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A New Species of Mudskipper, Periophthalmus takita (Teleostei: Gobiidae:Oxudercinae), from Australia, with a Key to the GenusAuthor(s): Zeehan Jaafar and Helen K. LarsonSource: Zoological Science, 25(9):946-952. 2008.Published By: Zoological Society of JapanDOI: http://dx.doi.org/10.2108/zsj.25.946URL: http://www.bioone.org/doi/full/10.2108/zsj.25.946

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© 2008 Zoological Society of JapanZOOLOGICAL SCIENCE 25: 946–952 (2008)

A New Species of Mudskipper, Periophthalmus takita(Teleostei: Gobiidae: Oxudercinae), from Australia,

with a Key to the Genus

Zeehan Jaafar1* and Helen K. Larson2

1Department of Biological Sciences, National University of Singapore, Singapore 1175432Museum and Art Gallery of the Northern Territory GPO Box 4646,

Darwin NT 0801, Australia

A new species of mudskipper (Gobiidae: Oxudercinae) from northern Australia is described. This species was previously misidentified as P. novaeguineaensis Eggert, 1935. Periophthalmus takita sp. nov. can be distinguished from its congeners by the following suite of characters: modally VIII spines in the first dorsal fin; second dorsal and anal fins with I, 11–12 rays; shape and color pattern of the first and second dorsal fins; and the extent of fusion between the two innermost rays of the pelvic fin. A re-diagnosis of P. novaeguineaensis is provided, as well as a revised key to the genus Periophthalmus.

Key words: Periophthalmus, Oxudercinae, Gobiidae, mudskipper, new species

INTRODUCTION

The gobiid fish species Periophthalmus cantonensis novaeguineaensis was first described by Eggert (1935) based on specimens from Merauke, Irian Jaya, Indonesia. In his revision of the Oxudercinae, Murdy (1989) discovered that the type specimen of P. novaeguineaensis, deposited at Universität Tübingen, was destroyed during World War II. Murdy (1989) proceeded to designate a lectoype (ZMA 112.466) and 21 paralectotypes (ZMA 112.945) from south-ern Irian Jaya and treated Periophthalmus expeditioniumWhitley, 1953, from Queensland as its junior synonym.

Larson and Takita (2004) recognized three different spe-cies within P. novaeguineaensis as described by Murdy (1989). They described two of these species as new (P. darwini and P. murdyi) and re-diagnosed P. novaeguineaensis based on material from northern Australia. The lectotype of P. novaeguineaensis Eggert was not available for their study, but that they did see Murdy’s notes on the specimen. They also examined the holotype of P. expeditionium Whitley and considered it to be a separate species from what they considered to be P. novaeguineaensis.

Recent examinations of a number of Periophthalmus type specimens by the senior author revealed that Perioph-thalmus murdyi Larson and Takita, 2004, is in fact a junior synonym of P. novaeguineaensis. The species referred to as P. novaeguineaensis in Larson and Takita, and figured in Murdy (1989: Fig. 41, Pl. 2F), is here recognized as an undescribed species, known only from northern Australia. Periophthalmus expeditionum is shown to be a junior syn-onym of the true P. novaeguineaensis.

Here we describe the new species from northern Australia and re-diagnose P. novaeguineaensis. We also include a key to the genus Periophthalmus Bloch and Schneider, 1801, modified from Larson and Takita (2004) and Murdy (1989).

MATERIALS AND METHODS

Morphological measurements and meristic counts follow those of Murdy (1989) with one exception: pectoral fin length [PL] does not include the fin base. Measurements were made with vernier cal-lipers to the nearest 0.1 millimeter and are expressed as a percent-age of the standard length of the fish [SL] unless otherwise stated.

Counts of the second dorsal and anal fins distinguished between segmented and unsegmented rays, with the last two rays counted as a single element, as they share the ultimate pterygiophore. Pre-dorsal scales were counted in a straight line starting from the scales anterior to the first dorsal spine forward to posterior of the interor-bital. Counts of the longitudinal scale series follow the method used by Murdy (1989) but stopped at the hypural crease. Transverse scale counts forward were made from the anal fin origin anterodor-sally to the to first dorsal fin base. Specimens used in this study are lodged at the Museum and Art Gallery of the Northern Territory, Australia (NTM); Australian Museum, Sydney, Australia (AMS); Raf-fles Museum of Biodiversity Research, Singapore (ZRC); and Zoo-logical Museum of Amsterdam, The Netherlands (ZMA).

