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Three new species of Rainbowfishes (Melanotaeniidae) from the Birds Head Peninsula, West Papua Province, Indonesia.

Abstract

Three new species of melanotaeniid rainbowfishes are described from fresh waters of the Birds Head Peninsula, West Papua Province, Indonesia. Melanotaenia ericrobertsi is described from 41 specimens, 18.4-52.1 mm SL, from the upper Kladuk River system. It was first collected in 1982 and included among the type series of M. irianjaya. However, recent genetic evidence reveals it is a distinct species, closely related to Melanotaenia from nearby drainages including M ajamaruensis, M. boesemani, M fasinensis, and two additional new species described in this paper, M. laticlavia and M. multi radiata. Melanotaenia laticlavia is described on the basis of 6 specimens, 33.769.6 mm SL from Aifuf Creek, and M multiradiata from 23 specimens, 37.4-122.5 mm SL from Sisiah Creek. These Birds Head species are primarily distinguished on the basis of distinctive adult male colour patterns and cytochrome b genetic analysis. However, M. laticlavia exhibits two separate patches of vomerine teeth, an unusual melanotaeniid feature, and M multiradiata has a relatively high pectoral-fin ray count (> 90 % with 15 rays or more).

Zusammenfassung

Beschrieben werden drei neue Arten der Regenbogenfis-che, Melanotaeniidae, aus Stiggewassern der Vogelkopf-Hal-binsel, Provinz Westpapua, Indonesicn. Melanotaenia eri-crobertsi n. sp. wird auf der Grundlage von 41 Exemplaren mit einer Lange zwischen 18,4 und 52,1 mm SL vom [degrees]berm lOaduk-Flusssystem beschrieben. Sic wurden 1982 erstmals gesammelt und der Typenreihe M irianjaya zugeordnet. Doch haben neuere genetische Untersuchungen ergeben, dass es sich urn eine eigene Art handelt, die den Melano-taeniidae aus nahe gelegenen Einzugsgebieten nahe ver-wandt ist, darunter M ajamaruensis, M boesemani, M fasi-nensis sowie zwei weitere Arten, die hier ebenfalls beschrieben werden: M laticlavia and M multiradiata. Melanotaenia Laticlavia wird auf der Grundlage von 6 Exem-plaren mit 33,7-69,6 mm SL vom Aifuf Creek beschrieben, wahrend M rnultiradiata anhand von 23 Exemplaren mit 37,4-122,5 mm SL vom Sisiah Creek beschrieben wird. Die drei zentralen Anal auf der Vogelkopf-Halbinsel lassen sich vor allem durch gut erkennbare Farbmerkrnale bei den Man-nchen unterscheiden sowie durch eine genetische Analyse des Cytochrom b. Hinzu kommt, class M latickvia an zwei getrennten Stellen Gaumenzahne besitzt, was unter Regen-bogenfischen ungewohnlich ist, und dass M multi radiatus eine relativ hohe Zahl an Flossenstrahlen an der Brustflosse aufweist (>90 % mit 15 Flossenstrahlen oder mehr).

Resume

Trois nouvelles especes de Melanotaeniides sont decrices en provenance d'eaux douces de la Birds Head Peninsula, province de West Papua, Indonesie. Melanotaenia ericro-bertsi n. sp. est dealt sur base de 41 specimens, 18,4 - 52,1 mm de LS, du bassin superieur dc la Kladuk River. Il a ete collecte d'abord en 1982 et inclus parmi les varietes types de M. irianjaya. Toutefois, une evidence genetique recente revele que c'est une espece distincte, tres proche de Mela-notaenia de drainages voisins comprenant M. ajamaruensis, M. boesemani, M. fasinensis, et deux autres nouvelles es-peces decrites dans cet article, M. laticlavia et M multira-diata. Melanotaenia laticlavia est decrit sur base de 6 specimens, 33,7 - 69,6 mm de LS, de l'Aiful Creek et M. mul-tiradiata, sus base de 23 specimens, 37,4 - 122,5 mm de LS, de Sisiah Creek. Les especes du centre de la Birds Flead se distinguent cssentiellement par des patrons de coloration differents pour les males adultes et par ranalyse genetique du cytochrome b. Toutefois, M. laticlavia pos-sede deux groupes separes de dents vomerines, tine carac-teristique inhabituelle pour les Melanotaeniides, et M. multi radiata a un nombre relativement &eve de rayons a la pectorale (> 90% avec 15 rayons ou plus).

Sommario

Tre nuove specie di pesci arcobaleno sono descritte dalle acque dolci dalla Birds Head Peninsula, Provincia di Papua Occidentale, Indonesia. Melanotaenia ericrobertsi n. sp. e descritta sulla base di 41 esemplari, 18,4-52,1 mm SL, dal bacino superiore del flume Kladuk. E stata raccolta la prima volta nel 1982 ed era compresa tra le serie tipo di M irianjaya. Tuttavia, una recente analisi genetica ha rivelato che si tratta di una specie distinta, strettamente legata alle specie di Melanotaenia dei bacini idrogeografici limitrofi, tra cui M. ajamaruensis, M. boesemani, M fasinensis e le due nuove specie descritte in questo articolo, M. laticlavia e M multiradiata. Melanotaenia laticlavia e descritta sulla base di 6 esemplari, 33,7-69,6 mm SL da Aifuf Creek, e M. multiradiata da 23 esemplari, 37,4-122,5 mm SL da Sisiah Creek. Le specie provenienti dalle zone centrali della Birds Head sono distinte soprattutto sulla base della carat-teristica colorazione dei maschi adulti e dell'analisi geneti-ca del citocromo b. Turtavia, M. laticlavia presenta due aree separate di denti vomerini, una caratteristica insolita nei melanotenidi e M. multiradiatus ha un numero relati-vamente alto di raggi pettorali (> 90% con 15 raggi o pitt).

INTRODUCTION

Melanotaeniids are one of only four endemic families (with Neoceratodontidae [extant species only], Lepidogalwddae and Pseudomugilidae) entirely confined to the fresh waters of Australia-New Guinea. Rainbowfishes are small (usually < 10 cm SL), colourful fishes which occur in a wide range of lotic and lentic habitats from clear, rapidly flowing streams to semi-stagnant mud holes. While many species have quite narrow ranges, they are the most widespread family within Australia-New Guinea where they are often rhe most abundant fish present (Unmack et al. 2013). Rainbowfishes thrive in captivity and thanks to their often brilliant colours, small size, and ease of breeding, are highly prized by aquarists. Allen and Cross (1982) and Allen (1995) published well-illustrated, reviews of the family. The latter work included accounts of 59 species or subspecies of melanotaeniids and 15 species of the closely allied Pseudomugilidae. However, in the intervening years since 1995, a host of new species have been discovered, particularly in New Guinea (Allen 8c Renyaan 1996 and 1998; Allen 1996, 1997, 1998, and 2001; Allen & Unmack 2008 and 2012; Allen et al. 2008 and 2014; Allen tic Hadiaty 2011 and 2013; Kadarusman et al. 2010, 2011 and 2012; and McGuigan 2001). The family currently contains 83 species belonging to seven genera (Es-chmeyer 2014), but many new species can be expected with future exploration, particularly in remote sections of New Guinea. The present paper desscribes three new species from the Birds Head Peninsula of West Papua Province, Indonesia. Two of these species were collected by the first author in 1982 and 1999, but were not recognized as distinct until recent genetic samples were obtained. The third species was provided by H.-G. Evers and J. Christian, who sampled extensively across the southern portion of the peninsula during October 2013.

