcated; 1st anal-fin spine short, 2nd very robust, longest,

clearly longer than 3rd; dorsal and anal fins with high scaly

sheath; pectoral-fin tip not reaching to vertical at anus or

pelvic-fin tip when depressed; caudal fin slightly rounded.

Preserved colouration. Based on all type specimens: head and

body generally dark-brownish; 2 faint longitudinal stripes on

head and body.

Distribution. Hapalogenys filamentosus sp. nov. is currently

known only off Iloilo, Panay I., Philippines. Efforts by the first

author to collect other examples of the species from fish markets

and trawl catches throughout the Philippine Is. (Luzon,

Mindoro, Panay, Cebu and Mindanao Is.) have been in vain.

Apart from the type specimens, no examples are known to

exist in museum collections.

Ecological note. The habitat of H. filamentosus sp. nov. is

likely to be similar to that of H. kishinouyei and H.

dampieriensis sp. nov.

Etymology. The name " filamentosus'’ refers to the filamentous

1st ray of the pelvic fin in this species.

Hapalogenys kishinouyei Smith and Pope, 1906

English name; Striped Velvetchin

Japanese Name: Shimasetodai

Figures IE, 2D-F, 3G-H

Hapalogenys kishinouyei Smith and Pope, 1906: 476, fig. 6 (type

locality: Urado, Kanagawa, Japan).-Jordan and Thompson, 1912: 554,

fig. 3 (Tokyo).-Izuka and Matsuura, 1920:150 (Takamatsu, Kagawa,

Japan).-Fowler, 1931: 269 [Urado, Kanagawa and Takamatsu, Kagawa,

Japan, Philippines (doubtful locality)]. -Kyushin et al.,1982: 102 (South

China Sea). -Akazaki, 1984: 173 (East Asian Shelf ).-Okamura et al.,

1985: 489, 678 (Okinawa Trough). -Chen et al., 1997: 102, fig. 306

(South China Sea).-Iwatsuki et al., 2000a; 133 (East Asian Shelf and

Australianspecimensnowref erred to //.4ampieneu5i5sp. nov. ).-Shimada,

2000: 841 [southern Japan, Philippines (doubtful locality), and north-

west Australian specimens now referred to H. dampieriensis sp.

nov.)]. -Randall and Lim, 2000: 619 (South China Sea).-Shinohara et al.,

2001: 326 (Tosa Bay, Japan).-Shimada, 2002: 841 [southern Japan,

Philippines (doubtful locality) and north-west Australian specimens now

referred to H. dampieriensis sp. nov.].-Youn, 2002: 339, 613 (Korea).

Type material. Holotype: USNM 55610, 81 mm SL, Tokyo, Japan (see

fig. 2D).

Non-type material. 45-393 mm SL, n= 39. MUFS 770, 235 mm

SL, Osaka Fish Market, Japan; MUFS 12316, 12343, 16^264 mm SL,

Meitsu, Nango, Miyazaki, Japan, set net, about 25 m; MUFS 12421,

338 mm SL, Ooshima L, Meitsu, Nango, Miyazaki, Japan, large set

net, about 50 m depth; MUFS 12589, 45 mm SL, off Kihachi, Miyazaki,

Japan; MUFS 12852-12854, 3; 242-268 mm SL, Meitsu, Nango,

Miyazaki, Japan, set net, about 50 m; MUFS 14041, 14286, 153-168

mm SL, Meitsu, Nango, Miyazaki, Japan; MUFS 14937, 145 mm SL,

Meitsu, Nango, Miyazaki, Japan; MUFS 16351, 262 mm SL, Meitsu,

Nango, Miyazaki, Japan; SFU 1325, 133 mm SL, Shanghai, China;

SFU Chingtao, China; SFU South China Sea; SFU 58-2573, 183 mm

SL, China; SFU 63-0092, 187 mm SL, China; SFU 3858, 147mm SL,

Taiwan Strait, China; URM-P 28055, 221 mm SL, East China Sea;

URM-P 28056, 207 mm SL, East China Sea near Taiwan.

Diagnosis. A species of Hapalogenys with the following

combination of characteristics: fleshy lower lip with dense cluster

of very short papillae (figs 3G-H), scaly posteriorly (on posterior

abdominal part of angular of lower jaws, fig. 3H); 10 pores on

and behind chin, including a single very small pore near

symphysis, plus 2 moderately-sized pores anteroventrally on

each dentary, a single large pore ventrally at midpoint of dentary,

and a single large pore ventrally at midpoint of angular (2

posteriormost pores sometimes slit-like) (see Gloerfelt-Tarp and

Kailola, 1984:197, fig. 2); no scales on maxilla (fig. 3G); 5 broad

distinct longitudinal stripes, 1st from front of 1st dorsal-fin spine

along dorsal midline, 2nd from nape to base of mid dorsal-fin soft

rays, 3rd from eye to last dorsal-fin ray base, 4th from preopercular

flange, through base of pectoral fin, to lower caudal peduncle,

Yukio Iwatsuki and Barry C. Russell

last from isthmus to base of anal spinous fin; the 1st, 4th and 5th

stripes lost in specimens larger than about 250 mm SL (figs IE,

2D-F) and their width below base of 5th and 6th dorsal-fin spines

clearly greater than pupil diameter at all sizes; soft rayed portions

of dorsal and anal fins somewhat rounded posteriorly and

somewhat angulated posteriorly, respectively (figs IE, 2D-E);

pelvic-fin tip extending slightly beyond anus but clearly not

reaching to base of 1st anal-fin spine when depressed (figs IE,

2D-E); 44-47 pored lateral-line scales; a procumbent spine-like

process (exposed tip of 1st pterygiophore) at origin of dorsal.

Description. Counts and proportional measurements as

percentage of SL of the holotype and 5 paratypes are given in

Table 1. Data for the holotype are presented first, followed by

non-type material (if different) in parentheses. Characters

given in the diagnosis are not repeated.

Body deep, 49% of SL (44-53% of SL), relatively

compressed, covered with ctenoid scales, ctenii free on

posterior margin of exposed area; lateral line continuous until

on hypural plate; orbit diameter clearly larger than interorbital

space; suborbital depth clearly less than eye diameter; head

covered with scales, extending almost to anterior nostrils; chin

and ventral part of urohyal naked; cheek and opercular bones

fully scaled; jaws with bands of pointed conical teeth,

outermost teeth generally much enlarged but no distinct

canines; teeth absent on vomer and palatines; preopercle

serrate; upper opercle with 2 short spines posteriorly; single

notched dorsal fin; 4th dorsal-fin spine longest; 1st dorsal- and

anal-fin soft rays not strongly bifurcated; 1st anal-fin spine

short, 2nd anal fin longest, clearly longer than 3rd; dorsal and

anal fins with high scaly sheath; pectoral-fin tip not reaching

to vertical at anus or pelvic-fin tip when depressed; caudal fin

generally truncate.

Live colouration. Based on colour photographs of specimens

(MUES 20896, 14286): head and body generally dark-brownish,

slightly lighter on jaws; dorsal, anal and pectoral fins dark-

brownish (similar to body); pelvic-fin rays and membranes

black; pelvic-fin spine dark-brownish; iris brownish-golden

(orange-golden); lips, lowest parts of cheek and preopercle, and

interopercle pale-brownish.

Preserved colouration. Based on the holotype and non-type

specimens: head and body generally dark-brownish; 5

longitudinal stripes on body; pelvic-fin rays with black

membrane, pelvic-fin spine pale-brown; iris, lips, ventral

portion of cheek and preopercle, and interopercle brown.

Distribution. Hapalogenys kishinouyei is currently known

only from the East Asian Shelf (fig. 5).

Biological note. The largest specimen of Hapalogenys

kishinouyei studied was 556 mm SL, collected from a set net

catch at Meitsu fish market, Nango, Miyazaki, Japan on 14 Nov

1994 (specimen not kept). This species commonly reaches

150-300 mm SL.

Remarks. Although Fowler, 1931 reported Hapalogenys

kishinouyei from the Philippines and adjacent regions, the

species was originally based on specimens from Urado,

Tokyo, and Takamatsu, Kagawa, Japan. Despite subsequent

reports including the Philippines in the species’ distribution

(Iwatsuki et al., 2000a; Shimada, 2000, 2002), attempts by us

to locate H. kishinouyei specimens in fish markets and

museum collections over the past ten years have been

unsuccessful in documenting any Philippine occurrences of

this species. This strongly suggests that the species is not

distributed in the Philippines. However, Chen et al., 1997

provided a fine colour plate of the species from the South

China Sea and van Quan (unpublished document) listed the

species from northern Vietnam.

Hapalogenys merguiensis Iwatsuki, Satapoomin and Amaoka,

2000

New English name: Mergui Velvetchin

Figures IF, 4A-B

Hapalogenys merguiensis Iwatsuki, Satapoomin and Amaoka,

2000a: 133, fig. lA-C (type locality: Mergui Basin, southern Myanmar

Sea, Andaman Sea).

Type material. Holotype: HUMZ 90021, 242 mm SL, Mergui Basin,

southern Myanmar Sea (11°27'N, 97 °16'E). Paratypes (174-242 mm

SL, n=6): AMS I. 22739-001, 192 mm SL, southern Andaman Sea, off

Satun Province near Phuket, W coast of southern Thailand; HUMZ

90022, 178 mm SL, same data as holotype; MULS 15800, 177 mm SL,

Taninthayi Coast, southern Myanmar Sea; PMBC 10985, 199 mm SL,

off Satun Province near Phuket, W coast of southern Thailand, southern

Andaman Sea; PMBC 10986, 179 mm SL, southern Andaman Sea, off

Satun Province near Phuket, W coast of southern Thailand.

Non-type material. HUMZ 33397, 172 mm SL, Andaman Sea.

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster of

very short papillae and barbels (figs 4A-B), slightly scaly

posteriorly (fig. 4B); 10 pores on and behind chin, including a

single very small pore near symphysis, plus 2 moderately-sized

pores anteroventrally on each dentary, a single large pore

ventrally, at midpoint of dentary, and a single large pore ventrally

at midpoint of dentary and angular (2 posteriormost pores

sometimes slit-like) (see Gloerfelt-Tarp and Kailola, 1984:197,

fig. 2); no scales on maxilla (fig. 3A); 2 indistinct oblique dark

bands, 1st descending from nape to behind pectoral fin, and 2nd

from base of anterior 2nd or 3rd dorsal-fin spine and soft dorsal-

fin base, curving backwards through lateral line to soft anal fin

and caudal peduncle (1 whitish or indistinct pale brown oblique

band, curving backward and becoming wider, from base of first

3 dorsal-fin spines, to anus and spinous anal-fin base) (fig. IF);

soft rayed portions of dorsal and anal fins rounded posteriorly

and somewhat truncate posteriorly, respectively (fig. IF); pelvic-

fin tip not reaching to 1st anal-fin spine when depressed (fig. IF);

39-42 pored lateral-line scales; a procumbent spine-like process

(exposed tip of 1st pterygiophore) at origin of dorsal fin.

Description. A detailed description was given by Iwatsuki et

al., 2000a and is not repeated here.

Distribution. Hapalogenys merguiensis is currently known

only from the Andaman Sea, in depths from about 80-180 m

(see Iwatsuki et al., 2000a).

Revision of the genus Hapalogenys

Hapalogenys nigripinnis Schlegel in Temminck and Schlegel,

1843

New English name: Short Barbeled Velvetchin

Japanese name: Higesoridai

Figures IG, 4C-D

Pogonias nigripinnis Schlegel in Temminck and Schlegel, 1843:

59, pi. 25 (type locality: Nagasaki Bay, Nagasaki, Japan).

Hapalogenys nitens Richardson, 1844b: 84, pi. 43, figs 1-2 (type

locality: Canton, China). -Richardson, 1844a: 463 (Macao, China).

Hapalogenys aculeatus Nystrdm, 1887: 10 (type locality: Japan).

Hapalogenys guentheri Matsubara, 1933 (originally gUntheri):

86, fig. 6 (type locality: Pusan, South Korea).

Type material. Lectotype: RMNH D282, 292 mm SL, Nagasaki Bay,

Nagasaki, Japan. Paralectotpes: RMNH D283 (dried), 245 mm SL,

RMNH D284 (dried), 206 mm SL, RMNH 722, 240 mm SL, RMNH

D2216 (dried), 347 mm SL, ZMB 8121 (dried), 198 mm SL, all as

Nagasaki, Japan.

Non-type material. BMNH 1968.3.11.1 {\\o\o\y^t oi Hapalogenys

nitens), 107 mm SL, Canton, China; LAKU 51028-51029 (holotype

and paratype of Hapalogenys guentheri, [originally gUntheri}), 2: 205

mm SL and 138 mm SL, respectively, Pusan, Korea, holotype coll,

date unknown, paratype coll. 30 Apr. 1927; ZMUU 275 (holotype of

Hapalogenys aculeatus), 320 mm SL, Nagasaki, Japan. 43 other

specimens examined (53-404 mm SL) are listed in Iwatsuki and

Nakabo, 2005.

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster of

very short papillae and barbels (figs 4C-D), scaly posteriorly

(on posterior abdominal part of angular of lower jaws, fig. 4D);

10 pores on and behind chin, including a single very small pore

near symphysis (usually not apparent owing to dense covering

papillae), plus 2 moderately-sized pores anteroventrally on each

dentary, a single large pore ventrally at midpoint of dentary, and

single large pore ventrally at articulation of dentary and angular

(2 posteriormost pores usually slit-like concave traces or slit-

like) (see figs 4C-D; Gloerfelt-Tarp and Kailola, 1984: 197, fig.

2); scales on maxilla (fig. 4C); body often with 2 indistinct

oblique dark bands, 1st descending from nape to behind pectoral

fin and running to posterior part of soft anal-fin rays, 2nd

descending from base of anterior 3rd or 4th dorsal-fin spines

and soft dorsal-fin base, curving backwards through lateral line

to upper part of caudal peduncle (fig. IG, often uniformly dark

or pale-brown); soft rayed portions of dorsal and anal fins

strongly rounded posteriorly and slightly angulated posteriorly,

respectively (fig. IG); pelvic-fin tip not reaching to base of 1st

anal-fin spine when depressed (fig. IG); 45-48 (rarely 44 or 49)

pored lateral-line scales; a procumbent spine-like process

(exposed tip of 1st pterygiophore) almost completely hidden by

predorsal scales at origin of dorsal fin.

Description. A detailed description was given by Iwatsuki et

ah, 2000a and is not repeated here.

Distribution. Hapalogenys nigripinnis is currently known only

from the East Asian Shelf in depths less than 100 m (see

Iwatsuki and Nakabo, 2005; see fig. 6). The species has not

been confirmed as occurring around the Ryukyu or Ogasawara

Is., or southern Japan.

Remarks. The status of H. aculeatus Nystrdm, 1887, H. nitens

Richardson, 1844b and H. guentheri Matsubara, 1933

(originally H. guntheri) as junior synonyms of H. nigripinnis

Schlegel in Temminck and Schlegel, 1843, was discussed by

Iwatsuki and Nakabo, 2005.

Hapalogenys sennin Iwatsuki and Nakabo, 2005

New English name: Long Barbeled Velvetchin

Japanese name: Higedai

Figures IH, 3E-F

Hapalogenys sennin Iwatsuki and Nakabo, 2005: 861 (type

locality: Meitsu, Miyazaki, Japan).

Type material. Holotype: MULS 11649 (fig. IH), 232 mm SL, Meitsu,

Nango, Miyazaki, Kyushu L, Japan (31°31.9'N, 131°23.5'E), set net, less

than 30 m depth. Paratypes (n=13, all from Japan): LAKU 38698, 234

mm SL, Oki L, Shimane; LAKU 85960, 227 mm SL, Tateyama, Chiba,

Boso Peninsula; LRLM 3715, Shima-cho, Shima-gun, Mie; KPM-NI 49,

208 mm SL, Kanagawa; MULS 2086, 176 mm SL, Miyazaki fish market,

Miyazaki, Japan; MULS 7149, 68 mm SL, mouth of the Kaeda R.,

Miyazaki, Japan; MULS 11678, 207 mm SL, female, Meitsu, Nango,

Miyazaki; MULS 14627, 79 mm SL, mouth of the Kaeda R., Miyazaki;

MULS 16060, 248 mm SL, Shirahama, Wakayama; MULS 20810, 194

mm SL, Moroyose, Hyogo, Sea of Japan; MULS 21573, 296 mm SL,

female, Meitsu, Nango, Miyazaki; MULS 22226, 46 mm SL, Hitotsuba

Inlet, Ooyodo R., Miyazaki; NSMT-P 60236, 174 mm SL, Suzaki, Chiba;

OMNH-P 2682, 183 mm SL, Moroyose, Hyogo, Sea of Japan, set net.

Non-type material. 21 specimens examined (49-296 mm SL, all

from Japan) are listed in Iwatsuki and Nakabo, 2005.

Diagnosis. A species of Hapalogenys with the following

combination of characters: fleshy lower lip with dense cluster of

long and short papillae and barbels on chin (figs 4E-F), not scaly

posteriorly (on posterior abdominal part of each angular of lower

jaws, fig. 4F); 10 pores on and behind chin (only 4 slit-like pores

on angular on each lower jaw in young and then becoming

covered in adults; fig. 4F); no scales on maxilla (fig. 4E); 2 faint

oblique dark bands on body, 1st descending from nape to behind

pectoral fin and then running to posterior part of soft anal -fin rays,

and 2nd from base of anterior 3rd or 4th dorsal-fin spines and soft

dorsal-fin base, curving backward through lateral line to upper

part of caudal peduncle, or often becoming uniform dark-brown

on body (fig. 4E-H of Iwatsuki and Nakabo, 2005); soft rayed

portions of dorsal and anal fins strongly angulated posteriorly

(fig. IH, anal more strong angulated than dorsal); pelvic-fin tip

not reaching to base of 1st anal -fin spine when depressed (fig.

IH); 44-45 (rarely 43 or 46) pored lateral -line scales; a

procumbent spine-like process (exposed tip of 1st pterygiophore)

at origin of dorsal fin, sometimes hidden by predorsal scales.

Description. A detailed description was given by Iwatsuki and

Nakabo, 2005 and is not repeated here.

Distribution. Hapalogenys sennin is currently known only

from the southern part of Japan (excluding the Ryukyu and

Ogasawara Is.) and is possibly endemic to the Japanese region

(fig. 6; see Iwatsuki and Nakabo, 2005).

Ecological notes. Collection data indicate that habitat of

Hapalogenys sennin is restricted to river mouths and coastal

rocky-sandy bottoms in depths of less than 50 m. Detailed

habitat features are given in Iwatsuki and Nakabo, 2005.

Yukio Iwatsuki and Barry C. Russell

Figure 4. Head (left) and lower jaw ventral view (right) of 3 species, Hapalogenys merguiensis (A-B), H. nigripinnis (C-D) and H. sennin (E-F).

A-B), HUMZ 33397, 172 mm SL, Andaman Sea; C-D), ZMUU 275, holotype of H. aculeatus, 320 mm SL, Nagasaki, Japan; E-F), MUFS

11649, holotype, 232 mm SL, Meitsu, Nango, Miyazaki, Japan. Arrow indicates squamation on maxilla.

Revision of the genus Hapalogenys

Discussion

Iwatsuki et al., 2000a demonstrated that Hapalogenys

maculatus Richardson, 1846, from Canton, China, should be

considered as a ''nomen dubium"\ because the seven presently

known valid species of Hapalogenys all lack the round spots

on the upper half of the body, tail and vertical fins that

supposedly characterise H. maculatus. The description of H.

maculatus is highly suggestive of Plectorhinchus cinctus

Schlegel in Temminck and Schlegel, 1843, a haemulid (Iwatsuki

et al. 2000a).

Other nominal species previously placed in Hapalogenys

include: Hapalogenys meyenii Peters (synonymized under

Parapristipoma trilineatum Thunberg by Iwatsuki et al.,

2000b); Hapalogenys petersi Day (subsequently removed to

the genus Dinoperca, and assigned to a new family

Dinopercidae, by Heemstra and Hecht, 1986); and

Hapalogenys pictus Tortonese (transferred to Plectorhinchus

by Smith, 1962). In addition, the holotype (ZMB 10179, 215

mm SL) of H. atlanticus Reichenow from Chinchoxo (=

Chichoua), Gabon, West Africa, was shown to be a senior

synonym of Centrarchops chapini Fowler by Heemstra and

Iwatsuki (in press). Hapalogenys nigripinnis Schlegel in

Temminck and Schlegel, 1843 was shown to be a senior

synonym of H. nitens Richardson, 1844b, H. aculeatus

Nystrdm, 1887 and 77. guentheri Matsubara, 1933, by Iwatsuki

and Nakabo, 2005. Accordingly, we recognise seven valid

Hapalogenys species, including two new species: H. analis,

H. dampieriensis sp. nov., H. filamentosus sp. nov., H.

kishinouyei, H. merguiensis, H. nigripinnis and H. sennin

(figs lA-H and 1 3).

Table 3 includes selected characters of the above seven

Hapalogenys species. All have the following combination of

characters: dorsal-fin rays X or XI, 13-17, anal-fin rays III,

nine or ten, relatively deep body, a procumbent spine-like

process (exposed tip of first pterygiophore) at origin of dorsal

fin, ten pores (total) on and behind chin, including a single

very small pore near symphysis plus four pores on each lower

jaw, and a cluster of very short dense barbels and/or papillae

on the fleshy lower lip (figs 3-4; Iwatsuki et al., 2000a).

Hapalogenys analis differs from other congeners in having

five to seven alternating whitish and dark-brown bands on the

head and body (fig. lA), a black margin posteriorly on the

dorsal-, anal- and caudal-fin soft rays (fig. lA), black membrane

on the spinous dorsal- and anal-fin (until first anal-fin ray), and

pelvic-fin rays (fig. lA), and well-developed papillae and

barbels on the fleshy lower lip (figs lA, 3A-B). Other congeners

lack these characters (figs IB-H). Hapalogenys nigripinnis

and H. sennin differ from other Hapalogenys species in having

vertical and longitudinal stripes on the body (fig. IB-H),

higher counts of pored lateral-line scales (usually 44-48 vs.

44-45; Table 3). In addition, H. sennin has extremely well-

developed papillae and barbels on the fleshy lower lip with a

dense cluster on the chin (fig. 4F). Hapalogenys nigripinnis

differs from the latter in having scales on the maxilla (vs.

absent in H. sennim, fig. 4C, E) and rounded soft dorsal fin

posteriorly (vs. strongly angulated in H. sennim, fig. IG, IH).

Hapalogenys dampieriensis sp. nov., H. filamentosus sp.

nov. and H. kishinouyei are similar to each other in overall

appearance and are accordingly identified as the "Hapalogenys

kishinouyei complex”, a species group defined by having two

to five longitudinal stripes on the body. However, H.

dampieriensis sp. nov. differs from the two other species in

having four narrow dark longitudinal stripes, two of which are

lost in adults (figs 2A-C in 65-158 mm SL and fig. IB in about

270 mm SL). In contrast, H. kishinouyei has five (only two in

adults) broad dark longitudinal stripes on the body (figs 2D-F

in 81-168 mm SL and fig. IE in 266 mm SL), and H.

filamentosus has two faint narrow longitudinal body stripes,

the filamentous pelvic-fin ray almost reaching to or slightly

beyond base of first anal-fin spine when depressed (vs. slightly

beyond anus but not reaching to first anal-fin spine base in H.

dampieriensis and H. kishinouyei-, figs 1C, D). Furthermore,

H. filamentosus differs from H. dampieriensis sp. nov. in

having the posteriormost angle of the jaw reaching to a vertical

through the centre of the eye (vs. the jaw reaching slightly

beyond a vertical through the anteriormost eye membrane in

H. dampieriensis sp. nov. and H. kishinouyei in similar sized

specimens of 130-160 mm SL, fig. 1C and figs 2B-C, 2E-F).

Randall, 1981 reported many marine antitropical and anti-

equatorial species from the Indo-Pacific, and the distribution

of the genus Hapalogenys itself also seems to be generally

anti-tropical (figs 5-6; Randall, 1981). Hapalogenys analis,

H. kishinouyei, H. nigripinnis and H. sennin have sympatric

distributions and occur essentially in the same areas, viz.

“East Asian Shelf”, except off the Ryukyu and Ogasawara Is.

However, within this general area the collection data indicate

they occupy different niches.

Hapalogenys kishinouyei is distributed in depths less than

200 m along the Pacific coast of Japan from Boso Peninsula,

Honshu L, through the southernmost Shikoku and Kyushu Is.;

the East China Shelf slope from Senkaku I. to southern

Kyushu; and in the South China Sea. Hapalogenys analis is

distributed in the East China Sea (including western Japan)

and South China Sea in shallow coastal waters, in depths less

than 100 m. Hapalogenys nigripinnis is densely distributed

off southern Japan (except the Ryukyu and Ogasawara Is.),

southern Korea, East China Sea, Taiwan Strait and Hong

Kong, in depths less than 50 m (Iwatsuki and Nakabo, 2005).

H. sennin is generally distributed off southern Japan (except

the Ryukyu and Ogasawara Is.) and inhabits shallow coastal

rocky and sandy shores in depths less than about 30 m during

spring and autumn (unknown in winter; Iwatsuki and Nakabo,

2005). By comparison, H. dampieriensis sp. nov., H.

filamentosus sp. nov. and H. merguiensis have allopatric

distributions and occur off north-western Australia, the

Philippines (off Iloilo, Panay L), and in the Andaman Sea,

respectively.

Comparative material examined. Centrarchops chapini: ZMB

10179 (holotype of Hapalogenys atlanticus), 215 mm SL,

Chinchoxo (=Chichoua), Gabon, west Africa Parapristipoma

trilineatum: ZMB 1050 (holotype of Hapalogenys meyenii),

340 mm SL, Manila, Philippines.

Yukio Iwatsuki and Barry C. Russell

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Revision of the genus Hapalogenys

Acknowledgments

The authors are very grateful to M. McGrouther and H. Motomura

(AMS), A.-M. Hine, J. Maclaine and P. Campbell (BMNH),

D. Catania, T. Iwamoto and W. N. Eschmeyer (CAS),A. Graham,

P. Last, and G. Yearsley (CSIRO), T. Nakabo (FAKU), S. Kimura

(FRLM), K. Nakaya, M. Yabe and K. Amaoka (HUMZ), C. Li

and X. Li (lOCAS), H. Senou (KPM), P Pruvost and G. Duhamel

(MNHN), K. Matsuura and G. Shinohara (NSMT), H. Larson

and R. Williams (NTM), K. Hatooka (OMNH), U. Satapoomin

and S. Bussarawit (PMBC), J. Johnson (QM), M. J. P van Oijen

and J. van Egmond (RMNH), H.-L. Wu (SFU), T. Yoshino

(URM), S. Jewett, J. Williams, L. Palmer, V. G. Springer, and

S. Raredon (USNM), G. Moore and B. Hutchins (WAM),

P Bartsch, H.-J. Paepke and C. Lamour (ZMB), M. Eriksson and

B. Femholm (ZMUU) and K. K. P Lim (ZRC) for loans and

facilities made available during the authors’ visits.

We also thank I. Akagawa for translation of French articles

into Japanese, T. Suzuki (Amagasaki High School, Hyogo,

Japan) and H. Senou (KPM) for use of photographs of

Hapalogenys nigripinnis, andT. Yoshino (URM) for suggestions

regarding the treatment of historical references. Finally, we

thank G. S. Hardy, Ngunguru, New Zealand, for critical

comments on the manuscript, and for help with English. Lastly,

we are very grateful for many helpful suggestions to improve

the manuscript from two anonymous reviewers and the guest

editors. This study was supported in part by grants, awarded to

the first author, by the Ministry of Education, Science, Sports

and Culture, Japan (1364069 and 16570079).

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Memoirs of Museum Victoria 63(1): 47-56 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Two new species of Parapercis (Perciformes: Pinguipedidae) from north-eastern

Australia, and rediscovery of Parapercis colemani Randall & Francis, 1993

Jeffrey W. Johnson

Ichthyology, Queensland Museum, PO Box 3300, South Brisbane, Q 4101, Australia (jeff.johnson@qm.qld.gov.au)

Abstract Johnson, J.W. 2006. Two new species of Parapercis (Perciformes: Pinguipedidae) from north-eastern Australia, and

rediscovery of Parapercis colemani Randall & Francis, 1993. Memoirs of Museum Victoria 63(1): 47-56.

Two new species of pinguipedid fishes, Parapercis flavolabiata and P. sexlorata, are described. The former is known

from 20 specimens collected by epibenthic sled and demersal trawl from within the Great Barrier Reef, Australia, between

Anzac Reefs and Lady Musgrave Island, in depths from 45 to 79 m. It is most similar to Parapercis multiplicata Randall,

1984 and P. colemani Randall & Francis, 1993 in appearance, but is unique among the genus in having a combination of

dorsal rays V, 21, anal rays I, 17-18, lateral-line scales 51-55, palatines edentate, angle of subopercle produced and with

small spinules, 9-10 abdominal and 20 caudal vertebrae, and colouration including six broad red saddles across the back,

a purplish-red line from the anterior edge of each orbit around the snout, and a series of yellow-edged red spots on the

postorbital part of the head and nape. Parapercis sexlorata is described from seven specimens trawled off central eastern

Australia between Cape Moreton, Queensland and North Solitary Island, New South Wales in 86 to 137 m. It is easily

distinguished from its closest congeners with palatine teeth and dark transverse bars, P. macrophthalma Pietschmann,

1911 and P. muronis Tanaka, 1918, by having four rather than five dorsal-fin spines and six rather than five transverse bars.

Parapercis colemani Randall & Francis, 1993 was previously known only from the holotype, collected in 1.5 m depth in

the lagoon of Norfolk Island. Additional specimens collected on the NORFANZ survey in the Tasman Sea indicate that

the species usually inhabits much deeper habitats and occurs over a wider geographic area. New localities and meristic

data are reported for the additional material.

Keywords Pinguipedidae, Parapercis, new species, Queensland, Great Barrier Reef, Australia

Introduction

Parapercis Bleeker, 1863 currently includes about 54

recognised species, including 51 from the Indo-west Pacific

and single representatives in the south-eastern Pacific, south-

eastern Atlantic and north-eastern Atlantic (Cantwell, 1964;

Randall and McCosker, 2002; Randall, 2003; Randall and

Yamakawa, 2006; this study). While shallow reef-dwelling

species are relatively well known, those found on soft-bottom

habitats in depths in excess of 50 m have been more poorly

sampled and elucidated. A study of pinguipedid fishes in

Australian ichthyological collections has revealed a number of

undescribed species taken during demersal trawl surveys. Two

of these species are described herein.

Parapercis colemani Randall & Francis, 1993 was described

from a single female collected in a shallow sandy bay at Norfolk

I. The authors indicated that they were unable to locate further

specimens in the region despite expending considerable effort,

and surmised that the species may have been a waif from a

breeding population elsewhere, possibly in relatively deep

water. In 2003 a consortium of Australian, New Zealand and

French scientific organisations collaborated to investigate the

biodiversity of benthic communities on the Norfolk Ridge and

Lord Howe Rise of the Tasman Sea on the NORFANZ cruise.

Specimens of an unidentified species of Parapercis collected at

two widely separated localities during this cruise are here

identified as P. colemani, confirming these assumptions. New

localities, meristic data, and other details at variance with the

original description are reported for the material.

Methods and materials

Institutional acronyms for types and comparative material are

those of Leviton et ah, 1985. Lengths of specimens are given

as standard length (SL), being the distance from the front of

the upper lip to the base of the caudal fin (posterior end of the

hypural plate). The abbreviation HL is used for head length.

Lateral-line scale counts are to the base of the caudal fin, and

do not include several additional pored scales on the fin. Gill-

raker counts include all rudiments. Vertebral counts were made

from radiographs. Where assessed, sex was determined by

examination of gonads through a small incision in the abdomen.

Jeffrey W. Johnson

Measurements were taken with dial calipers, to the nearest 0.5

mm for SL and the nearest 0.1 mm for all other measurements,

with the aid of a stereo microscope where necessary. Methods

for counts and measurements otherwise follow Randall, 2003.

Where different, values for paratypes follow those of the

holotype in parentheses. Specimens of P. flavolabiata were

collected by epibenthic sled and P. sexlorata by trawl, except

where indicated otherwise. Meristic and morphometric details

for the new species are presented in Tables 1-2.

Comparative material examined

Parapercis macrophthalma ZMB 16160, holotype, 107.5 mm,

Takao, Taiwan, Dr. Haberer.

Parapercis muronis BSKU 36390, 126 mm, Tosa Bay, Japan, 6

Feb 1982; BSKU 37293, 122 mm, Tosa Bay, Japan, 27 Apr 1982.

Parapercis flavolabiata sp. nov.

Yellowlip Grubfish

Figures 1-2, 5; Tables 1-2.

