A new species of damselfish (Chrysiptera: Pomacentridae) from Papua New Guinea and eastern Indonesia.
The pomacentrid genus Chrysiptera Swainson, 1839, contains small, often brightly coloured fishes that are common inhabitants of coral and rocky reefs in the Indo-Pacific region. Allen (1991) recognized 25 species, but an additional six species have since been described (Allen & Adrim 1992; Randall 1994; Allen & Rajasuriya 1995; Allen 1999; Allen & Bailey 2002; Allen & Erdmann 2008). The members of this genus (formerly known as Glyphidodontops Bleeker, 1877) are relatively elongate (2.1-2.7 in SL for most species, although the complex of species treated herein usually 1.9-2.2 in SL) pomacentrids, with smooth preopercular and suborbital margins, and most species possess biserial dentition. Based on preliminary genetic studies it appears the group is polyphyletic and probably composed of several genera. However, we prefer to recognise a single genus pending comprehensive genetic studies of intrafamilial relationships.
The present paper describes a new species that has been previously misidentified (e.g. Allen 1991) as C. parasema (Fowler, 1918). However, Allen & Erdmann (2008) demonstrated there are at least three distinctive species confused under this name: the true C. parasema ranging from the Ryukyu Islands southward to the Philippines and western Indonesia, C. giti Allen & Erdmann, 2008, from south-western New Guinea and the Togean Islands, Indonesia, and an undescribed species from northern New Guinea and far north Sulawesi, which is described herein.
MATERIALS AND METHODS
Lengths of specimens are given as standard length (SL) measured from the anterior end of the upper lip to the base of the caudal fin (posterior edge of hypural plate); head length (HL) is measured from the same anterior point to the posterior edge of the opercle flap; body depth is the maximum depth taken vertically between the belly and base of the dorsal spines; body width is the maximum width just posterior to the gill opening; snout length is measured from the anterior end of the upper lip to the anterior edge of the eye; orbit diameter is the horizontal fleshy diameter, and interorbital width the least fleshy width; upper jaw length is taken from the front of the upper lip to the posterior end of the maxilla; caudal peduncle depth is the least depth, and caudal peduncle length is the horizontal distance between verticals at the rear base of the anal fin and the caudal fin base; lengths of fin spines and rays are measured to their extreme bases (not from the point where the ray or spine emerges from the basal scaly sheath); caudal fin length is the horizontal length from the posterior edge of the hypural plate to a vertical at the tip of the longest ray; caudal concavity is the horizontal distance between verticals at the tips of the shortest and longest rays; pectoral fin length is the length of the longest ray; pelvic fin length is measured from the base of the pelvic spine to the filamentous tip of the longest soft ray; pectoral ray counts include the small splint-like, uppermost rudimentary ray; only the tube-bearing anterior lateral-line scales are counted; a separate count is given for the deeply pitted scales occurring in a continuous series midlaterally on the caudal peduncle; the decimal figure "1.5" appearing in the scale row count above the lateral line refers to a complete scale (1) and small truncated scale (0.5) at the base of the dorsal fin; gill raker counts include all rudiments and are presented as separate counts for the upper and lower limbs as well as a combined count; the last fin ray element of the dorsal and anal fins is usually branched near the base and is counted as a single ray.
Counts and proportions for the holotype are followed by the range for paratypes in parentheses. Proportional measurements expressed in thousandths of the SL are provided in Table I. Counts for dorsal rays, anal rays, pectoral rays, total gill rakers on the first arch, and tubed lateral-line scales are presented in Table II. Type specimens are deposited at the Australian Museum, Sydney (AMS), Pusat Penelitian dan Pengembangan Oseanologi, Jakarta, Indonesia (NCIP), National Museum of Natural History, Washington, D.C. (USNM), and the Western Australian Museum, Perth (WAM).