L Recent phylogenetic evidence (Unmack et al. 2013) indicates that Melanotaenia contains three main lineages, western, northern and southern, correlated respectively with the following major biogeographic regions: the Birds Head Peninsula of western New Guinea, northern New Guinea, and the combined southern New Guinea/Australian region. The Birds Head region and adjacent Birds Neck of West Papua (Fig. 1) has proved a particularly fertile area for recent discoveries. Allen (1990) provided a summary of the seven species known to inhabit the Birds Head mainland at that time. A total of 21 species are currently known from the region, including the offshore islands of the Raja Ampat Group (Allen et al. 2014). The present paper describes three additional new Melanotaenia from the Birds Head Peninsula.

MATERIALS AND METHODS

Counts and measurements that appear in parentheses refer to the range for paratypes if different from the holotype. Type specimens are deposited at Museum Zoologicum Bogoriense, Cibinong, Java, Indonesia (MZB), National Museum of Natural History, Washington, D.C. (USNM), and Western Australian Museum, Perth (WAM). In addition, we have examined relevant type specimens at Nat-uralis Biodiversity Center, Leiden (RMNH).

Comparative material examined: Melanotaenia aja-maruensis--RMNH 28068 (holotype), 78.0 mm SL, Ayamaru Lakes region, Birds Head Peninsula, approximately 1[degrees]21'S, 132[degrees]16'E; RMNH 28069 (paratypes), 46 specimens, 25.5-63.7 mm SL, collected with holotype; WAM P26792-001 (paratypes), 6 specimens, 43.8-67.7 mm SL. M boe-semani--RMNH 28061 (holotype), 66.3 mm SL, Ayamaru Lakes region, Birds Head Peninsula, approximately 1[degrees]21'S, 132[degrees]16'E; RMNH 28062 (paratypes), 27 specimens, 35.0-63.0 mm SL, collected with holotype; WAM P. 26791-001 (paratypes), 3 specimens, 49.5-63.4 mm SL. M fasinensis--MZB 17700 (holotype), Ween Village, Birds Head Peninsula, 1[degrees]13.856'S, 131[degrees]58.186'E; MZB 17701 (paratypes), 4 specimens, 91.0-120.2 mm SL, collected with holotype. M irianjaya - MZB 4952 (holotype), 50.0 mm SL, Fruata, Bomberai Peninsula, approximately 2[degrees]59'S, 133[degrees] 32'E; MZB 4953, 50 specimens, 20.0-58.0 mm SL, collected with holotype; WAN1 P.29960-001, 53 specimens, 32.0-79.0 mm SL, near Bintuni, Birds Head Peninsula, approximately 2[degrees]05'S, 133[degrees] 30'E.

The methods of counting and measuring are as follows: dorsal and anal rays--the last ray of the anal and second dorsal fins is divided at the base and counted as a single ray; lateral scales--number of scales in horizontal row from upper edge of pectoral-fin base to caudal-fin base, excluding the small scales posterior to the hypural junction; transverse scales--number of scales in vertical row (excluding small truncated scales along base of fins) between anal-fin origin and base of first dorsal fin; predorsal scales--number of scales along midline of nape in front of first dorsal fin; cheek scales--total number of scales covering suborbital and preoper-culum; standard length (SL)--measured from tip of upper lip to caudal-fin base; head length (HL)--measured from tip of upper lip to upper rear edge of gill opening; cau.dal peduncle depth is least depth and caudal peduncle length is measured between two vertical lines, one passing through base of last anal ray and the other through caudal-fin base; caudal concavity--horizontal distance between verticals at tips .of shortest and longest rays.

Rainbowfish species used to generate sequence data are shown in Table I. We sequenced the mitochondrial cytochrome b (cytb) gene and used GARLI to obtain the best Maximum Likelihood tree and 1000 bootstrap replicates. Methods for obtaining DNA sequence data and their analyses follows Allen & Unmack (2012) and Allen et al. (2014) except where noted as follows: the model of sequence evolution TrN+G was the best one identified by ModelTest 3.7 (Posada & Crandall, 1998), we used attach-mentspertaxon = 58 and trees were rooted with the two "Waigeo" group species (M catherinae and M synergos). GenBank accession numbers are provided in Table I for all sequences included in this study.

Table I. Rainbowfish species from the Birds Head region of Papua
Barat Province used in the phylogenetic analysis including locality
data, the number of individuals examined and their GenBank
accession number. In the locality Field the code AS indicates
fish were sourced from rainbowfish aquarium hobbyists.

Species                            Locality         N    GenBank #

Melanotaenia ajamaruensis    Soum Ck, Birds Head    4  KM211354-5.1

Melanotaenia ammeri          crib to Arguni Bay,    4    KC133615.1
                             Birds Ncck

Melanotaenia angfa           AS, Yakati R, Birds    3    KC133616.1
                             Ncck

Melanotaenia arfakensis      AS, Prafi R. Birds     2    KC133617.1
                             Head

Melanotaenia batanta         Warmon Ck, Batanra Is  3    JQ282008.1

Melanotaenia boesemani       AS, Birds Head         3   KC 133618.1

Melanotaenia catherinae I    Kali Raja, Waigeo Is   3   KC 133629.1

Melanotaenia catherinae II   Wei Sam Ck, Waigeo Is  1   KC 133630.1

Melanotaenia ericrobertsi    Suswa village,         2   KC 133628.1
                             Karabara R, Birds
                             Head

Melanotaenia fasinensis      Sawiat Ck, Birds Head       KM211357.1

Melanotaenia flavipinnis     Ifaupan Ck, Misool Is  2  KF954094-5.1

Melanotaenia fredericki I    AS, Birds Head         2    KC133621.1

Melanotaenia fredericki II   AS, Sth of Sorong,     2    KC133619.1
                             Birds Head

Melanotaenia kokasensis      Kokas village,         7   KC 133623.1
                             Bomberai, Birds Neck

Melanotaenia laticlavia      AifiifCk, Birds Head   2  KM211358-9.1

Melanotaenia misoolensis I   Wai lama, Misool Is    5   KC 133624.1

Melanotaenia misoolensis II  Wai Tama, Misool Is    2   KC 133625.1

Melanotaenia multiradiata    Sisiah Ck, Birds Head  2  KM211360-1.1

Melanotaenia parva           AS, L Kurumoi, Birds   4   KC 133626.1
                             Neck

Melanotaenia salawati        Kali Dokrur, Salawati  9    KC133620.1
                             Is

Melanotaenia sp. Rawarra     Scbyar R, Birds Head   2   KC 133627.1

Melanotaenia sneideri        small ck in Kumawa     3      KM211356
                             Mnts, Birds Neck

Melanotaenia synergos I      Wai Bin Ck, Batanta    2    KC133631.1
                             Is

Melanotaenia synergos II     AS, Batanta Is         2   KC 133632.1


Melanotaenia ericrobertsi, n. sp.