Material examined. Holotype. QM 1. 37762, male, 84 mm, east of

Mystery Cay, Qld, 21°22.5'S, 152°07.5'E, 55 m, C. Bartlett on FRY

Lady Basten, 19 Nov 2005.

Paratypes: (N = 16) AMS 1.43796-001, 3: 39-85 mm, NW of Reef

21-253, Qld, 21°20.1'S, 152°15.9'E, 60 m, C. Bartlett on FRY Lady

Basten, 20 Nov 2005; CSIRO H.6303-01, 79 mm, SE of Anzac Reefs,

Qld, 18°47.rS, 147°57.3'E, trawl 79.2 m,D. Gledhill onPPN Gwendoline

May, 13 Dec 2003; QM 1. 36139, 3: 55-83 mm, west of Riptide Cay,

Qld, 21°14.rS 151°45'E, 45 m, C. Bartlett on ERY Lady Basten, 25 May

2005; QM 1. 36598, 43 mm, NE of Eulalie Reef, Qld, 19°06.3'S,

148°06.9'E, 55 m, C. Bartlett on ERY Lady Basten, 9 Sept 2004; QM 1.

36632, 91 mm, west of Stucco Reef, Qld, 19°33.3'S, 149°31'E, 77 m, C.

Bartlett on ERY Lady Basten, 11 Sept 2004; QM 1.37763, 2: 49 mm,

east of North Reef, Qld, 23°ll.rS, 152°00.9'E, 65 m, C. Bartlett on ERY

Lady Basten, 9 Nov 2005; QM 1.37764, 3: 35-53 mm, north of Joist

Reef, Qld, 19°26. 1'S, 149°40.5'E, 74 m, C. Bartlett on ERY Lady Basten,

26 Nov 2005; QM 1.37765, 48 mm, NE of Hyde Reef, Qld, 19°42.3'S,

150°10.5'E, 70 m, C. Bartlett on ERY Lady Basten, 26 Nov 2005; QM

1.37766, 67 mm, SE of Lady Musgrave L, Qld, 23°54.9'S, 152°30.9'E,

52 m, C. Bartlett on ERY Lady Basten, 5 Nov 2005.

Other material: BPBM 40411, 59 mm, north of Hyde Reef, Qld,

19°39.9'S, 150°04.5'E, 77 m, C. Bartlett on PRY Lady Basten,

26/11/2005; CSIRO H.6329-01, 83 mm, WSW of Mystery Cay, Qld,

21°24.9'S, 151°52.5'E, trawl 67 m, D. Gledhill on PRY Gwendoline

May, 27/11/2005; QM 1.37767, 82 mm, NE of Lavers Cay, Qld,

21°ll.rS, 152°07.5'E, trawl 61 m, D. Gledhill on PRY Gwendoline

May, 3 Dec 2005.

Diagnosis. Dorsal-fin rays V, 21; anal-fin rays I, 17-18 (usually

I, 17); pectoral-fin rays 16-18 (usually 17); lateral -line scales

51-55; gill rakers 3-5-1-10-11; predorsal scales 8-9; scales on

cheek cycloid, in about 11 horizontal rows; usually 8 canine

teeth in outer row at front of lower jaw; vomer with a crescentic

row of 6-10 robust conical teeth; palatines edentate; angle of

subopercle produced, with about 7 small spinules; 9-10

abdominal and 20 caudal vertebrae; colouration including 6

broad red saddles across the back, a purplish red to violet line

from the anterior edge of each orbit around the snout, a series

of yellow-edged red spots on the postorbital part of the head

and nape, and a row of dark-red spots along soft dorsal fin.

Description. Dorsal-fin rays V, 21; anal-fin rays I, 17 (I, 17-18,

only 2 with I, 18); all dorsal- and anal-fin rays branched, last to

base; pectoral-fin rays 17 (16-18, 2 with 16, 1 with 18), upper

ray unbranched, others including lowermost branched; pelvic-

fin rays I, 5; branched caudal-fin rays 14 (14-15); lateral-line

scales 53 (51-55), plus 2-3 smaller scales on caudal-fin base;

scales above lateral line to origin of dorsal fin 5 (5-6), to base

of anterior soft rays of dorsal fin 31 / 2 ; scales below lateral line in

an oblique row to origin of anal fin 11 (11-12); circumpeduncular

scales 24; predorsal scales 9 (8-9), extending just anterior to a

vertical from preopercular margin; horizontal row of scales

from preorbital across cheek to edge of preopercle 19; gill

rakers on 1st arch 3-1-11, total 14 (3-5-1-10-11=13-16);

branchiostegal rays 6; vertebrae 9-1-20 (one paratype with

9 +20, 2 with 10+20).

Body depth 6.15 (5.7-6.3) in SL; body subcylindrical,

greatest width 0.95 (0.90-0.95) in body depth; head length

3.35 (3. 1-3.4) in SL; snout pointed, its length 3.6 (3.55-4.0) in

HL; orbit diameter 3.45 (3.35-3.7) in HL; eyes directed more

laterally than dorsally, bony interorbital space narrow, 8.05

(8.0-11.2) in HL; caudal-peduncle depth 3.3 (3.15-3.6) in HL;

caudal-peduncle length 2.95 (2.85-3.5) in HL.

Mouth slightly oblique, lower jaw protruding slightly, with

curved canine teeth at front of each jaw slightly projecting and

visible when mouth is closed; upper jaw extending to a vertical

at anterior margin of pupil, upper jaw length 2.5 (2.3-2.65) in

HL; upper jaw with 22 (20-23) outer curved canines, first 8 or

9 clearly the largest, of these 8th or 9th the largest, those

following gradually reducing in size posteriorly, broad inner

band of villiform teeth anteriorly, narrowing gradually to form

only 2 rows at rear of jaw; front of lower jaw on each side with

4 (3-4) enlarged curved canines in distinctly separate outer

row (6 paratypes with total of 8, 3 with 7, and 7 with 6), tooth

nearest symphysis smallest, the others gradually increasing in

size laterally, last in the series largest, broad inner band of

villiform teeth extending posteriorly from symphysis to side

of jaw just posterior to largest tooth in outer row, next 6 (5-9)

teeth in single row, enlarged and gradually increasing in size

posteriorly, then followed by single row of 14 (13-15) smaller

slightly curved conical teeth. Vomer with crescentic row of 9

(6-10) robust conical teeth, medial teeth largest, usually

several smaller additional teeth posteriorly; palatines edentate.

Tongue spatulate with broadly rounded tip.

Gill membranes united with broad free fold, not attached to

isthmus. Gill rakers short, the longest about one-third length of

longest gill filament on first gill arch. Anterior nostril small,

inconspicuous, situated in front of centre of eye, slightly more

than half distance to tip of snout, with membranous posterior

flap, the latter folded around near base to form partial tube, flap

usually lying flat against snout in preserved specimens. Posterior

nostril slightly more than half distance from snout to anterior

margin of eye, dorsoposterior to, and about twice width of

anterior nostril, its opening simple, aperture slightly oval-shaped;

intemarial distance about 1.5 times width of posterior nostril.

Opercle with distinctly exposed, robust, pointed spine,

angle of subopercle expanded and armed with about 7 small

spinules; preopercle and remainder of subopercle entire, the

margins naked and broadly rounded.

Two new species of Parapercis

Figure 1. Holotype of Parapercis flavolabiata, QM 1.37762, 84 mm SL, male.

Lateral line continuous, ascending smoothly from opercle

to below 2nd or 3rd soft dorsal-fin ray, then approximately

following contour of back; scales ctenoid, except for those on

nape, cheeks, breast, midline of belly, and some anteriorly on

opercle and posteriorly on base of pectoral fins, those on

middle of sides with about 36 cteni; scales on cheek extending

forward approximately to a line between middle of eye and

posterior tip of maxilla; no scales on dorsal, anal or pelvic

fins; 2 or 3 rows of small cycloid scales on base of pectoral-fin

rays; elongate ctenoid scales densely arranged on proximal

two-thirds of caudal fin.

Origin of dorsal fin at or just posterior to a vertical from tip

of opercular flap, the predorsal length 3.25 (3.1-3.35) in SL; 1st

dorsal-fin spine shortest, 1 1.35 (9.05-12.85) in HL; 2nd and 3rd

dorsal-fin spines progressively longer; 4th dorsal-fin spine

longest, 3.8 (3.3^.25) in HL; 5th dorsal-fin spine intermediate

in height between 2nd and 3rd spines; membrane from 5th spine

to 1st soft ray moderately notched, attached at about two-thirds

height of 5th dorsal-fin spine; longest dorsal-fin soft ray the

19th (19th-20th), 1.95 (1.95-2.25) in HL; origin of anal fin

below base of 5th to 6th soft dorsal-fin ray, preanal length 2.1

(1.95-2.1) in SL; anal-fin spine slender, closely attached to 1st

soft ray, 5.7 (4.95-7.45) in HL; longest soft anal-fin ray the

15th (15th or 16th), 2.35 (2.15-2.5) in HL; caudal fin emarginate,

in males 2nd and 3rd branched ray from upper margin produced

to form a short filament, and about 10th-13th rays slightly

produced to form a small rounded lobe, length of caudal fin

without filament 4.75 (3. 85^.7) in SL; pectoral fins rounded,

10th ray usually longest, 4.7 (4.55-5.05) in SL, shorter than

pelvic fins; origin of pelvic fins in advance of upper base of

pectoral fins and approximately on a vertical from upper corner

of operculum, prepelvic length 3.8 (3.5-3.85) in SL; pelvic-fin

spine closely attached to 1st soft ray, its termination fleshy and

difficult to accurately determine; 4th soft pelvic-fin ray longest,

reaching to base of 2nd soft anal-fin ray (origin of anal fin to

base of 3rd soft ray), 4.1 (3 .7^.25) in SL.

Colour in alcohol. Head, body and fins mostly uniformly pale

yellowish-brown. Some faint dark saddles dorsally above

lateral line persisting in larger specimens.

Colour fresh. Male holotype pale pinkish-white above on body,

with 6 broad inverted triangular red saddles, extending almost

to ventral midline on each side. Secondary narrow vertical red

bars between the 3rd and 4th, 4th and 5th and 5th and 6th

saddles, and on the caudal peduncle extending ventrally from

lateral line to a similar level. A small red blotch also in the same

series, just below pectoral fin base. Pigmentation of bars and

saddles darker blood-red at their lower apex. Lower sides, belly

and breast shading to pearly white. Head mostly flesh-pink.

Opercle below opercular spine and adjacent upper four-fifths of

pectoral-fin base pale yellow. A short diffuse oblique red and

yellow bar (red anteriorly, yellow posteriorly) from tip of

maxilla to about half distance to middle of eye. A distinct

purplish-red to violet line broadly but faintly margined in pale

orange-yellow from middle of anterior edge of orbit, curving

around snout to meet orbit on opposite side. Chin and anterior

half of upper and lower lips diffusely flushed with red. Posterior

half of upper lip bright yellow. Purple-pink reflective band

across anterior part of interorbital, followed posteriorly by a

small pale-yellow blotch adjacent to inner edge of each eye.

Transverse row of 3 yellow-edged red spots across occipital

region, lowermost on each side behind upper third of eye, the

upper central spot on dorsal midline. Four additional yellow-

edged red spots in a crescentic row on nape immediately behind

head, upper 2 directly behind inner margin of orbit, lowermost

on anterodorsal edge of post-temporal. Area from upper corner

of operculum to posterodorsal part of head diffusely washed in

yellow. Posteroventral margin of orbit with faint purple-pink

reflective line underlying the orbital membrane. Dorsal and

ventral margins of iris yellow, followed by a broad purplish-red

area extending to upper and lower edges of pupil respectively,

remainder of iris behind and in front of the pupil yellow. Distal

third of spinous dorsal fin membrane pale translucent yellow.

Jeffrey W. Johnson

Figure 2. Holotype of Parapercis flavolabiata, QM 1.37762, 84 mm SL, male. (A) Lateral view of head (B) Dorsal view of head.

Two new species of Parapercis

Table 1. Selected meristic and morphological values for type specimens of Parapercis flavolabiata and P. sexlorata (measurements as percentage

of standard length).

P. flavolabiata P. sexlorata

Holotype

QM 1.37762

Paratypes

(n=16)

Holotype

QM 1.33274

Paratypes

(n = 6)

Standard length (mm)

84.0

35.0-91.0

120.0

73.5-95.0

Dorsal-fin rays

V, 21

IV, 23

Anal-fin rays

I, 17

I, 17 - 1, 18

Pectoral-fin rays

16-18

Gill rakers

3 + 11

3-5 + 10-11

6 + 9

4-6 + 8-10

Lateral-line scales

51-55

53-57

Lower jaw teeth (outer row)

4 + 4

3-4 + 3-4

3 + 3

Vertebrae (abdominal -i- caudal)

9 + 20

9-10 + 20

10 + 22

9 + 23 (2)

Body depth

16.3

15.9-17.5

19.3

15.6-18.1

Body width

17.4

17.4-18.5

18.3

18.0-19.8

Head length

29.8

29.2-32.5

26.8

25.6-27.9

Snout length

8.2

7.8-8.5

5.7

5.8-6.3

Orbit diameter

8.6

8.4-9.6

7.9

8.4-9.5

Interorbital width

3.7

2.7-3.8

2.3

1.6-2.2

Preorbital depth

4.3

3.7-4.7

3.9

3.1-3.8

Upper jaw length

12.0

11.9-13.6

12.1

11.2-12.8

Predorsal length

30.8

29.7-32.2

30.8

29.0-31.2

Preanal length

47.9

47.6-51.6

45.2

46.0-48.7

Prepelvic length

26.3

26.1-28.4

24.7

24.1-26.0

Caudal-peduncle depth

9.0

9.0-9.5

10.0

9.3-10.1

Caudal-peduncle length

10.1

9.1-10.2

7.8

7.6-8.9

Dorsal-fin base

62.0

59.1-62.5

67.3

61.5-65.3

First dorsal-fin spine length

2.6

2.4-33

4.1

2.6-4.4

Second dorsal-fin spine length

5.4

3.9-4.9

5.2

3.7-5.7

Third dorsal-fin spine length

7.1

6.4-7.7

5.7

5.1-6.0

Fourth dorsal-fin spine length

7.9

7.6-9.2

5.9

5.7-7. 1

Fifth dorsal-fin spine length

5.7

5.2-1.5

Longest dorsal-fin ray

15.4

14.0-15.7

15.0

13.5-15.3

Anal-fin base

42.3

40.7-43.0

45.6

43.9-46.8

Anal-fin spine length

5.2

4.3-5.8

3.8

4.6-5.4

Longest anal-fin ray

12.7

12.5-13.6

12.9

13.1-13.4

Caudal-fin length

21.1

21.4-25.9

17.2

19.1-21.5

Pectoral-fin length

21.3

19.8-21.9

20.6

20.1-22.3

Pelvic-fin length

24.5

23.4-26.9

20.4

22.0-23.7

Table 2. Frequency of lateral-line scales and gill rakers in type specimens of Parapercis flavolabiata and P. sexlorata and in type and non-type

material of P. colemani (* denotes holotype).

Lateral-line scales

51 52 53 5 4 55 56 57 58

P. flavolabiata 1 5 5* 5 1 -

P. colemani - - - - 1 2 1 P

P. sexlorata - - 1 1 2 2* 1 -

Gill rakers

Upper

Lower

Total

P. flavolabiata

P. colemani

P. sexlorata

4>K

Jeffrey W. Johnson

Basal two-thirds of membrane between 1st and 2nd dorsal

spines blood-red, that between other spines mainly transparent,

but with numerous tiny white dots. Soft dorsal fin with numerous

tiny white dots, translucent yellow on basal and distal 3rd, but

with central 3rd transparent. Distinctive small dark-red spot in

transparent area just above midheight of fin following each ray.

Anal fin with basal half translucent whitish and distal half pale

translucent yellow. Caudal fin with a broad oblique blood-red

band covering about one-third of fin anterodorsally, but

extending to full length of fin ventrally, followed by a central

white area, and a broad pale-yellow margin distally. Narrow

bright-yellow edge to anterodorsal half of caudal fin. Centre of

fin with about 4 small darker red smudge-like spots interspersed

with dusky melanophores. Pectoral fins transparent. Pelvic fins

white, with a faint yellowish blush proximally. Tongue pinkish-

white. Peritoneum white.

Distribution. Demersal trawl grounds between Anzac Reefs,

18°47.1'S, 147°57.3'E and Lady Musgrave I., 23°54.9'S,

152°30.9'E, Great Barrier Reef, Qld, in depths from 45 to 79.2 m.

(fig. 4).

Etymology. Erom the latin flavo for yellow and labiata for

lipped, in reference to the distinctive yellow posterior half of

the upper lip.

Remarks. Parapercis flavolabiata is most similar in colouration

to P. multiplicata Randall, 1984, sharing with that species a series

of reddish saddles across the back and a row of dark-red spots

along the soft dorsal fin, however it may easily be distinguished

by the presence of only six saddles (rather than eight), dark

marking in spinous dorsal fin confined to first membrane (rather

than anterior three), presence of a bright-yellow upper lip,

purplish-red line across the snout, and series of yellow-edged red

spots on the postorbital region of the head (all absent), fewer

pectoral fin rays (16-18, usually 17 versus 14—16, usually 15),

lower lateral-line scale count (51-55 versus 56-58) and fewer

predorsal scales (8-9 versus 10-12). Parapercis colemani

Randall & Erancis, 1993 has similar colouration and meristics to

P. flavolabiata, but has five reddish saddles that only cover the

upper third of the body and a series of ten separate vertical yellow

bars below (versus six red saddles continuous to lower sides and

interspersed posteriorly with independent shorter red bars), lacks

the distinctive head markings of P. flavolabiata, and has a higher

lateral-line scale count (55-58 versus 51-55). Parapercis

punctulata Cuvier, 1829 and P. signata Randall, 1984 are also

somewhat similar in colouration, but have dark markings and

spots on anal fin (absent in P. flavolabiata), differ markedly in the

colouration of the head and spinous dorsal fin (membranes of the

latter with black blotch covering entire base), and have more

lateral-line scales (55-57 versus 51-55).

This species was taken during a wide-ranging study of

benthic inter-reef fishes and invertebrates across the entire

Australian Great Barrier Reef region. Most prior surveys of this

region have been more restricted geographically and have

concentrated on areas fished commercially for penaeid prawns

and scallops, spp, using variously adapted commercial

otter trawl gear and scallop dredges. This survey utilised small

but fairly conventional trawl gear, but employed a specialised

epibenthic sled with a diameter of 1.5 m and stretched mesh size

of 25 mm. Almost 1200 tows of 200 m were undertaken with

this apparatus. It was responsible for the capture of numerous

poorly known benthic fish species and accounted for all but

three of the specimens collected of Parapercis flavolabiata.

Parapercis sexlorata sp. nov.

Sixstrap Grubfish

Eigures 3-5; Tables 1-2.

Material examined. Holotype. QM 1. 33274, 120 mm, east of Cape

Moreton, Qld, 27°03.58'S, 153°31.57'E, 86 m, Qld Fisheries Service,

14 Mar 2001.

Paratypes: (N = 6) AMS 1.22873-004, 91 mm, SE of Cape Byron,

NSW, 28°43'S, 153°49'Eto 28°39'S, 153°50'E, 128-137 m, K. Graham

onYRNKapala, 1 Nov 1978; AMS 1.31484-006, 86 mm, off Brunswick

Heads, NSW, 28°27'S, 153°50'E, 119-137 m, K. Graham on FRV

Kapala, 16 Feb 1991; AMS 1.37977-002, 2: 73.5-93 mm, off Sandon

Bluffs, NSW, 29°40'S, 153°40.6'E, 107-109 m, K. Graham on FRV

Trader Horn, 8 Jul 1999; AMS 1.40445-004, 95 mm, off Sandon

Bluffs, NSW, 29°39'S, 153°4rE to 29°42'S, 153°40'E, 108-117 m, K.

Graham on FRV Trader Horn, 5 Jul 1999; QM 1. 37579, 84.5 mm,

ENE of North Solitary L, NSW, 29°54'S, 153°36'E, 102 m, K. Graham

on FRV Kapala, 2 Aug 1978.

Diagnosis. Dorsal-fin rays IV, 23; anal-fin rays I, 19; pectoral-

fin rays 19; lateral-line scales 53-57; gill rakers 4-6-1-8-10;

predorsal scales 10-11; scales on cheek ctenoid, in about 9

horizontal rows below middle of eye; 6 canine teeth in outer

row at front of lower jaw; vomer with a crescentic outer row of

6-7 robust conical teeth and several smaller teeth behind;

palatines with a single row of about 7 robust conical teeth;

angle of subopercle produced, with several feeble spinules; 9-

10 abdominal and 22-23 caudal vertebrae; head and body pink,

with 6 narrow oblique posteroventrally-directed black bars

across the back, 1st extending to lateral line, 2nd to middle of

sides, others to lower 3rd of body; a small black spot in upper

corner of caudal-fin base; scales above lateral line in area

between bars and spot with dusky margins; spinous dorsal fin

membrane dusky; upper lip yellowish.

Description. Dorsal-fin rays IV, 23; anal-fin rays I, 19; all

dorsal- and anal-fin rays branched, last to base; pectoral-fin

rays 19, upper ray unbranched, others including lowermost

branched; pelvic-fin rays I, 5; branched caudal-fin rays 15;

lateral-line scales 56 (53-57), plus 2-3 smaller scales on

caudal-fin base; scales above lateral line to origin of dorsal fin

41/2 ( 41 / 2 - 51 / 2 ), to base of anterior soft rays of dorsal fin 3/2 (3/2-

41 / 2 ); scales below lateral line in a oblique row to origin of anal

fin 14 (13-14); circumpeduncular scales 24; predorsal scales 11

(10-11), extending forward just posterior to a vertical from

preopercular margin; horizontal row of scales from preorbital

across cheek to edge of preopercle about 19; gill rakers on 1st

arch 6-1-9, total 15 (4-6-1-8-10=12-16); branchiostegal rays 6;

vertebrae 10-1-22 (2 paratypes with 9-1-23).

Body depth 5.15 (5.5-6.4) in SL, more robust in larger

specimens; body subcylindrical, greatest width 1.05 (0.80-1.0)

in body depth, depth equal to or greater than width only in

larger specimens; head length 3.75 (3. 6-3.9) in SL; snout

bluntly rounded, its length 4.7 (4.1-4.65) in HE; orbit diameter

Two new species of Parapercis

Figure 3. Holotype of Parapercis sexlorata, QM 1.33274, 120 mm SL, male.

% ^

Figure 4. Paratype of Parapercis sexlorata, QM 1.37579, 84.5 mm SL (Photo: K. Graham).

3.4 (2.75-3.1) in HL; eyes directed as much dorsally as

laterally, bony interorbital space narrow, 11.9 (11.65-15.7) in

HL; caudal-peduncle depth 2.7 (2. 6-3.0) in HL; caudal-

peduncle length 3.4 (3.0-3.65) in HL.

Mouth slightly oblique, jaws terminal, curved canine teeth

at front of each jaw slightly projecting, but concealed by lips

when mouth is closed; upper jaw extending to a vertical at

posterior margin of pupil, upper jaw length 2.2 (2.05-2.5) in

HL; upper jaw with 19-21 outer curved canines, first 10 (8-10)

larger than those following, of these either 1st or 9th usually

the largest, 11th and subsequent teeth gradually reducing in

size posteriorly, broad inner band of villiform teeth anteriorly,

narrowing gradually but still forming several rows at rear of

jaw; front of lower jaw on each side with 3 enlarged curved

canines in distinctly separate outer row, tooth nearest

symphysis smallest, 3rd in the series largest; broad inner band

of villiform teeth extending posteriorly from symphysis to

side of jaw 4-5 rows posterior to largest tooth in outer row,

next 2 teeth abruptly larger, then followed by single row of

15-16 smaller slightly curved conical teeth. Vomer with

Jeffrey W. Johnson

1H*

★

19"-

. 1 ■ *

12*

2tr

28*

- — -

' .1

" |i ^

V - \

....... ^

Figure 5. Distribution of Parapercis flavolabiata and P.

sexlorata based on specimens examined.

crescentic outer row of about 7 robust conical teeth, medial

teeth largest, several smaller adjacent teeth posteriorly;

palatines with row of about 7 robust conical teeth and several

closely spaced additional teeth near midsection of row. Tongue

spatulate with broadly rounded tip, covered with numerous

minute papillae.

Gill membranes united with broad free fold, not attached to

isthmus. Gill rakers short, the longest about one-sixth length of

longest gill filament on 1st gill arch. Anterior nostril moderately

large, situated in front of centre of eye, about half distance to tip

of snout, with membranous posterior flap, the latter folded

around near base to form partial tube. Posterior nostril about half

distance from anterior nostril to anterior margin of eye, dorso-

posterior to, and slightly wider than anterior nostril, its opening

simple and slightly raised, aperture slightly oval-shaped;

intemarial distance about twice width of posterior nostril.

Opercle with distinctly exposed, robust, pointed spine,

angle of subopercle expanded, with several feeble spinules;

preopercle and remainder of subopercle entire, the margins

naked and broadly rounded.

Lateral line continuous, ascending from opercle to below

2nd or 3rd dorsal spine, then approximately following contour

of back; scales ctenoid, except for some on shoulder region and

pectoral-fin rays and those on nape, breast and midline of belly,

scales on middle of sides with about 40-50 cteni; scales on

cheek extending forward approximately to a line between middle

of eye and midlength of maxilla; no scales on dorsal, anal or

pelvic fins; small elongate scales on pectoral-fin rays, some rows

covering proximal half of fin; elongate ctenoid scales densely

arranged on proximal three-fourths of caudal fin.

Origin of dorsal fin distinctly posterior to a vertical from tip

of opercular flap, the predorsal length 3.25 (3.2-3.45) in SL; 1st

dorsal-fin spine shortest, 6.55 (6.0-10.5) in HL; subsequent

spines progressively longer; 4th (3rd or 4th) dorsal-fin spine the

strongest, 4th always the longest, 4.5 (3.75^.55) in HL;

membrane from 4th spine to 1st soft ray moderately notched,

attached at about two-thirds height of 4th dorsal-fin spine;

longest dorsal-fin soft ray the 17th (17th-20th), 1.8 (1.75-1.9)

in HL; origin of anal fin below base of 4th or 5th soft dorsal-fin

ray, preanal length 2.2 (2.05-2.2) in SL; anal-fin spine slender,

closely attached to 1st soft ray, 7.15 (4.95-5.65) in HL; longest

soft anal-fin ray the 16th (16th or 17th), 2.05 (1.9-2.05) in HL;

caudal fin rounded, length of caudal fin 5.85 (4.65-5.25) in SL;

pectoral fins rounded to very bluntly pointed, 10th ray usually

longest, 4.85 (4.5-5. 0) in SL, slightly longer than pelvic fins

(shorter than pelvic fins in all smaller paratypes); origin of pelvic

fins slightly in advance of upper base of pectoral fins and on a

vertical just anterior to tip of operculum, prepelvic length 4.05

(3.85^.15) in SL; pelvic-fin spine closely attached to 1st soft

ray, its termination fleshy and difficult to accurately determine;

4th soft pelvic-fin ray longest, reaching midway between vent

and anal fin origin (from midway between vent and anal-fin

origin to base of 1st anal-fin soft ray), 4.9 (4.2^.55) in SL.

Colour in alcohol. Holotype, QM 1.33274 (fig. 3) pale yellowish

brown, with 6 narrow mostly oblique posteroventrally-directed

black bars across the back and upper sides, 1st extending from

immediately anterior to origin of spinous dorsal fin to lateral

line, 2nd from 2nd soft dorsal-fin ray to middle of sides, others

from 7th, 12th, 17th, and 22nd soft dorsal-fin rays to lower 3rd

of body. Small black spot in upper corner of caudal-fin base.

Scales above lateral line in area between each transverse bar

and between last bar and caudal spot with dusky margins,

producing a reticulate pattern. A narrow pale border around

each transverse bar. Lower sides, belly, breast and snout to

preopercle pale yellowish-brown. Upper part of opercle and

upper postorbital region to rear of interorbital dark-brown,

punctuated above margin of preopercle with a central pale area.

A narrow curved brown line extending ventrally behind eye

from postorbital area to just below lower margin of eye. Midline

of nape with a broad band of scales with dusky margins,

extending posteriorly from origin of scaled area to 1st transverse

black bar anterior to dorsal-fin origin. Spinous dorsal-fin

membrane diffusely dusky. Transverse black bars of upper

body produced slightly on to base of adjacent soft dorsal-fin

rays and membrane, soft dorsal fin otherwise uniformly pale.

Middle section of inner pelvic-fin rays dusky to black. Anal fin

uniformly pale. Caudal fin with 7 contrasting narrow dark

wavy vertical lines.

Two new species of Parapercis

Figure 6. Parapercis colemani, NMV A.251 12-003, 84 mm SL (Photo: NORFANZ Founding parties).

Colour fresh. Based on colour photograph of paratype, QM

1.37579 (fig. 4), pale-pink on head and body, with 6 transverse

black bars and a black spot on upper corner of caudal-fin base.

Each bar produced ventrally in dusky-yellow. Belly and breast

shading ventrally to pinkish-white. Scales above lateral line in

area between each transverse bar and between last bar and

caudal spot with distinct dusky margins, producing a reticulate

pattern. A narrow pink border, devoid of dusky-edged scales,

around each transverse bar. Much of pectoral-fin base,

postorbital and opercle diffusely dusky, infused with yellow.

Upper lip dark orange-yellow, lower lip white. Spinous dorsal

fin dusky grey. Soft dorsal-fin rays orange anteriorly, some

wavy oblique orange lines visible posteriorly, membrane

mostly translucent. Inner pelvic-fin rays dusky at midsection,

whitish at base, yellowish distally. Anal fin faintly orange.

Caudal fin mauve with about 7 narrow wavy vertical orange-

yellow lines.

Distribution. Scattered on demersal trawl grounds between Cape

Moreton, Qld, 27°03.58'S, 153°31.57'E and ENE of North

Solitary I., NSW, 29°54'S, 153°36'E, in depths from 86 to 137 m

(fig. 5).

Etymology. Erom the latin sex for 6 and lorata for strap, in

reference to the 6 distinctive narrow black strap-like bars across

the back of this species.

Remarks. Among currently recognised species, Parapercis

sexlorata is most similar to P. macrophthalma Pietschmann,

1911 and P. muronis Tanaka, 1918, sharing with those taxa

palatine teeth and a series of black bars across the back and

upper sides. However it may easily be distinguished by the

dorsal-fin ray count of IV, 23 versus V, 23-24, six versus eight

canine teeth in outer row of lower jaw, and six versus five

transverse bars. This species appears to be quite rare, with only

seven specimens captured in a region that has been surveyed

quite intensively by demersal trawl on various occasions since

the late 1970s.

New records of Parapercis colemani Randall & Erancis, 1993

Parapercis colemani Randall & Erancis, 1993 was described

from a single female collected in 1 .5 m depth in a sandy inshore

lagoon at Norfolk I. There have been no published records or

additional material reported since the original description.

Three specimens of an unidentified species of Parapercis were

collected in 2003 during the NOREANZ cruise, one from the

Lord Howe Rise and two from the Northern Norfolk Ridge,

Tasman Sea region, in depths of 89-113 m. These are identified

here as conspecific with P. colemani. The new records indicate

that the species usually inhabits much deeper habitats over a

considerably wider geographic area than was previously

inferred from the type locality. It is difficult to draw conclusions

on the abundance of the species from NOREANZ results, as

few trawls were undertaken in the depth range that specimens

were caught (over 90% of trawls and sleds were in depths

greater than 200 m).

The new material has identical dorsal-, anal- and pectoral-

fin ray counts, and similar dental formulae to the holotype.

Several other counts and features at slight variance to the

original description are reported below. The lateral-line scale

and gill raker counts are slightly expanded (Table 2). Pelvic

fins of the holotype are described as “not reaching anus” and

their length 1.7 in HL. However in the new material they reach

to between the base of the first to third soft rays of the anal fin,

and are 1.2-1. 3 in HL. The holotype is damaged, with the

right pelvic fin cut off at the base and missing, and the left one

missing the distal portion of the third to fifth rays. Erom

proportional measurements given for the type and a current

measurement of the damaged left fin, the pelvic fin appears to

have been measured to the tip of the longest intact (second)

ray, rather than to the tip of the longer fourth ray. It is therefore

difficult to accurately compare the pelvic fin length and reach

of the type versus new non-type material. However, in the new

material the second pelvic-fin ray reaches between two scales

prior to the anus and the midpoint of the anus, whereas in the

Jeffrey W. Johnson

type it reaches to about four scales prior to the anus, indicating

that the type had slightly shorter pelvic fins overall. The female

holotype has the second to fourth caudal-fin rays only slightly

prolonged, the fin length to tip of upper lobe 1.4 in HL.

Although the new specimens are all larger than the type, those

with intact fins have these rays distinctly prolonged, the fin

length 1.1-1. 2 in HL.

