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A new deepwater species of the genus Apogonichthyoides (Apogonidae) from the Timor Sea, north-western Australia.

INTRODUCTION

Relatively few cardinalfish are known from the deeper zones of the continental shelf (~61 m, 200 ft) and down the continental slope (~290 m, 951 ft). Most collections of deepwater apogonids have been taken by some type of trawl or trap. Some exceptional catches have been made by hook and line, divers using mixed gas rebreathers or research submarines. Apogon gouldi Smith-Vaniz, 1977, taken by submersible between 262 and 277 m (Smith-Vaniz et al. 1999) is the deepest dwelling representative of the apogonid fauna in the Atlantic Ocean. An Indo-west Pacific species, Ostorhinchus gularis (Fraser & Lachner, 1984), captured at 290 m by trawl, is the deepest-dwelling apogonid known to date. Other deep-dwelling apogonids likely await discovery by science.

The new species, part of a collection examined at Museum Victoria, Melbourne, Australia, was taken in a beam trawl on the bottom in 95 m with five other apogonids. The trawl contained a significant amount of reddish algae that may have cushioned these apogonids from extensive loss of scales, broken spines and other bodily damage and may have helped to retain them in the trawl. Specimens of the new species were retained as vouchers for DNA tissue samples at Museum Victoria and the Commonwealth Scientific and Industrial Research Organization, Hobart, Australia. This new description brings Apogonichthyoides Smith, 1949, as treated by Fraser & Allen (2010) to 20 species.

METHODS

Methods for making counts and measurements are given in Fraser (2005). In the dorsal spine formula, a spine inside parentheses (I) shows that this spine is hidden below the skin. Morphometric proportions as % of standard length (SL) are given for the holotype followed by the ranges for paratypes (Table I). Variation of meristic characters is presented in parentheses for paratypes. Standard lengths in millimeters are given in material examined lists unless otherwise noted. One specimen was cleared and stained for bone and cartilage using the method of Taylor and Van Dyke (1985). The descriptive formulae of Springer and Smith-Vaniz (2008) were followed for axil skeleton, supraneurals and pterygiophore insertion patterns (Table II). The last pterygiophore in both dorsal and anal fins is curved backwards, not obviously between neural and hemal spines. These last pterygiophores were counted as if they were in the next gap between spines. Gill raker counts were broken down into rudiments and rakers by arch. Oral incubation of eggs was checked by examining the mouth cavity of specimens with buccal enlargement. All digital photographs were processed through Adobe Photoshop CS4 software. Other figures were obtained from Wild camera lucida tracings, scanned on a flatbed scanner and completed in Photoshop. Acronyms used to identify collections from museums follow Leviton et al. (1985) and Fricke & Eschmeyer (2009).

Apogonichthyoides miniatus n. sp.

(Figures 1-4; Tables I-II)

[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]
Table I. Proportions as a percent of standard length for
Apogonlchthyoldes mlnlatus. The proportions for holotype
are followed by parentheses for paratypes.

Body proportions percent of standard length  A. miniatus N=15

greatest body depth                              46.3 (41-45)

head length                                      43.7 (42-43)

eye diameter                                     13.4 (13-15)

snout length                                      9.4 (10-12)

bony interorbital width                          13.1 (11-12)

upper jaw length                                 21.1 (20-23)

caudal peduncle depth                            15.7 (15-16)

caudal peduncle length                           21.4 (19-24)

1st dorsal fin spine length                         4.6 (3-6)

2nd dorsal fin spine length                      11.7 (11-13)

3rd dorsal fin spine length                      23.1 (21-23)

4th dorsa fin spine length                       22.6 (18-21)

9th dorsal fin spine length                      15.7 (14-17)

1st anal fin spine length                           4.0 (4-5)

2nd anal fin spine length                        16.0 (14-17)

pectoral fin length                              29.7 (23-28)

pelvic fin length                                30.0 (25-29)

Table II. Supraneurals and pterygiophore insertion pattern for the
cleared and stained Apogonlchthyoldes mlnlatus. 0 = supraneural,
/ = vertebral spines, 1, 2, 3,4 = number of pterygiophores between
spines, - = no pterygiophore, c = caudal vertebral complex.