Table I. Proportional measurements of selected type specimens of Chrysiptera arnazae as percentage of the standard length. Character Holotype Paratype Paratype Paratype Paratype NCIP WAM NCIP USNM WAM 6334 P.32921 6335 391578 P.32913 Standard 37.8 39.1 36.1 34.0 32.7 length (mm) Body depth 19.5 20.9 16.8 16.3 17.5 Body width 7.5 7.9 6.6 6.3 6.8 Head length 11.9 13.0 12.3 11.3 10.7 Snout length 3.3 4.2 3.3 3.0 2.8 Orbit 4.4 4.8 4.3 3.9 4.2 diameter Interorbital 4.1 4.8 4.0 3.6 3.8 width Caudal 5.8 6.0 5.7 5.3 5.2 peduncle depth Caudal 4.7 5.1 4.5 4.8 4.2 peduncle length Predorsal 3.6 3.9 3.5 3.3 3.3 length Preanal 14.5 14.6 13.6 13.0 13.9 length Prepelvic 23.7 24.9 24.0 22.2 21.6 length Length dorsal 14.1 15.3 14.2 13.3 12.7 fin base Length anal 23.7 24.7 21.5 20.4 20.3 fin base Length 10.2 10.0 8.8 8.6 8.9 pectoral fin Length pelvic 11.8 12.1 11.0 11.3 10.4 fin Length pelvic 12.0 11.7 10.6 9.8 11.1 spine Length 1st 7.1 7.5 6.5 6.6 6.4 dorsal spine Length 2nd 3.0 3.8 2.7 2.2 3.1 dorsal spine Length 6th 7.9 7.9 7.1 7.1 7.0 dorsal spine Length 5.9 6.5 5.6 5.5 6.3 longest dorsal ray Length 1st 9.6 9.0 8.7 8.3 8.2 anal spine Length 2nd 3.4 4.2 3.6 3.2 2.8 anal spine Length 7.6 8.3 7.3 7.1 6.7 longest anal ray Length caudal 9.2 8.9 9.0 8.3 8.5 fin Caudal 10.5 9.0 10.6 8.5 10.4 concavity Character Paratype USNM 391578 Standard 28.0 length (mm) Body depth 14.3 Body width 5.5 Head length 9.9 Snout length 2.6 Orbit 4.2 diameter Interorbital 3.2 width Caudal 4.1 peduncle depth Caudal 3.6 peduncle length Predorsal 3.0 length Preanal 11.4 length Prepelvic 18.6 length Length dorsal 11.5 fin base Length anal 17.9 fin base Length 7.2 pectoral fin Length pelvic 9.0 fin Length pelvic 8.1 spine Length 1st 5.4 dorsal spine Length 2nd 2.4 dorsal spine Length 6th 5.6 dorsal spine Length 4.1 longest dorsal ray Length 1st 7.1 anal spine Length 2nd 2.4 anal spine Length 5.5 longest anal ray Length caudal 6.9 fin Caudal 7.9 concavity Table II. Frequency of counts in soft dorsal rays, soft ana rays, pectoral rays, total gill rakers on first arch, and tubec lateral-line scales of Chryslptera arnazae. Counts for pectoral rays and lateral-line scales were usually recorded for both sides of each individual (n = 18). Dorsal Anal Pectoral rays rays rays 9 10 10 11 13 14 15 2 16 1 17 4 25 4 Gill Lateral-line rakers scales 22 23 24 11 12 13 14 15 4 12 2 3 7 10 12 4
We also conducted a genetic analysis of the new species and its relatives. Multiple tissue samples of C. springeri, C. hemicyanena, C. giti, C. parasema, and the new taxon were collected from various locations across the West Pacific. Genomic DNA was extracted using 10% Chelex (Walsh et al 1991), then mtDNA control region DNA sequence data was collected by PCR amplification using primers CRA and CRE (Lee et al 1995) followed by sequencing on an ABI 3730 using BigDye 3.0 terminator chemistry (Applied Biosystems) following protocols described in Drew et al. 2008. Forward and reverse sequences were proofread in Sequencher (GeneCodes, Corp), then aligned in TCoffee (Poirot et al 2003) with subsequent alignment by eye. Unweighted parsimony analysis was conducted via the heuristic search option in PAUP* 4.0b10 (Swofford 2002) with 1000 bootstrap replicates to assess clade support. Maximum likelihood analyses were conducted in PAUP* 4.0b10 (Swofford 2002) employing a heuristic search with model parameters determined from ModelTest Server 3.8 (Posada 2006) with 100 boostrap replicates to assess clade support.