Colouration of the type, based on the original description

and figures, and direct comparison of the preserved specimen,

is almost identical with the new material (fig. 6), with the

following exception. The type has a longitudinal row of eight

small vague dark spots on the upper side (dusky orange-red in

fresh specimen). Although these still persist in the preserved

type specimen, they are lacking in the other material. In all

other details, including the formation of the five dark saddles

above, ten yellow bars below, faint midlateral white stripe,

reddish blotch below eye, series of small transverse dark

markings across the top of the head and postorbital, and fin

colouration, they are entirely consistent.

Material examined. Parapercis colemani AMS 1.33434-001,

holotype, female, 82 mm, Norfolk L, Emily Bay, eastern side, 29°04'S,

167°57'E, spear, 1.5 m, J.E. Randall, 17 Eeb 1991; CSIRO H.6028-02,

female, 107 mm, Tasman Sea, Lord Howe Rise, SE of Lord Howe L,

31°49'S, 159°20'E, benthic sled, 89 m, R Last, A. Graham and D.

Gledhill on RV Tangaroa, 22 May 2003; NMNZ P.11712, 87 mm,

Norfolk I.,R. Bell, 4 Aug 1913; NMVA.25112-003, males?, 2: 83-84

mm, Tasman Sea, North Norfolk Ridge, 28°54.43'S, 167°40.54'E,

beam trawl, 111-113 m, D. Bray on RV Tangaroa, 15 May 2003.

Acknowledgements

Many thanks to D. Gledhill, CSIRO Marine Research and C.

Bartlett, Queensland Museum for provision of fresh specimens

and photographs of P.flavolabiata. K. Graham, NSW Fisheries

collected most of the type material of P. sexlorata and gave

permission for reproduction of his colour photograph of this

species. R Bartsch, Humboldt University, Berlin and H. Endo,

Kochi University, Japan kindly made available on loan the

holotype of P. macrophthalma and specimens of P. muronis.

respectively. M. McGrouther, Australian Museum, A. Graham,

CSIRO Marine Research, D. Bray, Museum Victoria and A.

Stewart, Museum of New Zealand assisted with loan material

and collection data.

References

Bleeker, P. 1863. Onzieme notice sur la faune ichthyologique de Pile

deTernate. Nederlands Tijdschrift Dierkunde 1: 228-238.

Cantwell, G.E. 1964. A revision of the genus Parapercis, family

Mugiloididae. Pacific Science 28(3): 239-280.

Cuvier, G. 1829. Histoire naturelle des poissons. Tome troisieme. Suite

du Livre troisieme. Des percoides a dorsale unique a sept rayons

branchiaux et a dents en velours on en cardes. Histoire Naturelle

des Poissons 3: i-xxviii + 2 pp. 500 pp.

Leviton, A.E., Gibbs, R.H., Jr., Heal, E. and Dawson, C.E.. 1985.

Standards in herpetology and ichthyology: part 1. Standard

symbolic codes for institutional resource collections in herpetology

and ichthyology. Copeia 1985: 802-832.

Retschmann, V. 1911. Ueber Neopercis macrophthalma n. sp. und

Heterognathodon doederleini, Ishikawa, zwei fische aus Eormosa.

Annalen des K. K. Naturhistorischen Hofimuseums Wein 25: 431^35.

Randall, J.E. 1984. Two new Indo-Pacific mugiloidid fishes of the genus

Parapercis. Freshwater and Marine Aquarium 7(12): 41^9.

Randall, J.E. 2003. Review of the sandperches of the Parapercis

cylindrica complex (Perciformes: Pinguipedidae), with description

of two new species from the western Pacific. Bishop Museum

Occasional Papers 72: 1-19.

Randall, J.E. and Erancis, M.P 1993. Parapercis colemani, a new

pinguipedid fish from Norfolk L, south-western Pacific Ocean.

New Zealand Journal of Marine and Freshwater Research 27:

209-214.

Randall, J.E. and McCosker, J.E. 2002. Parapercis lata, a new species

of sandperch (Perciformes: Pinguipedidae) from the central Pacific.

Proceedings of the California Academy of Sciences 53(8): 87-93.

Randall, J.E. and Yamakawa, T. 2006. Parapercis phenax from Japan

and P. banoni from the southeast Atlantic, new species of

pinguipedid fishes previously identified as P. roseoviridis.

Zoological Studies 45(1): 1-10.

Tanaka, S. 1918. Twelve new species of Japanese fishes. Dobutsugaku

Zasshi [Zoological Magazine, Tokyo] SQ{?)56y. 223-221.

Memoirs of Museum Victoria 63(1): 57-64 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Simipercis trispinosa^ a new genus and species of sandperch (Perciformes:

Pinguipedidae) from eastern Australia

Jeffrey W. Johnson^ and John E. Randaff^

^Manager Ichthyology, Queensland Museum, PO Box 3300, South Brisbane, Q 4101, Australia (jeff.johnson@qm.qld.gov.

au) (author for correspondence)

^Senior Ichthyologist, Bishop Museum, 1525 Bernice St., Honolulu, Hawai’i, 96817-2704, USA (jackr@hawaii.rr.com)

Abstract Johnson, J.W. and Randall, J.E. 2006. Simipercis trispinosa, a new genus and species of sandperch (Perciformes:

Pinguipedidae) from eastern Australia. Memoirs of Museum Victoria 63(1): 57-64.

Simipercis trispinosa is described as a new genus and species of pinguipedid fish from 93 specimens collected mostly

by demersal trawl from the continental shelf of eastern Australia between Swain Reefs, Queensland and Broken Bay, New

South Wales in depths from 51 to 170 m. The new genus is most closely related to Parapercis Bleeker, 1863, but is unique

among pinguipedid fishes in having a combination of vomer and palatines edentate, dorsal spines 3, no distinct exposed,

pointed opercular spine, preopercle and subopercle entire, ten abdominal and 20-21 caudal vertebrae, low naked fleshy

nuchal crest, interorbital and suborbital with large ctenoid scales extending forward to near anterior margin of eye, and

head and body moderately compressed.

Keywords Pinguipedidae, Simipercis, Parapercis, new genus, new species, Queensland, New South Wales, Australia

Introduction

Rosa and Rosa, 1998 provided a diagnosis and definition of the

Pinguipedidae, describing three primary osteological

synapomorphies of the family: hyomandibular with oblique

crest (Gosline, 1968), intercalar with sharp posteriorly directed

process, and sphenotic with elongate, anteriorly directed

process. Imamura and Matsuura, 2003 redefined the family

using 14 derived characteristics, only one of which was used

by Rosa and Rosa (hyomandibular with oblique crest). They

also confirmed that Cheimarrichthys von Haast, 1874 should

be excluded from the Pinguipedidae and placed in its own

family, Cheirmarrichthyidae.

The Pinguipedidae currently contains five genera: the

monotypic Kochichthys Kamohara, 1961 from Japan and

Taiwan, Pinguipes Cuvier, 1829 with one species from each of

the south-eastern Pacific and south-western Atlantic, the

monotypic Prolatilus Gill, 1865 from the south-eastern Pacific,

Pseudopercis Miranda-Ribeiro, 1903 with two species from

the south-eastern Atlantic, and Parapercis Bleeker, 1863 with

about 54 recognised species, including 51 from the Indo-west

Pacific and single representatives in the south-eastern Pacific,

south-eastern Atlantic and north-eastern Atlantic (Rosa and

Rosa, 1998; Randall and McCosker, 2002; Imamura and

Matsuura, 2003; Randall, 2003; Randall and Yamakawa, 2006;

this study).

The first author recently examined all known pinguipedid

specimens throughout Australian ichthyological collections to

validate identifications for a project on demersal continental

shelf and slope fishes. During these studies two nominal species

of Parapercis, P. naevosa Serventy, 1937 and P. stricticeps De

Vis, 1884, regarded by Cantwell, 1964 as junior synonyms of P.

allporti (Gunther, 1876) and P. xanthozona (Bleeker, 1849)

respectively, were discovered to be valid species. In addition, a

number of undescribed species collected by demersal trawl

were also identified. Most of these currently await description,

but the most distinctive of the latter is described here.

Methods and materials

Institutional acronyms for types and comparative material are

those of Leviton et al, 1985. Lengths of specimens are given as

standard length (SL), being the distance from the front of the

upper lip to the base of the caudal fin (posterior end of the

hypural plate). The abbreviation HL is used for head length.

Lateral-line scale counts are to the base of the caudal fin, and do

not include several additional pored scales on the fin. Gill-raker

counts include all rudiments. Vertebral counts and descriptions

of osteological features were made from radiographs and skeletal

preparations. Methods for counts and measurements otherwise

follow Randall, 2003. Where different, values for paratypes

follow those of the holotype in parentheses. Specimens were

Jeffrey W. Johnson and John E. Randall

collected by trawl, except where indicated otherwise. Meristic

and morphometric details for the new species are presented in

Tables 1-2. Material examined for detailed comparison of

opercular osteology is listed. However an additional 30 unlisted

species of Parapercis were examined externally for character

states of the opercular bones.

Comparative material examined

Parapercis binivirgata QM 1.37220, 146 mm, SE of Cape

Moreton, 27°03'S, 153°31'E, 86 m, 14 Mar 2001 (skeleton).

Parapercis nebulosa QM 1.36874, 113 mm, Hervey Bay,

25°03.5'S, 152°46.6'E, 16.6 m, 1 Jul 2002; QM 1.36875, 152 mm,

same data (skeletons).

Parapercis sp.3 (undescribed) QM 1.36876, 118 mm, east of South

Stradbroke L, 27°48.8'S, 153°49.7'E, 161-165 m, 24 Jul 2002 (skeleton).

Simipercis gen. nov.

Type species. Simipercis trispinosa sp. nov.

Diagnosis. The type species of Simipercis differs from all other

genera of the family Pinguipedidae in having 3 dorsal spines, a

low naked fleshy nuchal crest (fig. Id), and dorsal angle of

opercular bone broad, not forming a distinct exposed, pointed

spine (figs 2a-d), and a combination of the following features:

vomer and palatines edentate, subopercle and preopercle entire,

10 abdominal and 20-21 caudal vertebrae, scales including

those on head and pelvic region large and ctenoid, interorbital

with a single row of large scales, extending to, or just beyond

anterior margin of eye, suborbital scales reaching a vertical

between anterior margin of pupil and anterior margin of eye,

and head and body moderately compressed (body width 1.3-

1.55 in depth in adults).

Remarks The new genus is most closely related to Parapercis

Bleeker, 1863, with similar vertebral counts and dental

formulae, but is unique among pinguipedid fishes in having

three dorsal spines {Kochichthys 2, Parapercis 4-6, Pinguipes

6-7, Prolatilus 4, Pseudopercis 4-7), a low naked fleshy nuchal

crest (no fleshy crest known in other genera), and dorsoposterior

angle of opercular bone not forming an exposed, distinctly

pointed spine (figs 2a-d). The dorsoposterior angle of the

opercular bone is relatively broad and narrowly truncate

posteriorly, with a minute shallow notch. The posteroventral

edge of the notch forms a feeble, bluntly triangular point which

is hidden by the skin in undamaged specimens. A horizontal

ridge on the inner surface of the opercular bone is visible

externally under light, and is situated well below the dorsal

margin of the opercle. The ridge terminates slightly before the

posterior tip of opercular bone. In other genera there is at least

one exposed, robust, distinctly pointed opercular spine, and in

Parapercis, the horizontal ridge on the inner face of the

opercular bone is situated very close to (rather than well below)

its dorsal margin, and extends to the tip, reinforcing the

opercular spine (figs 3a-f).

Simipercis also differs from other pinguipedid genera by the

combination of: vomer and palatines edentate (only Prolatilus

lacks vomerine teeth, only Prolatilus and some Parapercis lack

palatine teeth); margin of subopercle entire (usually at least some

small spines or spinules in other genera); 10 abdominal and 20-21

caudal vertebrae {Parapercis 9-10+18-22, Kochichthys 10+19,

Pinguipes 1 5-16+20-2 1 , Prolatilus 1 5-1 6+20-2 1 , Pseudopercis

16-18+20-22); large interorbital scales extending to, or just

beyond, anterior margin of eye (only Kochichthys, Prolatilus and

Parapercis haackei have interorbital scales); cheek and suborbital

with large ctenoid scales, extending to a vertical between anterior

margin of pupil and anterior margin of eye (variously squamate

or naked, usually with small cycloid scales on cheek, suborbital

with small scales extending to below middle of eye or more

posteriorly in other genera); and head and body moderately

compressed (generally cylindrical or subcylindrical anteriorly in

other genera). A detailed examination of osteological and

myological features was not carried out on Simipercis; however

it agrees well with the familial diagnosis and description of Rosa

and Rosa, 1998 and with additional features of the family defined

by Imamura and Matsuura, 2003 that were examined (eg. unique

crest on the hyomandibular present, posterior margin of opercular

bone moderately concave, six branchiostegal rays, lowermost

ray of pectoral fin branched, 15 branched caudal-fin rays).

Etymology. From the Latin similis, meaning similar or like, in

reference to the pinguipedid genus Percis Bloch and Schneider,

1801 (the latter being preoccupied by the agonid genus Percis

Scopoli, 1777).

Simipercis trispinosa sp. nov.

Threespine Grubfish

Figures la-e, 2a-d, 4; Tables 1-2.

Material examined. Holotype. QM 1. 32697, 114 mm, east of Noosa, Qld,

26°24'S, 153°39'E, 104 m, Queensland Fisheries Service, 22 Jun 2001.

Paratypes: (N = 92) AMS 1.31473-002, 130 mm, off lluka, NSW,

29°24'S, 153°35'E, 67-77 m, K. Graham on FRV Kapala, 6 May 1990;

AMS 1.32121-004, 120 mm, off Newcastle, NSW, 32°53'S, 152°01'E,

73-79 m, K. Graham on FRV Kapala, 1 Jun 1990; AMS 1.32209-002, 2:

66.5-116 mm, off Newcastle, NSW, 32°54'S, 151°57'E, 6A-70 m, K.

Graham on FRV Kapala, 23 May 1990; AMS 1.32217-001, 5: 105.5-

135 mm, off Newcastle, NSW, 32°53'S, 152°02'E, 73-75 m, K. Graham

on FRV Kapala, 29 Aug 1991; AMS 1.33510-004, 97 mm, E of Clarence

R., NSW, 29°26'S, 153°34'E, 6^68 m, K. Graham on FRV Kapala, 2

Apr 1992; AMS 1.37355-011, 6: 39-109 mm, E of Swain Reefs, Qld,

22°23.49'S, 153°04.48'E, 170 m, trap, J. Lowry and K. Dempsey on FV

Seadar Bay, 8 Sep 1995; AMS 1.37572-001, 105 mm, south-east of

Evans Head, NSW, 29°13'S, 153°31'E, 51-53 m, K. Graham on FRV

Kapala, 15 Jun 1995; AMS 1.37587-008, 100 mm, E of Swain Reefs,

Qld, 22°23.49'S, 153°04.48'E, 138 m, J. Lowry and K. Dempsey on FV

Seadar Bay, 8 Sep 1995; AMS 1.37600-030, 2: 70-118 mm, E of Swain

Reefs, Qld, 22°28.34'S, 152°59.45'E, 139 m, trap, J. Lowry and K.

Dempsey on FV Seadar Bay, 8 Sep 1995; AMS 1.38271-001, 2: 58-75

mm, off Newcastle, NSW, 32°54'S, 151°58'E, 67-72 m, K. Graham on

FRV Kapala, 6 Apr 1995; AMS 1.40462-001, 2: 87.5-121 mm, south-E

of Sandon Bluffs, NSW, 29°40'S, 153°28'E, 55-59 m, K. Graham on FV

Trader Horn, 6 Sep 1999; AMS 1.43651-002, 76 mm, E of Manly, NSW,

33°36.5'S, 151°29.3'Eto 33°40.5'S, 151°26.8'E, 69-80 m, K. Graham on

FVKirrawana, 10 Aug 2005; BMNH 2005.6.2.1, 114mm,Eof Coolum,

Qld, 26°30.6'S, 153°35.2'E, 102.3 m, Queensland Fisheries Service, 21

Jun 2001; BPBM 37231, 4: 118-125 mm, off Newcastle, NSW,

32°53.5'E, 151°59.5'E, 6^75 m, K. Graham on FRV Kapala, 6 Mar

1991; CAS 222272, 116 mm, off Newcastle, NSW, 32°53.5'E,

151°59.5'E, 67.7-71.3 m, K. Graham, 6 Mar 1991; CSIRO H.6247-01,

112 mm, off southern end of Swain Reefs, Qld, 22°26.1'S, 152°41.1'E,

Simipercis trispinosa, a new genus and species of Sandperch (Perciformes: Pinguipedidae) from eastern Australia

Table 1. Selected meristic and morphological values for type specimens of Simipercis trispinosa (measurements as percentage of standard length).

Holotype

QM 1.32697

Paratypes (n = 92)

(range)

Standard length (mm)

114

39.5 - 135

Dorsal-fin rays

III, 24

III, 24-25

Anal-fin rays

I, 18

I, 17-19

Pectoral-fin rays

19-21

Gill rakers

4+ 11 = 15

3-6 + 9-13 = 12-18

Lateral-line scales

46-54

Upper jaw teeth (outer row)

27 + 27

24-29 + 24-29

Lower jaw teeth (outer row)

4 + 4

4-5 + 4-5

Vertebrae (abdominal -i- caudal)

10 + 21

10 + 20 (2)-21 (10)

Body depth

20.7

19.0 - 24.0

Body width

15.3

14.4 - 17.2

Head length

25.9

24.9 - 27.8

Snout length

5.0

4.0 - 6.3

Orbit diameter

9.2

7.4 - 10.2

Interorbital width

2.7

2.2 -2.9

Preorbital depth

3.8

2.8 -4.0

Upper jaw length

10.4

9.8-11.1

Predorsal length

25.0

24.3 - 26.2

Preanal length

43.0

41.3-46.4

Prepelvic length

24.6

22.8 - 25.4

Caudal-peduncle depth

9.1

8.5 - 9.7

Caudal-peduncle length

10.0

8.2-11.1

Dorsal-fin base

64.6

64.3-70.0

First dorsal-fin spine length

4.2

3.3 -4.9

Second dorsal-fin spine length

5.4

5.0 -6.2

Third dorsal-fin spine length

6.9

6.1 -8.3

Longest dorsal-fin ray

21.3

17.5 - 23.6

Anal-fin base

48.7

46.0-49.8

Anal-fin spine length

7.3

6.0 -8.8

Longest anal-fin ray

14.4

11.9-15.1

Caudal-fin length

21.2

19.2 - 25.4

Pectoral-fin length

23.2

20.8-23.9

Pelvic-fin length

22.9

18.4-26.2

Table 2. Frequency of lateral-line scales and gill rakers in type specimens of Simipercis trispinosa.

Lateral-line scales

Gill rakers

Upper

Lower

Total

4 5

17 18

50 28

1 1

Jeffrey W. Johnson and John E. Randall

112 m, RV Gwendoline May, 23 Apr 2004; NSMT P.70863, 105.5 mm,

E of Coolum, Qld, 26°30.6'S, 153°35.2'E, 102.3 m, Queensland Eisheries

Service, 21 Jun 2001; QM 1.22042, 4: 82-116 mm, E of Swain Reefs,

Qld, 22°06'S, 153°02'E, 150 m, Queensland Eisheries Service, 28 Aug

1983; QM 1.33181, 96 mm, E of Coolum, Qld, 26°32'S, 153°39'E, 123

m, Queensland Eisheries Service, 8 Aug 2001; QM 1.33182, 98.5 mm, E

of Noosa, Qld, 26°28'S, 153°39'E, 112.3 m, Queensland Eisheries

Service, 8 May 2001; QM 1.33183, 25: 59.5-123 mm, E of Coolum,

Qld, 26°30.6'S, 153°35.2'E, 102.3 m, Queensland Eisheries Service, 21

Jun 2001; QM 1.33333, 3: 80-96.5 mm, E of Point Cartwright, Qld,

26°39'S, 153°35'E, 112 m, Queensland Eisheries Service, 14 Sep 2001;

QM 1.33994, 4: 6^82 mm, E of Noosa, Qld, 26°20'S, 153°46'E, 110 m,

Queensland Eisheries Service, 19 Jul 2002; QM 1.33995, 16: 65-118

mm, E of Noosa, Qld, 26°25'S, 153°40'E, 119 m, Queensland Eisheries

Service, 19 Jul 2002; QM 1.34142, 2: 87.5-115 mm, E of Noosa, Qld,

26°24'S, 153°41'E, 98 m, Queensland Eisheries Service, 17 May 2001;

QM 1.36879, 2: 113-118 mm, E of Noosa, 26°22.2'S, 153°42.4'E, 115-

119 m, Queensland Eisheries Service, 19 Jul 2002; USNM 383404 128

mm, off Newcastle, NSW, 32°53.5'E, 151°59.5'E, 67.7-71.3 m, K.

Graham, 6 Mar. 1991.

Other material: (spirit specimens) AMS E.2963, 78 mm, 21 km

north-east of North Reef, Qld, 23°07'S, 152°05'E, 128 m, EIS

Endeavour, 1909; AMS 1.32199-003, 56 mm, off Angourie, NSW,

29°28'S, 153°33'E, 64-66 m, K. Graham on ERV Kapala, 20 Nov 1990;

AMS 1.33577-001, 107 mm, off Clarence R., NSW, 29°29'S, 153°33'E,

66-70 m, K. Graham on ERV Kapala, 19 Mar 1992; AMS 1.37978-003,

59 mm, off Sandon Bluffs, NSW, 29°44'S, 153°26'E, 55-61 m, K.

Graham on EV Trader Horn, 1 Jul 1999; AMS 1.39897-003, 112 mm,

off Sandon Bluffs, NSW, 29°44'S, 153°26'E, 55-62 m, K. Graham on

EV Trader Horn, 8 Jul 1999; QM 1.36878, 6: 63-118 mm, E of Peregian

Beach, 26°25.9'S, 153°45.2'E, 132-134 m, Queensland Eisheries

Service, 18 Jul 2002; QM 1.37224, 12: 7^121 mm, E of Coolum, Qld,

26°30.6'S, 153°35.2'E, 102 m, Queensland Eisheries Service, 21 Jun

2001. (skeletal specimens) QM 1.36870, 79 mm, QM 1.36871, 124 mm,

QM 1.36872, 113 mm, QM 1.36873, 100 mm, all E of Coolum, Qld,

26°32.9'S, 153°36.3'E, 119-120 m, Queensland Eisheries Service, 20

Jul 2002; QM 1.37223, 109 mm, E of Coolum, Qld, 26°30.6'S,

153°35.2'E, 102 m, Queensland Eisheries Service, 21 Jun 2001.

Diagnosis. See generic diagnosis.

Eigure 1. Type specimens of Simipercis trispinosa. (A) Holotype, QM 1.32697, 114 mm SL, male (B) Paratype, BPBM 37231, 125 mm SL, male

(C) Paratype, AMS 1.32217-001, 115 mm SL, male (photo: K. Graham) (D) Paratype, CSIRO H. 6247-01, 112 mm SL, male, dorsal view of nape,

showing fleshy nuchal crest (photo: D. Gledhill) (E) Paratype, AMS 1.43651-002, 76 mm SL, female (photo: S. Humphries).

Simipercis trispinosa, a new genus and species of Sandperch (Perciformes: Pinguipedidae) from eastern Australia

Description. Dorsal-fin rays III, 24 (3 with III, 25 rays); anal-fin

rays I, 18 (3 with I, 17 and 1 with I, 19); all dorsal- and anal-fin

rays branched, last to base; pectoral-fin rays 20 (19-21, 30 with

19, 5 with 21), upper ray unbranched, others including lowermost

branched; pelvic-fin rays I, 5; branched caudal-fin rays 15;

lateral-line scales 50 (46-54), plus 3^ smaller scales on caudal-

fin base; scales above lateral line to origin of dorsal fin 3, to base

of anterior soft rays of dorsal fin 21/2; scales below lateral line in

an oblique row to origin of anal fin 10(10-11); circumpeduncular

scales 19 (19-20); predorsal scales about 9 (8-10) to posterior

margin of eye, separated along dorsal midline by a low fleshy

nuchal crest and not in well-defined rows of equally-sized scales,

continuing in a single row of a further 8 (8-9) large scales from

posterior margin of eye through interorbital space to anterior

margin of eye, or just forward of this point; origin of nuchal

crest about 3 scales behind posterior margin of eye, termination

about 21/2 scales anterior to dorsal-fin origin; horizontal row of

scales from preorbital across cheek to edge of preopercle 14; gill

rakers on 1st arch 4-1-11, total 15 (3-6-1-9-13=12-18);

branchiostegal rays 6; vertebrae 10-1-21 (20-21).

Figure 2. Operculaof Simipercis trispinosa, showing poorly developed

points at dorsoposterior angle, and low position of inner horizontally-

aligned opercular ridge. (A) QM 1.36872, 113 mm SL, outer surface

(B) QM 1.36872, 113 mm SL, inner surface (C) QM 1.36871, 124 mm

SL, outer surface (D) QM 1.36871, 124 mm SL, inner surface.

Figure 3. Opercula of selected species of Parapercis, showing strong,

well-developed spine at dorsal angle and high position of inner

horizontally-aligned opercular ridge. (A) P. binivirgata, QM 1.37221,

146 mm SL, outer surface (B) P. binivirgata, QM 1.37221, 146 mm

SL, inner surface (C) P. nebulosa, QM 1.36875, 152 mm SL, outer

surface (D) P. nebulosa, QM 1.36875, 152 mm SL, outer surface (E)

P. “sp. 3”, QM I. 36876, 118 mm SL, outer surface (F) P. “sp. 3”, QM

I. 36876, 118 mm SL, inner surface.

Jeffrey W. Johnson and John E. Randall

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Figure 4. Distribution of Simipercis trispinosa based on specimens

examined.

Body depth 4.85 (4.15-5.25) in SL; body moderately

compressed, greatest width 1.35 (1.15-1.55) in body depth,

juveniles subcylindrical, but specimens becoming increasingly

compressed with age (1.3-1.55 in specimens greater than 90

mm SL); head length 3.85 (3. 6-4.0) in SL, proportionately

longest in adults; snout blunt, its length 5.2 (3.9-6.4) in HL;

orbit diameter 2.8 (2. 6-3. 5) in HL; eyes directed more laterally

than dorsally, bony interorbital space narrow, 9.5 (8.6-12.0) in

HL; caudal-peduncle depth 2.85 (2.65-3.15) in HL; caudal-

peduncle length 2.6 (2.35-3.2) in HL.

Mouth slightly oblique, terminal, with curved canine teeth

at front of lower jaw slightly projecting and visible when mouth

is closed; upper jaw extending to a vertical between anterior

margin and middle of pupil, upper jaw length 2.5 (2.35-2.6) in

HL; upper jaw with 27 (24-29) outer curved canines, closely

and evenly spaced, 2nd or 3rd from symphysis largest, but not

distinctly larger than those following, gradually reducing in

size posteriorly, broad inner band of villiform teeth anteriorly,

narrowing gradually to form single row at rear of jaw; front of

lower jaw on each side with 4 (4-5, usually 5) enlarged curved

canines in distinctly separate outer row, tooth nearest

symphysis smallest, the others gradually increasing in size

laterally, 4th usually largest (occasionally 5th), broad inner

band of villiform teeth extending posteriorly from symphysis

to side of jaw 12-18 rows back, next 5-7 teeth enlarged and in

single row (of these, 4-5 posterior-most teeth largest and

strongly recurved), then followed by single row of 12-17

smaller slightly curved conical teeth. Total number of teeth in

each jaw generally slightly more in adults than in juveniles.

Vomer and palatines edentate. Tongue spatulate with broadly

rounded tip, its surface covered with tiny papillae.

Gill membranes united with broad free fold, not attached

to isthmus. Gill rakers short, the longest about one -third length

of longest gill filament on 1st gill arch. Anterior nostril small,

inconspicuous, situated in front of centre of eye about one-

third distance to tip of snout, with membranous posterior flap,

the latter folded around near base to form partial tube, flap

usually lying flat against snout in preserved specimens.

Posterior nostril close to anterior margin of eye, dorsoposterior

to, and about 3 times width of anterior nostril, its opening

simple, anterior edge thin and slightly expanded, making

aperture slightly oval-shaped; internarial distance about twice

width of posterior nostril.

Opercle with no exposed pointed spine; dorsoposterior

angle of opercular bone narrowly truncate with minute shallow

notch, posteroventral corner of notch broadly and bluntly

triangular, but hidden by skin and scales or barely exposed;

subopercle and preopercle entire, the margins broadly rounded

and slightly overlapped by large ctenoid scales.

Lateral line continuous, ascending fairly abruptly from

opercle to below origin of soft dorsal fin, then approximately

following contour of back; all scales ctenoid, except for those

on pelvic fins and distally on pectoral fins, those on middle of

sides with about 50 cteni; scales on cheek extending forward

to a vertical between anterior margin of eye and anterior

margin of pupil; no scales on dorsal and anal fins; small

elongate cycloid scales on basal 3rd of pelvic fins; small scales

on basal 4th of pectoral fins, ctenoid proximally and cycloid

Simipercis trispinosa, a new genus and species of Sandperch (Perciformes: Pinguipedidae) from eastern Australia

distally; proportionately larger ctenoid scales densely arranged

on basal two-thirds of caudal fin.

Origin of dorsal fin at a vertical just anterior to tip of

opercular flap, the predorsal length 4.0 (3. 8^.1) in SL; 1st

dorsal-fin spine shortest, 6.15 (5. 1-7.6) in HL; 3rd dorsal-fin

spine longest, 3.75 (3. 1^.25) in HL; membrane from 3rd

spine to 1st soft ray moderately notched, attached at about

four-fifths height of 3rd dorsal-fin spine; longest dorsal-fin soft

ray the 19th (18th-20th), 1.2 (1.2-1.45) in HL; origin of anal

fin below base of 7th to 8th soft dorsal-fin ray, preanal length

2.35 (2.15-2.4) in SL; anal-fin spine slender, closely attached

to 1st soft ray, 3.55 (3.05-3.7) in HL; longest soft anal-fin ray

the 15th (15th or 16th), 1.8 (1. 7-2.1) in HL; caudal fin truncate

to slightly rounded, but in males 3rd branched ray from upper

margin filamentous, and about 10th-13th rays slightly produced

to form a small lobe, length of caudal fin without filament 4.7

(3.95-5.2) in SL; pectoral fins rounded, 12th ray usually

longest, 4.3 (4.2^.8) in SL, longer than pelvic fins in adults,

shorter than pelvic fins in juveniles; origin of pelvic fins in

advance of upper base of pectoral fins and slightly anterior to a

vertical from dorsal-fin origin, prepelvic length 4.05 (3. 95^.4)

in SL; pelvic-fin spine closely attached to 1st soft ray, its

termination very fleshy and difficult to accurately determine;

4th soft pelvic-fin ray longest, reaching almost to base of 2nd

soft anal-fin ray (origin of anal fin to base of 3rd soft ray), 4.35

(3.8-5.45) in SL.

Colour in alcohol. Head, body and fins mostly uniformly pale

yellowish brown. Margins of scales on upper part of body

faintly greyish. Narrow naked area of nuchal crest silvery-white

(gradually fading in preservative), interspersed variably with

dusky melanophores.

Colour fresh. Male holotype pale rose-pink, with numerous

vague narrow oblique posteroventrally-directed yellow bands,

shading to pearly-white on lower part of opercle, pectoral fin

base, breast and belly. Oblique pale yellow bar from lower edge

of eye across cheek. Dorsal margin of eye yellow, remainder of

iris silvery-white, variably washed with pale red above and

posteriorly. Nuchal crest on predorsal midline silvery -white,

with numerous scattered dusky melanophores (nuchal crest

pale golden-pearl, surrounded basally with diffuse red in

paratype CSIRO H. 6247-01, fig. Id). 1st to 3rd (1st and 2nd in

some paratypes) dorsal-fin rays crimson in males, dorsal-fin

spines, remaining rays and dorsal-fin membrane pale yellow to

semitransparent. Anal-fin membrane chalky-white on basal

three-fourths, crimson distally in males. Caudal fin translucent

greyish with 4-5 irregular wavy diagonal lemon-yellow bars,

lower margin bright-yellow with a crimson flash submarginally

on outer half of fin. Some paratypes with yellow and red

colouration interspersed along lower margin of fin, and with a

lesser 2nd red flash immediately above, near distal edge of fin.

Pectoral fins translucent. Proximal half of pelvic fins white,

yellowish distally (some larger paratypes with outer half of fin

faintly pale red-pink). Peritoneum silvery-black.

Distribution. Demersal trawl grounds between Swain Reefs,

Qld (22°06'S, 153°02'E) and Manly, NSW (33°40'S, 151°26'E),

at depths of 51-170 m (fig. 4).