                                     Insertion pattern

A.        dorsal      0/0/0-1/2/1/1/1/1/1/1/2/3/2/1/-/-/-/-/-/-/-/-/-/c
mlnlatus  anal    1 / 4 / 2 / 2 / 1 / - / - / - / - / - / - / - / - / c
10+14


Holotype: NMV A29729-003, 35.0 mm SL, 46.1 mm TL, 10.5 km NNE of Cartier Island, Timor Sea, L30 transect, 12[degrees]26'42"S to 123[degrees]36'03"E, to 12[degrees]26'58"S 123[degrees]36'35"E. RV Southern Surveyor, SS0507/188, beam trawl, 7 July 2006, 95 m, digital color photograph on board.

Paratypes: NMV A29729-039, (8, 15.2-34.4) same data as holotype. NMV A29729-054, (1, 33.0), same data as holotype, cleared and counter stained. NTM S.16659-001, (1, 31.3), same data as holotype. WAM P.33240-001, (1, 35.1), same data as holotype. CSIRO H 6597-04, (1, 30.9), same data as holotype, digital color photograph, DNA tissue as BW-A4844, GT2004. CSIRO H 6597-05, (11, 19.0-36.5) same data as holotype; DNA tissue (3, 19.0, 31.0, 36.5), DNA BW-A4845-4847, GT2008-2010. BPBM 41014, (1, 31.0), same data as holotype. CAS 229329, (1, 30.0) same data as holotype. USNM 398149, (1, 34.5), same data as holotype.

Comparative material:

Apogonichthyoides atripes - NMV3175 (1, 59.9), Australia, Queensland, 12[degrees]03'17"S 143[degrees]15'10"E; NMV R5961 (1, 71.9), Australia, Queensland, 19[degrees]58'S 148[degrees]41'E.

Apogonichthyoides brevicaudatus - syntypes of Apogon brevicaudatus, ZMA 101.127 (2, 35.4-100.7) Indonesia, Aru Is., Jedan I. AMS I.22831-027 (1, 80), Western Australia, Northwest Shelf, 140 km west of Port Hedland; AMS I.20771-016 (10, 55-93), Queensland, Cape York, E of Captain Billy Creek; AMS I.15557-133 (5, 74-84), Queensland, Gulf of Carpentaria; QM I.11202 (1,92), Queensland, 16[degrees]54'S 139[degrees]37'E, 11 m; USNM 329155 (1, 92), Northern Territory, Drysdale I., 11[degrees]33'30"S 135[degrees]52'30"E; WAM P.5854 (1, 72), Western Australia, Shark Bay, July 1963; WAM P 1101 (1, 71), Western Australia, Shark Bay; WAM P.14226 (1, 81), Northern Territory, Darwin, 9 September 1965.

Apogonichthyoides chrysurus - syntypes of Apogon chrysurus, AMS I.1799-1801 (3, 51.0-54.9), Australia, Lord Howe Island, 1888. USNM 213338 (3, 31-38), Queensland, One Tree I., 11 December 1966, X-ray.

Apogonichthyoides miniatus - AMS I.45000-001 (3), same data as holotype (not seen).

Archamia fucata - USNM 52203, (1, 57), cleared and stained.

Foa albimaculosa - holotype of Apogon albimaculosus, AMS IB.8347, (1, 65.0), Australia, Queens-land, near Gloucester Island, 20[degrees]01'S 148[degrees]15'E, September 1967, trawl, 18.3 m. Paratypes of Apogon albimaculosus, AMS I.15557-287 (1, 56); Gulf of Carpentaria, 17[degrees]25'S 140[degrees]10'E, Rama Station 410, 27 November 1963, 10.1 m. AMS I.16232-001 (2, 58-63), 3 miles N of Magnetic Island, Townsville, 19[degrees]10'S 146[degrees]50'E, June 1969, 21.9 m. AMS I.16895-001 (1, 60), Papua New Guinea, N of Yule I, 8[degrees]49'S 146[degrees]31'E, trawl, 13 February 1971, 36.6 m. AMS I.16896-001 (1, 55), Papua New Guinea, N of Yule I., 13 November 1970; 33 m.

Holapogon maximus - syntypes of Apogon maximus, BMNH 1889.4.15.21-27 (2, 171-198), Muscat, Oman.