Chrysiptera arnazae n. sp.
(Figs 1-3; Tables I-III)
Table III. Comparison of diagnostic colour pattern differences of members of the "hemlcyanea complex" of Chryslptera species. Species Ground Anal fin Pelvic fin colour colour colour C. arnazae blue & yellow & yellow * yellow blue C. cymatilis blue blue blue C. giti blue & yellow blue yellow C. hemicyanea blue & yellow yellow yellow C. oxycephala pale green white white C. parasema blue & blue blue yellow C. pricei blue & white white white C. sinclairi blue blue blue C. springeri blue blue blue C. species A blue blue blue * pelvic fins of C. arnazae rarely one-half to two-thirds blue.
Chrysiptera parasema (non Fowler). Allen, 1991: 97 (in part; underwater photo of individual from Madang, Papua New Guinea).
Chrysiptera parasema (non Fowler). Allen et al. 2003: 74 (in part; underwater photo).
Chrysiptera cf. parasema. Kuiter & Tonozuka 2001: 396 (underwater photo of individual from Maumere, Flores).
Holotype. NCIP 6334, 37.8 mm SL, Pulau Lemon, 00[degrees]53.395'S, 134[degrees]04.782'E, near Manokwari, Papua Barat Province, Indonesia, 6-10 m, clove oil and hand net, M. Erdmann, 23 October 2007.
Paratypes: AMS I.44560-001, 3 specimens, 22.6-30.5 mm SL, Hayne Harbour Lagoon, 02[degrees]03.048'S, 147[degrees]25.532'E, Los Negros Island, Manus, Papua New Guinea, 8 m, clove oil and net, G. Allen, 26 October 2006; NCIP 6335, 2 specimens, 32.7-35.5 mm SL, collected with holotype; USNM 391578, 3 specimens, 28.0-34.9 mm SL, Saripa Bay, 00[degrees]07.002'S, 130[degrees]21.866'E, Waigeo, Raja Ampat Islands, Papua Barat Province, Indonesia, 10-12 m, clove oil, M. Erdmann, 22 August, 2007; WAM P. 27827-026, 29.2 mm SL, Hayne Harbour Lagoon, 02[degrees]03.048'S, 147[degrees]25.532'E, Los Negros Island, Manus, Papua New Guinea, 7-9 m, rotenone, G. Allen & R. Knight, 6 October 1982; WAM P.29624-088, 4 specimens, 29.2-36.8 mm SL, Kranket Lagoon, 05[degrees]11.724' S, 145[degrees]44.641'E, Madang, Papua New Guinea, hand nets, 5-10 m, G. Allen, 15 October 1987; WAM P.31144-004, 3 specimens, 21.2-29.3 mm SL, Susan's Reef, approximately 05[degrees]18'S, 150[degrees]08'E, Kimbe Bay, West New Britain Province, Papua New Guinea, rotenone, 7-8 m, G. Allen, 20 April 1996; WAM P.32921-001, 39.1 mm SL, collected with holotype; WAM P.32913-001, 36.1 mm SL, collected with USNM paratypes.
Diagnosis: A species of the pomacentrid genus Chrysiptera with the following combination of characters: dorsal rays XIII,10 (rarely 9); anal rays II,11 (rarely 10); pectoral rays usually 14 (occasionally 13 or 15); gill rakers on first branchial arch 6-8 + 14-17 (usually 7 + 16), total rakers 22-24; tubed lateral line scales 11-15 (usually 12-14); colour in life brilliant blue except abruptly yellow behind vertical line extending from last dorsal spines to about anterior half of anal fin; pelvic fins usually yellow with narrow blue anterior margin (rarely with one-half to two-thirds of fin blue).