Susvva Rainbovvfish

(Figs 2-4; Tables II & V)

Holotype. MZB 22114 (formerly WAM P.27868001, paratype of M irianjaya), female, 52.1 mm SL, Auk River, near Suswa Village, 0[degrees]56.472'S, 132[degrees]17.729'E, about 120 km east of Sorong, Birds Head Peninsula, Papua Barat Province, Indonesia, 0-1.5 m, seine net, G. R. Allen & H. Bleher, 18 November 1982.

Paratypes. MZB 22115 (collected with holotype, also formerly part of WAM P.27868-001, paratypes of M irianjaya), 10 specimens, 22.734.3 mm SL; WAM P. 27868-001, 30 specimens, 18.4-40.3 mm SL.

Diagnosis: A species of melanotaeniid rainbow-fish distinguished by the following combination of characters: dorsal rays IV to VI + 1, 1 3- 16 (usually V + 1,14 or 15); anal rays 1,23-27 (usually 25 or 26); pectoral rays 13-15 (usually 14 or 15); lateral scales 37-39 (usually 38); transverse scales 9 or 10; predorsal scales 17-19; circumpeduncular scales 12 or 13; total gill rakers on first arch 15-19 (usually 17 or 18); total scales covering preoperculum 1216 (mean 14.1); origins of first dorsal and anal fins about level; greatest body depth of adult male about 2.7-3.1 in SL; colour of adult male in life generally mauve to pale yellowish with orange stripe between each scale row on side of body; blackish zone on lower two-thirds of caudal peduncle and adjacent body above last few anal rays; blackish patch immediately behind pectoral-fin base; pelvic and median fins reddish with narrow white margin on dorsal fins.

Description: Dorsal rays VI + 1,15 (IV-VI + 1,1316); anal rays 1,26 (1,23-27); pectoral rays 15(13- 15); pelvic rays 1,5; branched caudal rays 15; lateral scales 37 (37-39); transverse scales 10 (9 or 10); predorsal scales 17 (17-19); cheek scales 15(12- 16); gill rakers on first branchial arch 3 + 13 (2-4 + 14 or 15), total gill rakers on first arch 15(16-19).

Body depth 3.6 (3.8-4.3 in paratypes, 2.7-3.1 in adult males from photographs) in SL, head length 4.0 (3.6-4.0) in SL; greatest width of body 2.3 (2.1-2.6) in greatest body depth; snout length 3.4 (3.0-3.4) in HL; eye diameter 2.9 (2.4-2.9) in HL; interorbital width 2.7 (2.7-3.0) in HL; depth of caudal peduncle 2.3 (2.2-2.8) in HL; length of caudal peduncle 1.4 (1.3-1.6) in HL.

Jaws about equal, oblique, premaxilla with an abrupt bend between the anterior horizontal portion and lateral part; maxilla ends below about anterior edge of pupil; maxillary length 2.9 (2.9-3.3) in HL; lips thin; teeth conical with slightly curved tips, extending on to outer surface of lips; teeth of upper jaw in 6-9 irregular rows anteriorly, reduced to 1-2 rows posteriorly, where they are exposed when mouth is closed; teeth in lower jaw in about 6-8 irregular rows anteriorly, reduced to 1-2 rows posteriorly; vomerine teeth poorly developed, holotype with a few scattered teeth in narrow band; palatine teeth absent.

Scales of body cycloid, relatively large, and arranged in regular horizontal rows; scale margins weakly crenulate; predorsal scales extending forward to rear half of interorbital space; preopercle with 2 scale rows between its posterior angle and eye; scales absent on preorbital.

Predorsal length 2.1 (2.0-2.1) in SL; preanal length 2.0 (2.0-2.2) in SL; prepelvic length 2.9 (2.6-2.9) in SL; length of second-dorsal fin base 4.6 (4.0-4.9) in SL; length of anal-fin base 2.6 (2.6-2.8) in SL.

First dorsal-fin origin about equal with anal-fin origin; longest spines of first dorsal fin 1.5 (2.12.9) in HL, its depressed tip reaching slightly beyond origin of second dorsal fin; longest ray (generally anteriormost) of second dorsal fin 1.8 (1.72.0) in HL, depressed posterior rays extending less than one-half length of caudal peduncle; longest anal ray (generally middle rays in type specimens, but anterior rays in adult males as determined by photographs) 2.0 (1.8-2.1) in HL; length of pelvic fins 1.5 (1.7-1.9), fin tips when depressed reaching from slightly beyond base of anal-fin origin in most paratypes to base of second soft anal ray in holotype; length of pectoral fins 1.3 (1.3-1.7) in HL; length of caudal fin 1.0 (1.0-1.1) in HL; caudal fin moderately forked, caudal concavity 3.8 (3.2-4.8) in head length.

Colour in life (Figs 2 & 4-upper): upper part of head and predorsal region greenish brown; body generally mauve to yellowish, usually with narrow orange stripe between each horizontal scale row; body scales with narrow brown margins, imparting overall network appearance; blackish patch, mainly covering first 3-4 mid-lateral scales behind pectoral-fin base; prominent blackish zone occupying lower two-thirds of caudal peduncle and adjacent body above last few anal rays; head with broad black stripe behind eye, extending to upper rear margin of operculum, usually with silvery yellow or reddish spot immediately below; remainder of operculum and adjacent cheek dusky greyish; pelvic and median fins reddish with narrow white margin on dorsal fins; submarginal blackish stripe present on second dorsal fin of adults, but often poorly developed; pectoral fins translucent. Young specimens, approximately 5 cm total length, are similar, but generally pinkish overall rather than yellowish.

Colour of holotype in alcohol (Fig. 3): generally light brown dorsally, grading to pale yellowish tan on lower sides; scale margins with fine, brown margins, imparting network appearance; faint, grey mid-lateral stripe on middle of side, more distinct on posterior third of body, including caudal peduncle; pelvic and median fins grey; pectoral fins translucent. Paratypes are similar in colour with the dark mid-lateral stripe showing variable intensity.

Sexual dimorphism: Not apparent among type specimens, which are invariably females or juveniles. However, photographs of captive adults (Figs 2 & 4-upper) indicate typical Melanotaenia sexual differences consisting of a deeper body (32.4-37.6 % of SL, calculated from photographs), taller first dorsal fin, and longer soft dorsal and anal rays. Males generally have the longest anal rays at the beginning of the fin compared to the middle of the fin in females.