Etymology. Erom the Latin tres meaning three, and spina

meaning thorn or spine, in reference to the 3 dorsal-fin spines, a

character state not found in any other species of Pinguipedidae.

Remarks. Although it has a distributional range of at least 11

degrees latitude, Simipercis trispinosa has been trawled from

only several relatively small areas of the east Australian coast

(fig. 4). There appears to be a strong correlation between

increased depth of capture and decreasing latitude across the

range of the species. Specimens from near the northern range

extremity, in the southern Great Barrier Reef region, occur in

considerably greater depths (112-170 m) than those at the

southern end, in New South Wales (51-80 m). Those centrally

located, off the Sunshine Coast in southern Queensland were

recorded from intermediate depths (98-123 m). Most trawl

catches of S. trispinosa have included several males in excess of

110 mm SL with prominent fin markings and well-developed

caudal-fin filaments (maximum recorded size 135 mm SL),

among a larger group of smaller females, ranging from about

60-100 mm SL. The largest female examined with ripe gonads

measured 100 mm SL. Nuchal crests were noted in specimens of

all sizes and are not related to age, gender or sexual maturity.

Acknowledgements

We thank D. Roy and M. Tonks, Queensland Department of

Primary Industries and Eisheries, for provision of fresh

specimens; M. McGrouther, Australian Museum, for sending

material on loan; D. Gledhill, CSIRO Marine Research, K.

Graham, NSW Eisheries and M. McGrouther, for making

available photographs of the new species; and H. Imamura,

Hokkaido University Museum, for kindly providing vertebral

counts of Kochichthys and Pseudopercis specimens that he

had examined.

References

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branchiaux et a dents en velours on en cardes. Histoire Naturelle

des Poissons 3: i-xxviii + 2 pp. 500 pp.

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537-547.

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Academy of Natural Sciences of Philadelphia 17: 66-68.

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of the United States National Museum 124: 1-78.

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relationships of the family Pinguipedidae (Teleostei: Perciformes).

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Niem, V.H. (eds.) FAO Species Identification Guide for Fishery

Purposes. The Living Marine Resources of the Western Central

Pacific. Vol. 6. Bony Fishes, part 4 (Labridae to Latimeriidae),

estuarine crocodiles, sea turtles, sea snakes and marine mammals,

pp. 3501-3510.

Randall, J.E. 2003. Review of the sandperches of the Parapercis

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Memoirs of Museum Victoria 63(1): 65-75 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Notoraja hirticauda, a new species of skate (Chondrichthyes: Rajoidei) from the

south-eastern Indian Ocean

Peter R. LasP and John D. McEachran^

'CSIRO, Marine Laboratories, G.P.O. Box 1538, Hobart, Tasmania 7001, Australia (peter.last@csiro.au)

^Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, TX 77843, U.S.A

(j-mceachran@neo.tamu.edu)

Abstract Last P.R. and McEachran J.D. 2006. Notoraja hirticauda, a new species of skate (Chondrichthyes: Rajoidei) from the

south-eastern Indian Ocean. Memoirs of Museum Victoria 63(1): 65-75.

A new skate, Notoraja hirticauda sp. nov., is described from off central Western Australia in the south-eastern Indian

Ocean. It is distinguished from its congeners in some proportional measurements, the distributions of dermal denticles and

bristle-like thorns, the absence of oronasal pits, and several skeletal structures. The new species has a relatively long snout,

multiple irregular rows of short thorns on the dorsal and lateral surfaces of the tail, and dermal denticles on the ventral

surface of the tail. Its rostral cartilage, which is longer than the nasobasal length, has appendices that are subtriangular in

cross-section posteriorly and abut the rostral shaft. Within Notoraja, N. hirticauda most closely resembles N. subtilispinosa

Stehmann from the Philippines and Indonesia.

Keywords Chondrichthyes, Rajidae, new species, skate, Notoraja hirticauda, south-eastern Indian Ocean

Introduction

The highly diverse and endemic rich skate faunas of Australia

and New Zealand are represented by at least 62 species, of which

all but two appear to be restricted to the Australasian region

(Last and Yearsley, 2002). Similarly, of the 40 or so Australian

species (Last and Stevens, 1994), only one also occurs off New

Zealand. Several new species have been identified from recent

exploratory, trawl-fishing catches in Australian seas but most of

these remain undescribed. For example, the rajoid fauna off the

western coast of Australia consists of at least 14 species of which

only three have been described (Last and Stevens, 1994). Many

of these appear to have narrow depth and geographic ranges.

Among the least well-known skates, at both specific and

supraspecific levels within the suborder, are Australian members

of the Notoraja-Xike, skates.

McEachran and Miyake (1990) presented a cladogram

dividing rajoids into two major clades: Group I consisting of

nine supraspecific taxa including the pavorajine subgroups

Pavoraja Whitley, Notoraja Ishiyama and a recently described

genus, Brochiraja (Last and McEachran, 2006; formerly as an

unnamed taxon “Subgenus A” of Pavoraja, sensu McEachran,

1 984); and Group II consisting of 1 1 supraspecific taxa including

“typical” rajine subgroups such as Raja and Dipturus. Group I

skates are well represented in the western Pacific with at least

30 species occurring off Australasia (Last and Yearsley, 2002).

However, relationships within the clade, which includes the

pavorajine subgroups and Insentiraja Yearsley and Last, are not

fully understood. McEachran and Last (1994) have suggested

that Brochiraja (as Subgenus A) is a subgroup of Notoraja

rather than Pavoraja, as originally suggested by McEachran

(1984). However, elucidation of the relationships within this

generic complex is dependent on descriptions of other members

within the clade. Notoraja was resurrected by Stehmann (1989)

and presently contains four valid nominal species, N. laxipella

(Yearsley & Last) and N. ochroderma McEachran and Last

from north-eastern Australia, N. tobitukai (Hiyama) from Japan,

and N. subtilispinosa Stehmann from the Philippines and

Indonesia, although other species are known to occur in the

Indo-west Pacific (McEachran and Last, 1994). A new species

of Notoraja is described below based on material from the

continental slope off Western Australia.

Several different methods for taking external measurements

of skates have been proposed. This has caused considerable

confusion in the past, so our methods are explained here in

some detail. We used the point-to-point approach proposed by

Hubbs and Ishiyama (1968), but raw data are finally expressed

as proportions of total length rather than disc width. The preoral

length is defined as the measurement from the snout tip to the

anterior median edge of the upper jaw rather than to the mouth

Peter R. Last and John D. McEachran

proper. Mouth width is the greatest dimension across the tooth

band of the upper jaw rather than visible width of the upper jaw

between the inner margins of integument (Yearsley and Last,

1992). Spiracular measurements refer to distances around the

main depression and do not include the anterior secondary

depression. The maximum and minimum widths of the nasal

curtain refer to the greatest dimensions across the curtain

(usually across the lobes) and the width between the innermost

insertions of the curtain immediately before the mouth

respectively. The nasal lobe dimension is the greatest width of

the lobe from the insertion of the curtain to its lateral margin.

Pelvic-fin lobe lengths are taken from the point of articulation

of the anterior lobe as described by McEachran and Stehmann

(1984) and Stehmann (1985) rather than from the haemal spine

(Hubbs and Ishiyama, 1968). The width of the pelvic-fin base is

the minimum distance between these points of articulation.

Measurements to the cloaca originate from near its posterior

margin (i.e. approximated from the origin of the first haemal

spine) rather than at its midpoint. Skeletal measurements and

counts follow Hubbs and Ishiyama (1968) and McEachran and

Compagno (1979, 1982).

Some mature paratypes were dissected to reveal the

structure of the neurocranium, scapulocoracoid and clasper;

skeletal elements were highlighted with Alcian blue eight GS

cartilage stain. All specimens were radiographed to verify

anatomical dissections, to count vertebrae, teeth and pectoral-

fin radials, and to determine the shape of the pelvic girdle.

Specimens examined in this study were obtained from the

Australian National Eish Collection, Commonwealth Scientific

and Industrial Research Organisation, Hobart (CSIRO).

Notoraja hirticauda sp. nov.

Eigures 1-8, Tables 1-3

Notoraja sp. C — Last and Stevens, 1994: 312, figs 33, 34.5, PI 50

(all fig. captions incorrect, as N. sp B).

Material examined. Holotype. Australia: CSIRO H1816-02, 446 mm

TL, mature male, west of Shark Bay, Western Australia, 26°36'S,

112°09'E, 760 m, 28 December 1989, RV “Akebono Maru”, sta. 41.

Paratypes (n=8): Australia: CSIRO CA2820, 312 mm TL, female,

CSIRO CA2821, 264 mm TL, female, north of Nickol Bay, Western

Australia, 18°38'S, 116°54'E, 600 m, 7 Apr 1982, LRV “Soela”, Cr. 2,

sta. 30; CSIRO CA2822, 241 mm TL, female, CSIRO CA2823, 277

mm TL, female, north of Nickol Bay, Western Australia, 18°41'S,

116°45'E, 596 m, 6 Apr 1982, LRV “Soela”, Cr. 2, sta. 28; CSIRO

CA2824, 361 mm TL, immature male, north of Nickol Bay, Western

Australia, 18°43'S, 116°33'E, 612 m, 7 Apr 1982, LRV “Soela”, Cr. 2,

sta. 29; CSIRO H1816-03, 448 mm TL, mature male, captured with

holotype; CSIRO HI822-0I, 372 mm TL, female, west of Shark Bay,

Western Australia, 26°44'S, 112°19'E, 735 m, 28 Dec 1989, RV

“Akebono Maru”, sta. 40; CSIRO H2574-06, 428 mm TL, female,

west of Lreycinet Estuary, Western Australia, 26°38'S, 112°30'E, 500

m, 30 Jan 1991, LRV “Southern Surveyor”, Cr. 1, sta. 37.

Diagnosis. A small species of Notoraja with the following

combination of characteristics: preoral snout 13.5-15.2% TL;

preorbital snout 12.3-13.8% TL; mutiple irregular rows of

small, bristle-like thorns and large dermal denticles present on

both surfaces of tail; single preorbital thorn present; no oronasal

pits; rostral cartilage about 1.25 in nasobasal length; rostral

appendices about 62% of nasobasal length, subtriangular in

cross-section posteriorly and evenly abutting rostral shaft;

pectoral radials 61-66; trunk vertebrae 24-27, precaudal tail

vertebrae 71-76; and both surfaces of body pale.

Description. Disc heart-shaped, 1.11 times as broad as long in

holotype (1.11-1.16 times in paratypes); maximum angle in

front of spiracles 73-75° in adult males, 93-97° in females and

juveniles; anterior margin initially straight, slightly more convex

in females and immatures, moderately concave opposite

spiracles in mature male holotype; posterior margin and comers

broadly convex (figs 1,2). Snout to axis of greatest width 61.5%

(57.7-60.9%) of disc length. Snout tip with prominent fleshy

and laterally flattened apical process; process rather broad-based

and almost continuous with anterior profile of disc in mature

males, more lobe-like in immatures and adult females. Pelvic

fins deeply incised, anterior lobe moderately long, narrow at

base and tapering to blunt point; posterior lobe relatively short,

with strongly convex lateral margins; anterior lobe length 95.7%

(93-113%) of posterior lobe. Tail narrow at base, tapering

gradually posteriorly, very slender toward tip; moderately

depressed over length, width 1.42 (1.44—1.72) times height at

axil of pelvic fin; slightly convex on ventral surface; lateral skin

folds originating well behind posterior pelvic-fin lobes, extending

to below mid-length of epichordal caudal-fin lobe and broadening

distally (only slightly narrower than height of epichordal lobe);

length of tail from rear of cloaca 1.40 (1.40-1.50) times distance

from tip of snout to rear of cloaca; predorsal tail length 78.1%

(78.2-81.4%) of tail length; width at midlength of tail 2.06

(1.99-2.72) in orbit diameter; width at axils of pelvic fins 0.80

(0.85-1.00) in orbit diameter. Dorsal fins of similar shape and

size; rather short and moderately tall with evenly convex anterior

margin; posterior margin straight or slightly convex; tip pointed

or acutely rounded; fins separated by short interspace. Epichordal

caudal-fin lobe relatively well developed, separated by short

interspace from and distinctly longer than 2nd dorsal-fin base;

hypochordal caudal lobe very low, originating near end of lateral

fold, not confluent with epichordal lobe. Preorbital snout length

4.02 (3.15-3.50 in females and immatures) times orbit diameter;

preoral snout length 2.29 (1.98-2.35) times intemarial distance.

Orbit diameter 1.17 (1.11-1.45) times interorbital distance; 1.87

(2.02-3.08) times length of spiracles. Lateral nasal fold expanded

slightly, with weak triangular process along lateral margin,

posterior margin smooth or with a weak fringe; nasal curtain

moderately well developed, broadly rounded and weakly fringed

(often hardly detectable) along posterior margin. Oronasal pits

absent. Upper and lower jaws slightly arched on either side of

symphysis; upper jaw of holotype indented at symphysis, not

indented in females and immatures. Teeth in 36 (35^0) series in

upper jaw, 34 (32-39) series in lower jaw; plate-like with short

cusps in females, cusps acutely pointed in males, arranged

quincuncially. Distance between 1st gill slits 1.72 (1.63-1.91)

times distance between nostrils; distance between 5th gill slits

1.07 (1.04—1.27) times distance between nostrils; length of 1st

gill slit 6.7 (4.7-8.4) times mouth width.

Dorsal surface of disc, posterior lobe of pelvic fin, dorsal

fins, epichordal lobe of caudal fin, and lateral skin folds densely

New skate from the south-eastern Indian Ocean

Table 1. Morphometric data for the holotype of Notoraja hirticauda sp. nov., with ranges and means for paratypes. Measurements are expressed

as percentages of total length

Holotype

CSIRO

H1816-02

Paratypes

n=8

Min

Max

Mean

Total length (mm)

446

241

448

Disc width

48.7

47.3

51.3

48.7

Disc length

43.7

41.8

45.2

43.0

Head length (dorsal)

17.8

16.9

17.7

17.4

Head length (ventral)

25.1

23.1

25.7

24.3

Snout length (preorbital)

13.8

12.3

13.2

13.0

Snout length (preoral)

15.2

13.5

15.0

14.6

Prenasal length

11.1

10.1

11.0

10.5

Snout to maximum width

26.9

24.5

26.6

25.7

Orbit diameter

3.3

3.8

4.2

3.9

Distance between orbits

2.8

2.7

3.5

3.0

Orbit and spiracle length

4.3

4.2

5.0

4.5

Spiracle length

1.8

1.3

2.1

1.7

Distance between spiracles

5.4

6.3

5.7

Mouth width

7.2

5.7

6.6

6.3

Nare to mouth

4.3

3.6

4.7

4.2

Distance between nostrils

6.1

6.0

6.4

6.2

Width of first gill slit

1.0

0.8

1.2

1.0

Width of third gill slit

0.8

0.9

1.3

1.1

Width of fifth gill slit

0.7

0.9

1.1

0.9

Distance between first gill slits

10.6

10.4

11.7

11.1

Distance between fifth gill slits

6.5

6.4

7.8

7.2

Length of anterior pelvic-fin lobe

13.7

13.1

15.0

14.4

Length of posterior pelvic-fin lobe

12.2

12.3

14.5

13.4

Tail width at axil of pelvic fins

4.1

4.0

4.5

4.2

Tail height at axil of pelvic fins

2.9

2.5

3.1

2.7

Tail width at tips of pelvic fins

3.9

3.2

3.9

3.6

Tail height at tips of pelvic fins

2.4

2.1

2.6

2.3

Width of tail across its midpoint

1.6

1.5

2.0

1.7

Distance— snout to cloaca

41.7

40.1

42.4

41.0

Distance— cloaca to 1st dorsal fin

45.3

45.6

48.4

46.8

Distance— cloaca to 2nd dorsal fin

50.0

50.2

52.2

50.8

Distance— cloaca to caudal-fin origin

54.5

54.0

56.0

55.0

Distance— cloaca to caudal-fin tip

58.3

58.4

61.0

59.5

Number of tooth rows (upper jaw)

38.2

Number of tooth rows (lower jaw)

35.6

Number of trunk vertebrae

25.4

Number of precaudal tail vertebrae

73.0

Number of pectoral-fin radials

64.2

Peter R. Last and John D. McEachran

Table 2. Morphometric data fotr the neurocranium of paratype (CSIRO H1822-01) of Notoraja hirticauda sp. nov. Measurements are expressed

as percentages of nasobasal length

Neurocranium

% length

Nasobasal length (mm)

31.4

Cranial length

72.8

231.8

Rostral cartilage length

39.3

125.2

Rostral cartilage width

1.6

5.2

Prefontanelle length

33.1

105.4

Cranial width

39.0

124.2

Interorbital width

10.1

32.3

Rostral base

3.8

12.0

Anterior fontanelle length

14.4

45.9

Anterior fontanelle width

3.9

12.5

Rostral appendix length

19.3

61.5

Rostral appendix width

5.4

17.2

Rostral cleft length

10.8

34.4

Cranial height

7.4

23.5

Width across otic capsules

19.0

60.5

Width of basal plate

9.0

28.7

Greatest width of nasal aperture

16.4

52.1

Least width of nasal aperture

7.0

22.4

Internasal width

3.9

12.5

Table 3. Morphometric data for the scapulocoracoid of paratype (CSIRO H2574-06) of Notoraja hirticauda sp. nov. Measurements are expressed

as percentages of scapulocoracoid length

Scapulocoracoid

% length

Scapulocoracoid length (mm)

22.7

Scapulocoracoid height

17.2

75.5

Premesocondyle

9.0

39.6

Postmesocondyle

13.7

60.4

Postdorsal fenestra length

6.5

28.8

Postdorsal fenestra height

5.2

23.0

Anterior fenestra length

2.9

13.0

Anterior fenestra combined height

6.5

28.8

Base length

20.6

90.6

Anterior corner height

16.4

71.9

Posterior corner height

12.8

56.1

New skate from the south-eastern Indian Ocean

Figure 1. Notoraja hirticauda sp. nov. Holotype (CSIRO H1816-02, 446 mm TL, male), a. Dorsal view; b. Ventral view.

Peter R. Last and John D. McEachran

fig. 4a

Figure 3. Squamation on the mid-tail of Notoraja hirticauda sp.

nov. Holotype (CSIRO H1816-02, 446 mm TL, male), a. Dorsal

view; b. Lateral view.

fig. 4b

fig. 4c

Figure 4. Neurocranium of Notoraja hirticauda sp. nov. Paratype (CSIRO H1822-01, mature female, 372 mm TL: a. dorsal view; b. lateral view;

c. posterior view), fig. abbreviations: ac-anterior cerebral vein foramen, af-anterior fontanelle, antc-antorbital condyle, bf-basal fenestra, end-

endolymphatic foramen, es-efferent spiracular artery foramen, hf-hyomandibular facet, into-intercerebral vein foramen, ja-jugal arch, IbX-

lateralis branch of vagus nerve foramen, obf-otic branch of facial nerve foramen, oc-occipital condyle, of-orbital fissure, one-orbitonasal canal

foramen, os-optic stalk, peri-perilymphatic nerve foramen, pf-posterior fontanelle, poc-preorbital canal, postp-postorbital process, prep-

preorbital process, prof-profundus nerve foramen, pterp-pterotic process, ra-rostral appendix, rb-rostral base, rn-rostral node, rs-rostral shaft,

Il-optic nerve foramen, Ill-oculomotor nerve foramen, Vll-hyomandibular branch of facial nerve foramen, IX-glossopharyngeal nerve

foramen, X-vagus nerve foramen.

New skate from the south-eastern Indian Ocean

Figure 5. Lateral view of left scapulocoracoid of Notoraja hirticauda sp. nov. Paratype (CSIRO H2574-06, mature female, 428 mm TL). fig.

abbreviations: ab-anterior bridge, ac-anterior corner, adf-anterodorsal fenestra, avf-anteroventral fenestra, msc-mesocondyle, mtc-metacondyle,

pdf-postdorsal fenestra, prc-procondyle, pvf-postventral foramina, rc-rear corner, scp-scapular process.

Figure 6. Ventral view of pelvic girdle of Notoraja hirticauda sp. nov.

a. Holotype, (CSIRO H1816-02, male, 446 mm TL); b. Paratype

(CSIRO H2574-06, female, 428 mm TL). fig. abbreviations: ilp-iliac

process, obf-obturator foramina, prep-prepelvic process.

Figure 7. Lateral view, partially expanded, of right clasper of Notoraja

hirticauda sp. nov. Holotype (CSIRO H1816-02, 446 mm TL, male),

fig. abbreviations: cf-cleft, hy-hypopyle, pj-projection, pr-

pseudorhipidion, rh-rhipidion, sl-slit, sp-spike.

Peter R. Last and John D. McEachran

fig. 8a

Figure 8. Right clasper cartilages of Notoraja hirticauda sp. nov. Holotype (CSIRO H1816-02, 446 mm TL, male): a) lateral view; b) dorsal

view; c) ventral view. fig. abbreviations: atr2-accessory terminal 2 cartilage, ax-axial cartilage, dmg-dorsal marginal cartilage, dtrl-dorsal

terminal 1 cartilage, dtr2-dorsal terminal 2 cartilage, tb-terminal bridge, vmg-ventral marginal cartilage, vtr-ventral terminal cartilage.

New skate from the south-eastern Indian Ocean

covered with fine, more or less evenly spaced dermal denticles;

thorns (fig. 3) small, delicate, confined to tail and preorbit, and

alar regions of mature males. Denticle cusps on disc very

slender, erect and slightly recurved; bases quadriradial, radii

arranged more or less at right angles, anterior radius generally

shortest (sometimes barely detectable). Both surfaces of tail

densely covered with denticles and small, bristle-like thorns;

thorns variable in size, scattered, interspersed with denticles;

median thorns in an almost linear series, other thorns distributed

randomly or in irregular series; largest thorns long-based,

somewhat compressed, sharply pointed distally, upright and

triangular or with narrow recurved tips; most numerous dorsally

on centre of tail; thorns on ventral surface of tail generally

smaller than those dorsally but much larger than denticles of

disc. Single, rather short preorbital thorn present. Alar thorns

strongly recurved and directed postero-medially, central group

in indistinct rows and merging with a cluster of smaller thorns

near anterior outer margin of pectoral fin; holotype with about

5-6 central rows, each with 1-4 thorns (mature male paratype

with about 9-12 rows, each with 1-2 thorns); all thorns non-

depressible, obliquely oriented to surface of disc, not fitting into

slits in integument. Claspers, anterior lobes of pelvic fins, and

ventral surfaces of disc and posterior pelvic-fin lobes naked;

integument of ventral surface rather delicate.

Skeletal morphology. Neurocranium with nasobasal length

20.7% of distance from snout to posterior margin of cloaca and

80.5% of cranial width. Rostral shaft slender, of moderate length,

about 125% of nasobasal length; rostral base 12% of nasobasal

length (fig. 4, Table 2); rostral appendices long, flattened

anteriorly, subtriangular in cross-section and abutting (rather

than fused to) rostral shaft posteriorly; propterygia of pectoral

girdle reaching rostral node; nasal capsules very large, ovoid, set

at about 26° to transverse axis of neurocranium; foramen for

profundus nerve near antero-lateral margin of nasal capsules;

preorbital process poorly developed; postorbital process poorly

developed and barely distinct from pterotic process; anterior

fontanelle very narrow, with acutely rounded apex and bilobed

posterior margin, extending to leading edge of nasal capsule;

posterior fontanelle indistinct (not visible in radiographs and

only detectable after staining and dissection), irregularly notched

due to broad margin of acellular perichondrial tissue, fontanelle

covered with dense fibrous connective tissue; basal fenestra

kidney-shaped; anterior cerebral vein foramen antero-dorsal to

optic nerve foramen and above orbito-nasal canal foramina;

oculomotor nerve foramen dorsal to optic stalk; jugal arches

weak; basal plate and intemasal plate relatively narrow, about

28.7% and 12.5% of nasobasal length respectively.

Scapulocoracoid of female moderately short, longer than

deep, tapering slightly posteriorly; lateral face expanded

slightly between mesocondyle and metacondyle (fig. 5, Table

3); anterior bridge wide, thin, strongly depressed; anterior

dorsal fenestra slightly larger than anterior ventral fenestra;

scapular process elevated above antero-dorsal margin; anterior

corner weakly angular, antero-lateral margin very slightly

convex; rear corner not extended distally; mesocondyle slightly

depressed; postdorsal fenestra moderately expanded, oval; 3

postventral foramina, central smallest.

Pelvic girdle with short prepelvic processes (fig. 6); iliac

processes moderately well developed, strongly recurved;

ischiopubic region relatively narrower in males than in females,

with straight anterior margin in males and weakly convex

margin in females, but with strongly concave posterior margins

in both sexes; iliac region with 2-3 obturatorial foramina.

Claspers long, very slender, proximal to and not expanded at

origin of glans (fig. 7); without dermal denticles or pseudo-

siphon; glans weakly depressed, moderately expandable. Inner

dorsal lobe with pseudorhipidion extending from level of

hypopyle and medial to slit, to about distal quarter of glans;

continuing distally almost to glans margin as a fold of integument;

lobe with deep cleft lateral to pseudorhipidion; slit proximal to

cleft; spur and flag absent. Ventral lobe without shield; pela well

developed, extending from level of hypopyle into distal half of

glans; projection slightly curved laterally, extending for almost

half length of glans from beneath pela to its pointed, naked distal

tip near glans margin; spike disc-shaped with sharp, naked

lateral and distal margins, ventral to projection, partly covered

by sentina but disto-lateral margin visible. Axial cartilage with a

slender conical tip; extending well beyond other cartilages (fig.

8). Dorsal marginal cartilage not expanded distally; with a

truncate distal border (with the dorsal terminal 2 cartilage) and a

long, slender, distomedial extension forming the pseudorhipidion.

Dorsal terminal 1 cartilage relatively narrow and band-like with

irregular anterior and posterior margins; located at about

midlength of pela and enveloping axial cartilage. Dorsal terminal

2 cartilage rather broad, almost spatulate distally; connected to

medial aspect of axial cartilage by well-developed terminal

bridge and loosely connected to tip of axial by connective tissue.

Ventral marginal cartilage not flared laterally at level of

hypopyle; extending almost to distal margin of accessory

terminal 2 (as component projection). Ventral terminal cartilage

J-shaped; proximo-medial condyle articulating with ventral

marginal cartilage, disto-medial extension articulating with tip

of axial cartilage; inner limb seated near articulation of accessory

terminal 2 and ventral marginal cartilages; acute proximal tip

not forming component eperon; disto-lateral margin strongly

curved inward and not forming component shield. Accessory

terminal 1 cartilage lacking, possibly fused with ventral terminal

cartilage to form component projection. Accessory terminal 2

cartilage with large disc-shaped distal extension forming

component spike; articulating with postero-medial margin of

ventral marginal cartilage.

Colour in preservative. Dorsal surface straw-coloured to pale

greyish-yellow, minute melanophores scattered widely over

most of surface; ventral surface whitish with translucent skin,

palest near margins of disc, minute melanophores lightly

scattered over tail. In life, paler, creamish to white above,

pinkish over body cavity and head; eyes black; tail brilliant

white; skin totally transparent ventrally, viscera distinct.

Size. A small skate attaining at least 448 mm TL and 222 mm

disc width; males maturing larger than 360 mm TL.

Distribution. Known from the south-eastern Indian Ocean on

the central western sector of the Australian continental slope

(between Shark and Nickol Bays, Western Australia), in depths

of 590-760 m. Apparent stenobathic and narrow geographic

Peter R. Last and John D. McEachran

ranges may not be an artefact; trawl data from depths shallower

and deeper, and in areas adjacent the geographic range, included

other sympatric rajoid species, but excuded N. hirticauda.

Etymology. A combination of the Latin hirtus (rough or bristly)

and cauda (tail) in allusion to the dense coverage of fine

denticles on both surfaces of the tail. Known by the vernacular

‘ghost skate’.

Remarks. Notoraja hirticauda fits the diagnosis of Notoraja as

defined by Stehmann (1989) and McEachran and Last (1994),

except that the tail is unusually thorny (compared to thorns

absent or in a medial row in other Notoraja species), the rostral

cartilage (including node) is distinctly longer than the nasobasal

length (rather than about equal in length), and the rostral

appendices are subtriangular posteriorly and abut the rostral

shaft (rather than plate-like and running closely parallel to the

shaft). Notoraja hirticauda is further distinguished from its

nominal congeners A. laxipella,N. ochroderma,N . subtilispinosa,

and A. tobitukai, in proportional measurements, lacking oronasal

pits, coloration, and in some skeletal structures. Of these species,

Notoraja hirticauda exhibits greatest overall similarity to A.

subtilispinosa. It is also superficially similar to A. laxipella but

has tail thorns (otherwise absent), a well-developed anterior

bridge in the scapulocoracoid (rather than thin or absent), a

robust terminal bridge in the clasper (rather than incomplete),

and fewer precaudal vertebrae. The relatively long snout of A.

hirticauda (preorbital snout length 12.4—13.7%) is more typical

of A. laxipella (13.4—14.7% TL) than the oth&r Notoraja species

(10.8-12.5% TL). A longer snout is also evident from the

prenasal length (10.1-11.0% vs. 8.9-10% TL in these other

Notoraja species) and in the shape of the neurocranium (rostral

cartilage length 125% in A. hirticauda vs. 88-92% of nasobasal

length in A. tobitukai and A. subtilispinosa). Notoraja

ochroderma has a relatively long rostral cartilage (ie. 104% of

nasobasal length; McEachran and Last, 1994) but it is still much

shorter than that of A. hirticauda. Notoraja ochroderma and A.

hirticauda (combined orbit and spiracle lengths 3. 5^.7% TL),

and A. hirticauda (4.2-5% TL), have smaller orbits and spiracles

than A. tobitukai and A. subtilispinosa (5-5.7% TL).

Members of the genus Notoraja also exhibit significant

interspecific variation in the mouth and tail sizes, the pelvic-fin

shape, and the gill-slit size and interspacing. Notoraja hirticauda

appears to have smaller gill slits (eg. first gill-slit width 0.8-1 .2%

vs. 1.2-L5% TL) and narrower intergill distances (eg. width

between first gill slits 10.4—11.7% vs. 11.5-13% TL) than A.

subtilispinosa. The anterior pelvic-fin lobe varies greatly from

slightly shorter in mature males (anterior 93-96% of posterior

lobe) to slightly longer than the posterior lobe in most females

(typically 104—110%) of A. hirticauda. The anterior lobe is

subequal or longer in A. subtilispinosa and A. tobitukai (anterior

100-117% in posterior lobe) and shorter than the posterior lobe

in A. ochroderma (anterior 76-90% in posterior lobe). Notoraja

hirticauda and A. tobitukai (tail width at pelvic-fin tips 3 .2-3 .9%

TL) have a slightly broader tail than A. subtilispinosa and A.

laxipella (width at pelvic-fin tips 2.2-3. 1% TL). The four

Notoraja species appear to have larger mouths than A. laxipella

{5.1-1. 1 vs. 4.8-5.5).

The pectoral-fin radial and trunk vertebral counts of A.

hirticauda closely resemble those of A. subtilispinosa, A.

tobitukai and A. laxipella (ie. 24-27 and 61-67 for vertebral

and radial counts respectively). Both counts are higher for A.

ochroderma (30-32 and 87-89 for vertebral and radial counts

respectively). Similarly, the scapulocoracoid of A. hirticauda is

more similar to those of A. subtilispinosa, A. tobitukai and A.

laxipella than to that of A. ochroderma which has a relatively

more elongate scapulocoracoid with a very large, horizontally

expanded post-dorsal fenestra. Notoraja hirticauda has three

postventral foramina like female specimens of A. subtilispinosa

and A. laxipella examined to date (four in A. tobitukai and five

in A. ochroderma). The pelvic girdle of A. hirticauda most

closely resembles that of A. tobitukai. In both species, the

anterior contour of the ischiopubic bar is slightly convex and

the prepelvic processes are short and rather robust. The

prepelvic processes of A. subtilispinosa and A. ochroderma

are much more slender and the anterior contour of the

ischiopubic bar of the latter is straight (McEachran and Last,

1994). As discussed above, the neurocranium of A. hirticauda,

like A. laxipella, has a relatively long rostral shaft and

appendices compared to other species of Notoraja but the

presence of jugal arches is shared by A. laxipella and all

members of the genus except A. ochroderma.

The fontanelle structure of the neurocranium of A.

hirticauda differs from the normal condition found in skates. A

posterior fontanelle is well developed in most rajoids. Stehmann

(1989) could not detect either anterior or posterior fontanelles

in the neurocranium in radiographs of the holotype of A.

subtilispinosa, and Ishihara and Stehmann (1990) observed

only an anterior fontanelle in radiographs of a second specimen

from Indonesia. Similarly, only the anterior fontanelle was

detectable in X-rays of the neurocrania of A. hirticauda types,

although an ill-defined, posterior aperture overlain with smooth

uncalcified cartilage and dense connective tissue was evident

from dissection. A similar, poorly-defined posterior fontanelle

exits in A. laxipella. In comparison, the posterior fontanelles of

other related Australasian skates (ie Notoraja ochroderma,

Irolita, Pavoraja and Brochiraja) are sharply defined in

radiographs (Last and McEachran, unpubl.).