Description: Dorsal rays XIII,10 (one paratype with XIII,9); anal rays II,11 (one paratype with III,10); pectoral rays 14 (13-15); gill rakers on first branchial arch 7 + 16 (6-8 + 14-17), total rakers 23 (22-24); lateral line scales with tubes 14/15 (11-15); scales in lateral series from upper rear margin of opercle to base of caudal fin 27; scales above lateral line to base of middle dorsal spines 1.5; scales below lateral line to anus 9.
Body depth 1.9 (1.9-2.2) in SL; maximum body width 2.6 (2.3-2.7) in depth; HL contained 3.2 (2.8-3.2) in SL; snout 3.6 (3.1-4.1), eye 2.7 (2.4-3.1), interorbital space 2.9 (2.7-3.7), least depth of caudal peduncle 2.1 (2.1- 2.5), length of caudal peduncle 2.5 (2.1-2.8), all in HL.
Mouth terminal, oblique, jaws forming an angle of about 30[degrees] to horizontal axis of head and body; maxillary reaching to vertical through anterior edge of eye; teeth of jaws biserial, those of outer row more or less incisiform with flattened tips, upper jaw with about 34 (32-34) teeth and lower jaw with about 38 (34-38) teeth in outer rows, largest about one-third diameter of pupil in height; secondary row of slender buttress teeth behind those of outer row in the spaces between them; single nasal opening on each side of snout; nostril with low fleshy rim; preorbital and suborbital relatively narrow, greatest depth of suborbital about one-half pupil diameter, ventral margin smooth; margin of preopercle smooth, without any denticulations; margin of opercular series smooth except blunt, flattened spine on upper portion near angle.
Scales of head and body finely ctenoid; preorbital, suborbital, snout tip, lips, chin and isthmus naked; pair of primary transverse scale rows on cheek with row of smaller scales along lower margin; dorsal and anal fins with basal scaly sheath; basal half of caudal fin covered by scales; pectoral fins covered by scales only at base; axillary scale cluster between base of pelvic fins about two-thirds length of pelvic spine.
Tubed lateral line scales ending below posterior spines of dorsal fin; pits or pores present on 5 (4-6) scales immediately posterior to last tubed scale; continuous series of 6 (6-8) pored or pitted scales mid-laterally on caudal peduncle to caudal base.
Origin of dorsal fin at level of third tubed scale of lateral line; spines of dorsal fin gradually increasing in length to sixth or seventh spine, remaining spines slightly decreasing in length; membrane between spines deeply incised; first dorsal spine 4.0 (3.3-5.1), seventh dorsal spine 1.5 (1.5-1.8), last dorsal spine 2.0 (1.7-2.6), longest soft dorsal ray 1.2 (1.3-1.5), all in HL; length of dorsal fin base 1.6 (1.6-1.7) in SL; first anal spine 3.5 (3.0-4.2), second anal spine 1.6 (1.5-1.8), longest soft anal ray 1.3 (1.3-1.5), all in HL; base of anal fin 2.3 (2.3-2.6) in base of dorsal fin; caudal fin emarginate with rounded lobes, its length 3.6 (3.1-4.3) in SL; pectoral fin reaching vertical through origin of anal fin, longest ray 3.2 (3.0-3.5) in SL; filamentous tips of pelvic fins reaching slightly beyond origin of anal fin when undamaged (tips broken off in holotype and most paratypes), longest ray 3.2 (2.8-3.6) in SL.