Remarks: Within western New Guinea there is a complex of Me/anotaenia species with similar morphology and colouration (herein called the Aya-maru complex). A number of these species occur in the region near Lake Ayamaru, e.g., M ajamaruensis, M boesemani, and M fasinensis, but others such as M alfakensis, M irianjaya, M veoliae and M wanoma occur more broadly across the Birds Head region (Fig. 1). This group also contains the three new species described herein: M ericrobertsi, M laticlavia, and M multiradiata. Most of these species are part of the same genetic clade within the western lineage, although some such as M lati-clavia are more distantly related genetically (Fig. 5). These fishes are generally similar in appearance and characterised by relatively high dorsal and anal fin ray counts, and a live colour pattern that features a dark submarginal stripe and outer white margin on the second dorsal fin.

Two of the new taxa, M ericrobertsi and M multi-radiata, were previously confused with M irianjaya, which was described on the basis of 170 specimens collected by G. R. Allen (Allen 1985) at widely scattered locations on the Bomberai and Birds Head peninsulas of West Papua, but now believed to occur mainly on the Bomberai Peninsula (Fig. 1). Based on general similarity in colour pattern, morphometric proportions, and meristic features the specimens were believed to represent a single, slightly variable species. Moreover, without recourse to current genetic methods, it did not seem unusual for a rain-bovvfish to have such a broad distribution, encompassing numerous river systems. However, an emerging pattern based on recent genetic results (Unmack et al. 2013) reveals that many of the former widespread species are likely divisible into two or more taxa with individual taxa frequently being restricted to one or a few river basins.

Melanotaenia irianjaya was originally described on the basis of relatively small individuals (largest 58 mm SL), which did not show typical adult colouration, including that of the adult male, which is particularly diagnostic. Thanks to their eventual introduction to the aquarium hobby, photographs and tissue samples were obtained of live adults from the type locality (Fruata, West Papua) and from Suswa, where 41 paratypes were collected by G. R. Allen in 1982. Photographs of live adults (Fig. 4) and DNA results (Fig. 5) from the two locations revealed important species-level differences between these two populations and prompted our description of the Suswa fish.

Colour differences include the presence of pronounced blackish zones on the anterior and posterior, mid-lateral parts of the body in M ericrobertsi, and much narrower white margin on the second dorsal CI n in comparison to M irianjaya. Photographs (Fig. 4) also reveal a difference in the shape of the anal fm of mature males, a feature that is not evident in the relatively small type specimens of both species. The anal fin of M irianjaya is more rounded, with the tallest rays in the middle of the fin in contrast to that of M ericrobertsi, which has the longest rays anteriorly, a condition that is unusual among male members of the genus, which usually have the longest anal rays either centrally or posteriorly.

I - L Genetic results (Fig. 5) indicate that the new species is actually more closely related to M fasi-nensis (Fig. 10C), which occurs further downstream in the same catchment. These species appear to be nearly morphological twins with no reliable morphometric or meristic features to distinguish them. However, colour pattern and genetic differences are diagnostic. The two prominent blackish zones on the body of M ericrobertsi and lack of this feature in M fasinensis is particularly useful for separating them, at least in adult fish. Additional differences between this species and the other two new taxa are provided in the remarks sections for M laticlavia and M multi radiata.

Zoogeography and habitat: The new species is known only from the vicinity of Suswa Village (Fig. 1), which is situated in the foothills of the Tamrau Mountains at an elevation of about 176 m, approximately 120 km east of the city of Sorong. The type locality (Fig. 6) is part of the extensive Kladuk River system and is approximately 215 km upstream from the Karabra Estuary on the south coast of the Birds Head Peninsula. The stream, which ranged from about 15 to 30 m in width at the type locality, was flowing through primary and secondary rainforest habitat over gravel and sand substrate with minimal aquatic vegetation. Rainbowfish were mainly concentrated around log debris. Temperature and pH values of 27-28[degrees] C and 7.5 were recorded at the time of collection.

Etymology: The new species is named ericrobertsi in honour of Eric Roberts, a pilot with Associated Mission Aviation (AMA), Papua Province, Indonesia. Eric is an aquarium fish enthusiast who collected live specimens and is responsible for the introduction of this species to the aquarium hobby.

Melanotaenia laticlavia, n. sp.

Aifuf Rainbowfish (Figs 7-8 & IOD; Tables III & V)

Holotype. MZB 22116, male, 69.6 mm SL, Aifuf Creek, 1[degrees]19.613'S, 132'35.415'E, about 43 km east of Lake Ayamaru, Birds Head Peninsula, Papua Barat Province, Indonesia, 0-1.0 m, seine net, H. G. Evers and J. Christian, 6 October 2013. Paratypes. MZB 22117 (collected with holorype), 3 specimens, females, 33.7-36.1 mm SL; WAM P.34025-001, 2 specimens, 57.0-61.6 mm SL.

Diagnosis: A species of melanotaeniid rainbow-fish distinguished by the following combination of characters: dorsal rays IV-VI + 1,14-17; anal rays 1,24-26 (usually 24); pectoral rays 14 or 15 (usually 14); lateral scales 36-37 (usually 37), transverse scales 9 or 10; predorsal scales 16-18 (usually 17); circumpeduncular scales 13; total gill rakers on first arch 16 or 17 (usually 16); total scales covering preopercul um 13-19 (adults with 16-19) ; adult with two separate patches of vomerine teeth; origin of first dorsal fin about level with anal-fin origin; greatest body depth of adult male 2.8-3.0 in SL; colour in life generally olive green to yellowish with faint orange stripe between each horizontal scale row; mid-lateral dark stripe often present, broader and more prominent on posterior half of body; dorsal and anal fins with dark submarginal stripe and pale outer margin; dorsal and ventral margins of caudal fin narrowly blackish.

Description: Dorsal rays V + 1,14 (IV-VI + 1,1517); anal rays 1,24 (1,24-26); pectoral rays 14 (14 or 15); pelvic rays 1,5; branched caudal rays 15; lateral scales 36 (37); transverse scales 9 (10); predor-sal scales 18 (16-17); cheek scales 16 (13-19); gill rakers on first branchial arch 2 + 14 (2 + 14 or 15), total gill rakers on first arch 16 (16 or 17).

Body depth 2.9 (2.8- 3.0, 3.4-3.5 in juveniles) in SL, head length 3.8 (3.5-3.7) in SL; greatest width of body 2.7 (2.5-2.6, 2.1-2.2 in juveniles) in greatest body depth; snout length 2.9 (3.2-4.1) in HL; eye diameter 3.2 (2.6-3.0) in HL; interorbital width 2.8 (2.7-2.9) in HL; depth of caudal peduncle 2.4 (2.4-2.7) in HL; length of caudal peduncle 1.6 (1.5-2.0) in HL.

Jaws about equal, oblique, premaxilla with an abrupt bend between the anterior horizontal portion and lateral part; maxilla ends below about anterior edge of pupil or slightly anterior to this point; maxillary length 2.6 (2.7-3.4) in HL; lips thin; teeth conical with slightly curved tips, extending on to outer surface of lips; teeth of upper jaw in 6-7 irregular rows anteriorly, reduced to 1-2 rows posteriorly, where they are exposed when mouth is closed; teeth in lower jaw in about 12 irregular rows anteriorly, reduced to 1-2 rows posteriorly; vomerine teeth in 2 well-separated patches, with about 10-12 teeth in each patch of holotype; palatine teeth absent.