SYSTEMATICS

Periophthalmus takita nov. sp.(Figs. 1–5, Tables 1, 2)

Periophthalmus novaeguineaensis Murdy 1989: 42, Fig. 41, Pl. 2F (in part).Periophthalmus novaeguineaensis, Larson and Takita, 2004: 181–183, Figs. 7E, 7F, 9 (in part).

Materials ExaminedHOLOTYPE: NTM S10418-006, Cameron’s Beach,

Shoal Bay, Northern Territory, 26 February 1982, 65.2 mm.

* Corresponding author. Phone: +65-65162969;Fax : +65-67792486;E-mail : dbszj@nus.edu.sg

doi:10.2108/zsj.25.946

New Species of Mudskipper 947

PARATYPES: NTM S10418-004, data as for holotype, nine specimens (30.5–55.0 mm); ZRC 50769, data as for holo-type; three specimens (36.8–53.9 mm); AMS I.25515-001, Port Hedland, Six Mile Creek, Western Australia, seven specimens (35.8–64.6 mm); NTM S10410-003, Ludmilla Creek, Darwin, Northern Territory, 29 December 1981, four specimens (35.6–44.7 mm); NTM S14637-032, south end of Field Island, Northern Territory, 27 May 1998, six specimens (21.4–60.1 mm).

Fig. 1. Holotype of Periophthalmus takita, NTM S.10418-006, 65.2 mm SL, Cameron’s Beach, Northern Territory, Australia.

Fig. 2. Paratype of Periophthalmus takita, NTM S.10418-004, 55.4 mm SL, Cameron’s Beach, Northern Territory, Australia.

Fig. 3. First dorsal fin of Periophthalmus takita, paratype, NTM S.10418-004, 55.4 mm SL, Cameron’s Beach, Northern Territory, Australia.

Fig. 4. Second dorsal fin of Periophthalmus takita, paratype, NTM S.10418-004, 55.4 mm SL, Cameron’s Beach, Northern Territory, Australia.

Fig. 5. Pelvic fin of Periophthalmus takita, holotype, NTM S.10418-006, 65.2 mm SL, Cameron’s Beach, Northern Territory, Australia.

Table 1. Meristic data for Periophthalmus takita (n=30).

HolotypeNTM S10418-006

Range

Fin-ray countsFirst dorsal fin IX VII to XISecond dorsal fin I,12 I,11–12Anal fin I,12 I,10–12Pectoral fin 13 13–15Segmented caudal-fin rays 16 16–18

Scale countsLateral row 64 58–73Transverse rows back 17 16–22Transverse rows front 17 17–24Predorsal 23 22–31

Table 2. Morphometric data for Periophthalmus takita (n=30).

HolotypeNTM S10418-006

Range Mean±SD

In % SLHead length 28.1 23.3–29.3 27.4±1.3Head depth 14.0 12.8–16.5 14.3±1.1Body depth 10.9 9.3–13.8 11.0±2.5Predorsal length 25.0 19.8–26.8 24.8±1.5Length of D1 base 17.5 16.8–21.8 18.2±4.0Length of D2 base 23.2 21.3–26.3 24.3±1.3Gap between D1 & D2 7.4 2.5–7.6 5.4±3.2Length end D2 to caudal fin 15.2 10.3–19.3 14.4±1.9Anal fin base 21.2 18.3–21.9 20.1±1.0Caudal peduncle depth 8.4 5.0–8.7 7.3±0.8Caudal fin length 22.1 19.2–28.9 23.7±2.5Pectoral base height 6.4 4 .5–7.4 6.2±0.7Pectoral fin length 19.6 15.6–27.0 18.9±4.7Pelvic fin length 14.7 11.0–15.0 13.1±1.1