Neamia octospina - WAM P.28022-021 (1, 39), Rowley Shoals, Clerke Reef, Western Australia, 17[degrees]18'S 119[degrees]22'E, 4 August 1983. WAM 28032.009 (2, 37-42), Rowley Shoals, Clerke Reef, Western Australia 17[degrees]18'S 119[degrees]22'E, 12 August 1983. AMS I. 33720-017 (1, 20.9), Ashmore Reef, Western Australia.

Quinca mirifica - UF 173741, (1, 51), Western Australia, aquarium reared, 12 October 2006, cleared and stained.

Vincentia badia - holotype, WAM P.25999.006, (1, 69.7), Western Australia, Rockingham, 29 January 1978.

Vincentia conspersa - syntype of Vincentia waterhousii, NMV 51865 (1, 72.4), Gulf of St. Vincent, Australia.

Vincentia punctata - syntype of Apogon punctatus, AMS I.19302-001 (1, 91.7), King Georges Sound, 35[degrees]03'S 117[degrees]58'E; 1878.

Diagnosis: An apogonine with nine visible dorsal spines, reddish head and body (pale in alcohol), dark spot on body in life (and in alcohol) above pectoral fin, 16 pectoral-fin rays, 7-8 well-developed gill rakers, 2-3 rows of palatine teeth and a rounded caudal fin.

Description: Range of proportions (as percentage of standard length) in Table I. Holotype data given first followed by that of paratypes in parentheses. Internal osteology from cleared and counter-stained paratype.

Dorsal fin IX spines as VIII-I,9 or (VII(I)-I,9), third spine longest and strongest, eighth spine shortest; sixth radial closely associated with pterygiophore supporting eighth spine (Fig. 2) but with a gap to free eighth spine; anal fin II,8; pectoral fin rays 16-16, two upper and three lower rays unbranched; pelvic fin I,5; longest procurrent caudal rays segmented, principal caudal rays 1+8-7+1; pored lateral line scales 23; transverse scale rows above lateral line 2; transverse scale rows below lateral line missing (6); median predorsal scales 2; cir median predorsal scales 2; circumpeduncular scale rows missing (12 as 5+2+5); total gill rakers and rudiments 14 (14-15), well-developed 7 (7-8), upper arch 2+1, lower arch 6+5 (6-7+5-6); second arch with tooth patches and rudiments; elongate interarcual cartilage extending from uncinate process of first epibranchial to lateral tip of second pharyngobranchial; first epibrachial with long, thin cartilage ending with ossified first pharyngobranchial.

Villiform teeth in band on premaxilla and dentary; 2-3 (1-2) rows on vomer; 2-3 rows on palatine; none on ectopterygoid, endopterygoid or basihyal.

Vertebrae 10+14; eight epineurals on first eight vertebrae; eight ribs on third to tenth vertebrae, last rib expanded; interhaemal gap 1+4 (Table I); five free hypurals (Fig. 3); one pair of slender uroneurals; three epurals; a free parhypural; three thin supraneurals, generally straight; two supernumerary spines on first dorsal pterygiophore; no procumbent spines (spurs); thin Y-shaped basisphenoid, free from parasphenoid and prootics; neurocranium with frontal tabs similar to Vincentia Castelnau, 1872 (Fraser 1972: pl 31); supramaxilla absent; posttemporal smooth or with 1-3 small serrae on posterior margin; preopercle with 2-3 small, widely spaced serrae near angle, ridge smooth; infraorbitals smooth; infraorbital shelf present on third bone; ring of scleral cartilage, no ossification present; 7 branchiostegals; ceratohyal notched, suture smooth with epihyal; urohyal with short anterior process.

Scales ctenoid on cheek, subopercle, missing on opercle, ctenoid on nape, with ctenoid or cycloid predorsal scales, ctenoid on breast, behind pectoral, abdomen and rest of body; pelvic fin base with two large scales, first ctenoid, second elongate and cycloid; no developed axillary scale; pored lateral line scales complete, second scale with one pore above and one below main canal (Fig. 4); last pored lateral line scale not elongate and with one pore above and one below main canal.