Scales of body cycloid, relatively large, and arranged in regular horizontal rows; scale margins weakly crenulate; predorsal scales extending forward to rear half of interorbital space; preopercle with 2-3 scale rows between its posterior angle and eye; scales absent on preorbital.

Predorsal length 2.1 (2.0-2.2) in SL; preanal length 2.0 (1.9-2.1) in SL; prepelvic length 2.6 (2.6-2.7) in SL; length of second-dorsal fin base 3.8 (3.8-4.3) in SL; length of anal-fin base 2.5 (2.4-2.5, 2.7-3.0 in juveniles) in SL.

First dorsal-fin origin about half eye diameter behind level of anal-fin origin; longest spines of first dorsal fin 1.9 (1.5-1.6, 2.4-2.6 in juveniles) in HL, its depressed tip reaching origin of second dorsal fin in juveniles and reaching to about base of second soft ray in mature males; longest ray (generally anteriormost in juveniles and penultimate in males) of second dorsal fin 1.8 (2.0-2.5) in HL, depressed posterior rays extending less than one-half length of caudal peduncle in juveniles and nearly full length of caudal peduncle in mature males; longest (sixth to ninth rays in males and females) anal rays 1.9 (2.1, 2.5-2.8 in juveniles) in HL; pelvic fin tips when depressed reaching to base of second or third soft anal fin ray in mature adults; length of pelvic fins 1.4 (1.4-1.5, 1.8-2.0 in juveniles); length of pectoral fins 1.4 (1.4-1.5) in HL; length of caudal fin 1.2 (1.1-1.3) in HL; caudal fin moderately forked, caudal concavity 3.3 (3.1-5.3) in head length.

Table III. Table II. Proportional measurements of selected type
specimens of Melanotaenia ericrobertsi expressed as percentage
of the standard length.

              Holotype  Paratype  Paratype  Paratype  Paratype
                MZB       WAM       WAM       MZB       MZB
               22116    P.34025   P.34025   22117      22117

Sex               male      male      male  juvenile  juvenile

Standard          69.6      61.6      57.0    36.0      36.1
length (mm)

Body depth        34.3      35.2      33.7    28.9      29.1

Body width        12.6      14.1      12.8    13.6      13.9

Head length       26.1      26.8      27.7    27.2      28.8

Snout length       9.1       8.4       8.8     7.8       8.3

Maxillary          9.9      10.1       9.8     9.4      10.2
length

Eye diameter       8.2       9.4       9.1    10.6      11.1

Bony               9.5       9.6      10.2     9.4      10.0
interorbital
width

Depdi of          10.9      11.0      10.9    10.8      10.8
caudal
peduncle

Length of         16.8      15.7      14.2    18.6      16.6
caudal
peduncle

Predorsal         48.7      49.2      51.1    46.4      48.5
distance

I'reanaJ          50.3      48.1      49.8    50.3      51.5
distance

Prepelvic         38.1      36.7      38.2    38.3      38.2
distance

2(nd)             26.6      26.0      26.3    26.1      26.3
dorsal-fin
base

Anal-fin          40.5      39.9      42.1    33.6      34.6
base

Pectoral-fin      19.0      19.2      19.3    19.4      19.7
length

Pelvic fin        18.1      19.2      18.4    15.3      14.7
length

Longest ray       14.1      18.0      17.7    10.3      11.9
l(sl) dorsal
fin

Longest ray       14.9      13.3      12.6    13.1      11.4
2(nd) dorsal
fin

Longest anal      14.1      12.5      13.5    11.1      10.5
ray

Caudal-fin        21.7      21.4      22.5    23.9      23.5
length

caudal             8.0       6.0       5.3     8.9       7.8
concavity

              Paratype
                MZB
               22117

Sex           juvenile

Standard        33.7
length (mm)

Body depth      29.4

Body width      13.4

Head length     27.9

Snout length     6.8

Maxillary        8.3
length

Eye diameter    10.1

Bony             9.5
interorbital
width

Depdi of        10.7
caudal
peduncle

Length of       16.3
caudal
peduncle

Predorsal       47.2
distance

I'reanaJ        51.9
distance

Prepelvic       39.2
distance

2(nd)           23.1
dorsal-fin
base

Anal-fin        36.5
base

Pectoral-fin    19.0
length

Pelvic fin      15.1
length

Longest ray     11.0
l(sl) dorsal
fin

Longest ray     11.3
2(nd) dorsal
fin

Longest anal    10.1
ray

Caudal-fin      23.1
length

caudal           5.6
concavity


CY Colour in life (Figs 7 & 10D): generally olive green to bronzy yellow, often darker dorsally, with darker scale margins imparting network appearance; narrow orange stripe between each horizontal scale row on side of body; frequently with dark blue to blackish mid-lateral stripe on side of body, often interrupted and usually narrower on anterior half and much broader (up to 3 scales wide) posteriorly; breast yellow orange in adult male, whitish in female; both sexes usually with broad blue-grey zone, tapering nearly to a point posteriorly, on lower side from below pectoral-fin base to above middle of anal fin; dorsal and anal fins pale orange to vivid orange in adult males, pale yellow to bluish in females; second dorsal and anal fins with dark brown to blackish submarginal stripe and pale blue outer margin; caudal fin translucent with dusky greyish rays and narrow blackish dorsal and ventral margins; pelvic fins orange in adult males and whitish in females; pectoral fins translucent; iris mainly golden yellow; broad blackish stripe across uppermost part of operculum and vivid red spot (about half pupil size) immediately below.

Colour of holotype in alcohol (Fig. 8): head brownish dorsally, grading to whitish ventrally; operculum with blackish stripe crossing uppermost portion with silvery patch immediately below; light brown on back, grading to whitish ventrally; each scale of side with thin brown margins imparting overall network appearance; blackish mid-lateral stripe on side from pectoral-fin base to caudal-fin base, poorly developed and interrupted on anterior half, but distinctly wider (covering up to 3 scales) on posterior half; second dorsal, anal, pelvic and caudal fins generally dusky grey with blackish sub-marginal stripe (more prominent on dorsal fin) and narrow whitish margin on dorsal and anal; caudal fin with narrow, blackish upper and lower margins; pectoral fins translucent. The two adult male paratypes (WANI) exhibit similar markings, but are overall much darker with thicker dark scale margins and numerous microscopic melanophores on the lower portion of the sides. Juvenile paratypes (33.7-36.1 mm SL) are yellowish tan, grading to white ventrally. The largest juvenile has a well-developed, dark mid-lateral stripe, which is only visible on the posterior portion of the body in the smaller juveniles. A remnant of the blue-grey zone on the lower side, mentioned in the live colour description, is also evident in the largest juvenile.

Sexual dimorphism (based on photographs because no adult female specimens were available for comparison): males appear to have a deeper body compared to females which is typical for the genus. The greatest depth of the three adult male types is 33.7-35.2% of SL compared with about 30% of SL for the female shown in Fig. 7. In addition, the greater height of the first dorsal fin (significantly overlapping the dorsal fin origin) and last soft dorsal rays (extending for most of caudal peduncle length) are typical male features of the genus. Finally, the dark upper and lower margins of the caudal fin are more vivid in males compared to females.