In % pelvic fin length

Extent of fusion of innermost pelvic rays 59.4 41.5–75.6 57.4±9.7

Z. Jaafar and H. K. Larson948

DiagnosisFirst dorsal fin spines modally VIII; second dorsal fin

rays modally I, 12; anal fin rays modally I, 11; caudal fin modally with 17 segmented rays; lateral scales 58–73; pre-dorsal scales 22–31. Pelvic fin with prominent frenum; sparse, small brown spots on dorsal surface. Inner rays of pelvic fin fused for half but never more than three-quarters of their length. Head and body pale grey, whitish ventrally; small, rounded, blackish spots on cheeks, pectoral fin base, and anterior part of body; characteristic round orange spots along lower half of head and body (less numerous posteri-orly). First dorsal fin broadly rounded, grey to black, with very thin, white margin restricted to tips of spines. Second dorsal fin whitish grey with narrow pale marginal stripe, two broad black bands, one sub-marginal and one medial, and a basal row of black spots. Anal fin pale orange to creamy. Pectoral fins pale with fine dusky markings mostly following fin rays. Caudal fin dusky to pinkish grey.

DescriptionTables 1 and 2 present meristic and morphometric data;

Figs. 1 and 2 are photographs of the holotype and one of the paratypes, respectively. Larson and Takita (2004: Fig. 9) illustrated a freshly dead specimen (as P. novaeguineaen-sis).

A robust Periophthalmus with compressed body and somewhat cylindrical head. Background color of head and body pale grey to light greyish-brown, becoming cream-colored ventrally; rounded blackish to dark grey spots present on cheeks, pectoral fin base, and anterior portion of body; characteristic round orange spots (becoming brown in preservative) along lower half of head and body, becoming less numerous posteriorly on body. Short vertical, silvery white lines along lower half of body, mostly post-abdominal; lines usually persist in preserved specimens. Branchiostegal membranes pale to dusky. Six dark brown irregular saddle-like bars crossing dorsum, coursing anteroventrally: first bar just anterior to first dorsal fin origin, second at middle of first dorsal fin, third (may be indistinct) at second dorsal fin ori-gin, fourth at middle of second dorsal fin, fifth at rear of sec-ond dorsal fin, and sixth at origin of caudal procurrent rays. First dorsal fin (Fig. 3) tall and broadly rounded; first dorsal fin elements never reaching second dorsal fin when depressed; fin reddish grey, grey basally in life, becoming black to grey upon preservation; thin whitish to creamy mar-ginal stripe (may be restricted to tips of spines); fin color uni-form in some specimens but in others becoming darker dis-tally. Second dorsal fin (Fig. 4) with narrow white marginal stripe, often restricted to tip of spines; two broad black bands, located sub-marginally and medially; and basal row of black blotches, which may appear to merge and form a basal line. Pelvic fins with rounded tips and prominent fre-num; innermost rays united for about half to three-quarters of their length. Anal fin yellow to orange in life; unmarked in preservation. Caudal fin dusky grey to pinkish grey, yellow-ish white along ventral margin. Pectoral fins with margins of fin rays finely spotted with grey; hyaline upon preservation. Pelvic fin (Fig. 5) yellowish to greyish-orange in life; hyaline in preservation, with occasional small, sparse, brown spots on dorsal surface.

EtymologyThis species is named for our colleague Dr. Toru Takita,

in recognition of his contributions to the knowledge of mud-skipper ecology, used as a noun in apposition.

DistributionCurrently known only from the Northern Territory and

Western Australia.

RemarksThe re-diagnosis of P. novaeguineaensis provided by

Larson and Takita (2004) and the figures in Murdy identified as P. novaeguineaensis (Murdy, 1989: Fig. 41, Pl. 2F) refer to this species. Murdy included some material of this spe-cies in his account of P. novaeguineaensis, including the fol-lowing specimens: NTM S.10418-004, Shoal Bay; NTM S.10410-003, Ludmilla Creek, Darwin; NTM S.10421-002, Elizabeth River mouth, Darwin Harbour; NTM S.10426-002, Buffalo Creek (Murdy, 1989; Larson and Takita, 2004). Due to the confusion involved with these two species, we list in Table 5 the identity of various specimen lots used in older and the current treatments. However, we were unable to examine all of the specimens used in Murdy’s study.

Periophthalmus takita is similar to P. novemradiatus Eggert, 1935, in having two dark bands on the second dor-sal fin, but differs in the following; VIII spines modally in the first dorsal fin (versus IX in P. novemradiatus), a larger gap between the first and second dorsal fins (versus 0.6–1.6%), 22–31 predorsal scales (versus 18–24), six saddle-like bars across dorsum (versus eight bars) and first spine of first dorsal fin not elongate in males (versus first spine elongate in males).