Head with many small pores, in poor condition in all specimens; anterior end of supraorbital canal as broad slit pore on tip of snout at edge of upper lip, large pore slightly above and medial to posterior nare, large pore over eye; lachrymal with large anterior pore near flat posterior nare opening, two large ventral slit pores along edge of lachrymal; anterior portion of dentary with dentary (anterior) and mental (ventral) pores, large articular pore; supratemporal canal with posterior canals extending almost to base of first dorsal fin over anterior predorsal scale.

Anterior naris tubular, posterior naris flat.

Caudal fin rounded; second dorsal and anal fin with rounded edge.

Post-mortem colors: See Figure 1A. Holotype: generally reddish on head, body and fins; head reddish with pale broad cheek mark below eye, iris brownish, snout, nape and preopercle reddish, opercle with silvery area; reddish body with nearly round darkish spot above pectoral fin and just behind opercular membrane, smaller than eye and larger than pupil; three faint, small, midline areas of melanophores evenly spaced behind body spot, lighter cast of reddish to yellowish areas on side becoming wide-spread on side of abdomen at and below base of pectoral fin to anal fin; first dorsal fin with darkish membranes between third and sixth dorsal spine forming spot-like marking, faint reddish to yellowish membranes below darkish area through eighth dorsal spine; second dorsal spine yellowish proximally and yellowish to pale distally with broad reddish area inbetween; pelvic fins with alternating pale yellowish and reddish markings on pelvic spine, pelvic rays reddish with blackish tips; anal fin pale yellowish proximally and reddish distally; caudal fin with pale area proximally adjacent to wide reddish band and pale distally.

Preserved color pattern: See Figure 1B. Holotype: head with faint cheek bar composed of melanophores, faint bar from upper eye edge posteriorly; darkish spot distally in first dorsal fin crossing membrane of third spine to part of sixth spine, mostly clear proximally, few melanophores in membranes between second and third spines and near base between fourth and fifth spines; second dorsal and caudal fins without melanophores, anal fin with distal melanophores on several fin rays; pelvic fin rays with distal melanophores on membranes and fin rays; large darkish spot above pectoral fin and just behind opercular membrane larger than iris and smaller than eye diameter on body, edging of lower half of spot round, upper half quasi-triangular; scattered melanophores extending dorsally as inverted quasi-triangular pattern, partially including lateral line scales; faint scattered bar-like melanophores on body behind spot, slightly more dense near midline, followed posteriorly by two faint, mostly mid-line melanophore patterns, one on body, one on midcaudal peduncle. Paratypes: all with similar patterns, large body spot above pectoral fin and just behind opercular membrane oval or slightly irregular on dorsal half with smaller mid-line melanophores in three patches on body and caudal peduncle; cheek mark as single or multiple lines of melanophores; faint bar from upper eye edge posteriorly; darkish spot distally in first dorsal fin crossing membrane of third spine to part of sixth spine, mostly clear proximally, few melanophores in membranes between second and third spines and near base between fourth and fifth spines; second dorsal and caudal fins without melanophores, anal fin with or without distal melanophores on several fin rays; pelvic fin rays with distal melanophores on membranes and fin rays; peritoneum pale, intestine pale, stomach blackish.

Etymology: The Latin word miniatus meaning bright red or cinnabar-red for the general coloration of the head, body and fins.

Distribution: Known only from 95 m, one collection in shelf waters of the Timor Sea, north-western Australia.

Remarks: Specimens were captured in a beam trawl which was towed for about 1.1 km on a course of about 117[degrees] true. Information gathered from a video transect across the path of station 188 showed sandy sediments, initially with pebbles interspersed with patches of bare sand and then outcrops of rocks with large sponges. Other summary data from the haul: target on hard ground, very good haul, 3-4 bins coral, sponges, urchins. Abundant red algae caught at this trawl station may have provided habitat cover for the new species. Photographs of five other apogonids were taken at the same station. These include Jaydia cf. poecilopterus (Cuvier in Cuvier & Valenciennes, 1828), Apogon cf. doryssa Jordan & Seale, 1906, Ostorhinchus cheni (Hayashi, 1990), Ostorhinchus franssedai (Allen, Kuiter & Randall, 1994), and Siphamia sp. All are probably new record depths for these species, except for Ostorhinchus cheni (WAM P.24796 at 142 m).