The rostral appendices are considered to be important

evolutionary characteristics (McEachran, 1984: McEachran

and Miyake, 1990). Three Notoraja species (A. tobitukai, A.

subtilispinosa and A. ochroderma) are considered to possess

plate-like rostral appendices (Stehmann, 1989; McEachran and

Last, 1994). These structures are thickened, almost triangular in

cross-section distally in A. hirticauda, and more closely

resemble the subconical appendices found in A. laxipella and

Brochiraja. Eurther inspection of the neurocranium of A.

laxipella revealed a relatively thinner, rostral cartilage separated

slightly from its appendices (rather than continuously abutting

them as in A. hirticauda).

The external form of the clasper of A. hirticauda is consistent

with the general type found in Brochiraja and Notoraja, except

for A. ochroderma, which has an eperon and a shield formed by

the sharp proximal margin of the ventral terminal cartilage

(McEachran and Last, 1994). The dorsal terminal 1 cartilage,

small in A. hirticauda, is absent in A. ochroderma and sometimes

absent in A. tobitukai. McEachran and Last (1994) suggested

that the “rhipidion” of Notoraja, Pavoraja and Irolita is

structurally different and probably not homologous with the

New skate from the south-eastern Indian Ocean

rhipidion of Ishiyama (1958, 1967), Stehmann (1970) and

Hulley (1972), that occurs in the majority of skate taxa in Group

II (McEachran and Miyake, 1990). Ishiyama (1958, 1967)

applied the terms pent and pela to this structure but since pent

has been proposed for a separate component, the “rhipidion” of

Group I should be called the pela to avoid further confusion.

Within Notoraja, only N. ochroderma and N. hirticauda are

whitish on both dorsal and ventral surfaces. N. ochroderma

possesses oronasal pits which are lacking in all other Notoraja

species. The dorsal squamation of N. hirticauda, consisting of

fine, erect but slightly recurved denticles, is similar to

N. subtilispinosa, N. ochroderma and N. laxipella. Also, the tail

of N. hirticauda and N. ochroderma, unlike the other two

species, has irregular rows of small thorns. However, unlike N.

ochroderma, the ventral surface of the tails of N. hirticauda and

the other Notoraja species are covered in denticles.

Much of the pavorajine skate fauna of the Indo-Pacific

remains undescribed. Within the known fauna, N. hirticauda

appears to be much more similar to N. subtilispinosa and N.

laxipella than either N. ochroderma or N. tobitukai, and the

supraspecific structure of the Notoraja group needs further

investigation. McEachran and Last (1994) noted that N.

ochroderma did not fit the synapomorphy scheme of either

Notoraja or Pavoraja but tentatively placed it in Notoraja as a

conservative measure. Yearsley and Last (1992), who

demonstrated similarities between the subgenus Insentiraja and

Notoraja, temporarily placed the former in Pavoraja based on

McEachran’s earliest synapomorphy scheme (1984) and stressed

the need for a review of Indo-Pacific pavorajines. A study of the

relationships of Australian and New Zealand skates is now in

progress by McEachran and Last (unpubl.).

Acknowledgements

We thank M. Gomon, B. Ward and G. Yearsley for comments on

the manuscript. B. Hope helped with measurements. A. Graham

provided technical support, including radiography, specimen

curation and the checking of specimen data. T. Carter took

photographs of the types and G. Davis illustrated the skeletal

structures. Digital imagery and enhancement was completed by

L. Conboy.

References

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and description of skates (Rajidae). Copeia 1968(3): 483-491.

Hulley, P. A. 1972. The origin, interrelationships and distribution of

southern African Rajidae (Chondrichthyes, Batoidei). Ann. S. Ajr.

Mus. 60: 1-103.

Ishihara, H. and Stehmann, M. 1990. A second record of the deep-

water skate Notoraja subtilispinosa from the Flores Sea,

Indonesia. Jap. J. Ichthyol. 37: 182-186.

Ishiyama, R. 1958. Studies on the rajid fishes (Rajidae) found in the

waters around Japan. J. Shimonoseki Coll. Fish. 7: 1-394.

Ishiyama, R. 1967. Fauna Japonica: Rajidae (Pisces). Biogeographical

Society of Japan: Tokyo, 84 pp.

Last, P. R. and McEachran J. D. 2006. New softnose skate genus

Brochiraja from New Zealand (Rajidae: Arhynchobatinae) with

description of four new species. N. Z. J. Mar. Freshwater Res. 40:

65-90.

Last, P. R. and Stevens, J. D. 1994. Sharks and rays of Australia.

CSIRO: Melbourne, 513 pp.

Last, P. R. and Yearsley G. K. 2002. Zoogeography and relationships of

Australasian skates (Chondrichthyes: Rajidae). J. Biogeogr. 29:

1627-41.

McEachran, J. D. 1984. Anatomical investigations of the New Zealand

skates Bathyraja asperula and B. spinifera, with an evaluation of

their classification within the Rajoidei (Chondrichthyes). Copeia

1984: 45-58.

McEachran, J. D. and Compagno, L. J. V. 1979. A further description

of Gurgesiellafurvescens with comments on the interrelationships

of Gurgesiellidae and Pseudorajidae (Pisces, Rajoidei). Bull. Mar.

Sci. 29: 530-553.

McEachran, J. D. and Compagno, L. J. V. 1982. Interrelationships of

and within Breviraja based on anatomical structures (Pisces,

Rajoidei). Bull. Mar. Sci. 32: 399-425.

McEachran, J. D. and Last, P. R. 1994. New species of skate, Notoraja

ochroderma, from off Queensland, Australia, with comments on

the taxonomic limits of Notoraja. (Chondrichthyes, Rajoidei).

Copeia 1994: 413-421.

McEachran, J. D. and Miyake, T. 1990. Phylogenetic interrelationships

of skates: a working hypothesis (Chondrichthyes: Rajoidei) in:

Pratt Jr, H. L., Gruber, S. H. and Taniuchi, T. (eds). Elasmobranchs

as Living Resources: Advances intheBiology, Ecology, Systematics,

and the Status of the Fisheries. NOAA Technical Report 90.

McEachran, J. D. and Stehmann, M. 1984. A new species of skate.

Neoraja carolinensis, from off the southeastern United States

(Chondrichthyes, Rajoidei). Proc. Biol. Soc. Wash. 97: 724-735.

Stehmann, M. 1970. Vergleichend morphologische und anatomische

Untersuchungen zur Neuordnung der Systematik der

nordostatlantischen Rajidae. Arch. Fischwiss. 21: 73-164.

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“Walther Herwig” nach Sudamerika. LXIV. Bathyraja

papilionifera sp. n. (Pisces, Batoidea, Rajidae), eine weitere neue

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Memoirs of Museum Victoria 63(1): 77-90 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Nomenclature and distribution of the species of the porcupinefish family

Diodontidae (Pisces, Teleostei)

Jeffrey M. Leis

Ichthyology, Australian Museum, 6 College St, Sydney, NSW 2010, Australia (e-mail; jeffl@austmus.gov.au)

Abstract Leis, J.M. 2006. Nomenclature and distribution of the species of the porcupinefish family Diodontidae (Pisces, Teleostei).

Memoirs of Museum Victoria 63(1): 77-90.

The tetraodontiform fish family Diodontidae is widely distributed in tropical and temperate marine waters. The family

has more than 70 nominal species, over 60% of which were described in the 100 years following Linnaeus. As a

consequence, many descriptions are less than detailed, and many types are no longer extant, if they existed at all. The high

incidence of synonymy, the many ‘old’ descriptions and the wide geographical distributions of the species has led to a great

deal of confusion. The present study, based on examination of diodontid holdings in 29 major collections, and including

the extant types of all but two of the nominal species, attempts to clarify the nomenclature and distribution of the species

of the family. Although some species boundaries are not entirely clear, only 18 or 19 of the nominal species are herein

regarded as valid (one as a subspecies). Tentative assignments of species to genera maintain current usage. Final

assignments to genera must await a cladistic analysis of relationships within the family. Four species are circumtropical,

four species (plus a subspecies) are confined to the Atlantic and appear to form a species group, four species are widely

distributed in the tropical Indo-Pacific, two species are confined to tropical Australasia, and three are endemic to temperate

Australia. One species described from New Zealand either occurs also in Australia, or is a synonym of an Australian

species. Synonymies, a key to the recognized species and a table of the identities of nominal species are provided.

Keywords Tetraodontiformes, burrfish, Allomycterus, Chilomycterus, Cyclichthys, Dicotylichthys, Diodon, Lophodiodon, Tragulichthys

Introduction

The porcupinefish family Diodontidae contains about

19 species in seven or eight genera of warm to temperate seas.

There are about 75 nominal species in the family, and several

of the species have very wide distributions. The species are

conspicuous, readily captured, of unusual morphology, and

have been the focus of interest by naturalists from ancient

times. Several species have pelagic stages that reach large

sizes and differ in appearance from the demersal adults.

All this has led to a great deal of confusion as to the number

of species, their distributions and the correct names for

them. The purpose of this paper is to clarify a number of

nomenclatural issues and the distributions of the species.

Due to their wide distributions, the species are obvious targets

for molecular genetic studies, and it is reasonable to expect

that some taxa that are currently, on the basis of morphology

alone, considered to represent a single, widespread species

will be subdivided once genetic studies are undertaken.

Conversely, some Atlantic taxa have less than clear separations

and may eventually be considered conspecific. Therefore,

it is important to lay some groundwork for these expected

future studies.

This paperlists the senior synonymofeachmorphologically-

defined species, followed by the junior synonyms. Brief

justifications for synonymies are provided, as are descriptions

of the distribution of the species based on material examined

or on identifiable literature records. In addition to the key

provided here, regional keys to the species, and illustrations of

them, can be found in Leis (1986, ref. 5686 - western Indian

Ocean); Leis (2001, ref. 26318 - western central Pacific Ocean:

this key also covers all species in the eastern Indian Ocean);

Allen and Robertson (1994, ref. 22193 - eastern Pacific Ocean);

Leis (2003, ref. 27121 - western central Atlantic Ocean); Leis

(in press, eastern central Atlantic Ocean). The key in Leis

(2001, ref. 26318) includes all Indo-Pacific species and genera

recognized herein, except the two temperate Australasian

species Diodon nicthemerus Cuvier and Allomycterus pilatus

Whitley (for these, see Kuiter, 1993, ref. 23929, or Gomon et

ah, 1994, ref. 22532 ).

Materials and methods

Abbreviations of fins are as follows: D, dorsal; A, anal; P,

pectoral; C, caudal. The spines mentioned are the dermal spines

(i.e., modified scales): fins of diodontids lack spines. These

Jeffrey M Leis

dermal spines have subdermal bases (or roots) that have either

two approximately opposing bases upon which the exposed

spine pivots when it elevates, or three (occasionally four)

broadly more or less equidistantly spaced bases that render the

exposed spine immobile. The exposed portion of the spine

varies in length and shape, but erectile spines are generally

round in cross-section, whereas fixed spines can vary from

round to compressed in cross-section. See Leis (1978, ref. 5529;

1986, ref. 5686; or 2001, ref. 26318) for more information on

spine morphology. Behind the massive beak-like jaws of

diodontids is a grinding, or trituration, plate formed by the

fused premaxillae and dentaries. This plate is often armed with

transverse plates of teeth, called trituration teeth.

Specific information on types is included only if it

supplements or corrects information in Eschmeyer (2005; 17

Oct 2005 version). To keep the literature cited list to a manageable

length, it includes only references not included in Eschmeyer’s

(2005) on-line database (http://www.calacademy.org/research/

ichthyology/catalog/fishcatmain.asp). The Eschmeyer reference

number is included with the text citation. Information on Diodon

is included in Leis (1978, ref. 5529) and in Leis and Bauchot

(1984, ref. 12539). Eor Diodon, only information on species

described since 1978 and information updating species

distributions is included here.

I examined specimens of diodontids from the following

institutions (codes after Leviton et ah, 1985, ref. 9683): AMS,

ANSP (loans based on holdings list), BMNH, BPBM, CAS,

CSIRO, BAKU, EMNH (loans based on holdings list), ERSKU,

LACM, MCZ, MNHN, NMNZ, NMV, NSMT, NTM, QM,

RMNH, ROM, RUSI, SAMA, SIO, SME, UA, USNM, WAM,

ZMA, ZMB, ZMUC. Distributions are based primarily on

museum specimens examined, but are supplemented with

reliable literature accounts.

Results

Family Diodontidae

Diagnosis. Small to medium-sized fishes to 1 m in length,

commonly 20-50 cm. Body wide and capable of great inflation,

covered with massive spines that may be quite long; spines with

large bases, or roots, under the skin; long spines usually erectile

and two-rooted, short spines usually fixed in erect position by

their three-rooted bases. Head broad and blunt; gill opening a

relatively small, vertical slit immediately before pectoral-fin

base; nasal organ usually in small tubes located in front of

large eyes; mouth large, wide and terminal; teeth fused to form

a strong, beak-like crushing structure without a median suture

dividing the upper and lower jaws into left and right halves.

Dorsal and anal fins without spines, set far back on body, and,

like caudal fin, generally rounded; most fin rays branched;

bases of fins often thick and fleshy; no pelvic fins. Lateral line

inconspicuous. No normal scales.

Genera. There is no generally agreed-upon allocation of species

to the nominal genera, nor is there any cladistic analysis of the

family or any subset of it. Most authors recognize Diodon (five

species, revised by Leis, 1978, ref. 5529) for species in which

nearly all the dermal spines are erectile. Eour monotypic Indo-

Pacific genera, three of which are confined to Australasia,

contain species that have a mixture of fixed and erectile spines

-Allomycterus, Dicotylichthys, Lophodiodon and Tragulichthys

- are usually recognized (see Gomon, 1994, ref. 22532 ; Leis,

2001, ref. 26318) and are in this paper. Chilomycterus sensu lato

(about ten species with nearly all dermal spines fixed and

immovable) has been more problematical. Tyler (1980, ref.

4477) recognized three groupings of Chilomycterus: 1) Atlantic

Chilomycterus ’ (five species confined to the Atlantic, and called

by him antennatus, antillarum, mauretanicus, schoepfii and

spinosus)-, 2) what I call herein ‘Circumtropical Chilomycterus’

(a circumtropical group considered by Tyler to consist of four

species called by him affinis, atinga, reticulatus and tigrinus)\

and 3) Tndo-Pacific Chilomycterus’ (considered by Tyler to

consist only of orbicularis). I agree with Tyler (1980, ref. 4477)

that nominal species in each of these three groups are

morphologically more similar to each other than they are to

species in the other groups. The type species of Chilomycterus

is Diodon reticulatus Linnaeus (1758, ref. 2787); thus, if these

groupings prove to be valid at the generic level, the circumtropical

group becomes Chilomycterus, and I use it in that sense herein.

Cyclichthys typically is used for several Indo-Pacific species

(including two not mentioned by Tyler [1980, ref. 4477]), and

the type of Cyclichthys is the Indo-Pacific orbicularis; thus, the

Indo-Pacific grouping can be considered Cyclichthys for the

purposes of this paper. The Atlantic group of species, regarded

by Tyler to be the most phylogenetically basal, is nearly always

included in Chilomycterus. If these Atlantic species were

removed from Chilomycterus, the generic name available for

them is Lyosphaera, based on the unique pelagic stage found in

at least some members of this group. Unfortunately, the identity

of the type species is not clear (on the basis of distribution,

schoepfii seems most likely). Lyosphaera has not been used as a

generic name for these five Atlantic species, and until a full

cladistic analysis is performed on the group, its use is not

recommended. Eor the purposes of this paper, I use Tyler’s term,

‘Atlantic Chilomycterus”, to identify this grouping.

Chilomycterus (ex Bibron) Brisout de Barneville, 1846 (sensu

stricto)

Chilomycterus (ex Bibron) Brisout de Barneville, 1846 (type

species, Diodon reticulatus Linnaeus)

Cyanichthys Kaup, 1855 (type species is D. coeruleus [non-D.

caeruleus Quoy and Gaimard] Kaup = D. reticulatus Linnaeus, 1758)

Diagnosis. All spines fixed, with long subdermal bases but

short or absent external spines (relatively smaller in larger

individuals); some spines on top of head with 4 bases; 10 C

rays; 21-23 vertebrae; heavy jaw teeth, but trituration teeth

few; no tentacles; nostril in adult an open, cup-shaped organ

with reticulations; 1 or more spines wholly on dorsal surface of

caudal peduncle; fins spotted; no large blotches on dorsal

surface of head or trunk. Some additional osteological

characters are given by Tyler (1980, ref. 4477).

The type species of Chilomycterus is Diodon reticulatus

Linnaeus (1758, ref. 2787). Cyanichthys coeruleus Kaup (1855,

ref. 2571) was based on an unregistered BMNH specimen of

43 mm SL (see also Gunther, 1870, ref. 1995 ). Although Kaup

Species of porcupinefish

Table 1. Nominal diodontid species and their current identity. Allomycterus jaculiferus McCulloch is included because of confusion about its

identity. Bibliographic details can be found in Eschmeyer (2005).

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Jeffrey M Leis

identified this specimen as D. coeruleus Quoy and Gaimard

(an alternate spelling of caeruleus), it is clearly a juvenile,

pelagic specimen of Chilomycterus reticulatus (Linnaeus).

Kaup (1855, ref. 2571) asserted the specimen was from New

Guinea, but according to Gunther (1870, ref. 1995) the locality

of the specimen “was never known at the British Museum”. A

single species (see below).

Chilomycterus reticulatus (Linnaeus, 1758)

Diodon reticulatus Linnaeus, 1758: 334 (India)

Diodon echinatusl Linnaeus, 1758; 335 (India)

Diodon tigrinus Cuvier, 1818: 127, pi. 6 (Moluccas)*

Chilomycterus affinis Gunther, 1870: 314 (unknown locality)*

Chilomycterus californiensis Eigenmann, 1891: 1133 (San Pedro,

California)*

Euchilomycterus quadradicatus Waite, 1900; 208 (Lord Howe 1.)*

Chilomycterus lissogenys Gtinther, 1910: 476, pi. 179 (Hawaii)

Chilomycterus galapagosensis Klausewitz, 1958: 82, fig. 7

(Galapagos Is.)*

*extant type

Based on examination of 55 museum specimens, including

extant types, from throughout its range (circumglobal in warm

waters), I can find no morphological differences among the

different nominal species or among geographic locations. There

is variation in colour pattern similar to that found in the

circumtropical Diodon holocanthus (see Leis, 1978, ref. 5529)

but, like that species, it is not obviously geographically based.

Spotting on the body is variable, although usually present to

some degree, but smaller spots are present on at least some, and

usually all, of the fins. The pelagic juvenile phase has a distinctly

different colour phase from the benthic adult. The species

remains pelagic to about 200 mm SL, thus providing ample

opportunity for dispersal and maintenance of genetic continuity.

Therefore, I regard Tyler’s group of ‘circumglobal Chilomycterus'

to consist of a single species. The rationale for calling this

species Chilomycterus reticulatus (Linnaeus) follows.

Diodon atringa Linnaeus (1758, ref. 2787) is frequently

regarded as a synonym (often the senior synonym) of this

species. This name is often misspelled atinga by authors. This is

clearly incorrect: Linnaeus used the spelling atringa in both his

tenth and twelfth editions. Nelson et al. (2004, ref. 27807)

recently explained why atringa is correct; and Eschmeyer (2005)

used atringa. To avoid confusion with D. atinga Bloch (1785,

ref. 4866), a synonym of D. hystrix, I herein use the spelling

atringa for the Linnaean species, regardless of the spelling used

by any subsequent author. Unfortunately, D. atringa Linnaeus is

not identifiable. There is no type, and Linnaeus’ description

could apply to any of several species of Chilomycterus or

Cyclichthys (including C. reticulatus, C. antennatus or C.

spinosus), and the same is true of Artedi (1738), the only source

cited by Linnaeus, and the pre-Linnaean authors cited by Artedi.

Artedi, 1738 mentioned that the fins of his “Ostracion bidens

sphaericus. . .”, upon which Linnaeus based his D. atringa, were

spotted, but large individuals of C. antennatus (Cuvier, 1816,

ref. 993 ) also have spotted fins (see below), so this is not

diagnostic, as is often assumed. Linnaeus (1766, ref. 2786) cited

a plate in Seba (1759, ref. 18716) that represents either C.

reticulatus or C. antennatus. Brisout de Bameville (1846, ref.

296) was the first author to express a clear opinion, and

considered D. atringa Linnaeus to be synonymous with D. orbe

Lacepede (1798, ref. 2708). The latter was based on a specimen

from Brazil - no longer extant - that is clearly identifiable as the

Atlantic Diodon spinosus Linnaeus (see below). Le Danois

(1959, ref. 12003) considered atringa Linnaeus to be

approximately equivalent to Tyler’s “Atlantic Chilomycterus"

group (which includes D. spinosus Linnaeus) with several

subspecies roughly equivalent to Tyler’s species.

In contrast, D. reticulatus Linnaeus (1758, ref 2787) is readily

identifiable. Linnaeus based his description on Artedi ’s

""Ostracion subrotudus . . ." . Atredi (1738) cited a Willughby

(1686) plate of “Orbis muricatus and reticulatus” that is clearly

identifiable as reticulatus by its colour, general morphology,

spine distribution and spine shape. The name reticulatus has been

in regular use as a senior synonym (in addition to the nine

1870-1926 references listed by Fowler, 1936, ref. 6546 and the

>30 post-1985 references listed by Eschmeyer, [2005]: Lowe,

1844, ref. 2833; Brisout de Barenville, 1846, ref. 296; Gunther,

1870, ref. 1995; Poey, 1876, ref. 3510; Goode, 1876, ref. 1832,

1877, ref. 13360; Jordan and Gilbert, 1883, ref. 2476; Eigenmann,

1885; Poll, 1959, ref. 12014; Tyler, 1980, ref. 4477; Leis, 1981,

1984; Leis and Bauchot, 1984, ref. 12539). Some authors,

apparently following Jordan and Evermann (1898, ref. 2444,

have considered reticulatus to be a junior synonym of atringa,

but none have attempted to justify this view. It is clear from

Jordan and Evermann ’s description and key that they incorporated

more than one species in their concept of C. atringa, including at

least C. reticulatus and C. antennatus. Jordan and Evermann

(1898) described C. atinga as having dark dorsal blotches and a

‘supraocular cirrus’, both features that are lacking in C. reticulatus

(Linnaeus) and in Tyler’s ‘circumtropical Chilomycterus' , but

present in species of the ‘Atlantic Chilomycterus' group.

There is a great deal of confusion in the literature as to just

what constitutes C. reticulatus and C. atringa. For much of the

19th century, most authors accepted Bloch’s (1785, ref. 21381)

concept of D. atinga {-D. hystrix Linnaeus, 1758, ref. 2787),

and although Brisout de Bameville (1846, ref. 296) pointed

out that this was in error, the use of D. atinga sensu Bloch

persisted for some years. Seret and Opic (1981) stated without

reasons that reticulatus was a synonym of C. atringa

(Linnaeus), but their illustration of C. atringa shows what

appears to be C. antennatus (Cuvier) (Seret and Opic kindly

provided unpublished dorsal and lateral views of the specimen

that strengthen this opinion). Similarly, Tortonese (1973, ref.

7192), without comment, listed reticulatus as a junior synonym

of atringa, but, later, Tortonese (in Whitehead et al., 1986, ref.

13677) illustrated as C. atringa a specimen of the eastern

Atlantic C. spinosus mauretanicus (Le Danois), but with

spotted fins, a feature I have not observed in the latter species.

Smith-Vaniz et al., 1999 (ref. 25013) listed C. atringa

(Linnaeus) as occurring in Bermuda, but had seen no

specimens, stating that their listing was based on Goode’s

(1876, ref. 1832; 1877, ref. 13360) and Gunther’s (1870, ref.

1995) records of C. reticulatus. Unfortunately, this leads to

ambiguity because, Smith-Vaniz et al. (1999, ref. 25013) could

be interpreted as considering reticulatus a synonym of atinga,

or as considering that the other authors misidentified their

Species of porcupinefish

specimens. Some other workers (e.g., Fowler, 1936, ref. 6546;

Lozano Rey, 1952) have included reticulatus of authors in

their synonymies of atringa, but not reticulatus Linnaeus

(1758, ref 2787), implying that they questioned others’ concept

of reticulatus rather than that they considered reticulatus

Linnaeus to be a junior synonym of atringa.

In more recent years, a view has developed among some

American workers that Atlantic individuals of this taxon are

C. atringa, whereas the Indo-Pacific individuals are either C.

affinis (Robins et ah, 1991, ref. 14237) or C. reticulatus (Nelson

et al., 2004, ref. 27807), but, again, no justification for this or

means of distinguishing the two nominal species has ever been

presented. Tyler (1980, ref. 4477) tentatively recognized four

species in this complex that have, based on his material

examined and text, different distributions: C. atringa (western

Atlantic); C. reticulatus (eastern Atlantic and Indo-Pacific); C.

tigrinus (western Indian Ocean); and C. affinis (Eastern

Pacific). However, Tyler (1980, ref. 4477) said that C. tigrinus

may be the young of C. reticulatus (I agree). So confusion

about the identity and distribution of these species continues.

In summary, D. atringa Linnaeus is unidentifiable, and the

post-Linnaean use of the name by various authors has been

inconsistent as to what species was being included: at least

four species and two multi-species groups have been identified

as D. atringa by various authors at various times. In spite of

the use of C. atringa (usually spelled atinga) by several

authors, the name should be regarded as a nomen dubium, and

not used. Diodon reticulatus Linnaeus is clearly identifiable,

and the use of the name has been remarkably consistent: it

should be used for this species.

Diodon echinatus Linnaeus (1758, ref 2787) is seemingly

equivalent to his Chilomycterus reticulatus (see Leis and

Randall, 1982). Linnaeus’ (1758, ref 2787) description and the

Marcgrave plate to which Artedi (1738) referred could apply to

any Chilomycterus or Cyclichthys species. Linnaeus (1766, ref.

2786) referred to a Seba (1759, ref. 18716) figure that is clearly

Diodon hystrix. Gronow (1854, ref. 6828), in his account of

''Holocanthus echinatus'\ cited a Seba (1759, ref. 18716) figure

that is either Chilomycterus reticulatus or C. antennatus, and a

Willughby (1686) figure that clearly represents C. reticulatus.

The holotype of Diodon tigrinus Cuvier (1818, ref. 18059)

is a specimen in the pelagic colour phase of C. reticulatus. The

species was recognized as a synonym of C. reticulatus as long

ago as Brisout de Barneville (1846, ref. 296).

Chilomycterus affinis Gunther (1870, ref. 1995) was based

on a specimen of unknown locality that is dried and thickly

varnished. The holotype has minimal spotting on the body, and

the spines, particularly on the head, are distorted by the

taxidermy and insertion of large, blue glass eyes. However,

there is nothing outside of the range of C. reticulatus variability

in this specimen. In the absence of any locality information, it

is unclear why most authors regarded this as a Pacific species.

Chilomycterus californiensis was described by Eigenmann

(1891, ref. 18744) on the basis of a specimen that he initially

did not obtain from the fisherman who captured it “on account

of the unreasonable price asked for it”. However, the fish was

subsequently “procured by the National Museum”, and

Eigenmann (1892) redescribed and figured it. Therefore,

USNM 43860 is in fact the holotype, in spite of Eigenmann’s

statement in the original 1891 description that “I did not obtain

it”. The holotype is in the pelagic colour phase of C.

reticulatus.

Euchilomycterus quadradicatus Waite (1900, ref. 4558)

from Lord Howe I. was based on a dried specimen - apparently

a beach wash-up - subsequently preserved in ethanol and in

poor condition. Although not figured by Waite, Whitley (1952)

illustrated the holotype (with some artistic license) clearly

showing the caudal-peduncle spine and four-rooted spines on

the head that in combination are diagnostic of Chilomycterus

reticulatus.

Chilomycterus lissogenys Gunther (1910, ref. 14460) was

based on an illustration by Garrett of a Hawaiian fish. Although

Garrett omitted some of the spines on the side of the head, he

clearly showed the spine on the caudal peduncle that is

characteristic of C. reticulatus. The illustration showed relatively

few spots on the body, but heavily spotted fins, a condition well

within the range of colour variation in this species.

The description and photo of Chilomycterus galapagosensis

Klausewitz (1958, ref. 12080) are clearly that of C. reticulatus.

The description of the nostrils alone is diagnostic. Klausewitz

distinguished his new species from C. atringa, which he

described as having a supraorbital cirrus and large dorsal

blotches (presumably based on the description of Jordan and

Evermann [1898, ref. 2244], which was based on more than

one species), by its lack of these two characteristics. He

distinguished it from C. californiensis by colour, but the latter

is in pelagic-phase colour, whereas C. galapagosensis has

typical, spotted demersal colour.

Distribution. Circumglobal in warm temperate to tropical waters:

W Atlantic - 39°N to 24°S

E Atlantic - Madeira (and possibly to Portugal) and Cape

Blanco to Angola

W Indian Ocean - South Africa to Tanzania and Reunion,

Seychelles and Mauritius

E Indian Ocean - Broome, Western Australia to Bali and

Timor

W and central Pacific - Japan to Lord Howe I. and northern

New Zealand, to Tuamotos to Hawaii (and in the east Pacific

barrier)

E Pacific - San Pedro, California to Chile, Galapagos and

Revillagigedos

Occurrences of this species are patchy, and many are based

on pelagic juveniles: in particular, adults are unknown from

broad areas of the Indo-Pacific. Pelagic juveniles are frequently

found poleward of the distribution of adults in areas of strong,

poleward currents.

If future work indicates that C. reticulatus contains more

than one geographically distinct species, several names are

available for Indo-Pacific populations, but no name is clearly

based on Atlantic material. Most of the extant types are either

dried or fixed in alcohol, so it may be possible to obtain genetic

data from them that could be helpful. Unfortunately, there are

no Linnaean types that might assist in this regard, and

Linnaeus’ usage of ‘habitat in India’ cannot be taken at face

value in most cases.

Jeffrey M Leis

Cyclichthys Kaup, 1855

Cyclichthys Kaup, 1855 (type species Diodon orbicularis Bloch)

Diagnosis. All but 1 or 2 spines fixed; all spines with 3 bases,

except in C. spilostylus which has some spines on top of head

with 4 bases; 9 C rays; 19-20 vertebrae; no tentacles in adults;

nostril in adult a short tube with 2 openings; no spines wholly on

dorsal surface of caudal peduncle; no fins spotted; no large

blotches on dorsal surface. Some additional osteological characters

are given by Tyler (1980, ref. 4477) for C. orbicularis.

The type species of this genus is C. orbicularis (Bloch,

1785, ref. 21381). Kaup (1855, ref. 2571) included two species

in his Cyclichthys - orbicularis Bloch, and cornutus Kaup - but

designated neither as type species for the genus. Subsequently,

Bleeker (1865, ref. 416) was apparently the first to designate a

type species for Cyclichthys and chose orbicularis Bloch

(Eschmeyer, 2005). Eras er-B runner (1943, ref. 1495) used

Cyclichthys as a subgenus of Chilomycterus . He did not

consider C. hardenbergi, but included C. orbicularis, the

“Atlantic Chilomycterus” species, and C. echinatus non-

Linnaeus (= C. spilostylus) in his concept of the subgenus

Cyclichthys. Tyler (1980, ref. 4477) pointed out that C.

orbicularis had osteological differences from the other

diodontids he studied, and placed the species in a group on its

own. However, he was not able to examine specimens of C.

spilostylus or C. hardenbergi. Based on external morphology,

it appears that C. orbicularis differs from other species that

have been included in Cyclichthys by Eras er-B runner (1943,

ref. 1495), and there is merit in Tyler’s placement. If this were

done, then a new genus would probably have to be described

for hardenbergi and spilostylus, as they do not appear to be

monophyletic with the “Atlantic Chilomycterus” species.

Pending a cladistic analysis of relationships in the family, I

recognize three species in Cyclichthys, which has been standard

practice in recent years.

Cyclichthys orbicularis (Bloch, 1785)

Diodon orbicularis Bloch, 1785: 73, pi. 127 (Jamaica?, Cape of

Good Hope & Moluccas)*?

Diodon caeruleus Quoy and Gaimard, 1824: 201, pi. 65 (fig. 5)

(North of New Guinea, 142°E, at the Equator)*

Chilomycterus parcomaculatus von Bonde, 1923: 38, pi. 9 (fig. 2)

(Natal, South Africa)

* extant type

Based on examination of 88 lots from throughout the range,

including the extant types. There has been no real question as

to the identity of this wide-spread and common species.