Periophthalmus novaeguineaensis Eggert, 1935(Figs. 6–12, Tables 3, 4)

Periophthalmus cantonensis novaeguineaensis Eggert, 1935: 61, Fig. 2, Pl. 3, fig. 12.Periophthalmus expeditionium Whitley, 1953: 127, Fig. 2.Periophthalmus murdyi Larson and Takita, 2004: 178, Figs. 3–6, 7C, D, 8D–G, Tables 3, 4.

Material ExaminedPeriophthalmus cantonensis novaeguineaensis. Lecto-

type, ZMA 112.466, New Guinea, Merauke-Fluss, 41.9 mm; paralectotype, ZMA 112.945, New Guinea, Merauke-Fluss, in part, 18 of 21 specimens (31.5–46.7 mm).

Periophthalmus expeditionium. Holotype, AMS I.6195, Karumba, Queensland, Australia, 39.6 mm.

Periophthalmus murdyi. Holotype, NTM S.11193-005, northeast of Harrison Dam, Adelaide River, Northern Territory, 38 mm. Paratypes: NTM S.11193-004, Barramundi Farm, Adelaide River, Northern Territory, 4 October 1993, six speci-mens (34.6–43.7 mm); NTM S.14024-013, small creek north side of Roper River, Northern Territory, 8 September 1994, six specimens (32.1–43.6 mm); NTM S.15800-001, mudflats under jetty in Derby, Western Australia, 19 July 2002, six spec-imens (33.3–44.3 mm); NTM S.14467-006, mud and gravel pools on rock bar at Smith’s Landing, East Alligator River, Northern Territory, 3 June 1997, nine specimens (32.0–42.0 mm); NTM S.10426-002, upstream Buffalo Creek, Northern Territory, 21 April 1982, 17 specimens (32.0–41.0 mm). Other

New Species of Mudskipper 949

material, AMS I.5552-016, Gulf of Carpentaria, Norman River, Queensland, Australia, two specimens (35.8–39.4 mm).

DiagnosisFirst dorsal fin spines modally XI; second dorsal fin rays

modally I, 12; anal fin rays modally I, 11; caudal fin modally with 17 segmented rays; lateral scales 71–110; predorsal scales 25–32. Inner rays of pelvic fin always united for more than three-quarters of their length. First dorsal fin red-brown or black, with broad, black submarginal stripe and white dis-tal margin; posterior edge concave, giving triangular shape to fin. Second dorsal fin with whitish margin and one broad, black sub-marginal band and basal row of black spots. Usu-ally no markings on anal, pectoral, or caudal fins; small brown spots or dusky pigment along length of fin or fin rays in some individuals. Pelvic fin with prominent frenum, heavily pigmented on dorsal surface, and with narrow, yel-lowish to whitish marginal band.

DescriptionTables 3 and 4 present morphometric and meristic data,

respectively; Figs. 6–8 are photographs of the lectotype of P. novaeguineaensis, and holotypes of P. expeditioniumand P. murdyi, respectively. Larson and Takita (2004: 178–181, Figs. 3–6) provided a full description in their account of P. murdyi as well as color photographs of live and freshly dead specimens.

Basal color of head and body light brown to fawn, becoming cream-colored ventrally. Seven dark brown saddle-like bars on body crossing dorsum (persistent in pre-servative) and three or four irregular lines or thin bands crossing dorsal surface of head; markings on dorsum of head variable in intensity and may be absent in some spec-imens. Bars across dorsum on body coursing anteroven-trally: first bar commencing anterior to first dorsal fin origin, second at middle of first dorsal fin, third at end of the first dorsal fin, fourth after origin of second dorsal fin, fifth at rear of second dorsal fin, sixth at origin of caudal procurrent rays, and seventh at hypural crease. Side of head with two or three dark, irregular lines or narrow bands; most con-spicuous line running posterodorsally to front end of upper maxilla and may branch at preopercle, resulting in reticu-lated pattern. When alive, lower half of head and body with very small, bluish-white spots and short, vertical white lines present on flanks; latter persisting in preserved specimens. First dorsal fin (Fig. 9) triangular, dorsal fin elements never

Table 3. Meristic data for Periophthalmus novaeguineaensis(n=21).