Apogonichthyoides miniatus has nine visible dorsal spines. The eighth spine has lost articulation with its radial. A few other species of apogonids have, or variably have, nine visible dorsal spines, deeply notched as VIII-I,9, the eighth spine much reduced in length: Foa albimaculosa (Kailola, 1976), Apogonichthyoides chrysurus (Ogilby, 1889), Holapogon maximus (Boulenger, 1888), Lepidamia species, Neamia octospina Smith & Radcliffe in Radcliffe, 1912), Vincentia badia Allen, 1987, V. conspersa (Klunzinger, 1872) and V. punctata (Klunzinger, 1879). Most cardinal fish genera and species have lost a visible eighth spine, either hidden under the skin as an extremely short spine, a hidden nubbin no longer spine-like, or complete loss of the ossified spine (Fraser 1972: pl. 8).

Caudal skeleton elements of Apogonichthyoides miniatus were the plesiomorphic set of ossified elements (Fraser 1972: table 4). The single pair of uroneurals were very thin and not mirror images, one lacked the anterior hook and was in two pieces.

Apogonichthyoides miniatus lacked scleral ossicles in the eye. A cartilaginous ring was present. These bones have been overlooked in previous apogonid studies. Franz-Odendaal (2008) noted that 94% of the families examined had a consistent ossicle number. Cardinalfish fall in the variable 6% of families with species having or lacking a pair of ossicles. Variation in ossicle size occurs among apogonids. For example, Quinca mirifica Mees, 1966, has small ossicles when compared with a similar sized Archamia fucata (Cantor, 1849). Franz-Odendaal (2008) suggests that there may be a relationship between swimming activity and the presence or absence of ossicles. Cardinalfish may be an excellent family with which to test this hypothesis.

The head-pore pattern of Apogonichthyoides miniatus is similar to that described by Fraser & Allen (2010) for A. umbratilis except that: 1) there is a large anterior pore slightly dorsal and medial to the posterior naris present in A. miniatus in addition to the large pore over the eye and the lateral slit pore on the tip of the snout present in both species; 2) Apogonichthyoides miniatus has a greater posterior expansion of supertemporal canal branches closer to the origin of the first dorsal spine than in A umbratilis.

Apogonichthyoides miniatus, a member of the nigripinnis group (which contains 11 species) by virtue of the large body spot, might be confused with A. atripes (Ogilby, 1916). The 15 mm SL specimen of Apogonichthyoides miniatus has a preserved color pattern like that of the adults without any evidence of bars on the body, but had a distally darker anal rays and the distal portion of the caudal fin. Kuiter & Kozawa (2001: 57) show figures of juvenile to adult Apogonichthyoides atripes. The 30 mm SL juvenile has a broad basicaudal bar, an ocellus, completely dark anal and first dorsal fins, darkish body, a pale caudal peduncle and the head is uniform except for a narrow cheek mark, all characters unlike any Apogonichthyoides miniatus of similar size. Adult Apogonichthyoides atripes have well-developed serrated opercular edges, are dark brownish except for the distal part of the soft dorsal fin and the caudal fin which are pale or pale yellowish, and they lack the short darkish post ocular bar extending above the posttemporal. Small Foa albimaculosa with a visible eighth dorsal spine and complete lateral line are less likely to be confused with Apogonichthyoides miniatus, having 18-19 pectoralfin rays, dark spots distally in the yellowishrimmed anal and second dorsal fins, alternating yellowish and blackish marks present in the caudal, anal and second dorsal fins, alternating yellowish and blackish marks present on the spine of the second dorsal fin, second anal spine, pelvic spine and first two dorsal spines, the iris with alternating dark and light marks as spokes radiating from the pupil, alternating dark and light bars on lips, and a short darkish post-ocular bar extending above the posttemporal (Kuiter & Kozawa 2001: 65). Neither of these two species has any evidence of red coloration in life nor has a pale body with a darkish spot and pale second dorsal and anal fins in alcohol, all characters of Apogonichthyoides miniatus. In alcohol, Apogonichthyoides atripes has a darkish body spot present along with a basicaudal bar, darkish dorsal and anal fins while Foa albimaculosa lacks the darkish body spot and has darkish dorsal and anal fins

A female (32.2 SL) had ova ~0.4 mm diameter. One mouth-brooding male, 34.4 SL, has eggs ~0.7 mm diameter in its mouth. The largest specimen was 36.5 SL, 47.0 mm TL.