Remarks: Live colour pattern of adult males (Figs 7 and 10D) and DNA analysis (Fig. 5) provide the best means of separating Melanotaenia from western New Guinea. However, M. laticlavia is distinguished from other members of the Ayamaru complex, and all other Melanotaenia examined to date, in having two separate patches of vomerine teeth (Fig. 11A) in adults. Most members of the genus possess a single medial patch (Fig. 11B) or in the case of a few species from outside this region, may lack these teeth entirely. However, this feature is most obvious in large adults and is often lacking or poorly developed in juveniles and subadults.

This species- (as well as M ericrobertsi and M multi radiata) is further separable from M aja-maruensis and M boesemani on the basis of 9 or 10 transverse scale rows versus only 7 or 8 for the latter species. It also lacks the pair of diffuse, dusky grey bars on the ventral, anterior body, as well as the dusky blue-grey colour on the lower head and breast region, characteristic of adult males of these two species (Fig. 10A & B). Melanotaenia laticlavia is also distinguished by the dark upper and lower caudal-fin margins (also shared by M irianjaya), the possession of a dark submarginal stripe and prominent white outer margin on both dorsal and anal fins (only on the dorsal fin of other Ayamaru complex species), broad blue grey zone on lower side (tapering porteriorly), and a red spot on the operculum, which although common in the genus, is either rudimentary or absent on other Ayamaru complex species.

Zoogeography and habitat: The new species is known only from Aifuf Creek in the Birds Head region of West Papua (Fig. 1). The type locality (Fig. 11) consists of a narrow (about 1-2 m wide), relatively shallow (to about 0.5-1.0 m depth) stream with very gradual gradients flowing through second growth forest. The type specimens were mainly collected over sand and gravel bottoms. The stream is located in the upper reaches of the Kamundan River system, one of several large rivers that flow into the mouth of Berau Bay at the southernmost portion of the Birds Head Peninsula. The type locality, at 154 m elevation, is situated about 108 km north of the river mouth or following the meandering path of the river it is approximately 220 km upstream from the sea.

Etymology: The new species is named laticlavia (Latin: having a broad stripe) with reference to the broad bluish-grey stripe along the lower side of the body.

Melanotaenia multiradiata, n. sp

Moswaren Rainbowfish

(Figs 10E& 12-14; Tables IV & V)

Holotype. MZB 22118, male, 122.5 mm SL, Sisiah Creek, 1[degrees]27.784'S, 132[degrees]14.546'E, about 25 km east of Tern inabuan, Birds Head Peninsula, Papua Barat Province, Indonesia, 0-1.5 m, seine net, G. R. Allen, M. Allen & S. Renyaan, 5 September 1999. Paratypes. MZB 22119 (collected with holotype), 7 specimens, 37.4-83.7 mm SL; USNM 427109, 5 specimens, 40.7-97.3 mm SL; WAM P.31569-001, 10 specimens, 37.4-97.5 mm SL.

Diagnosis: A species of melanotaeniid rainbow-fish distinguished by the following combination of characters: dorsal rays IV-VI + 1,14-17 (usually V + 1,15); anal rays 1,23-26 (rarely 23); pectoral rays 14-17 (usually 15 or 16); lateral scales 37-38; transverse scales 9; predorsal scales 17-19; circumpedun-cular scales 12 or 13; total gill rakers on first arch 16-19 (usually 17 or 18); total scales covering pre-operculum 13-17 (mean 15.8); origin of first dorsal fin usually about eye diameter behind level of anal-fin origin, but occasionally level of dorsal and anal-fin origins about even; greatest body depth of adult male 2.7-3.2 in SL; colour in life generally bluish green to yellowish, grading to white ventrally; mid-lateral dark stripe usually present, broader and more prominent on posterior half of body; dorsal fins with dark submarginal stripe and pale outer margin.

Description: Dorsal rays V + 1,13 (IV-VI + 1,1416); anal rays 1,24 (1,23-26); pectoral rays 14/15 (14-17); pelvic rays 1,5; branched caudal rays 15; lateral scales 38 (37 or 38); transverse scales 9; pre-dorsal scales 18 (17-19); cheek scales 16 (13-17); gill rakers on first branchial arch 3 + 14 (2-4 + 14 or 15), total gill rakers on first arch 17 (16-19).

Body depth 2.7 (2.7- 3.6, 3.6-40 in juveniles < 43 mm SL) in SL, head length 3.9 (3.5-4.0) in SL; greatest width of body 2.7 (2.2-2.8) in greatest body depth; snout length 2.7 (2.7-3.0) in HL; eye diameter 3.8 (3.2-3.8) in HL; interorbital width 2.9 (2.6-3.0) in HL; depth of caudal peduncle 2.3 (2.3-3.0) in HL; length of caudal peduncle 1.6 (1.4-1.8) in HL.

Table IV. Proporcional measurements of selected type specimens of
Melanotaenia multiradiata expressed as percentage of the standard
length.

                Holotype  Paratype  Paratype  Paratype  Paratype
                  MZB       WAM       WAM       MZB       MZB
                 22118    P.31569   P.31569    22119     22119

Sex              male     female      male      male    female

Standard         122.5      97.5      96.1      83.7     80.5
length (mm)

Body depth        37.6      30.6      37.4      35.2     31.8

Body width        14.0      13.2      14.3      14.0     13.3

Head length       25.6      25.2      26.4      28.3     26.3

Snout length       9.6       8.7       8.8       9.6      9.2

Maxillary          9.7       9.0       9.3       9.2      9.6
length

Eye diameter       6.7       6.6       7.3       8.1      7.2

Bony               8.9       9.5      10.2       9.7      9.1
interorhital
width

Depth of          11.1      10.2      11.4      10.9     10.1
caudal
peduncle

Length of         16.0      17.1      15.7      15.4     16.5
caudal
peduncle

Predorsal         50.9      50.5      52.4      49.8     49.9
distance

Preanal           46.7      49.3      48.0      48.3     52.3
distance

Prepelvic         35.6      36.5      38.1      37.5     39.1
distance

2(nd)             26.9      25.6      26.8      25.0     25.3
dorsal-fin
base

Anal-fin          45.8      39.1      44.1      44.3     38.5
base

Pectoral-fin      17.3      17.9      16.6      19.1     18.6
length

Pelvic-fin        19.5      18.5      18.7      19.4     18.9
length

Longest ray       14.6      13.1      14.6      17.1     14.5
I(st) dorsal
fin

Longest ray       14.1      10.9      15.6      17.3     12.0
2(nd) dorsal
fin

Longest anal      14.7      14.4      15.0      15.4     12.9
ray

Caudal-fin        19.6      19.3      21.4      21.0     21.2
length

Caudal             6.9       9.0       6.8       7.6      7.6
concavity

                Paratype  Paratype  Paratype
                  WAM       MZB       WAM
                P.31569    22119     P.31569

Sex               male    female    female

Standard          71.3     58.2      50.6
length (mm)