HolotypeZMA 112.466

Range

Fin-ray countsFirst dorsal fin X IX-XIVSecond dorsal fin I,12 I,12–13Anal fin I,10 I,10–11Pectoral fin 12 12–15Segmented caudal-fin rays 17 16–18

Scale countsLateral row 79 71–110Transverse rows back 18 17–28Transverse rows front 21 18–30Predorsal 27 25–32

Table 4. Morphometric data for Periophthalmus novaeguineaensis(n=21).

LectotypeZMA 112.466 Range Mean±SD

In % SLHead length 25.6 24.3–26.5 25.8±0.7Head depth 13.6 11.8–15.3 14.0±0.8Body depth 11.3 9.3–12.4 10.7±1.3Predorsal length 26.1 24.5–27.2 25.3±1.2Length of D1 base 19.1 14.0–21.7 18.5±2.2Length of D2 base 24.4 21.2–25.2 23.3±1.4Gap between D1 & D2 5.8 2.5–8.5 5.2±1.8Length end D2 to caudal fin 14.8 11.3–17.1 14.2±1.8Anal fin base 17.6 17.5–21.2 18.4±1.2Caudal peduncle depth 6.0 5.2–7.2 6.4±0.5Caudal fin length 21.1 19.2–24.6 20.7±5.1Pectoral base height 5.8 4.2–7.1 5.4±0.8Pectoral fin length 17.1 13.1–19.3 16.7±2.0Pelvic fin length 11.6 9.3–12.5 11.7±0.7

In % pelvic fin length

Extent of fusion of innermost pelvic rays 80.4 77.4–90.9 82.9±3.0

Fig. 6. Lectotype of Periophthalmus novaeguineaensis, ZMA 112.466, 41.9 mm SL, Merauke, New Guinea, Indonesia.

Fig. 7. Holotype of Periophthalmus expeditionium, AMS I.6195, 39.6 mm SL, Karumba, Australia.

Fig. 8. Holotype of Periophthalmus murdyi, NTM S.11193-004, 38.0 mm SL, Adelaide River, Northern Territory, Australia.

Z. Jaafar and H. K. Larson950

reaching second dorsal fin when depressed; fin uniformly black or grey in preservative, with broad, white marginal band. Second dorsal fin (Fig. 10) with one broad, black sub-marginal band and basal row of black spots. Anal, caudal, and pectoral fins hyaline, with occasional small brown spots or pigment along fin elements. Pelvic fin (Figs. 11, 12) rounded with prominent frenum; innermost rays united for more than three-quarters of their length. Pelvic fins brown to blackish anterodorsally, marginal band lacking pigment.

DistributionCurrently known from Australia and Indonesia.

RemarksPeriophthalmus novaeguineaensis is distinguished from

P. takita in having XI spines modally in the first dorsal (versus VIII), one dark band on the second dorsal fin (versus two), and the innermost rays of the pelvic frenum joined for more than three-quarters of the length (versus joined half to three-quarters).

Periophthalmus novaeguineaensis sensu Murdy (1989) consisted of three species: P. novaeguineaensis Eggert, 1935; P. darwini Larson and Takita, 2004; and P. takita nov. sp. (Table 5). Periophthalmus novaeguineaensis sensu Larson and Takita (2004) is the new species described above as Periophthalmus takita. Three specimens (33.8–35.6 mm SL) from the paralectotype series of P. novaeguineaensis(ZMA 112.945) were found to be P. darwini.

Key To Species of Periophthalmus Bloch & SchneiderThe key is modified from Larson and Takita (2004) and

Murdy (1989). Abbreviations used are as follows: D1, first dorsal fin; D2, second dorsal fin; A, anal fin; C, caudal fin, and LS, longitudinal scale counts.