Although Bloch’s (1785, ref. 21381) types might all be lost,

his plate showing the arrangement of spines, especially those

near the mouth, is diagnostic of this species. There is one specimen

of unknown origin in ZMB that may be a syntype of this species

(Paepke, 1999, ref. 24282), but definitive evidence is lacking.

The alleged type locality of Jamaica appears to be in error.

The holotype of Diodon caeruleus Quoy and Gaimard

(1824, ref. 3574) was described and figured, and the specimen

is extant (see Leis and Bauchot, 1984, ref. 12539 ), leaving no

doubt that it is conspecific with C. orbicularis (Bloch).

Chilomycterus parcomaculatus von Bonde (1923, ref. 521)

was based on a specimen that was ‘inadvertently destroyed’

(S.X. Kannemeyer, personal communication, 1/2/80), but the

description and figure are diagnostic.

Distribution. Indo-west Pacific

W Indian Ocean - Capetown, South Africa to Red Sea,

Oman and Persian Gulf, Maldives, Reunion.

E Indian Ocean - Shark Bay, Western Australia to Burma

W Pacific - southern Japan and Sea of Japan to Sydney,

Australia and east to Philippines and New Caledonia.

Cyclichthys hardenbergi (de Beaufort, 1939)

Chilomycterus hardenbergi de Beaufort, 1939: 33-34 (New

Guinea)*

*extant type

Based on examination of 21 museum specimens from throughout

the limited range, including the holotype. There are no real

questions as to the identity of this species: de Beaufort’s (1939,

ref. 17230) description is diagnostic, and the type is extant. This

species has one of the more limited ranges within the family.

Distribution. Indo-Pacific

North-western Australia to the west coast of Cape York,

and the south coast of New Guinea. Kailola (1975) also

recorded it from the Trobirand Islands.

Cyclichthys spilostylus (Leis and Randall, 1982)

Chilomycterus spilostylus Leis and Randall, 1982: 363, figs 1, 2

(Red Sea)*

* extant types

Based on examination of 23 museum specimens, including the

types. This species was mis-identified as Cyclichthys echinatus

(Linnaeus, 1758, ref. 2787) by some authors (see Leis and

Randall, 1982, ref. 8453 ), but echinatus is most likely a synonym

of Chilomycterus reticulatus (Linnaeus). Other than this, there

are no real questions as to the identity of this wide-ranging

species. The pelagic stage of this species has a tentacle emerging

from each spine. These tentacles are lost at settlement.

Distribution. Indo-Pacific

W Indian Ocean - Capetown, South Africa to Gulf of Elat,

Red Sea, Muscat to western India and Mauritius (also a

Mediterranean record from Israel by Golani (1993), presumably

via Suez Canal)

E Indian Ocean - Northwest Cape, Western Australia to

Bali

W Pacific - Southern Japan to Hong Kong, Philippines,

New Caledonia, and northern Great Barrier Reef.

E Pacific - Galapagos (single record, including photograph,

by Humann [1997], repeated by Grove and Lavenberg [1997],

ref. 24023).

“Atlantic Chilomycterus^’

Lyosphaera Evermann and Kendall, 1898: 131 (type species

Lyospharea globosa Evermann and Kendall, possibly = Diodon

schoepfii Walbaum, 1792)

Species of porcupinefish

Atinga Le Danois, 1954: 2356. (type species Diodon atringa

Linnaeus - see Eschmeyer, 2005, and below).

Diagnosis. All spines fixed; all spines with 3 bases; 9 C rays;

19-20 vertebrae; tentacles present on lower jaw and usually

over eye; nostril in adult a short tube with 2 openings; no spines

wholly on dorsal surface of caudal peduncle; no fins spotted

(except in large C. antennatus)\ large blotches present on dorsal

surface. Some additional osteological characters are given by

Tyler (1980, ref. 4477). At least 2 of the species of this group

share the 'Dyosphaera'' larval stage (antennatus and the type

species of Lyosphaera), and others may do the same.

Although the genus Lyosphaera Evermann and Kendall

(1898, ref. 1281) is available as a generic name for this group, I

recommend against its use until a cladistic analysis of the

‘Atlantic Chilomycterus” species and their relationship to other

diodontids is undertaken. Lyosphaera has never been used in

this way, and the identity of the type species is unclear (although

most likely to be schoepfii based on distribution). The species

upon which Atinga Le Danois (1954, ref. 6451) is based is

unclear. The type species, D. atringa Linnaeus (1758, ref. 2787),

is not identifiable (see above under C. reticulatus). It is clear that

Le Danois’ (1954, ref. 6451; 1959, ref. 12003; 1962, 21440)

concept of atinga included a species of the “Atlantic

Chilomycterus'' group, although which species is unclear as her

illustrations of Atinga atinga atinga in the 1954 paper are of C.

antillarum (identified as male) and C. spinosus mauretanicus

(identified as female). In view of this confusion about the identity

of the type species, use of Atinga Le Danois (1954: ref. 6451) is

not recommended. It has been little used since its description.

The “Atlantic Chilomycterus'' is a group of similar species

previously recognized in various ways by Gunther (1870, ref.

1995), Le Danois (1959, ref. 12003) and Tyler (1980, ref. 4477).

See above regarding the generic status of these species.

Chilomycterus antennatus is the only member of this group

that I can separate on morphological grounds; principally, the

development of the fleshy tentacles over the eye. It also has a

colour pattern that differs more from the other species of the

“Atlantic Chilomycterus'' group than they do from each other.

The other four taxa differ only in colour, and have largely non-

overlapping distributions. In all but the case of the very similar

forms, C. spinosus (Linnaeus, 1758, ref. 2787) and C.

mauretanicus (Le Danois, 1954, ref. 6451), the distributions

do at least seem to come into contact. In contrast, the latter

two taxa occur only on opposite sides of the Atlantic and they

have only very minor differences in colouration. Hence, I treat

these two as subspecies: Chilomycterus spinosis spinosus and

Chilomycterus spinosus mauretanicus. In some cases, colour

patterns do exhibit intermediacy. Chilomycterus schoepfii

adults have a distinctive lined pigment pattern, but the youngest

C. schoepfii have a colour pattern not unlike that of C. spinosus

(dark background with lighter, diffuse spotting), and at

intermediate sizes, the dark background may have shrunk to a

mesh-like pattern with expanded lighter centres similar to that

of C. antillarum. Similarly, in northern South America, a

colour pattern with elements of both C. spinosus and C.

antillarum is present. Examination of the genetics of these

“Atlantic Chilomycterus" species would be very interesting.

Chilomycterus spinosus spinosus (Linnaeus, 1758)

Diodon spinosus Linnaeus, 1758: 335 (India)

Diodon orbe Lacepede, 1798: 124, pi. 3 (Rio de Janeiro)

Cyclichthys cornutus Kaup, 1855: 231? (unknown locality)*

Tetrodon torosus Larranaga, 1923: 390? (Uruguay)

*extant type

Based on examination of 24 lots (50-200 mm) from throughout

the range, and the extant type of C. cornutus Kaup (BMNH

1849.1.15.36).

Linnaeus (1758, ref. 2787) based his description of Diodon

spinosus on Artedi (1738), who cited an illustration by

Willughby (1686). This information is sufficient to determine

that the species is one of the “Atlantic Chilomycterus" species,

but without any lines or small spots on the body. This eliminates

schoepfii, antennatus, antillarum and mauretanicus, leaving

spinosus as the unlined, unspotted species of this group.

Lacepede (1798, ref. 2708) provided a figure of D. orbe

that clearly shows the arrangement of spines and the diagnostic

dorsal blotches and lack of small spots or lines on the body.

This and the type locality leave no doubt that Diodon orbe is

conspecific with C. spinosus (Linnaeus).

The type of Cyclichthys cornutus Kaup (1855, ref. 2571) is

a small, stuffed specimen of unknown origin with a thick coat

of varnish, but the spine arrangement and presence of a

supraocular tentacle show that it is clearly a species of the

“Atlantic Chilomycterus" group. The visible colour pattern

best fits C. spinosus (Linnaeus).

Tetrodon torosus Larranaga (1923 : 390 ref. 22561: not seen

by me) has been regarded as a synonym of Chilomycterus spinosus

(Linnaeus) since 1925 (Devincenzi, 1925, ref. 20322, see

Eschmeyer, 2005) and I am unaware of any subsequent use of the

name. If the synonymy of Devincenzi is correct, Uruguay would

represent the southernmost record of C. spinosus spinosus.

As noted above, eastern Atlantic specimens of C. spinosus

have oblique, irregular lines laterally on the trunk and head

that are lacking in western Atlantic specimens. Therefore, I

have recognized the western Atlantic population as the

nominate subspecies and the eastern Atlantic population as C.

spinosus mauretanicus (Le Danois) (see below).

Distribution. Western Atlantic

Erom northern coast of South America (Surinam and

British Guiana) to Rio de Janeiro, Brazil.

Chilomycterus spinosus mauretanicus (Le Danois, 1954)

Atinga atinga mauretanicus Le Danois, 1954: 2354 (Mauritania,

Gulf of Guinea)*

* extant types

Based on examination of 21 lots from throughout the range,

including the syntypes. This nominal species is considered a

subspecies of C. spinosus (Linnaeus) because only minor

colour differences separate it from its western Atlantic

counterpart. Le Danois (1954, ref. 6451) briefly described this

nominal species, apparently inadvertently, in a paper on sexual

dimorphism in diodontids, then redescribed it in 1959 (ref.

12003) and provided more information - some of it conflicting

- in 1962 (ref. 21440 ). See Leis and Bauchot (1984, ref. 12539)

for information on the status of the types.

Jeffrey M Leis

Distribution. Eastern Atlantic.

From central Angola to Canary Is. and perhaps Portugal.

Chilomycterus schoepfii (Walbaum, 1792)

Diodon schoepfii Walbaum, 1792: 601 (New York)

Diodon meulenii Walbaum, 1792: 602 (unknown locality)

Diodon geometricus var. lineatus Bloch and Schneider, 1801: 513

(New York)

Diodon maculato-striatus Mitchill, 1815: 470 (New York)

Diodon rivulatus Cuvier, 1818: 129, pi. 6 (unknown locality [New

York, USA according to Eschmeyer, 2005])*

Diodon nigrolineatus Ayres, 1842: 68 (Brookhaven, Long Island,

New York)

Diodon fuliginosus dcKay, 1842: 324, pi. 55 (fig. 181) (New York)

Diodon verrucosus dcKay (ex Mitchill), 1842: 325, pi. 55 (fig. 1)?

(New York)

Holocanthus areolatus Gronow in Gray, 1854: 27? (Cape of Good

Hope, South Africa?)

Chilomycterus pentodon Atkinson in Bryant, 1888: 18 (Beaufort,

North Carolina, USA)

* extant type.

Based on examination of 62 lots from throughout the range.

Unfortunately, I could locate types of only one of the ten nominal

species represented here.

Walbaum’s (1792, ref. 4572) description of D. schoepfii

mentions the diagnostic lined colour pattern of this species, as do

the descriptions of Diodon meulenii Walbaum (1792, ref. 4572),

Diodon geometricus var lineatus Bloch and Schneider (1801, ref.

471), Diodon maculato-striatus Mitchill (1815, ref. 13292),

Diodon rivulatus Cuvier (1818, ref. 18059), Diodon nigrolineatus

Ayres (1842, ref. 15926), Diodon fuliginosus deKay (1842, ref.

1098), Holocanthus areolatus Gronow in Gray (1854, ref. 6828)

and Chilomycterus pentodon Atkinson in Bryant (1888, ref.

13034), thus confirming their identification. The type locality of

South Africa for H. areolatus provided by Gronow introduces

some doubt, but this may well be an error, as there is no diodontid

species with a lined colour pattern in that area. Diodon

verrucosus deKay (ex Mitchill) (1842, ref. 1098) has a pigment

pattern similar to that of C. antillarum, but, apparently, C.

schoepfii passes through an early life-history phase with this

colour pattern, and the type locality of New York would seem to

eliminate the tropical C. antillarum, so I tentatively consider

verrucosus to be a synonym of C. schoepfii.

Distribution. Western North Atlantic

From Halifax, Nova Scotia (waif) to Belize (apparently

with a gap between southern Texas and Belize) on the mainland

and Cuba, Bermuda and Bahamas.

Chilomycterus antennatus (Cuvier, 1816)

Diodon antennatus Cuvier, 1816: 185, pi. 9 (unknown locality)*?

Chilomycterus briareos Metzelaar, 1919: 173, fig. 55 (Lesser

Antilles, St Eustatius)*

Lyosphaera digitalis Breder, 1927: 81, fig. 34 (locality unknown

Western North Atlantic or W Indies)*

*extant type.

Based on examination of 37 lots, including extant types (see

Leis and Bauchot, 1984, ref. 12539, for a discussion of the status

of the types of D. antennatus Cuvier [1816, ref. 993]). In spite of

assertions to the contrary, C. antennatus can have spotted fins.

Fin spotting in C. antennatus begins basally on all fins at about

50 mm SL. The caudal fin becomes mostly or entirely spotted

by 100-150 mm SL. Spotting on other fins seems variable, but

basal one -third to one -half of the P, D and A fins can be spotted

in specimens as small as 127 mm SL, whereas other specimens

as large as 200 mm may have spots only on the extreme base on

these fins. Because many ichthyologists have assumed that any

Chilomycterus with spotted fins is C. reticulatus (or one of its

synonyms), this has led to many misidentifications of C.

antennatus, and is probably the basis for Jordan and Evermann’s

(1998, ref. 2244) inclusion of what are apparently characteristics

of C. antennatus in their description of C. atringa.

Aside from colour differences, C. antennatus has larger

fleshy tentacles, particularly over the eye, than do the other

‘Atlantic Chilomycterus"" species. It clearly has a Lyosphaera

stage larva (Heck and Weinstein, 1978).

Cuvier’s (1816, ref. 993) description and figure were

diagnostic of the species, and what is probably the type is

extant in MNHN (see Leis and Bauchot, 1984, ref. 12539),

leaving no doubt about the identity of this distinctive species.

The description and figure of Chilomycterus briareos

Metzelaar (1919 ref. 2982) clearly refers to C antennatus, and

the type is extant. The fish has spots on the fins, particularly

on the caudal fin, which is common in larger individuals of C.

antennatus.

In contrast, Lyosphaera digitalis Breder (1927, ref. 635), is

the young "Lyosphaera stage’ of this species, virtually lacking

spines. Heck and Weinstein (1978) have documented the

transition of this distinctive "Lyosphaera stage’ to the juvenile

of C. antennatus.

Distribution. Western Atlantic (possibly eastern Atlantic).

W Atlantic - Key West Florida to Panama, Colombia and

Tobago, Bermuda, and throughout Caribbean and Antilles.

E Atlantic - no specimens, but see below.

There are persistent reports of C. antennatus from the

eastern Atlantic, but I have seen no specimens from this area.

Where published descriptions or illustrations of ""Chilomycterus

antennatus"" from the eastern Atlantic are diagnostic, they are

usually of C. spinosus mauretanicus , or in some cases C.

reticulatus. However, there is one published illustration of a

fish from Senegal that does appear to be C. antennatus, although

it is identified in the publication as C. atringa (Linnaeus) (Seret

and Opic, 1981). Unfortunately, the specimen was not retained

(B. Seret, personal communication). When I requested a

specimen for study, Seret, who was not in Senegal at the time,

kindly arranged for a colleague to send me one: it was C.

spinosus mauretanicus. Therefore, it is possible that C.

antennatus does occur rarely in the eastern Atlantic, most likely

as a waif from the west. Specimens are needed to confirm this.

Chilomycterus antillarum Jordan and Rutter, 1897

Diodon geometricus Bloch and Schneider, 1801: 513, pi. 96 (coast

of Brazil)

Chilomycterus antillarumiorddLndLud'RvhXex, 1897: 131 (Kingston,

Jamaica)*

Species of porcupinefish

* extant type

Based on examination of 41 specimens including extant types.

Although D. geometricus Bloch and Schneider (1801, ref. 471)

is an older name than C. antillarum Jordan and Rutter (1897,

ref. 10644), and the Bloch and Schneider figure clearly applies

to the same species, the name has been little used since its

description other than as a junior synonym of either spinosus or

schoepfii. Other than Paepke’s (1999, ref. 24282) catalogue of

Bloch types (unfortunately, the type of geometricus is lost), the

most recent correct use of geometricus was Gunther (1870,

ref.1995), and this for only one of his ‘varieties’ (i.e., beta). In

contrast, C. antillarum has been widely, almost universally,

used for this species (see Eschmeyer, 2005, for 13 publications

between 1983 and 2003; in addition, Bailey et al., 1960, ref.

27285; 1970, ref. 27286; Bdhlke and Chaplin, 1968, ref. 23150;

Randall, 1968; 1996; Tyler, 1977; 1980, ref.4477; Robins et al.,

1980, ref. 7111; 1991, ref. 14237; Lieske and Myers, 1994;

Cervigon, 1996, ref. 24489; Smith, 1997; Lyczkowski-Schultz,

et al., 2005). Because the senior synonym {geometricus) has

not been used as a valid name after 1899, and because the junior

name has been used in at least 25 works published by at least

ten authors over the last 50 years, this meets the criteria of

Articles 23.9.1 and 2 of ICZN (1999, ref. 26875), and prevailing

usage (of antillarum) must be maintained. In 2003, 1 suggested

that current usage of C. antillarum be maintained in the

interests of stability (Leis, 2003, ref. 27121), and here provide

evidence that ICZN criteria require this to be met.

Chilomycterus orbitosus Poey (1875: 69, ref. 18564) is

clearly a species of the “Atlantic Chilomycterus"" group, but

there is no known extant type. Poey’s brief description on

Cuban specimens seems to be based on a composite of C.

antillarum and C. schoepfii from Cuba, but perhaps best fits

the former. The name orbitosus has not been used since its

description, as far as I can ascertain. So, even if it could be

established that orbitosus Poey, 1875 and antillarum Jordan

and Rutter, 1897 are conspecific, I would recommend against

the use of the older orbitosus in the interests of stability.

Distribution. W Atlantic.

Florida, Bahamas and Cuba to Venezuela, Barbados and

Brazil

Some Brazilian specimens are intermediate in colour

between the crisp, dark hexagonal pattern typical of C.

antillarum and the dark background with vague lighter spots

typical of C. spinosus spinosus. The significance of this is

unknown, and further investigation is required.

Tragulichthys Whitley, 1931

Tragulichthys Whitley, 1931 (type species D. jaculiferus Cuvier)

Diagnosis. All spines fixed, except those in pectoral axil which

are by far the longest on the body; all spines except those in the

P axil with 3 bases; spines long to medium; 9 C rays; 19

vertebrae; no tentacles in adults; nostril in adult a short tube

with 2 openings, but may become bifurcate in larger individuals;

no spines wholly on dorsal surface of caudal peduncle, but

large spines extend over the peduncle nearly to the caudal-fin

base; no fins spotted; no large blotches on dorsal surface. Some

additional osteological characters are given by Tyler (1980, ref.

4477) as Diodon jaculiferus.

The type species of this monotypic genus is D. jaculiferus

Cuvier (1818, ref. 18059). Most of the spines are fixed in the

normal ‘burrfish’ manner, but those in the pectoral axil, which

are by far the longest on the body, are erectile. Some have

regarded Tragulichthys Whitley (1931, ref. 4673) as a synonym

or subgenus of Diodon (Fraser-Brunner, 1943, ref. 1495; Tyler,

1980, ref. 4477). But, until a full analysis of the phylogeny of the

family is forthcoming, it seems best to maintain current usage

and to recognize Tragulichthys at the generic level because the

only species has a number of morphological differences from the

five species normally included in Diodon.

Tragulichthys jaculiferus (Cuvier, 1818)

Diodon jaculiferus Cuvier, 1818: 130, pi. 7 (‘Indian Ocean via

Peron’)*

Chilomycterus grandoculis Ogilby, 1910; 19 (Moreton Bay,

Queensland)*

*extant type

Based on examination of 50 lots, including extant types, from

throughout the range. Aside from confusion regarding the

designation of a type species for Allomycterus (see below),

there have been few nomenclatural issues regarding this tropical

Australian species. References to this species from New

Zealand are of Allomycterus pilatus (see below): T. jaculiferus

does not occur in New Zealand.

Cuvier’s (1818, ref. 18059) description, figure, and the

extant type leave no doubt about the identity of this distinctive

species. The description of Chilomycterus grandoculis Ogilby

(1910, ref. 3288) details the diagnostic spination, and the extant

type makes it clear that it is conspecific with T. jaculiferus.

Distribution. Northern Australia.

From Derby, Western Australia to Darwin (including

Rowley Shoals) to Torres Strait and south to Moreton Bay,

Qld.

Dicotylichthys Kaup, 1855

Dicotlylichthys Kaup, 1855 (type species Dicotylichthys

punctulatus Kaup)

Atopomycterus Bleeker (ex Verreaux), 1865 [type species

Atopomycterus diversispinus Bleeker (ex Verreaux)]

Diagnosis. Spines on head and belly erectile, those on back and

sides fixed; fixed spines with 3 bases, erectile spines with 2

bases; spines long to medium; 9 C rays; 21 vertebrae; no tentacles;

nostril in adult bifid; no spines wholly on dorsal surface of caudal

peduncle, but large spines extend over the peduncle nearly to the

caudal-fin base; no fins spotted; no large blotches on dorsal

surface, but lateral bars present. Some additional osteological

characters are given by Tyler (1980, ref. 4477).

The type species of Dicotylichthys is D. punctulatus Kaup

(1855, ref. 2571). The sole species in this genus has erectile

spines on the head and belly, but fixed ones on the back and

sides. In contrast to the arrangement adopted here, some authors

follow Fraser-Brunner (1943, ref. 1495) and include all

diodontids that develop bifid nasal organs in Dicotylichthys.

Jeffrey M Leis

This would place in the same genus such disparate species as

pilatus with all fixed spines, punctulatus with a mixture of

erectile and fixed spines, and nicthemerus with all erectile

spines. However, Dicotylichthys is very similar to the monotypic

Lophodiodon (see below). Bleeker’s (1865, ref. 416) description

of Atopomycterus (based on an unpublished manuscript by

Verreaux held in MNHN) is brief, but fortunately the types of

A. diversispinus Bleeker are extant (see below), thus clearly

showing that Atopomycterus is a synonym of Dicotylichthys.

Dicotylichthys punctulatus Kaup, 1855

Dicotylichthys punctulatus Kaup, 1855: 230 (Cape of Good Hope,

South Africa and Mauritius, but these localities are apparently

incorrect, see below)*

Atopomycterus diversispinis Bleeker (ex Verreaux), 1865: 49

(Australia)*

Dicotylichthys myersi Ogilby, 1910: 18 (Moreton Bay, Queensland,

Australia)*

* extant type

Based on over 50 lots from throughout the range, including all

extant types. Kaup’s (1855, ref. 2571) description is not detailed,

but the extant specimens upon which he based his description

are all of this distinctive species.

Although Bleeker’s 1865 description of Atopomycterus

diversispinis is brief, and not detailed, the syntypes are extant

and readily identified as D. punctulatus Kaup (1855, ref. 2571;

see Leis and Bauchot, 1984, ref. 12539).

Dicotylichthys myersi Ogilby (1910 ref. 3288) was said by

Ogilby to differ from D. punctulatus by the relative size of the

abdominal spines, but the syntypes are well within the range

of relative spine size of D. punctulatus.

Distribution. South-eastern Australia.

From Moreton Bay, Qld to Bass Strait.

Kaup (1855, ref. 2571) reported that his type specimens came

from the Cape of Good Hope and Mauritius. Subsequently,

Gunther (1870, ref. 1995) reported that the only specimen in

BMNH identified as being from Mauritius was of questionable

locality, and that the sole specimen from the Cape of Good Hope

(which he identified as the ‘type’ of D. punctulatus) was

“presented by Sir A. Smith”. Smith was a medical doctor resident

in Cape Town who procured many specimens from passing

ships, and then provided them to the British Museum, where they

were generally assumed to have originated in Cape Town (Bass

et ah, 1975, ref. 7409). Thus, there is good reason to question the

locality data of Smith specimens (Bass et ah, 1975, ref. 7409) if

other evidence is inconsistent with them. Other than this BMNH

specimen, I have been unable to find any institution (including

RUSI) that has specimens of D. punctulatus from anywhere but

Australia, where it is abundant within its range. Thus, I conclude

that Sir A. Smith procured his specimen from a passing ship, not

from the Cape of Good Hope, and that this species is endemic to

south-eastern Australia.

Allomycterus McCulloch, 1921

Allomycterus McCulloch, 1921 (type species Allomycterus

jaculiferus [non-Cuvier] McCulloch = Allomycterus pilatus Whitley)

Diagnosis. All spines fixed except 1 or 2 in P axil; spines with 3

bases except erectile ones; spines long or short; 9 C rays; no

tentacles in adults; nostril in adult bifid; no spines wholly on

dorsal surface of caudal peduncle, but large spines that extend

over the peduncle nearly to the caudal-fin base; no fins spotted; no

large blotches on dorsal surface, but lateral bars may be present.

The type species of this genus is A. jaculiferus (non-

Cuvier) McCulloch. McCulloch (1921, ref. 2945), apparently

following Gunther’s (1870, ref. 1995) concept of jaculiferus,

provided an excellent illustration of the species he was

proposing as the type of his new genus. However, it was not D.

jaculiferus of Cuvier (1818, ref. 18059). Whitley (1931, ref.

4673) realized this, and also realized that McCulloch’s fish

was undescribed. Whitley therefore described as new

Allomycterus pilatus, and designated as his holotype the

specimen illustrated and described by McCulloch (1921). Not

surprisingly, this has caused some confusion.

Allomycterus pilatus Whitley, 1931

Allomycterus jaculiferus (non-Cuvier) McCulloch, 1921: 141, pi.

23 (fig. 2) (New South Wales, Australia)

Allomycterus pilatus Whitley, 1931: 125 (NSW, Australia)*

Allomycterus whitleyi Phillipps, 1932: 13, fig. 5 (New Zealand)*

*extant type

Based on 38 lots from throughout the range, including the

extant types. Confusion over the specific name of this species

is dealt with under the genus. There seem to be two forms of

this species, one with long, blade-like spines (A. whitleyi

form), and another with short, compressed spines. Both forms

occur off the Australian mainland, but I have seen only the

long-spine form from New Zealand, and the specimens with

the longest spines seem to be from New Zealand. These

differences are not obviously connected with sexual

dimorphism. Therefore, there may be two species of

Allomycterus, and a genetic study would be useful in clarifying

the situation. In addition, Kuiter (1993, ref. 23929) illustrates

two colour morphs among south-eastern Australian specimens

of A. pilatus, referring to deep-water and shallow-water forms.

The basis for the colour differences is unclear and should be

investigated. References to Allomycterus jaculiferus from

New Zealand are based on A. pilatus (see discussion under

Allomycterus).

Whitley’s (1931, ref. 4673) description and McCulloch’s

(1921, ref. 2945) illustration are clear, and could apply to no

other species. In addition, the holotype is extant.

Phillipps’ (1932, ref. 16393) A. whitleyi constitutes the long-

spined form from New Zealand, and although both holotype and

paratype are stuffed and distorted, they appear to differ from A.

pilatus only in the length and shape of the spines. Phillipps’

description contains two spellings of the specific name: two as

whitleyi and one as whiteleyi. Given the correct spelling of

Gilbert Whitley’s name (to whom the patronym refers), ''whiteleyi

is clearly a typographical error even though it appears before the

two uses of whitleyi within Phillips’ article.

Distribution. Southern Australia and New Zealand.

Rottnest I., WA, to Botany Bay, NSW, including Tas.;

Tasman Sea seamounts and ridges; and New Zealand.

Species of porcupinefish

Lophodiodon Fraser-Brunner, 1943

Lophodiodon Fraser-Brunner (type species Diodon calori

Bianconij

Diagnosis. Spines on head and belly erectile, those on back and

sides fixed; fixed spines with 3 bases, erectile spines with 2

bases; spines short to medium; anteriorly-pointing spines on

snout; 9 C rays; a small supraorbital tentacle in adults; nostril

in adult a short tube with 2 openings; no spines wholly on

dorsal surface of caudal peduncle, but large spines extend over

the peduncle nearly to the caudal-fin base; no fins spotted; no

large blotches on dorsal surface, but bars present laterally.

The type species of this genus is D. calori Bianconi (1854,

ref. 17949). The sole species in this genus has most spines on

head and belly erectile, and those on back and sides fixed. This

genus is similar in many ways to Dicotylichthys, differing

primarily in that the nasal organ in Dicotylichthys is bifid,

whereas in Lophodiodon, it is a hollow tube with two distinct

nostrils. There is reason to expect that two genera may

eventually be considered to be synonymous.

Lophodiodon calori (Bianconi, 1854)

Diodon calori Bianconi, 1854: 69 (Mozambique)

Lophodiodon nigropunctatus Smith, 1957: 222, fig. 4 (Port

Alfred, South Africa)*

*extant type

Based on 13 specimens from most of the range, including the

extant types. This species is widely distributed, but uncommon

in collections.

Although the name Diodon calori dates from Bianconi,

1854 (ref. 17949), the illustration of Diodon calori in Bianconi

(1855, ref. 295) is diagnostic for this species, with its large

number of short spines, four lateral bars and no dorsal

blotches.

Lophodiodon nigropunctatus Smith (1957, ref. 12171) was

based on juveniles (30-60 mm SL), and the apparent difference

in colour pattern with L. calori can be attributed to this. The

spination of Smith’s specimens is diagnostic.

Distribution. Indo-Pacific.

The 13 specimens I have examined are all from east Africa

and Seychelles, but the species is reliably reported from Oman,

Bali, Timor, the Australian Northwest Shelf, the South China

Sea and New Caledonia, and somewhat less reliably as the

similar D. punctulatus from New Guinea by Tortonese (1964,

ref. 9080) and Munro (1967, ref. 6844).

Diodon Linnaeus, 1758

This genus was revised by Leis (1978, ref. 5529) with additional

information on nomenclature and types in Leis and Bauchot

(1984, ref. 12539), and information contained there is not

repeated. Only information on Diodon species described since

1978 and on noteworthy new distributional information is

included here. Note that figs 9 and 17 of Leis (1978) were

switched (see 1979 errata facing p. 956, US Fishery Bulletin

76[4] ): fig. 9 labelled Diodon hystrix is actually D. holocanthus

and fig. 17 labelled Diodon holocanthus is actually D. hystrix.

Diodon eydouxii Brisout de Barneville, 1846

Diodon bertolettii de Lema, de Lucena, Saenger and de Oliveira,

1979: 35-38, figs 18-19 (Brazil)*

*extant type

Diodon bertolettii can readily be identified as a synonym of D.

eydouxii Brisout de Barneville based on its semi-lunate fins,

blue colour, fin-ray counts, and from the photographs provided

by de Lema et al. (1979, ref. 8836).

Leis (1978, ref. 5529) examined specimens of D. eydouxii

Brisout de Barneville (1846, ref. 296) from 19 scattered

localities in all warm oceans. I have now seen an additional 31

lots. These plus four acceptable literature records extend the

known distribution of this species.

Distribution. Pelagic, Atlantic, Indian and Pacific Oceans.

W Indian - Cape of Good Hope to Zanzibar

E Indian - only record is Andaman Sea

W Pacific - Indonesia to Okinawa

Central Pacific - from near Samoa to Hawaii

E Pacific - equator to 20°N plus a California record (Lea,

1998: from Los Angeles Harbour, but misidentified as C.

reticulatus [Linnaeus]).

W Atlantic - 28°S to 37°N

E Atlantic - 30°W is eastern-most specimen examined, but

there are apparently valid literature records from the Azores

(Azevedo, 2004) and from Spain (Crespo et al., 1987).

Diodon holocanthus Linnaeus, 1758

Diodon paraholocanthus, Kotthaus, 1979: 39, fig. 492 (Bab-el-

Mandeb, southern Red Sea)*

*extant type

Kotthaus (1979, ref. 8818) confused Diodon liturosus Shaw

with Diodon holocanthus Linnaeus, as is obvious from his

description and photograph (his fig. 491) of what he called D.

holocanthus. Then, having encountered the true D. holocanthus

in the north-western Indian Ocean, he described it as a new

species, D. paraholocanthus. The description and photo (his

fig. 492) of the holotype are entirely consistent with D.

holocanthus Linnaeus.

Leis (1978) examined 141 specimens of D. holocanthus

Linnaeus (1758, ref. 2787) from all warm oceans. I have now

seen more than 100 additional lots that extend the known

distribution of the species (see below). It is noteworthy that

there are still no records of D. holocanthus from the Pacific

Plate other than those reported by Leis (1978): Hawaii, Easter

and Pitcarin Is. Reference in Robertson et al. (2004) and

Mundy (2005, ref. 28379) to D. holocanthus occurring in the

Line Is. is incorrect (B.C. Mundy, personal communication).

Distribution. Circumtropical in Atlantic, Indian and Pacific

Oceans (except only peripherally on Pacific Plate).