Body depth        32.5     29.0      27.9

Body width        13.3     11.7      11.7

Head length       26.1     25.8      27.5

Snout length       9.0      8.9       9.1

Maxillary          9.1      9.1       9.1
length

Eye diameter       8.0      7.4       8.7

Bony               9.5      8.8       9.7
interorhital
width

Depth of          10.7     10.3       9.9
caudal
peduncle

Length of         17.7     19.1      17.8
caudal
peduncle

Predorsal         51.5     48.3      50.4
distance

Preanal           49.2     49.3      52.2
distance

Prepelvic         36.5     36.1      37.7
distance

2(nd)             24.4     22.2      21.9
dorsal-fin
base

Anal-fin          40.7     37.5      36.0
base

Pectoral-fin      19.1     18.4      16.8
length

Pelvic-fin        19.6     17.0      15.4
length

Longest ray       12.8     12.7      13.0
I(st) dorsal
fin

Longest ray       14.6     12.5      13.0
2(nd) dorsal
fin

Longest anal      14.2     11.3      10.3
ray

Caudal-fin        19.8     19.4      21.7
length

Caudal             4.8      4.3       7.5
concavity


Jaws about equal, oblique, premaxilla with an abrupt bend between the anterior horizontal portion and lateral part; maxilla ends below about anterior edge of pupil; maxillary length 2.6 (2.7-3.1) in HL; lips thin; teeth conical with slightly curved tips, extending on to outer surface of lips; teeth of upper jaw in 6-7 irregular rows anteriorly, reduced to 1-2 rows posteriorly, where they are exposed when mouth is closed; teeth in lower jaw in about 10 irregular rows anteriorly, reduced to 1-2 rows posteriorly; vomerine teeth in 2 separate patches (variable in paratypes, ranging from absent in smaller specimens to either a single narrow band or two separate patches); palatine teeth absent.

Scales of body cycloid, relatively large, and arranged in regular horizontal rows; scale margins weakly crenulate; predorsal scales extending forward to rear half of interorbital space; preopercle with 2-3 scale rows between its posterior angle and eye; scales absent on preorbital.

Predorsal length 2.0 (1.9-2.1) in SL; preanal length 2.1 (1.9-2.1) in SL; prepelvic length 2.8 (2.6-2.8) in SL; length of second-dorsal fin base 3.7 (3.7-4.6) in SL; length of anal-fin base 2.2 (2.3-2.8) in SL.

First dorsal-fin origin about half eye diameter behind level of anal-fin origin (paratypes variable, ranging from level with anal-fin origin to about eye diameter behind level of anal origin); longest spines of first dorsal fin 1.8 (1.5-1.6, 2.4-2.6 in juveniles) in HL, its depressed tip reaching slightly beyond origin of second dorsal fin in females and to about base of second or third soft ray in mature males; longest ray (generally anteriormost in females and juveniles, and penultimate in males) of second dorsal fin 1.8 (1.6-1.8 in adult males, 2.12.3 in males less than about 60 mm SL and females) in HL, depressed posterior rays extending about one-half length of caudal peduncle or less in females and juveniles, and nearly full length of caudal peduncle in mature males; anal rays of adult males more or less subequal except last few rays shorter than others, tallest rays 1.8 (1.7-1.8) in HL; rays of anterior half of female and juvenile anal fin taller than those of posterior half, tallest rays 1.8-2.7 in HL; length of pelvic fins 1.3 (1.31.8, fin tips when depressed reaching to base third or fourth soft anal fin ray in mature males and to first or second soft anal ray in females; length of pectoral fins 1.5 (1.4-1.6) in HL; length of caudal fin 1.3 (1.1-1.3) in HL; caudal fin moderately forked, caudal concavity 3.7 (2.8-6.0) in head length.

Table IV. Proporcional measurements of selected type specimens of
Melanotaenia multiradiata expressed as percentage of the standard
length.

                Holotype  Paratype  Paratype  Paratype  Paratype
                  MZB       WAM       WAM       MZB       MZB
                 22118    P.31569   P.31569    22119     22119

Sex              male     female      male      male    female

Standard         122.5      97.5      96.1      83.7     80.5
length (mm)

Body depth        37.6      30.6      37.4      35.2     31.8

Body width        14.0      13.2      14.3      14.0     13.3

Head length       25.6      25.2      26.4      28.3     26.3

Snout length       9.6       8.7       8.8       9.6      9.2

Maxillary          9.7       9.0       9.3       9.2      9.6
length

Eye diameter       6.7       6.6       7.3       8.1      7.2

Bony               8.9       9.5      10.2       9.7      9.1
interorhital
width

Depth of          11.1      10.2      11.4      10.9     10.1
caudal
peduncle

Length of         16.0      17.1      15.7      15.4     16.5
caudal
peduncle

Predorsal         50.9      50.5      52.4      49.8     49.9
distance

Preanal           46.7      49.3      48.0      48.3     52.3
distance

Prepelvic         35.6      36.5      38.1      37.5     39.1
distance

2(nd)             26.9      25.6      26.8      25.0     25.3
dorsal-fin
base

Anal-fin          45.8      39.1      44.1      44.3     38.5
base

Pectoral-fin      17.3      17.9      16.6      19.1     18.6
length

Pelvic-fin        19.5      18.5      18.7      19.4     18.9
length

Longest ray       14.6      13.1      14.6      17.1     14.5
I(st) dorsal
fin

Longest ray       14.1      10.9      15.6      17.3     12.0
2(nd) dorsal
fin

Longest anal      14.7      14.4      15.0      15.4     12.9
ray

Caudal-fin        19.6      19.3      21.4      21.0     21.2
length

Caudal             6.9       9.0       6.8       7.6      7.6
concavity

                Paratype  Paratype  Paratype
                  WAM       MZB       WAM
                P.31569    22119     P.31569

Sex               male    female    female

Standard          71.3     58.2      50.6
length (mm)