1a. Frenum uniting pelvic spines either prominent or vesti-gial. If vestigial, visible without magnification ............. 2

1b. Frenum absent, or visible only with magnification.... 14 2a. Innermost pelvic fin rays joined by membrane for the

entire length to form a rounded disk........................... 3 2b. Innermost pelvic fin rays not joined by membrane for

entire length................................................................. 5 3a. Distal margin of D1 straight; D2 and A greater than I,

10 and I, 9, respectively.............................................. 4 3b. Distal margin of D1 rounded; D1 uniformly dark brown

or black with white distal margin; D2 rays I, 9–11; A rays I, 8–11 (Thailand to Singapore) ....................................... P. walailakae Darumas and Tantichodok, 2002

4a. Whitish to bluish spots on head and body when alive; D2 rays I, 13 or 14 rays; A rays I, 12–14 (Sumatra to Peninsular Malaysia) ................................................................................... P. spilotus Murdy and Takita, 1999

4b. Orange spots on head and body when alive; D2 rays I,

Fig. 9. First dorsal fin of Periophthalmus novaeguineaensis, NTM S.14024-013, 39.6 mm SL, Roper River, Northern Territory, Australia.

Fig. 10. Second dorsal fin of Periophthalmus novaeguineaensis, NTM S.14024-013, 39.6 mm SL, Roper River, Northern Territory, Australia.

Fig. 11. Pelvic fin of Periophthalmus novaeguineaensis, Lecto-type, ZMA 112.466, 41.9 mm SL, Merauke, New Guinea, Indonesia.

Fig. 12. Pelvic fin of Periophthalmus novaeguineaensis, NTM S.14024-013, 39.6 mm SL, Roper River, Northern Territory, Australia.

New Species of Mudskipper 951

11 or 12; A rays I, 10–12 (India to Java) ............................................................P. chrysospilos Bleeker, 1852

5a. D1 greatly reduced in both sexes; gap between D1 and D2 always greater than D1 base; D1 with V spines modally (IV–VI) (north-western Australia) ................................................... P. darwini Larson and Takita, 2004

5b. Gap between D1 and D2 never greater than D1 base..................................................................................... 6

6a. D2 with two dark bands .............................................. 7 6b. D2 with one dark band................................................ 8 7a. D1 broadly rounded; D1 with VIII spines modally (VII–

XI); C segmented rays 17 modally (15–17); gap bet-ween D1 and D2 more than 2% of SL; pre-dorsal scales 22–31 (Australia)..................... P. takita sp. nov.

7b. D1 concave to straight; first spine in D1 greatly elon-gate in males; D1 with IX spines modally (VIII–X); C segmented rays 15 modally (13–15); gap between D1 and D2 less than 2% of SL; pre-dorsal scales 18–24 (India and northern Thailand).......................................................................P. novemradiatus (Hamilton, 1822)

8a. Height of D1 greater than body depth; D1 margin con-vex; D1 with prominent submarginal black band (Korea)...........................................................................

..............P. magnuspinnatus Lee, Choi and Ryu, 1995 8b. D1 height less than or equal to body depth; pattern and

color of D1 not as above............................................. 9 9a. D1 with no spots ....................................................... 10 9b. D1 with few or no spots ............................................ 1210a. D1 with numerous white spots; D1 with XI–XV spines;

pelvic frenum vestigial; LS typically more than 70 (east Africa to western Pacific) ......... P. kalolo Lesson, 1830

10b. D1 with XI or fewer spines; pelvic frenum prominent; LS typically fewer than 70......................................... 11

11a. D1 with orange spots turning black in preservative; D1 with VIII–XI spines; branchiostegal membrane heavily pigmented; ventral peritoneum densely black (Sumatra to Philippines) ....................... P. variabilis Eggert, 1935

11b. D1 with numerous white spots; branchiostegal mem-brane not pigmented; ventral peritoneum lightly pig-mented (Indonesia, Philippines) ............................................................................ P. malaccensis Eggert, 1935

12a. D1 plain gray or with few white spots basally; D1 with narrow black margin or row of black spots; D2 with one dark submarginal band (Arabian Gulf to Pakistan)................................................... P. waltoni Koumans, 1941

12b. D1 with no spots and distal white margin................. 13

Table 5. Periophthalmus specimens cited in Murdy (1989) and Larson and Takita (2004) and their present status.