W Indian - from Cape of Good Hope, South Africa to

Oman and Red Sea, Sri Lanka, Mascarenes, and Seychelles.

E Indian - Andaman Sea to Australia

W Pacific - west of Pacific Plate: Japan to New Caledonia

and Elizabeth and Middleton Reefs, Tasman Sea.

Jeffrey M Leis

In Australia, south to Ulladulla, NSW (36°S) off east coast,

and to Freemantle, WA (32°S) off west coast.

Central Pacific - Hawaii, Easter and Pitcarin Is. only.

E Pacific - southern California to Colombia

W Atlantic - Hudson Canyon (off New Jersey) to

Argentina.

E Atlantic - Liberia and Nigeria to northern Angola .

Diodon hystrix Linnaeus, 1758

Leis (1978) examined 43 specimens of Diodon hystrix Linnaeus

(1758, ref. 2787) from all warm oceans. I have now seen an

additional 80 lots that extend the documented distribution of

the species (see below).

Distribution. Circumtropical in Atlantic, Indian and Pacific

Oceans

W Indian Ocean - throughout the area from South Africa

(Tsitsikamma Coastal National Park) to the Red Sea, Sri

Lanka, and all major island groups.

Australia - south to Elizabeth and Middleton Reefs, Lord

Howe L, and northern NSW (29°S) off the east coast, and

Rowley Shoals on west coast.

W Pacific - New Caledonia and Kermadecs to Rotuma,

Pitcarin L, Hawaii and southern Japan.

E Pacific - Mexico to Chile

W Atlantic - 36°N to ca 20°S

Central Atlantic - Ascension and St Helena

E Atlantic - only 1 confirmed record at Fernando Po

Diodon liturosus Shaw, 1804

Leis, 1978 examined 30 specimens of Diodon liturosus Shaw

(1804, ref. 4015) primarily from the western Pacific. I have

now seen an additional 45 lots that extend the documented

distribution of the species (see below).

Distribution. Indo-west Pacific

W Indian Ocean - South Africa (Algoa Bay) to Oman and

southern Red Sea, Mascarenes, Seychelles, Laccadives and

Maldives.

E Indian Ocean - Phuket, Thailand to Ningaloo Reef,

W Pacific - from Maizuru, Japan to northern NSW,

Australia to New Caledonia to Society and Marshall Is.

Diodon nicthemerus Cuvier, 1818

Leis (1978) examined nine specimens of Diodon nicthemerus

Cuvier (1818, ref. 18059), all from southern Australia. Museums

in Australia contain large numbers of this species, and its

distribution is confirmed as being confined to the waters of

southern Australia. I have seen specimens from an area ranging

from Houtman Abrolhos Is., WA (28°S), to Nadgee, NSW

(37°S), although Kuiter (1993) reports D. nicthemerus as far

north as Seal Rocks (32°S). This is the most restricted

distribution of any species of Diodon.

Key to genera and species of the family Diodontidae

NB: in juveniles relative spine length and body colour generally

differ from those of adults

1. All body spines erectile and 2-rooted (except a few around

gill opening or dorsal-fin base) Diodon 10

— All or most body spines of back and sides fixed in an erect

position and 3 -rooted 2

L^on-Diodon

2. Indian and Pacific in distribution 3

— Atlantic in distribution

(NB: 1 Indo-Pacific species, Cyclichthys spilostylus, has

penetrated the eastern Mediterranean Sea through the

Suez Canal 14

Indo-Pacific non-Diodon

3. Spines on top of head and on belly erectile 4

— Spines on top of head and on belly fixed in an erect

position 5

4. 2 to 4 spines in the 1st row on the snout point toward the

mouth when not erect; no small, black spots scattered more

or less uniformly over head and trunk

Lophodiodon calori (Indo-west Pacific)

— All erectile spines point toward tail when depressed;

small, black spots scattered more or less uniformly over

head and trunk

Dicotylichthys punctulatus (south-east Australia)

5. A small spine or 2 wholly on the dorsal surface of the

caudal peduncle; normally 10 caudal rays; nasal organ of

adults an open ridged cup; adults with fins spotted

Chilomycterus reticulatus (circumtropical)

— No spines wholly on the caudal peduncle; normally 9

caudal rays; nasal organ of adults a short tube with either

2 openings or split at the end (not an open cup); no spots

on fins of adults 6

6. A set of 4 long fixed spines with their bases near the dorsal

and anal-fin bases - their pointed ends extend over the

caudal peduncle; a few spines in P axil erectile 7

— No especially long spines around dorsal and anal fin

bases; spines in P axil fixed 8

7. Very long spines (longer than rays of pectoral fin) in

pectoral-fin axil; 3-4 black spots (< eye) on sides of head

and trunk, none on back

Tragulichthys jaculiferus (tropical Australia)

— Spines of pectoral-fin axil not particularly elongate; some

eye-size dark spots on back generally associated with

spine bases Allomycterus pilatus

(temperate Australia, Tasman Sea and New Zealand)

8. Few black spots on body, those present at base of spines

dorsally and dorso-laterally; D, A and C fins with dusky

distal margin; only 2 spines over eye; 2 spines between

nostrils, 1 immediately adjacent to each nostril

Cyclichthys hardenbergi

(tropical Australia, southern New Guinea)

— Black spots in clusters dorsally and laterally, or associated

with spine bases laterally and ventrally; D, A and C fins

either clean or with faint, parallel bands; 3 spines over

eye; only 1 spine between nostrils, located medially 9

Species of porcupinefish

9. Spines few, 4 dorsally between pectoral-fin bases, 8 or 9

anterior to dorsal-fin base; a short, moveable spine near

corner of mouth; all spines on top of head with 3 bases;

black spots in clusters dorsally and dorsolaterally

Cyclichthys orbicularis (Indo-west Pacific)

— Spines more numerous, 5 or 6 dorsally between pectoral-

fin bases, 11 or 12 anterior to dorsal-fin base; no moveable

spines; some spines on top of head with 4 bases; black

spots at base of spines laterally and ventrally

Cyclichthys spilostylus (Indo-Pacific)

Diodon

10. None of spines wholly on caudal peduncle; body with

several large, dark dorsal or lateral blotches; no small,

dark spots on fins 11

— One or more small spines wholly on the dorsal surface of

caudal peduncle; body without large dorsal blotches; all fins

(anal sometimes excepted) heavily spotted 13

11. Temperate Australian waters only; no small, fixed, tri-

base spine immediately above gill opening; no small, flat

spines on the anterior border of the depression surrounding

the gill opening; 11 or fewer spines from lower jaw to

anus; adult colour pattern dominated by 4, large, lateral

bars, dorsum uniformly dark

Diodon nicthemerus (southern Australia).

— Tropical waters, with strays into warm temperate water; 1

or 2 small, fixed tri-base spines above gill opening; 3 or 4

small, flat spines forming the anterior border of depression

surrounding the gill opening; 12 or more spines from

lower jaw and anus; adult colour pattern dominated by

several large, dorsal blotches 12

12. Frontal spines obviously much shorter than spines

immediately behind pectoral fin; small downward-

pointing spine below anterior margin of eye; 17-22 spines

from lower jaw to anus; large dorsal blotches with distinct

pale border; blotch below eye not continuing over top of

head Diodon liturosus (Indo-Pacific)

— Frontal spines slightly shorter to much longer than spines

immediately behind the pectoral-fin base; small

downward-pointing spine below anterior margin of eye

absent (Indo-Pacific) or present (most Atlantic specimens);

12-15 spines from lower jaw to anus; dorsal blotches

without distinct pale border; blotch below eye continues

over interorbital in Indo-Pacific specimens

Diodon holocanthus (circumtropical)

13. Pectoral-fin rays 19-22; anal-fin rays 16-18; dorsal and

anal fins somewhat pointed to semilunate in adults;

relatively streamlined, head width of adults 3.3-4.0 in

standard length; 10-14 spines from lower jaw to anus; a

wholly pelagic species coloured dark-blue dorsally

Diodon eydouxii (circumtropical).

— Pectoral-fin rays 22-25 (rarely 21); anal-fin rays 14-16;

dorsal and anal fins rounded in adults; relatively robust,

head width of adults 2.4-3.3 in standard length; 14-19

spines from lower jaw to anus; juveniles (up to 20 cm)

pelagic and coloured blue dorsally, adults demersal and

coloured tan to brown

Diodon hystrix (circumtropical)

Atlantic non-Diodon

14. 1 or 2 small spines wholly on the dorsal surface of the

caudal peduncle; normally 10 caudal-fin rays; nasal organ

of adults, an open, ridged cup; adults with fins spotted; on

top of head some spines with 4 roots

Chilomycterus reticulatus (circumtropical)

— No spines wholly on the caudal peduncle; normally 9

caudal-fin rays; nasal organ of adults, a short hollow tube

with 2 openings; fins of adults usually without spots; all

spines with 3 roots ‘Atlantic Chilomycterus’’" 15

15. A large (ca. = eye diameter) tentacle above eye; colour

pattern dominated by large dorsal blotches and with small

spots scattered on back and sides, on fins only basally,

except on most or all of caudal fin from 10-15 cm standard

length, and on other fins from 20 cm

Chilomycterus antennatus (central-west Atlantic)

— Tentacles above eyes absent or small; no small spots on

fins or on back and sides; dorsal and lateral dark blotches

present 16

16. Network of hexagonal to circular black lines on back and

sides in adults

Chilomycterus antillarum (central-west Atlantic)

— Black lines on back and sides absent in adults, or if present,

wavy or approximately parallel - not intersecting to form

rings or polygons 17

17. Extensive series of dark-brown to black parallel lines

densely covering back and sides in adults

Chilomycterus schoepfii (western North Atlantic)

— No black lines on back; dorsal background dark with diffuse

lighter spots Chilomycterus spinosus 18

18. No black lines on sides of head or trunk; South American

in distribution Chilomycterus spinosus spinosus

(east coast South America)

— Irregular, approximately parallel black lines on sides of

head and trunk; eastern Atlantic in distribution

Chilomycterus spinosus mauretanicus

(west coast Africa)

Acknowledgements

I thank the curators and staff of the many Museums who have

allowed me access to their collections of diodontids over many

years and to many colleagues who provided information on

diodontids. W.N. Eschmeyer, CAS, and J.C. Tyler, USNM,

offered constructive comments on a draft of the manuscript.

S. Bullock provided editorial assistance. This research was

supported, over many years, by the Australian Museum, Sydney.

References

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in Eschmeyer, 2005, and are not repeated here.

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records of Fistularia petimba and Diodon eydouxii for the Azores,

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18(1): 8.

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living marine resources of the Eastern Central Atlantic, LAO,

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Memoirs of Museum Victoria 63(1): 91-96 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East

China Sea and off Western Australia

Tetsuji Nakabo^ Dianne J. Bray^ and Umeyoshi Yamada^

'Kyoto University Museum, Kyoto University, Kyoto 606-8501, Japan (nakabo@inet.museum.kyoto-u.ac.jp)

^Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (dbray@museum.vic.gov.au)

^ Hamada, Togitsu, Nishisonogi, Nagasaki Prefecture 851-2102, Japan

Abstract Nakabo T., Bray D. J. and Yamada U. 2006. A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East

China Sea and off Western Australia. Memoirs of Museum Victoria 63(1): 91-96.

Zenopsis stabilispinosa sp. nov. is described from the South China Sea, East China Sea and off Western Australia. It

differs from other species of Zenopsis in having the third anal-fin spine fused to its pterygiophore, seven dorsal-fin spines,

a short pelvic fin in adults, and 32 (13-1-19) vertebrae. A key to the species of Zenopsis is provided.

Keywords Zeidae, Zenopsis, new species. South China Sea, Western Australia

Introduction

The species of Zenopsis are characterized by having a deep

strongly compressed silvery body, scales absent from the body

except along the lateral line, concave dorsal head profile,

enlarged bucklers along most of the spinous dorsal-fin base and

along the soft dorsal- and anal-fin bases, three anal-fin spines,

six pelvic-fin elements, and narrow infraorbitals. Three species

of Zenopsis are currently recognized: Z. nebulosa Temminck

and Schlegel, 1845, Z. conchifer Lowe, 1852 and Z. oblonga

Parin, 1989. During surveys of demersal fishes in the South

China Sea and off Western Australia, ten specimens of Zenopsis

which differ significantly from these three Zenopsis species

were collected. They are described herein as a new species.

Material and methods

Museum abbreviations follow Leviton et ah, 1985, with the

exception of the following: SNFR - Seikai National Fisheries

Research Institute. Counts and measurements (Tables 1 and 2)

follow Nakabo, 2002a. Vertebral numbers and anal-fin

pterygiophore configurations were obtained from radiographs.

Zenopsis stabilispinosa sp. nov.

Figures 1-3A, 4, Tables 1-2

Zenopsis sp. nov. Bray, 1983: 106, fig. 16; Zenopsis sp. (off

Western Australia): Williams et al., 1996: 150 (off Western Australia),

Tyler et al., 2003: 2 (South China Sea and off Western Australia).

Material examined. Holotype. FAKU 64803 (307.2 mm SL), South

China Sea, 19°45.0'N, 114°04.0'E, 457-767 m, JAMARC (Japan

Marine Fishery Resource Research Center), 20 Jun, 1991.

Paratypes. FAKU 64804 (410.1 mm SL), South China Sea,

19°47.0'N, 114°05.0'E, 465-505 m, JAMARC, 21 Jun 1991; AMS

1.22826-004 (206.1 mm SL), North-west Shelf, Western Australia 210

km NW of Port Hedland, 18°44'S, 117°02'E, 396-406 m, J. R. Paxton,

CSIRO RV Soela, 13 Apr, 1982; AMS 1.31146-001 (2, 100.2-103.3

mmSL),off North-west Cape, Western Australia, 21°37.9'S, 113°59.3'E,

209-215 m, J. R. Paxton, CSIRO RV Southern Surveyor, 24 Jan, 1991;

AMS 1.31147-002 (3, 80.4-104.5 mm SL), off North-west Cape,

Western Australia, 21°44.5'S, 113°52.5'E, 290-320 m, J. R. Paxton,

CSIRO RV Southern Surveyor, 24 Jan, 1991; ASIZ P.0057609, off

Donggang, Pintung, Taiwan, 22 28 12 N, 120 25 48 E, K. T. Shao, 8

Oct, 1985; ASIZ P.0060011, fish market, Dahsi, Ylian, Taiwan, 24 57

00 N, 121 52 48 E, B. H. Gao, 10 Aug, 1997.

Diagnosis. Zenopsis stabilispinosa differs from its congeners in

the following combination of characters: dorsal fin with 7 spines;

anal fin with first 2 spines movable and 3rd spine immovable,

fused with its pterygiophore; pelvic fin short, 1.69-1.79 in HL,

in adult specimens greater than 300 mm SL; bony bucklers

along bases of spinous and soft-rayed portions of dorsal fin 3-

4- I-5-6; bucklers along ventral body margin anterior to pelvic fin

and between pelvic- and anal-fin origins, and along anal-fin base

5- 6-I-6-7-I-5-6 and vertebrae 32 (13 abdominal -i- 19 caudal).

Description, (values for paratypes in parentheses if different

from holotype). Dorsal fin VII, 26; anal fin III, 23; pectoral fin

rays 12 (12-13); gill rakers 3-1-9 (3^-i-8-10); dorsal bucklers

Tetsuji Nakabo, Dianne J. Bray and Umeyoshi Yamada

Figure 1. Zenopsis stabilispinosa sp. nov. Holotype, FAKU 64803 (307.2 mm SL).

Figure 2. Diagrammatic figure (A), and teeth on jaws and vomer (B) of the holotype of Zenopsis stabilispinosa sp. nov., FAKU 64803.

A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East China Sea and off Western Australia

Figure 3. Diagrammatic figures of anal-fin pterygiophores and spines

of 3 species of Zenopsis: A, Z. stabilispinosa, holotype, FAKU 64803

(307.2 mm SL); B, Z. nebulosa, FAKU 64805 (379 mm SL); C, Z.

conchifer, RUSI 14070 (135.5 mm SL). Scales indicate 10 mm.

Figured. Diagrammatic figure of ajuvenile of Zenopsis stabilispinosa,

AMS 1.31147-002 (80.4 mm SL), paratype.

4-1-6 (3^-i-5-6); ventral bucklers 6-I-6-I-5 (5-6-I-6-7-I-5-6);

vertebrae 13-1-19. Body deep, strongly compressed, scales absent

except along lateral line. Head strongly compressed, dorsal

profile slightly concave. Nostrils located just before eye; anterior

nostril rounded, posterior nostril oblong. Mouth large, strongly

oblique; upper jaw extremely protrusible. Teeth in upper jaw in

2 separate bands; anterior band with 3 irregular rows of small

caniform teeth, inner teeth larger than outer ones; posterior band

very narrow, long with minute caniform teeth. Lower jaw with 2

irregular rows of small caniform teeth; 2 teeth on inner row of

anteriormost part larger than other teeth. Vomer with 1 small

caniform tooth on right side, 2 small caniform teeth on left side

(with 0-3 caniform teeth on right side, 0-7 on left side; mode 2

on both sides). Palatine toothless. Tongue thick, long and pointed.

Gill-rakers on 1st gill-arch short. Pseudobranchiae well-

developed. Lateral line arched below spinous portion of dorsal

fin, following midline axis of body from below anterior one-third

of soft dorsal fin, to caudal-fin base. Caudal peduncle moderately

long, narrow. Dorsal-fin origin vertically to almost vertically

above posterior tip of operculum; anterior dorsal-fin spines long

with filamentous membranes; soft dorsal fin convex, rays

increasing in length to 18th ray. Anal fin origin near vertical

through notch between spinous and soft-rayed portions of dorsal

fin; 1st and 2nd anal-fin spines small, movable, 3rd spine

immovable (fused with pterygiophore); 2nd and 3rd anal-fin

spines and 1st anal-fin ray connected by membrane; soft anal fin

convex, rays increasing in length to about 16th ray. Pectoral fin

rounded, reaching to vertical below bases of 4th and 5th dorsal-

fin spines. Pelvic fin relatively short, 1.69 in HL(L79 in paratype

324.4 mm SL, but somewhat longer, 0.98-1 .26 in HL, in juvenile

paratypes 80.4—206.1 mm SL), origin anterior to vertical through

pectoral-fin base, fin not reaching anus when depressed

(extending to anus in paratypes less than 105 mm SL). Caudal fin

slightly emarginate.

4 (3-4) bony bucklers along base of spinous dorsal fin and

6 (5-6) bony bucklers along soft dorsal-fin base; each buckler

with a dorsal spine. 6 (5-6) bony bucklers anterior to pelvic-

fin base. 6 (6-7) bony bucklers with posterior spine between

origins of pelvic and anal fins. Bucklers absent from base of

anal-fin spines. 6 (5-6) bony bucklers, each with a vertical

spine, along base of soft anal fin.

Colour when fresh. Body silver (with many dark spots in juvenile

paratypes less than 105 mm SL). Dorsal margin of head and

body dark-brown from snout to caudal peduncle. Spinous dorsal

fin with a transverse broad dark-brown band on membrane; soft

dorsal fin dusky distally, white basally. Anal fin dusky distally,

white basally. Pectoral fin pale basally, dusky distally, with a

small dark-brown spot dorsally on fin base. Pelvic fin with 4

irregular black bands (6-7 dark bands in juvenile paratypes less

than 105 mm SL). Caudal fin with dark semicircular mark on

base and broad dark band on posterior margin.

Etymology. The specific name, stabilispinosa, refers to the

fused anal-fin spine.

We recommend that the standard name for this species be

the Fixed-spine Mirror Dory.

Tetsuji Nakabo, Dianne J. Bray and Umeyoshi Yamada

Table 1. Selected meristics for species of Zenopsis

Z. stabilispinosa

Z. nebulosa

Z. conchifer

Z. oblonga

Holotype

Paratypes

No. specimens

Dorsal fin

Vll, 26

Vll, 27

IX, 27

IX-X, 24-26

Vlll-X, 27-29

Anal fin

111, 23

111, 25-26

111, 25-27

Pectoral fin

12-13

11-13

Pelvic fin

Gill rakers (upper -i- lower)

3 + 9

3-4 + 8-10

3-4 + 7-9

2-3 + 8-10

Vertebrae (AV + CV)

32 (13 + 19)

35(13 + 22)

34-36(13 + 21-23)

Dorsal bony bucklers*

4 + 6

3-4 + 5-6

5-7 + 6-7

2-3 + 5

6-7 + 8-9

Ventral bony bucklers**

6 + 6 + 5

5-6 + 6-7 + 5-6

5-6 + 7-9 + 8-10

1-3 + 6-8 + 5-6

2-3 + 7-8 + 9-11

* along spinous and soft dorsal-fin bases

**anterior to pelvic fin + between pelvic fin and anal fin + along anal-fin base

Relationships. Zenopsis stabilispinosa differs from Z.

nebulosa, Z. conchifer and Z. oblonga in having the first two

anal-fin spines movable and a third fused to the pterygiophore

(versus three movable anal-fin spines in the other three species;

in some specimens of Z. conchifer the third anal-fin spine is

more or less fixed, but is not fused with the pterygiophore, fig.

3C). As the movable third anal-fin spine is based on the second

anal pterygiophore in the latter three species (fig. 3B), it is

likely that the spine -like posterior process projecting from the

third anal pterygiophore of Z. stabilispinosa represents the

third anal-fin spine fused to its pterygiophore (fig. 3A). Fused

anal-fin spines have previously been reported in two other

zeiform families: family Parazenidae - Cyttopsis rosea Lowe,

1843, and family Cyttidae - Cyttus novaezelandiae Arthur,

1885 and C. traversi Hutton, 1872 (Bray, 1983, Heemstra, 1980,

Tyler et ah, 2003).

Zenopsis stabilispinosa also differs from its congeners in

having seven dorsal-fin spines (versus eight to ten dorsal-fin

spines), 19 caudal vertebrae (versus 22-23 caudal vertebrae)

and a smaller pelvic fin, 1.69-1.79 in HL in specimens greater

than 300 mm SL of Z. stabilispinosa (versus almost the same

as or more than HL in other species).

Zenopsis stabilispinosa is similar to Z. nebulosa in having

five to six ventral bony bucklers anterior to the pelvic fin (versus

two to three bucklers in Z. conchifer and Z. oblonga). Zenopsis

stabilispinosa and Z. conchifer resemble Z. conchifer in lacking

ventral bony bucklers along the anal-fin spines (versus having

one to two ventral bony bucklers in Z. nebulosa and Z. oblonga).

Remarks. Opinion differs regarding whether the first pelvic-fin

element in Zenopsis is a spine or a ray. Here, we follow Tyler et

al., 2003 in regarding the first pelvic-fin element as a ray

because it is completely divided throughout its entire length,

despite being neither segmented nor branched.

Key to the species of Zenopsis

la) Dorsal-fin spines 7; 3rd anal-fin spine immovable (fused

to pterygiophore); caudal vertebrae 19

Z. stabilispinosa sp. nov.

lb) Dorsal-fin spines 8-10; 3rd anal-fin spine movable (not

fused to pterygiophore); caudal vertebrae 21-23 2

2a) Bony bucklers along spinous portion of dorsal-fin base

1-3; bucklers absent from base of anal-fin spines

Z. conchifer

2b) Bony bucklers along spinous dorsal-fin base 5-7; 1-2

bony bucklers along base of anal-fin spines 3

3a) Bony bucklers on ventral margin anterior to pelvic-fin

base 5-6 Z. nebulosa

3b) Bony bucklers on ventral margin anterior to pelvic-fin

base 2-3 Z. oblonga

Comparative material examined

Zenopsis conchifer. RUSI (J. L. B. Smith Institute of Ichthyology)

13801, (3, 65.6-135.8 mm SL), Kenya, Malindi, 3°04'S, 40°25'E,

R C. Heemstra, 17 Nov 1980; RUSI 14070 (5, 84.8-135.5 mm

SL), Kenya, Ras Ngomeni, 2°50'S, 40°34'E, P. C. Heemstra, 12

Dec 1980. Zenopsis nebulosa: FAKU 64805, SNFR 1068 (313-

381 mm SL), East China Sea, 30°49'9N, 128°56'2E, 355-379 m,

SNFR, 26 Oct 1991; SNFR 336, 1610, 1611, (86-152 mm SL),

East China Sea, 32°12'7N, 127°34'7E, 117 m, SNFR, 14 Oct

1982; SNFR 1066 (232 mm SL), East China Sea, 29°14'7N,

127°34'7E, 408 m, SNFR, 2 Oct 1989. Zenopsis oblonga: USNM

285048, E. Pacific, Nazca Ridge, 25°39'S, 85°37'W, 210 m

(paratype); USNM 353898, SE Pacific, Sala-Y-Gomez Ridge,

25°02S, 97.48°W, 330 m, N. Parin et ah, 3 May 1987.

A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East China Sea and off Western Australia

Table 2. Measurements of Zenopsis stabilispinosa sp. nov. Measurements are in mm; proportions (% SL) in parentheses.

Ol-

C--

o|-

Cl;

ol;

o|-

Cl;

IT)

Ol-

c/5

ol;

Ol-

(T)

C--

AMS

C--

£;

C--

oi-

C--

o|-

ol-

C--

ol;

Ol-

Clh

C--

Ol-

C--

o|-

ol;

ol-

o|-

-H

ol;

o|-

Ol-

259.1

T903

75.1

125.7

16.4

88.9

40.4

17.3

17.7

23.3

14.2

138.1

9.6

19.2

31.2

59.5

47.9

voz

14.0

9.3

8.3

17.3

Ol-

ol;

o|-

Ol-

o|-

^ §

ol;

o|-

Catalog no.

Total length

Standard length (SL)

Head length (HL)

Body depth

Body width

Predorsal length

Snout length

Eye diameter

Orbit diameter

Postorbital length

Interorbital length

Preanal length

Caudal peduncle depth

Caudal peduncle length

Pectoral-fin length

Pelvic-fin length

Caudal-fin length

1st dorsal-spine length

2nd dorsal-spine length

3rd dorsal-spine length

4th dorsal-spine length

5th dorsal-spine length

6th dorsal-spine length

7th dorsal-spine length

Longest dorsal-ray length

1st anal-spine length

2nd anal-spine length

3rd anal-spine length

Longest anal-ray length

Tetsuji Nakabo, Dianne J. Bray and Umeyoshi Yamada

Acknowledgements

We thank the staff of the Japan Marine Fishery Resource Research

Center (JAMARC) for donating the specimens of Z. stabilispinosa

to FAKU for this study. We are grateful to T. Irie and H. Horikawa,

Fisheries Research Agency, J. R. Paxton and J. M. Leis, Australian

Museum, Sydney and K.T. Shao, Academia Sinica, Taipei, for

the loan of specimens of Z. stabilispinosa, and to P. C. Heemstra

for loaning the specimens of Z. conchifer. Lastly, we thank J. C.

Tyler (USNM) for critically reviewing the manuscript.

References

Arthur, W. 1885. Notes on New Zealand fishes. Transactions of the

New Zealand Institute 17 (1884): 160-172, PI. 14.

Bray,D. J. 1983. Revision ofthe fish family Zeidae. 184 pp. Unpublished

Master’s thesis. School of Biological Sciences, Macquarie

University, Sydney, Australia.

Heemstra, P. C. 1980. A revision of the zeid fishes (Zeiformes: Zeidae)

of South Africa. Ichthyological Bulletin (41): 1-18.

Hutton, F. W. 1872. Fishes of New Zealand. Catalogue with diagnoses

ofthe species. Colonial Museum and Geological Survey Department.

Wellington. Fishes New Zealand: 1-93 + 95-133, Pis. 1-12.

Leviton, A. E., Gibbs, R. H. Jr., Heal, E. and Dawson, C. E. 1985.

Standards in herpetology and ichthyology. Part 1. Standard

symbolic codes for institutional resource collections in herpetology

and ichthyology. Copeia 1985 (3): 802-832.

Lowe, R. T. 1852. An account of fishes discovered or observed in

Madeira since the year 1842. Proceedings of the Zoological

Society of London 1850 (pt 18): 247-253.

Nakabo, T. 2002a. Introduction to Ichthyology, p.xxi-xlii in: Nakabo,

T. (ed.). Fishes of Japan with pictorial keys to the species, English

edition. Tokai University Press: Tokyo.

Parin, N. V. 1989. Zenopsis oblonga sp. nov. (Zeidae, Osteichthyes)

from the Nuska Ridge. Zoologicheskii Zhurnal 68(4): 150-153.

(in Russian with English summary).

Temminck, C. J. and Schlegel, H. 1845. Pisces, Parts 7-9: 113-172,

Pis. 1-143 + A, in: Fauna Japonica, sive descriptio animalium

quae in itinere per Japoniam suscepto annis 1823-30 collegit, notis

observationibus et adumbrationibus illustravit P. E de Siebold.

Tyler, J. C., O'Toole, B. and Winterbottom, R. 2003. Phylogeny of the

genera and families of zeiform fishes, with comments on their

relationships with tetraodontiforms and caproids. Smithsonian

Contributions to Zoology. 618: i-iv, 1-110.

Williams, A., Last, P. R., Gomon, M. F. and Paxton, J. R. 1996. Species

composition and checklist of the demersal ichthyofauna of the

continental slope off Western Australia (20-35°S). Records ofthe

Western Australian Museum, 18: 135-155.

Memoirs of Museum Victoria 63(1): 97-106 (2006)

iSSN 1447-2546 (Print) 1447-2554 (On-iine)

http://www.museum.vic.gov.au/memoirs/index.asp

Australian trevallies of the genus Pseudocaranx (Teleostei: Carangidae), with

description of a new species from Western Australia

William F. Smith-Vaniz and Howard L. Jelks

Abstract

Keywords

U. S. Geological Survey, 7920 NW 7P‘ Street, Gainesville, Florida 32653 USA (bill_smith-vaniz@usgs.gov)

Smith-Vaniz W.F. and Jelks H.L. 2006. Australian trevallies of the genus Pseudocaranx (Teleostei: Carangidae), with

description of a new species from Western Australia. Memoirs of Museum Victoria 63(1): 97-106.

Pseudocaranx dinjerra sp. nov. is described from Western Australia. It differs from the Australian endemic

Pseudocaranx wrighti in having the posterior margin of the upper jaw nearly vertical, lachrymal without scales, 19-21

versus 24-28 lower gill rakers and 53-66 versus 37-48 scales in the curved part of the lateral line. The new species is

distinguished from Australian populations of P. georgianus and Mentex” primarily by having a different combination of

meristic values including 14 versus 15 caudal vertebra, 19-21 versus 21-24 segmented anal-fin rays and 58-75 versus

72-95 total lateral-line scales. Diagnoses are given for P. wrighti and P. georgianus, and specimens from Queensland and

Lord Howe Island are discussed and provisionally identified as P. dentex.

Carangidae, Pseudocaranx, new species. Western Australia, antitropical distribution

Introduction

Carangid fishes of the genus Pseudocaranx'OiQQkex, 1863 occur

in subtropical and warm temperate regions of the world’s

oceans with single species, P. dentex Bloch and Schneider, 1801

and P. chilensis Guichenot, 1848, present in the Atlantic and

eastern Pacific oceans, respectively. In the Indo-west Pacific,

the genus consists of a complex of species for which 13 names

have been proposed, eight based on fish from Australia or New

Zealand. Except for the Australian endemic Pseudocaranx

wrighti Whitley, all of these species are very similar externally

and their scientific names have been inconsistently used and

frequently misapplied. Determining the taxonomic status of

some disjunct and sympatric populations (e.g., Yamaoka et ah,

1991; Masuda et ah, 1995) will probably require both

morphological and molecular data. Unraveling the evolutionary

history of these carangids is also challenging because they have

antitropical distributions that are not easily explained (Briggs,

1987) and is confounded by the Miocene origins of Indo-Pacific

coral reef fish biodiversity, which predates Pleistocene

glaciations (Read et ah, 2006).

In the course of acquiring vertebral counts from a large

series of Pseudocaranx from many localities, we observed that

all specimens from both sides of the Atlantic Ocean have 15

caudal vertebrae and those from many relatively isolated

locations in the Indo-Pacific consistently have either 14 or 15

caudal vertebrae. The discovery that Pseudocaranx species

from Western Australia include taxa with either 14 or 15 caudal

vertebrae and non-overlapping geographic distributions resulted

in a re-evaluation of the taxonomic status of species that Paxton

et ah, 1989 and Smith-Vaniz, 1999 had referred to collectively

as P. dentex. These fishes are herein recognized as P. georgianus

Cuvier and a new species P. dinjerra, respectively. Identification

of the Pseudocaranx species that occurs off southern Queensland

and at Lord Howe Island is unresolved and is here tentatively

referred to as Pseudocaranx sp. ''dentex'\

In their review of P. georgianus from temperate Australasian

waters, James and Stephenson, 1974 concluded that only one

species was represented but observed that some collections

from southern Australia contained two groups of fish each

having either 24 or 25 total vertebrae and different proportions

(body depth, snout/head lengths and eye diameter). Re-

examination of most of the specimens available for their study

revealed that all of those with 24 vertebrae are P. wrighti

(species not mentioned by them) and that those with 25 vertebrae

are P. georgianus. Although James and Stephenson, 1974 failed

to appreciate a number of additional characters that distinguish

these two species, they correctly reported that specimens from

Norfolk and Kermadec (Raoul) Is. consistently have 14 caudal

(24 total) vertebrae, as do most of those from North Cape, New

Zealand. Determination of the taxonomic status of these

extralimital fish is beyond the scope of the present study;

however, they definitely are neither P. dinjerra nor P. wrighti.