Colour in life (Figs 1 and 3): generally brilliant blue with vertical grey streak on each scale, becoming abruptly yellow posterior to vertical line extending from last dorsal spines to about anterior half of anal fin; broad blackish or dark grey stripe from snout tip to middle of anterior edge of eye and narrower black stripe below from upper lip to lower anterior corner of eye; iris blue with blackish or dark grey stripe along anterior and ventral margins, also blackish or dark grey stripe through middle of eye (continuation of upper snout stripe); narrow blackish bar usually present on single scale at upper opercular edge; spinous portion of dorsal fin brilliant blue; caudal fin yellow basally with translucent outer half; anal fin blue anteriorly and mostly yellow on remainder of fin; pelvic fins usually yellow with narrow blue anterior margin, rarely with expanded blue area covering one-half to two-thirds of fin (as in holotype); pectoral fins translucent with small blackish spot at base of uppermost 2-3 rays.
[FIGURE 1 OMITTED]
Colour in alcohol (Fig. 2): generally dark bluish grey, abruptly pale tan or whitish posterior to vertical line extending from last dorsal spines to about anterior half of anal fin; large bluish spots evident on centres of several cheek and opercle scales; spinous dorsal fin dark bluish grey; remaining fins translucent whitish except anterior half of anal fin dark grey; pelvic fins variable, anterior two-thirds dusky grey in holotype, but usually overall translucent whitish except narrow dark grey anterior margin.
[FIGURE 2 OMITTED]
The paratypes (WAM P.29624-088) from Madang, Papua New Guinea are similarly coloured, but exhibit a blue streak on most of the body scales, and a relatively large blue spot on each scale of the cheek and operculum. A dark grey stripe from the snout tip to the anterior edge of the eye is also evident.
Remarks: Chrysiptera arnazae belongs to a complex that contains nine other closely related species from the Indo-Australian Archipelago including C. cymatilis Allen, 1999, C. giti, C. hemicyanea Weber, 1913, C. oxycephala (Bleeker, 1877), C. parasema, C. pricei Allen & Adrim, 1992, C. sinclairi (Allen, 1987), C. springeri (Allen & Lubbock, 1976) and at least one additional undescribed species. The group is primarily restricted to the region that includes Indonesia, Philippines, New Guinea, and the Solomon Islands, ranging southward to offshore reefs of north-western Australia and northward to the Ryukyu Islands of Japan.
The members of the complex have a distinctive shape that is relatively deep-bodied (maximum depth 1.9-2.2 in SL) for the genus, a deeply incised spinous dorsal fin, and bright colouration largely consisting of blue or a combination of blue and yellow. Colour pattern differences (key to species below and Table II) in combination with gillraker and dorsal spine counts provide the best means of separation. The new species most closely resembles C. giti, C. hemicyanea, and C. parasema (Fig. 3). They are generally allopatric over most of the geographic range (Fig. 4), although C. hemicyanea cooccurs with C. giti at the Fakfak Peninsula of western New Guinea (Papua Barat Province, Indonesia), and C. arnazae is sympatric with C. parasema at eastern Flores in Indonesia. A study of genetic relationships (Barber et al., in preparation) within Chrysiptera reveals that the four species form a closely related, monophyletic clade.
[FIGURE 3 OMITTED]
[FIGURE 4 OMITTED]
Analysis of 389bp of mtDNA control region DNA sequence data from 21 samples representing six putative species of Chrysiptera revealed that C. arnzae is highly distinct from C. parasema, C. giti, C. hemicyanea, C. springeri and C. rollandi, forming a clade with 100% bootstrap support using both maximum parsimony and maximum likelihood (Fig. 5). Maximum intraspecific variation within C. arnazae was 2% uncorrected sequence divergence while minimum interspecific variation ranged to 12.7% uncorrected sequence divergence from C. giti from Fak Fak, Indonesia. While results suggest that C. arnazae is most clost related to C. giti, results also indicate that C. giti is polyphyletic, with samples from Amsterdam Island being a highly distinct sister clade to C. arnazae while C. giti from Fak Fak Indonesia, form a clade with C. hemicyaena and C. parasema, suggesting that further taxonomic study of this group is required.