ACKNOWLEDGEMENTS

A research fellowship from the Australian Museum, Sydney aided research on apogonids through Mote Marine Laboratory. Douglas Hoese, Jeffrey Leis, Mark McGrouther, Amanda Hay, Michelle Yerman and Sally Read of the Australian Museum provided assistance during my stay. Robert H. Robins of Florida Museum of Natural History, Gainesville, Florida, helped with museum loans from Australia. While in Australia, Dianne Bray and Martin Gomon of the National Museum Victoria, Melbourne, Helen Larson and Rex Williams of the Northern Territory Museum, Darwin, and Gerald Allen and Susan Morrison of the Western Australian Museum, Perth, provided assistance during my stay and helped with loans. Gerald Allen graciously discussed identities of a number of cardinal fishes. Alastair Graham of the Commonwealth Scientific and Industrial Research Organization, Hobart, Australia, provided information about the station, a color photograph and helped with a loan. Dianne Bray, who was onboard the research vessel, provided information about the station and a post-mortem color photograph. The survey SS05/2007 was undertaken on the Marine CSIRO Wealth from Oceans flagship with the assistance of the Commonwealth Department of the Environment, Water, Heritage and the Arts. Comments by two anonymous reviewers and the science editor were most useful.

Received: 19 March 2010 - Accepted: 14 April 2010

REFERENCES

FRANZ-ODENDAAL, T. A. 2008. Scleral ossicles of Teleostei: evolutionary and developmental trends. The Anatomical Record 291: 161-168.

FRASER, T. H. 1972. Comparative osteology of the shallow water cardinal fishes (Perciformes: Apogonidae) with references to the systematics and evolution of the family. Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology 34: 1-105.

FRASER, T. H. 2005. A review of the species in the Apogon fasciatus group with a description of a new species of cardinalfish from the Indo-West Pacific (Perciformes: Apogonidae). Zootaxa 924: 1-30.

FRASER, T. H. & ALLEN, G. R. 2010. Cardinalfish of the genus Apogonichthyoides Smith, 1949 (Apogonidae) with a description of a new species from the West-Pacific region. Zootaxa 2348: 40-56.

FRASER, T. H. & LACHNER, E. A. 1984. An unusual Indo-West Pacific cardinalfish of the genus Apogon (Teleostei: Apogonidae). Proceedings of the Biological Society of Washington 97: 632-636.

FRICKE, R. & ESCHMEYER, W. N. 2009. A guide to fish collections in the Catalog of Fishes database. On-line version updated 15 January 2010 http://research.calacad-emy.org/research/Ichthyology/catalog/fishcatmain.asp (accessed 2 February 2010).

KUITER, R. H. & KOZAWA, T. 2001. Apogonidae Pictorial Guide Fishes of the Indo-West Pacific. Zoonetics, Seaford, Australia 1-132.

LEVITON, A. E., GIBBS, JR., R. H., HEAL, E. & DAWSON, C. E. 1985. Standards in Herpetology and Ichthyology: Part I. Standard symbolic codes for institutional resource collections in Herpetology and Ichthyology. Copeia 1985(3): 802-832.

SMITH-VANIZ, W. F., COLLETTE, B. B. & LUCKHURST, B. E. 1999. Fishes of Bermuda: History, Zoogeography, annotated checklist, and identification keys. American Society of Ichthyologists and Herpetologists, Special Publication No. 4, I-x, 1-424.

SPRINGER, V. G. & SMITH-VANIZ, W. F. 2008. Supraneural and pterygiophore insertion patterns in carangid fishes, with description of a new Eocene carangid tribe, [dagger]Paratrachinotini, and a survey of anterior anal-fin pterygiophore insertion patterns in Acanthomorpha. Bulletin of the Biological Society of Washington 16: 1-73.

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Thomas H. Fraser

Mote Marine Laboratory, 1600 Ken Thompson Parkway, Sarasota, FL 34236-1096 USA. Email: cardinalfish@comcast.net
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Author:Fraser, Thomas H.
Publication:aqua: International Journal of Ichthyology
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Geographic Code:8AUST
Date:Jul 15, 2010
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