William F. Smith-Vaniz and Howard L. Jelks

Material and methods

Museum abbreviations follow Leviton et al., 1985. In the

material examined sections, specimen sizes are given as mm

fork length (FL) and cleared and stained specimens are indicated

as “C&S”; parenthetical expressions present number of

specimens, if more than one, followed by size range. Localities

are abbreviated and listed by major geographic areas. Scutes are

defined as scales that have a raised horizontal ridge on their

posterior margin with a small to moderate projecting spine

ending in a point not exceeding a 120° angle. All scutes are

counted, including those on the caudal-fin base. Pectoral-ray

counts do not include the dorsal-most spine -like element. Gill-

raker counts are from the first gill arch (usually on the right

side), with the raker at the angle included in the lower-limb

count; rudimentary gill rakers are defined as tubercles or short

rakers with the diameter of their bases greater than their height.

Measurements were analyzed using a sheared principal

component analysis (PCA) following Rohlf and Bookstein,

1987. The following 22 point-to-point measurements were taken

for the relatively few specimens > 200 mm FL: fork length (FL)

from tip of snout to tip of shortest median caudal-fin ray; snout

to origin of first dorsal fin (DIO); snout to origin of second

dorsal fin (D20); snout to origin of pelvic fin (P20); snout to

origin of first anal-fin spine (AlO); length of dorsal-fin base;

length of anal-fin base; DIO to P20; DIO to origin of second

dorsal fin (A20); D20 to A20; D20 to AlO; height of dorsal-

fin lobe; height of anal-fin lobe; pelvic fin length; pectoral fin

length; length of curved part of lateral line (CLL), measured as

a cord (straight-line distance) of the arch extending from the

upper edge of the opercle to its junction with the straight part;

length of straight part of lateral line (SLL), measured from its

junction with the curved part to its termination on the caudal-fin

base (end of last scute); head length from tip of snout to posterior

margin of the opercular flap; postorbital head length from

posterior margin of orbit to posterior margin of the opercular

flap; upper jaw length is from the snout tip to posterior end of

maxilla; eye diameter is the horizontal diameter of the orbit.

Pseudocaranx Bleeker

Pseudocaranx Bleeker, 1863: 82.

Type species. Scomber dentex Bloch and Schneider, 1801, by

subsequent designation of Fowler, 1936: 692.

Remarks. These fishes are presumed to comprise a monophyletic

group, but the most appropriate generic classification for them is

uncertain pending a well collaborated phytogeny of carangines.

Citula Cuvier, 1816 (type species Citula banksii Risso, 1820 [=

P. dentex] by subsequent monotypy) is an available senior

synonym of Pseudocaranx (type species Scomber dentex) but

this generic name has not been used for a nominal species of the

group since Risso’s description. In the interest of nomenclatural

stability, the junior name should continue to be used pending a

petition to the International Commission of Zoological

Nomenclature to conserve the more familiar generic name.

Other more recent synonyms of Pseudocaranx are Longirostrum

Wakiya, 1924, Usa (as a subgenus of Caranx) Whitley, 1927, and

Usacaranx Whitley, 1931. Within the Caranginae, Pseudocaranx

and Caranx equula Temminck and Schlegel, 1844, which Kijima

et al., 1986 and Gushiken, 1988 assign to the monotypic genus

Kaiwarinus Suzuki, 1962, are exceptional in having very poorly

developed inferior vertebral foramina, and may be a sister taxa.

Key to Australian species of Pseudocaranx

1. Posterior margin of upper jaw canted posteroventrally (fig.

lA); scales present on lachrymal; lower limb gill rakers

24-28; scales in curved portion of lateral line 37-48

(eastern Bass Strait to Exmouth Gulf, WA) P. wrighti

— Posterior margin of upper jaw nearly vertical (fig. IB); no

scales on lachrymal; lower limb gill rakers 19-23 (except

24-27 in fish from Qld and Lord Howe L, where P. wrighti

does not occur); scales in curved portion of lateral line

53-80 2

2. Segmented anal-fin rays 19-21; total lateral -line scales

58-75, rarely >71; caudal vertebrae 14 (Houtman Abrolhos

to North West Cape, WA) P. dinjerra

— Segmented anal-fin rays 21-24 (rarely 21); total lateral-line

scales 72-95, rarely <74; caudal vertebrae 15 3

3. Lower limb and total gill rakers 19-23 and 28-35 (rarely

35) , respectively (New Zealand and NSW to Lancelin L,

WA) P. georgianus

— Lower limb and total gill rakers 24—27 and 36^1 (rarely

36) , respectively (southern Qld, Australia and Lord Howe

I.) P. sp. ""dentex”

Pseudocaranx dinjerra sp. nov.

Figures 2, 3A, 4, 5; Tables 1-3

Pseudocaranx dentex (non Bloch and Schneider) in: Allen and

Swainston, 1988: 74, fig. 447 (misident., in part, brief descr.) in:

Hutchins, 1990: 270 (listed; Shark Bay); Hutchins, 1997: 247 (listed;

Houtman Abrolhos).

Material examined. Holotype. NMV A. 1962 (245), SW of Shark Bay,

25°28'S, 112°27'E, 25°19'S, 112°17'E; trawled in 131-139 m; M.F.

Gomon; sta. MFG-71; 4 Mar 1981.

Paratypes. 38 specimens, 77-230 mm FL. ANSP 148695 (2, 221-

230), off Cape Farquhar, 23°42'S, 113°0rE, 23°48'S, 112°58'E; trawled

in 156-160 m; M/V TM 71; 13 Sep 1979. WAM P.22338 (217), Cape

Cuvier, 24°13'S, 113°23'E; J. Penn; 29 Jul 1972. ANSP 148696 (20,

77-92) and USNM 385866 (15, 81-95.5), Houtman Abrolhos,

Hummock L, 28°48'S, 114°02'E, trawled in 43 m; Nov 1980.

Diagnosis. A species of Pseudocaranx with posterior margin of

upper jaw nearly vertical; lachrymal naked and expanded part

of maxilla only partially covered with scales; caudal vertebra

14; gill rakers 7-10 upper, 19-21 lower, 27-31 total; scales in

curved part of lateral line 53-66.

Description (values for holotype in parentheses). Dorsal fin rays

VIII-I, (25) 23-25; anal-fin rays II-I, (21) 19-21; pectoral-fin

rays (19) 18-20; vertebra 10 precaudal -i- 14 caudal; inferior

vertebral foramina on caudal vertebra 7 or 8-10; scales in

curved lateral line (61) 53-66; scales in straight LL (2) 2-11;

scutes in straight LL (30) 19-31; total scales in LL (63) 58-74;

total scales -i- scutes in LL (93) 86-99; developed gill rakers (8)

7-10 upper, (20) 19-21 lower, (28) 27-31 total, a single

rudimentary raker rarely present on either end of gill arch.

Australian trevallies of the genus Pseudocaranx

Figure 1. Head profiles and upper jaw shapes of Pseudocaranx: A,

P. wrighti, ANSP 135418, 169 mm FL, Cockburn Sound, WA; B, P.

georgianus, ANSP 138196, 200 mm FL, Cape Arid, WA.

Chest completely scaly; bases of dorsal and anal fins with

a wide scaly sheath anteriorly; lachrymal naked; dorsal 3rd to

half of expanded part of maxilla with a few embedded scales;

cheeks, preopercle, opercle and interopercle covered with

scales. Junction of curved and straight parts of lateral line

below segmented dorsal-fin rays (13) 12-13; length of curved

LL (0.57) 0.62-0.65 in straight LL; 1st dorsal-fin spines weak,

the 3rd spine longest and slightly longer than height of 2nd

dorsal-fin lobe; last dorsal- and anal-fin rays slightly longer

and more widely spaced than adjacent rays; 1st anal-fin

pterygiophore with anteroventral end short and bluntly

rounded; 2nd dorsal-fin lobe (3.1) 2. 8-3.1 in head length;

pectoral fin of holotype and larger paratypes (0.94) 0.91-0.94

in head length. Upper jaw (2.7) 2. 8-2.9 in head length, ending

slightly in front of anterior margin of eye (fig. 3A); adipose

eyelid weakly developed; lips slightly thickened and finely

papillose. Jaw teeth (difficult to see clearly without dissection)

in the holotype and 2 largest paratypes: upper jaw with a single

row (21 left, 22 right) 19-22 of small conical teeth and (0) 1

inner tooth near symphysis; lower jaw with an outer row (21

left, 22 right) 25-27 of small conical teeth and an irregular

inner row (11 left, 8 right) 13-18 of conical teeth on posterior

half of jaw. (Based on observed changes in dentition with

growth in other Pseudocaranx species, it is likely that an inner

row of dentary teeth is absent in individuals of P. dinjerra

larger than the holotype.) Vomerine tooth patch triangular-

shaped, without a median posterior extension, and sparsely

covered with small teeth.

Measurements of the holotype (in parentheses) and 2

paratypes, 221-230 mm, as percentages of FL: snout to DIO

(39) 41-42; snout to D20 (54) 56; snout to P20 (35) 34-35;

snout to A20 (57) 57; DIO to P20 (30) 30; DIO to A20 (39)

39-41; D20 to AlO (32) 33-35; D20 to A20 (32) 33-35; D2

base (35) 35-37; A2 base (31) 30-31; height dorsal-fin lobe

(10) 10-11; height anal-fin lobe (10) 9-10; pelvic-fin length

(15) 14-15; pectoral-fin length (33) 34; head length (31) 31-32;

postorbital head length (12) 12-13; snout length (12) 12; upper

jaw length (11) 11; eye diameter (7) 7-8; curved lateral-line

length (24) 24-25; and straight lateral-line length (24) 24-25.

Preserved coloration. Holotype and larger paratypes uniformly

pigmented, except spinous dorsal fin slightly dusky and opercle

with a prominent dark pupil-sized spot at level of pupil. Juveniles

with 7-9 dusky bands on body, extending ventrally from dorsum

and fading out on ventral half of sides. Bands widest dorsally

and equal or slightly wider than pale interspaces at mid-level of

side. Dark opercular spot, intense, smaller than pupil and slightly

vertically elongate. Inter-radial membranes of 1st dorsal fin

dusky, densely peppered with small melanophores.

Life coloration (from an underwater photograph, probably of

a subadult, from Shark Bay provided by J.B. Hutchins). Silvery

blue-green dorsally, fading to silvery with iridescence below,

faint mid-lateral yellow stripe from opercle to base of caudal fin;

faint yellow stripe on base of dorsal fin extending slightly onto

dorsum; prominent black spot on opercle at level of the pupil,

approximately diameter of pupil and vertically elongate.

Distribution. Endemic to WA (fig. 4). Definitely known from

Houtman Abrolhos (28°48'S) to off Cape Farquhar (23°42'S),

and reported (Allen and Swainston, 1988) to North-West Cape

(^21°47'S), but rare north of Shark Bay. Several photographs of

Pseudocaranx taken by J.B. Hutchins off Green Head, WA

(30°04'S) are tentatively identified as P. dinjerra based on the

tiny size of the opercular spot. We know of no collections of P.

dinjerra from well-sampled Rottnest I. (Hutchins and Pearce,

1994), located at 32°S, where both P. wrighti and georgianus

occur; but its occurrence there might be expected.

Dispersal of temperate and subtropical species along the

north-west coast of Australia is believed to be aided by counter-

currents flowing inshore of the southward-flowing warm Leeuwin

100

William F. Smith-Vaniz and Howard L. Jelks

Figure 2. Pseudocaranx dinjerra. Shark Bay, WA; photograph by J. B. Hutchins.

Current. Fluctuations of this current regimen - associated with

glacial and interglacial periods may have contributed to

reproductive isolation and eventual speciation of Western

Australian endemic subtropical species (Hutchins, 1994;

Hutchins, 2001a), presumably including the new Pseudocaranx.

Etymology. The trivial name dinjerra (west) is an Aboriginal

word (Anon, 1969), in reference to the Western Australian

endemic status of the species, and should be treated as an

appositional noun.

Remarks. This species is most similar to Pseudocaranx

georgianus but differs in having 14 caudal vertebra, and little

overlap in number of anal-fin rays (Table 1) and total lateral-line

scales (Table 3). Sheared PCA (fig. 5) revealed good separation

of the three analyzed taxa. Although 11 of the 15 specimens of

P. georgianus used in the analysis were obtained from the

Sydney Fish Market, according to Kailola et ah, 1993 the main

commercial fishery is located in New South Wales waters where

this species appears to be resident and non-migratory.

Pseudocaranx dinjerra has a relatively longer snout and upper

Jaw (PC2) and shorter straight lateral line (PC3) than P.

georgianus and P. sp. ''dentex'\ However, fork length (PCI)

accounted for 97.5% of the variation, while only 1.7% was

associated with PC2 and PC3. Differences were subtle, yet

consistent in the three groups.

Other than P. wrighti, the only other previously described

Indo-Pacific Pseudocaranx with 14 caudal vertebra is P. cheilio

Snyder, 1904, described from Honolulu, Hawaii. As mentioned

in the introduction, at many locations in the Indo-west Pacific

(including Hawaii and Easter Island) Pseudocaranx spp.

invariably have either 14 or 15 precaudal vertebrae. In contrast,

Yamaoka et ah, 1991 found two distinct genetic morphs (identified

by electrophoretic analysis) of “P. dentex” in Tosa Bay, Japan,

each with different vertebral counts. The two morphs had

strongly bimodal dorsal ray counts and the juveniles of one

morph also appeared to have more distinct narrow bands on the

body, leading these authors to strongly suspect that two species

were involved. Masuda et ah, 1995 also found significant mtDNA

differences between the same two sympatric Japanese

Pseudocaranx morphs. They implied that differences in

spawning and recruitment locations and associated water

temperatures may have affected the number of vertebrae.

Vertebral counts are intraspecifically very constant in all other

carangid genera (including 130-1- species), so the situation in

Pseudocaranx is very interesting if these morphs are not different

species. Neither of the Japanese studies considered gill raker

numbers, but our limited data indicate that they also differ

between these two morphs. Pseudocaranx cheilio from Hawaii

and the Japanese morph with 14 caudal vertebrae have higher

numbers of lower gill rakers, 27-30 versus 19-21 in P. dinjerra.

Australian trevallies of the genus Pseudocaranx

101

Figure 3. Heads of Pseudocaranx species: A, P. dinjerra, ANSP 148695, 230 mm FL, Cape Farquhar, WA; B, P. georgianus, USNM 385513, 275

mm FL, Sydney Fish Market; C, P. '"denteP\ CAS 16472, 404 mm FL, One Tree L; D, P. wrighti, WAM P.14007-18, 124 mm FL, Rottnest I.

Pseudocaranx georgianus Cuvier, 1833

Figures IB, 3B, 4, 5; Tables 1-3

Caranx georgianus Cuvier in Cuvier and Valenciennes, 1833: 85

(orig. descr. King George Sound, WA; syntypes MNHN 5854); Ogilby,

1893: 80, pi. 24 (descr.; biology; fisheries; edibility); James and

Stephenson, 1974: 402, fig. 2 (taxonomic status; synonymy; descr.

based, in part, on Pseudocaranx wrighti)-, Hutchins, 1979: 38, color

pi. 21 (Rottnest L; behaviour; distrib.); Smith-Vaniz et al., 1979: 12

(syntypes listed).

Caranx platessa Cuvier in Cuvier and Valenciennes, 1833: 84

(orig. descr. “La mer des Indes;” holotype MNHN 5856); Gunther,

1860: 440 (as first reviser, listed in synonymy of C. georgianus)-, James

and Stephenson, 1974: 402, fig. 3 (discussion of type locality, probably

Geographe Bay, WA; considered to be a synonym of C. georgianus)-,

Smith-Vaniz et al., 1979: 17 (holotype listed).

Caranx nobilis Macleay, 1881: 532 (orig. descr. Port Jackson,

NSW, Australia; holotype MAMU, apparently lost).

Usacaranx archeyi Griffin, 1932: 130, pi. 22 (orig. descr. North

side of Motuihi L, Hauraki Gulf, New Zealand; Holotype AIM 262).

Usacaranx georgianus. Roughley, 1951: 57, color pi. 22 (biology

and fisheries); Scott et al., 1974: 201, unnumb. fig. (desc.; distrib.).

Pseudocaranx dentex (non Bloch and Schneider). May and

Maxwell, 1986: 300, unnumb, color photo, (descr.; common name;

102

William F. Smith-Vaniz and Howard L. Jelks

Figure 4. Australian distributions of Pseudocaranx species based on

material examined.

distrib.); Okiyama, 1988: 474, unnumb. pi. (descr.; early stage); Paxton

et al., 1989: 582 (partial synonymy; distrib. in part, includes range of

P. dinjerra); Kuiter, 1993: 176, unnumb, color photos (descr.; distrib.;

in part, includes ranges of other spp.); Gomon et al., 1994: 584,

unnumb, color photos (descr.; distrib.; partial synonymy); Smith-

Vaniz, 1999: 2730 (descr.; synonymy; distrib. in part, includes ranges

of other spp.); Hutchins, 2001b: 33 (listed).

Diagnosis. A species of Pseudocaranx with posterior margin of

upper jaw nearly vertical (fig. IB); lachrymal naked and

expanded part of maxilla naked or with a few partially embedded

scales; caudal vertebra 15; gill rakers 8-13 upper, 19-23 lower,

28-35 total; scales in curved part of lateral line 55-78.

Distribution. Known from New Zealand and the southern half

of Australia from NSW to just north of Perth, WA, including

Rottnest I.

Remarks. Pseudocaranx dentex, described from Brazil, is the

oldest available name for any species of Pseudocaranx while P.

georgianus is the oldest name for a nominal species with an

Indo-Pacific type locality. The type specimens of both species

have 15 caudal vertebrae, as do all specimens from the Atlantic

and western Indian Oceans and some from Japan. Australian

specimens with the same vertebral count have fewer total gill

rakers (except those from Queensland and Lord Howe Island,

see Table 2) than do specimens from the latter localities, which

have 35^2 gill rakers. Unlike similarly sized specimens of P.

dentex from widely separate Atlantic localities, large adults

(>350 mm FL) from southern Australia and New Zealand

usually have a pronounced hump on their foreheads that is

correlated with hyperostosis of the supraoccipital bone, do not

have blunt snouts (see following remarks), and the dark opercular

spot, although variable in size and shape, is also usually diffuse

and noticeably larger than the pupil diameter. Because of these

differences, P. georgianus is here recognized as a valid species.

Pseudocaranx sp. “dentex”

Figures 3C, 4, 5; Tables 1-3

Material examined. Ill specimens, 45^95 mm FL. WAM P.14755

(170), 0.8 km E. Lancelin L, 31°33'S, 115°19'E. WAM P.14054 (187)

and WAM P.14762-3 (2, 122-132), Rottnest I. ANSP 148694 (234) and

ANSP 178913 (3, 170-186) Perth vicinity. AMNH 55688 (2, 187-200)

and ANSP 135419 (2, 145-139), Fremantle, Cockbum Sound. MNHN

5856 (217), holotype of Caranx platessa, “mer des Indies,” probably

Geographe Bay. WAM P.25064-004 (4, 45-90), Hardy Inlet. WAM

P.20253 (193), Broke Inlet. WAM P.21824-25 (2, 160-162), Cherryup,

34°58'S, 117°26'E. WAM P.20026 (219), Wilson Inlet. MNHN 5854 (2,

142-166), syntypes of Caranx georgianus. King George Sound. AMS

IA.653 (210), Albany, King George Sound. WAM P.21673 (193),

Brewer Bay, 34°24'S, 119°26'E. WAM P.21681 (153), Stokes Inlet.

ANSP 148693 (2, 102-174) and ANSP 148697 (7, 72-132), Duke of

Orleans Bay. ANSP 138196 (2, 193-200), Cape Arid. ANSP 138194

(189) and ANSP 138195 (2, 11^170), Isralite Bay. ANSP 138197

(192), Coffin Bay. ANSP 49316-18 (3, 181-185), Melbourne. ANSP

135420 (82 C&S), TMH D.197 (132), TMH D.530 (4, 90-108), TMH

D.531 (189), TMH D.774 (272), TMH D.830 (187), TMH D.1029 (67),

USNM 222104 (15, 140-227), Tasmania. BMNH 1896.6.17.49-52 (4,

135-495), Flinders L, 33°43'S, 134°31'E. AMS 1.19890-002 (3, 201-

263), Nadgee Nature Reserve, 37°26'S, 149°54'E. AMS 1.28734-002 (4,

191-210), Green Cape, Bittangabee Bay. ANSP 153525 (8, 156-180)

Jervis Bay. USNM 177016 (2, 255-294), New South Wales, no specific

locality. ANSP 138198 (2, 220-246), ANSP 147861 (2, 265-276),

ANSP153773 (6,315-378),USNM385513 (275), Sydney Fish Market.

ANSP 135421 (2, 94-95 C&S), CAS-SU 7433 (2, 149-150), CAS-SU

8321 (163) Sydney. ANSP 147826 (2, 144-162) 24 km E. Sydney

Harbour. AIM 262 (88), holotype of Usacaranx archeyi, N side of

Motuihi L, Hauraki Gulf, New Zealand.

? Scomber dentex'QXoch and Schneider, 1801: 30 (orig. descr.; Rio

de Janeiro, Brazil; holotype ZMB 14112).

? Usacaranx insulanorum Whitley, 1937: 223, pi. 13, fig. 2 (orig.

descr.; Elizabeth Reef; holotype lost).

Caranx nobilis. Grant, 1982: 302, color pi. 147 (brief descr., Qld).

Pseudocaranx dentex. Randall et al., 1990: 164 (misident. in part;

brief descr.).

Material examined. 38 specimens, 162-860 mm FL. Qld, Australia:

CAS 16472 (7, 404-432), One Tree L; QMB 1.13732 (321), Noosa,

26°23'S, 153°07'E; QMB 1.19416 (296), Cape Moreton. Lord Howe L:

AMS 1.5761-001 (673); AMS 1.7395-006 (18, 162-199); AMS 1.17178-

045 (195); AMS 1.17395-015 (2, 181-197); AMS 1.23674-001 (860);

CAS-SU 9158 (2, 170-198); BPBM 14833 (4, 145-210).

Remarks. Unlike P. georgianus from southern Australia and

New Zealand, no evidence of hyperostosis of the supraoccipital

is apparent in any of the large specimens available from One

Tree I. In addition to having more gill rakers (Table 2), adults

from off Queensland and Lord Howe Island do not have the

blunt snouts (fig. 3B) that are characteristic of many large

individuals from New Zealand (Ayling and Cox, 1982, pi. 19)

and southern Australia (Ogilby, 1893, pi. 24). As discussed

under “Remarks” for P. dinjerra, sheared PCA analysis

revealed consistent but subtle differences between all three

Pseudocaranx groupings (fig. 5).

The fish from Queensland and Lord Howe Island listed above

likely represent a fourth Australian Pseudocaranx species but

final determination requires additional study, especially

Australian trevallies of the genus Pseudocaranx

103

^ R dinjerra

O P. georgianus

A P sp. '"dentex"'

PC2

Figure 5. Sheared principal component analysis of 22 measurements

for 3 Pseudocaranx taxa. PC2 loadings were associated with snout

and upper jaw lengths, while PC3 was associated with straight

lateral line and anal-fin base lengths; see Material and methods

section for discussion of measurements used.

mitochondrial DNA analyses. James, 1980 reported limited

movement of tagged Pseudocaranx off north-east New Zealand,

so it is possible that fish from Queensland and Lord Howe Island

are isolated from those in southern Australia and New Zealand.

We are currently unable to distinguish them from P. dentex from

the Atlantic Ocean and South Africa, orfrom JapanPx^wt/ocarawv

sp. with 15 caudal vertebra. Two other antitropical Indo -Pacific

carangids, Seriola lalandi Valenciennes and ''Caranx'' equula,

also have similarly wide disjunct distributions, so this

zoogeographic pattern is not unique. If Atlantic Pseudocaranx

are indeed conspecific with those from Queensland offshore

reefs and Lord Howe Island, then the oldest available name for

them is P. dentex. If a different scientific name is required, one

possibility is Usacaranx insulanorum Whitley. That nominal

species was described (Whitley, 1937) from Elizabeth Reef

based on a single individual. Unfortunately, the holotype is not

extant (Eschmeyer, 1998) and no additional specimens are

available from the type locality that would allow detailed

comparisons. The original description included no vertebral

count but the reported high number of gill rakers (14-1-25) agrees

with specimens from One Tree and Lord Howe Islands.

Grant’s, 1982 photograph (colour pi. 147) of a postmortem

specimen from Noosa Heads, Queensland shows a fish with

mostly yellow dorsal and anal fins and a faint yellow mid-

lateral body stripe. The Pseudocaranx color photograph (Plate

IV-19) in Randall et ah, 1990 is misleading because it was

taken at Easter Island (Randall, pers. comm.).

Pseudocaranx wrighti Whitley, 1931

Eigures lA, 3D, 4; Tables 1-3

Usacaranx georgianus wrighti Whitley, 1931: 317 (orig. descr. 40

mi. west of Kingston, South Australia; Holotype AMS 1.10336).

Pseudocaranx wrighti. May and Maxwell, 1986; 301, unnumb,

photo, (descr.; common name; distrib.); Paxton et ah, 1989: 582

(synonymy; distrib.); Kuiter, 1993: 176, unnumb, color photos (descr.;

distrib.); Gomon et ah, 1994: 585, unnumb, color photo, (descr.;

distrib.); Hutchins, 1997: 247 (listed; Houtman Abrolhos); Hutchins,

2001b: 33 (listed).

Material examined. 125 specimens, 46-196 mm FL. CSIRO C.2751

(119), Exmouth Gulf; ANSP 134668 (4, 105-143), ANSP 135418 (8,

141-169), WAM P.14007-18 (11, 100-135), WAM P.14019-26 (8, 110-

144), WAM P.14043-53 (11, 119-150), WAM P.14055 (112), Rottnest I.

vicinity; ANSP 153537 (6, 150-176), ANSP 182762 (4, 172-196),

Perth vicinity; WAM P.27679 (182), Swan-Avon R.; AMS 1.10336

(122.5, holotype of Usacaranx georgianus wrighti), W of Kingston,

36°50'S, 139°20'E; AMNH 37652 (2, 169-181), Cockburn Sound,

Fremantle; ANSP 148691 (30 of 61, 46-159), Sepia Depression off

Garden L, 32°08'S, 115°37'E; SAMA 196 (2, 116-139), Doubtful I.

Bay; ANSP 145073 (186), Coffin Bay; SAMA 3110 (2, 120-122),

SAMA 3147 (144), off Port Lincoln; SAMA 4777 (145) Adelaide outer

harbour; SAMA 3930 (17, 97-125), St. Vincent Gulf; AMS 1.20194-

032 (9, 92-123), Investigator Strait; SAMA 3620 (164), Kangaroo L;

AMS 1.10397 (125), Flinders L, 33°45'S, 134°30'E; TMH D.535 (2,

94-103), Flinders L, 39°50'S, 148°00'E.

Diagnosis. A species of Pseudocaranx with posterior margin

of upper jaw canted posteroventrally (fig. lA); lachrymal and

expanded part of maxilla densely covered with scales; caudal

vertebra 14; gill rakers 10-15 upper, 24-28 lower, 35-43 total;

scales in curved part of lateral line 37-48.

Distribution. Endemic to Australia (fig. 4); eastern Bass Strait

extending W at least to Rottnest I., WA. A single collection of

P. wrighti from Exmouth Gulf (see material examined) at

approximately 22°S extends its distribution well into that of P.

dinjerra, if the record is not due to a specimen mix-up.

Remarks. This species differs from other Australian congeners

as indicated in the identification key and Table 3. It also differs

in having a well defined opercular spot that is nearly solid black

and approximately the diameter of the pupil (fig. 3D), in never

developing a yellow mid-lateral body stripe, and second dorsal

and anal fins transparent or dusky green, never with yellow

pigmentation. Pseudocaranx wrighti is the smallest species of

Pseudocaranx, rarely exceeding 200 mm EL.

Acknowledgements

We thank the curators and collection managers of the

ichthyological collections cited herein for specimen loans and

other courtesies during visits to their respective institutions.

Michael Walker, Western Australian Eisheries Department,

also provided specimens that were critical to this study. Thanks

are due to Sandra J. Raredon and Kyle Luckenbill for digital

specimen images, Barry J. Hutchins for providing color slides

of Australian Pseudocaranx, and both he and Rudie Kuiter for

sharing their observations on Pseudocaranx and patiently

answering our questions. Tim M. Berra provided help in

104

William F. Smith-Vaniz and Howard L. Jelks

Table 1. Frequency distributions of segmented fin rays and caudal vertebrae in Australian species of Pseudocaranx.

Dorsal-fin rays

Anal-fin

rays

Species

SD 18

wrighti

23.9

0.9 1

20.7

0.7

dinjerra

24.3

0.6

20.6

0.6

georgianus

sp. '"dentex”

26.4

0.8

22.7

0.8

Queensland

25.8

0.4

21.7

0.5

Lord Howe I.

25.7

0.7

22.0

0.5

Pectoral-fin

rays

Caudal vertebrae

Species

wrighti

18.1

0.4

113

dinjerra

18.7

0.5

georgianus

sp. '"dentex”

19.1

0.5

109

Queensland

20.1

0.3

Lord Howe I.

20.3

0.5

Table 2. Frequency distributions of developed gill rakers in Australian species of Pseudocaranx.

Upper gill rakers

Lower gill rakers

Species

7 8

wrighti

12.8

0.9

25.8

0.9

dinjerra

3 16

8.5

0.7

20.2

0.6

georgianus

sp. denteP"

10.1

0.9

21.9

0.8

Queensland

12.8

0.8

25.2

0.8

Lord Howe I.

12.7

0.6

25.8

0.6

Total

gill rakers

Species

wrighti

38.6

1.5

dinjerra

28.7

1.0

georgianus

sp. '"dentex”

32.0

1.4

Queensland

38.0

1.3

Lord Howe I.

38.7

1.0

selecting the scientific name for the new species. The first

author is also grateful for an Australian Museum Visiting

Museum Fellowship that provided much of the impetus for this

research. The manuscript benefited from comments of Martin

F. Gomon, Steven J. Walsh, and two anonymous reviewers.

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Memoirs of Museum Victoria

Volume 63 Number 1 2006

1 > A new species of Neamia (Perciformes: Apogonidae) from the West Pacific Ocean

Thomas H. Fraser and Gerald R. Allen

7 > Description of two new species of Nesogoblus (Pisces: Gobioidei: Gobiidae) from southern Australia

Douglass F. Floese and Helen K. Larson

15 > Description of a new species of dwarf Philypnodon (Teleostei: Gobioidei: Eleotridae) from

south-eastern Australia

Douglass F Hoese and Sally Reader

21 > Description of a new species of Heterocllnus (Blennoidei: Clinidae) from southern Australia

By Douglass. F Hoese and Denise S. Rennis

25 > Description of two new species of shore-eels (Gobiesocidae: Cheilobranchinae: Alabes) from

south-eastern Australia and Norfolk Island

Barry Hutchins

29 > Revision of the genus Hapalogenys (Teleostei: Perciformes) with two new species from the

Indo-West Pacific

Yuklo IwatsukI and Barry C. Russell

47 > Two new species of Parapercis (Perciformes: Pinguipedidae) from north-eastern Australia, and

rediscovery of Parapercis colemani Randall & Francis, 1993

Jeffrey W. Johnson

57 > SImIpercIs trispinosa, a new genus and species of sandperch (Perciformes: Pinguipedidae) from

eastern Australia

Jeffrey W. Johnson and John E. Randall

65 > Notoraja hirticauda, a new species of skate (Chondrichthyes: Rajoidei) from the south-eastern

Indian Ocean

Peter R. Last and John D. McEachran

77 > Nomenclature and distribution of the species of the porcupinefish family Diodontidae (Pisces,

Teleostei)

Jeffrey M. Lels

91 > A new species of Zenopsis (Zeiformes: Zeidae) from the South China Sea, East China Sea and

off Western Australia

TetsujI Nakabo, Dianne Bray and UmeyoshI Yamada

97 > Australian trevallies of the genus Pseudocaranx (Teleostei: Carangidae), with description of a

new species from Western Australia

William F Smith-Vaniz and Howard L Jelks

Contents