Key to Chrysiptera species in the Hemicyanea complex (note: in preserved specimens blue colour is generally dark grey to greyish-brown, and yellow is whitish to tan). 1a Colour pattern mainly brilliant blue in 2 life without yellow or whitish areas 1b. Colour pattern not primarily blue, either 5 a combination of blue and yellow, blue and white, or mainly pale greenish 2a. Dorsal spines usually 12 3 2b. Dorsal spines usually 13 4 3a. Most of scales immediately below C. species A dorsal-fin base, and on upper half of caudal peduncle blackish (Philippines and Sabah) 3a. Most of scales immediately below C. springeri dorsal-fin base, and on upper half of caudal peduncle blue (eastern Indonesia) 4a. Total gill rakers on first arch 22-27 C. cymatilis (Bismarck Archipelago and Milne Bay Province of Papua New Guinea and Solomon Is.) 4ba. Total gill rakers on first arch 31-34 C. sinclairi (Bismarck Archipelago of Papua New Guinea) 5a. Head and body primarily blue with 6 variable amounts of yellow colouration including caudal fin and peduncle 5b. Head and body either blue or light green 9 over most of body, grading to white ventrally 6a. Yellow colouration confined to caudal fin C. parasema and adjacent peduncle; anal fin entirely bluish (W. Indonesia, Sabah, and Philippines) 6b. Yellow colouration not confined to caudal 7 fin and adjacent peduncle; anal fin entirely or partly yellow 7a. Breast and lower one-fourth of body C. hemicyanea yellow (offshore reefs of NW Australia, S. Sulawesi, and W. New Guinea, and Kei Is.) 7b. Breast and lower one-fourth of body blue 8 8a. Pelvic fins almost always yellow (rarely C. arnazae one-half to two thirds blue), posterior portion of body abruptly yellow behind vertical line extending from last dorsal spines to about anterior fourth of anal fin (N. New Guinea, Halmahera, and N. Sulawesi) 8b. Pelvic fins blue, posterior portion of C.giti body abruptly yellow behind oblique line extending from upper caudal-fin base to anal fin origin (Fak Fak Peninsula, W. New Guinea and Togean Islands off northeastern Sulawesi) 9a. Head and body blue on dorsal two thirds, C. pricei white on lower third and most of caudal peduncle (Cenderawasih Bay, W. New Guinea) 9b. Head and body mainly pale green, grading C. oxycephala to white on lower third and most of caudal peduncle, most of body scales with 2-3 vertically aligned small blue spots (Solomon Islands and Palau to E. Indonesia and Philippines)
[FIGURE 5 OMITTED]
Distribution and habitat: C. arnazae is currently known only from the following locations (proceeding westward): Papua New Guinea (New Ireland, New Britain, Manus, and Madang), Papua Barat Province (Manokwari and Waigeo in the Raja Ampat Islands), Halmahera, Lembeh Strait, Sulawesi, and Maumere, Flores. Most of the present distribution appears to be closely correlated with insular fragments of the former South Caroline Arc, a prominent paleogeographic feature in the Pacific Basin. Over the past 20 million years these fragments, including New Ireland, New Britain, Manus, Waigeo, and large masses that gradually accreted to the northern New Guinea mainland, drifted westward as a result of subduction of the Pacific Plate at the northern margin of the Australian Plate (Hill & Hall 2003; Polhemus 2007). The presence of C. arnazae at Flores, Halmahera and the northern tip of Sulawesi is more difficult to account for, but is probably a more recent phenomenon facilitated by the relatively short distances being bridged by island "stepping stones".
The habitat consists of sheltered fringing reefs, usually in bays at depths between about 3-20 m. It generally occurs in small aggregations and is closely associated with branching formations of live coral, particularly Acropora spp. and the pocilloporid Seriatopora hystrix. The fish hover a short distance above the substratum, presumably feeding on zooplankton.
Etymology: This species is named arnazae to honour the second author's wife and best friend, Arnaz Mehta, who has selflessly supported his extensive field time in the Bird's Head region.