NTM Cat. No. Identification Source Present assignment

S.462 P. novaeguineaensis Murdy 1989 ? (number is an error)S.10408-002 P. novaeguineaensis Murdy 1989 P. takitaS.10410-003 P. novaeguineaensis Murdy 1989 P. takitaS.10410-003 P. novaeguineaensis Larson and Takita 2004 P. takitaS.10418-004 P. novaeguineaensis Murdy 1989 P. takitaS.10418-004 P. novaeguineaensis Larson and Takita 2004 P. takitaS.10421-002 P. novaeguineaensis Murdy 1989 P. takitaS.10421-002 P. novaeguineaensis Larson and Takita 2004 P. takitaS.14024-013 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.10426-002 P. novaeguineaensis Murdy 1989 P. novaeguineaensisS.10426-002 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.10429-025 P. novaeguineaensis Murdy 1989 2 spec. = P. takita; 1 spec. = P. darwiniS.10472-018 P. novaeguineaensis Murdy 1989 P. darwiniS.10554-001 P. novaeguineaensis Murdy 1989 P. darwiniS.10554-001 P. darwini Larson and Takita 2004 P. darwiniS.10554-004 P. darwini Larson and Takita 2004 P. darwiniS.10649-007 P. novaeguineaensis Murdy 1989 P. takitaS.10694-001 P. novaeguineaensis Murdy 1989 14 spec. = P. darwini, 27 spec. = P. sp. unident.S.10694-022 P. darwini Larson and Takita 2004 P. darwiniS.10718-056 P. novaeguineaensis Murdy 1989 P. takitaS.10989-002 P. novaeguineaensis Murdy 1989 P. takitaS.11114-002 P. novaeguineaensis Murdy 1989 P. novaeguineaensisS.11193-004 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.11193-005 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.11360-015 P. novaeguineaensis Murdy 1989 P. darwiniS.11360-015 P. darwini Larson and Takita 2004 P. darwiniS.13475-001 P. novaeguineaensis Larson and Takita 2004 P. takitaS.13496-003 P. novaeguineaensis Larson and Takita 2004 P. takitaS.14024-013 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.14400-006 P. darwini Larson and Takita 2004 P. darwiniS.14467-006 P. murdyi Larson and Takita 2004 P. novaeguineaensisS.14637-032 P. novaeguineaensis Larson and Takita 2004 P. takitaS.15492-002 P. novaeguineaensis Larson and Takita 2004 P. takitaS.15801-001 P. murdyi Larson and Takita 2004 P. novaeguineaensis

Z. Jaafar and H. K. Larson952

13a. A broad dark stripe at 2/3 of D2 height; D2 with basal row of dusky spots on interradial membranes; caudal fin narrow and pointed posteriorly (southern New Guinea to Australia) ..................................................................................... P. novaeguineaensis Eggert, 1935

13b. A broad dark stripe at mid D2; D2 with basal row of dusky spots on fin rays; caudal fin broad and rounded posteriorly (China to Japan).......................................................................................P. modestus Cantor, 1842

14a. D1 and D2 contiguous in adult males; D1 large and tri-angular in males, greatly reduced in females, barely perceptible in some; D2 lacking stripes (Papua New Guinea and Australia) ..............P. weberi Eggert, 1935

14b. D1 and D2 not contiguous in either adult males or females, D1 not reduced in females; D2 with a single dusky stripe ............................................................... 15

15a. D1 with white or black spots; LS typically fewer than 90................................................................................... 16

15b. D1 lacking or with few white spots posteriorly; LS typi-cally more than 90 (tropical western Africa) .........................................................P. barbarus (Linnaeus, 1766)

16a. D1 rounded, with prominent black spot posteriorly; D1 with not more than X spines (Malaysia, Indonesia, Singapore, Australia, Philippines) .................................................................................. P. gracilis Eggert, 1935

16b. D1 pointed and lacking black spot; D1 with more than X spines..................................................................... 17

17a. D1 with black stripe inframarginally; peritoneum densely black ventrally; LS 75 or more (western Indian Ocean to Oceania) .................................................................................. P. argentilineatus Valenciennes, 1837

17b. D1 with light brown stripe inframarginally; peritoneum lightly pigmented ventrally; LS 75 or fewer (Andaman Islands, Thailand, Indonesia, Australia, Philippines, Peninsular Malaysia) ..............P. minutus Eggert, 1935

ACKNOWLEDGMENTS

We thank Professor Peter Ng for his support and comments on this manuscript. We are grateful to Ronald Vonk, Mark McGrouther, Sally Reader, and Amanda Hay for the loan of specimens. This study was supported by grant R154-000-155-112 from the National University of Singapore to Professors Chou Loke Ming and Peter Ng.

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(Received September 5, 2007 / Accepted July 29, 2008)