We are grateful to Conservation International (CI), the Indonesian Department of Nature Conservation (PHKA), and especially to the Walton Family Foundation for sponsoring our taxonomic work on fishes of the Bird's Head region of western New Guinea. The first author is also indebted to The Nature Conservancy, especially staff member Alison Green, for supporting field work at New Britain, New Ireland, and Manus. Dan Polhemus provided valuable insight regarding the westward movement of Caroline Arc island fragments.
Received: 26 February 2010 - Accepted: 03 March 2010
ALLEN, G. R. 1991. Damselfishes of the world. Mergus Verlag, Melle, Germany, 271 pp.
ALLEN, G. R. 1999. Three new species of damselfishes (Pomacentridae) from Indonesia and eastern Papua New Guinea. Revue francaise d'Aquariologie 25 (3-4): 99-106.
ALLEN, G. R. 2009. Field guide to marine fishes of tropical Australia and south-east Asia. Fourth revised edition. Western Australian Museum, Perth, 287 pp.
ALLEN, G. R. & ADRIM, M. 1992. A new species of damselfish (Chrysiptera: Pomacentridae) from Irian Jaya, Indonesia. Records of the Western Australian Museum 16: 103-108.
ALLEN, G. R. & BAILEY, S. 2002. Chrysiptera albata, a new species of damselfish (Pomacentridae) from the Phoenix Islands, Central Pacific Ocean. Aqua, Journal of Ichthyology and Aquatic Biology 6 (1): 39-43.
ALLEN, G. R. & RAJASURIYA, A. 1995. Chrysiptera kuiteri, a new species of damselfish (Pomacentridae) from Indonesia and Sri Lanka. Records of the Western Australian Museum 17: 283-286.
DREW, J. A., KAUFMANN, L., ALLEN, G. R. & BARBER, P. H. 2008. Endemism and regonal color and genetic differences in five putatively cosmopolitan reef fishes. Conservation Biology 22: 965-976.
HILL, K. C. & HALL, R. 2003. Mesozoic-Cenozoic evolution of Australia's New Guinea margin in a west Pacific context. In: Evolution and Dynamics of the Australian Plate (Eds Hillis, R. R. & Muller, R. D.): 265-290. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372.
KUITER, R. H. & TONOZUKA, T. 2001. Indonesian Reef Fishes. Part 2, Zoonetics, Seaford, Victoria, Australia, pp. 304-622.
LEE, W. J., HOWELL, W. H. & KOCHER, T. D. 1995. Structure and evol ofution teleost mitochondrial control regions. Journal of Molecular Evolution 41: 54-66.
POLHEMUS, D. A. 2007. Tectonic geology of Papua. In: Ecology of Papua. Part One. (Eds Marshall, A. J. & Beehler, B. M.): 137-164. Periplus Editions, Singapore.
POITORT, O., O'TOOLE, E. & NOTREDAME, C. 2003. Tcoffee@igs: A web server for computing, evaluating and combining multiple sequence alignments. Nucleic Acids Research 31: 3503-3506.
POSADA, D. & CRANDALL, K. A. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817-818.
RANDALL, J. E. 1994. Two new damselfishes (Perciformes: Pomacentridae) from Arabian waters. Revue francaise d'Aquariologie 21 (1-2): 39-48.
SWOFFORD, D. L. 2002. PAUP. Phylogenetic analysis using parsimony (and other methods), version 4.0b10. Sinauer, Sunderland MA.
WALSH, P. S., METZGER, D. A. & HIGUCHI, R. 1991. Chelex-100 as a medium for simple extraction of DNA for PCR based typing from forensic material. Biotechniques 10: 506-513.
Gerald R. Allen (1), Mark V. Erdmann (2), and Paul H. Barber (3)
(1) Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: firstname.lastname@example.org
(2) Conservation International, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235 Indonesia
(3) Department of Ecology and Evolutionary Biology, University of California Los Angeles, 621 Charles E. Young Drive South, Los Angeles, CA 90095, USA
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|Author:||Allen, Gerald R.; Erdmann, Mark V.; Barber, Paul H.|
|Publication:||aqua: International Journal of Ichthyology|
|Date:||Apr 20, 2010|
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