Body depth        32.5     29.0      27.9

Body width        13.3     11.7      11.7

Head length       26.1     25.8      27.5

Snout length       9.0      8.9       9.1

Maxillary          9.1      9.1       9.1
length

Eye diameter       8.0      7.4       8.7

Bony               9.5      8.8       9.7
interorhital
width

Depth of          10.7     10.3       9.9
caudal
peduncle

Length of         17.7     19.1      17.8
caudal
peduncle

Predorsal         51.5     48.3      50.4
distance

Preanal           49.2     49.3      52.2
distance

Prepelvic         36.5     36.1      37.7
distance

2(nd)             24.4     22.2      21.9
dorsal-fin
base

Anal-fin          40.7     37.5      36.0
base

Pectoral-fin      19.1     18.4      16.8
length

Pelvic-fin        19.6     17.0      15.4
length

Longest ray       12.8     12.7      13.0
I(st) dorsal
fin

Longest ray       14.6     12.5      13.0
2(nd) dorsal
fin

Longest anal      14.2     11.3      10.3
ray

Caudal-fin        19.8     19.4      21.7
length

Caudal             4.8      4.3       7.5
concavity


Colour of holotype in life (Fig. 12): generally bluish green on dorsal portion of head and body, grading to whitish ventrally; body scales, including those of breast and ventral portion of side, with narrow blackish margins imparting overall network appearance; usually with prominent dark blue to blackish mid-lateral stripe from pectoral-fin base to caudal-fin base, less distinct and occupying single scale row on anterior half to two thirds of body, but more vivid and wider (covering 2-3 horizontal scale rows) on posterior body including caudal peduncle, although not clearly visible in Fig. 12 it was evident when first collected; head with broad blackish stripe from snout to upper rear margin of operculum, interrupted by eye; silvery spot on upper operculum immediately below aforementioned dark stripe, remainder of operculum and cheek yellowish to bronze; dorsal fins reddish with pronounced dark brown to blackish submarginal stripe and broad, white outer margin; anal fin bluish grey to yellow or reddish; caudal fin dusky grey with narrow, white posterior margin; pelvic fins dusky grey to whitish; pectoral fins translucent. Adult male paratypes generally similar, but sometimes with pronounced yellow area occupying 3-4 horizontal scale rows on middle of side (Fig. 10E). Females (Fig. 13) are similar to the pattern described for the holotype with adults generally displaying a conspicuous dark blue to blackish mid-lateral stripe.

Colour ofr holotype in alcohol (Fig. 14, upper): head and body brown dorsally, grading to yellowish tan on ventral half; blackish stripe crossing uppermost portion of operculum, continued on body as blackish mid-lateral stripe to caudal-fin base, less distinct anteriorly and consisting of broad vertical streak on each scale, forming continuous stripe on posterior third of body with maximum width of about two scale rows; pelvic and median fins dusky grey, second dorsal with blackish submarginal stripe and broad, white outer margin, also narrow white margin on caudal; pectoral fins translucent. Adult paratypes similar, although most adult specimens with more vivid, blackish mid-lateral stripe on body, particularly prominent on largest (97.5 mm SL) female (Fig. 14, lower). Juvenile paratypes (37.4-43.0 mm SL) are pale brown dorsally, grading to tan ventrally with a blackish mid-lateral stripe similar to that described For adults evident in the largest juvenile.

Sexual dimorphism: Although sample size was relatively small for most size classes, males typically have a deeper body compared to females. The average body depth of males greater than 80 mm SL is 35.2% of SL (range 31.5-37.6%, n = 8) and that of specimens from 50-79 mm SL is 31.2% of SL (range 29.6-32.5%, n = 2). In comparison females in these same size classes have average body depths of 31.2 (30.6-31.8%, n = 2) and 28.5 (27.9-29.0 %, n = 2) % of SL. The average body depth of juveniles, 37-43 mm SL was 26.4% SL (range 25.128.1 %, n = 6). Other male differences include the greater height of the first dorsal fin (significantly overlapping the dorsal fin origin) and last soft dorsal rays (extending for most of caudal peduncle length).

Remarks: M multiradiata is distinguished by its relatively high pectoral-fin ray count compared to other Ayamaru complex species with 91.3% of specimens having 15 or more pectoral rays (Table V). This value is significantly greater than 25.5% recorded for M ajamaruensis, 37.7% for M boese-mani, 40.0% for M. fasinensis, 12.5 % for M iri-anjaya and 16.7% for M laticlavia. Diagnostic colour pattern features of M multi radiata include a distinct white, posterior margin on the caudal fin and the possession of a well-developed and expanded mid-lateral, dark stripe on the posterior body (also shared with M ericrobertsi).

Zoogeography and habitat: The new species is known only from Sisiah Creek near Moswaren Village, which lies about 25 km east of the town of Teminabuan, near the south coast of the Birds Head region of West Papua (Fig. 1). The type locality (Fig. 15) consists of a 15-20 m wide stream with depths to about 2 m and slow to moderate flow through second growth forest. The type specimens were mainly collected over a limestone rock bottom, and were concentrated around log debris in the deeper sections. The stream was exceptionally clear at the time of collection with estimated underwater visibility of 20 m. The type locality is situated at an elevation of about 86 m, approximately 58 km upstream from where it eventually flows into Waromge Bay.

Etymology: The new species is named multi-radiata (Latin: many rays) with reference to the relatively high number of pectoral-fin rays.

Genetic results and discussion: A total of 70 individuals were sequenced from 19 species from the western lineage in the Birds Head region for the mtDNA cytb gene (Table I). Within species genetic diversity was low, with only six species (M aja-maruensis, M flavzinnis, M fredericki, M lati-clavia, M misooknsis, and M multi radiata) containing two haplotypes (the remaining 13 species each had only a single haplotype). Sequence analysis of the 29 OTUs (Table I) yielded 849 invariant characters, 39 variable but parsimony uninformative characters, and 253 parsimony informative characters. ML analysis recovered one tree with a likelihood score of -4260.229554 (Fig. 5). The relationships recovered were broadly congruent with the larger sequence dataset in Unmack et al. (2013). Rainbowfishes in the western lineage from the Birds Head region consist of three deeper clades. The most divergent is the "Waigeo" group which consists of the species M catherinae and M .synergos from Waigeo and Batanta islands. The remaining species separate in two groups, "Northern Birds Head" and the "Southern Birds Head" on the southern side of the Birds Head Peninsula and the Birds Neck region (Figs 1, 5). New samples included in this study reinforce the boundary between the "northern" and "southern Birds Head" groups. That is, rainbowfishes in drainages that enter up to the western edge of Berau Bay are part of a distinct clade ("southern Birds Head" group) relative to those in drainages west and north of this region ("northern Birds Head" group). This is likely influenced by low sea level drainage patterns which would potentially allow historical connections between drainages within Berau Bay and the adjacent Bintuni Bay (Fig. 1).

Several Birds Head rainbowfishes are separated by relatively small genetic divergences, such as M. aja-maruensis and M boesemani with a p-distance of 0.4%, M koktzsensis and M. sneideri by 0.6%, M ericrobertsi and M fasinensis by 1.3%. These values are similar to or less than variation within some Birds Head rainbowfishes such as M synergos I and 11 (1.2%), M fredericki I and 11 (1.0%), and M catherinae I and 11 (0.7%). However, most of these closely related species pairs in western New Guinea are located at different sites within the same drainage (except M. kokasensis and M sneiderz), whereas different populations of the same species are found in separate well-isolated drainages, which helps to explain their larger genetic divergences. Clearly though these closely-related species pairs have remained sufficiently isolated from one another despite their close geographic proximity to maintain these genetic divergences as well as having usually evolved differences in colouration, meristic counts and/or morphometric proportions. These differences beg the question as to whether intervening populations represent a series of intermediate variants, or whether species are separated by distinct boundaries due to differences in habitats or barriers. Only further sampling for each species pair can resolve that question. In the meantime, the combination of evidence supports their status as distinct species.

ACKNOWLEDGEMENTS

We are grateful for the generous support of the National Geographic Society who funded Gerald Allen's field work in 1999, which lead to the discovery of M multiradiata. We also thank Samuel Renyaan and Mark Allen, who participated in the 1999 expedition. We also acknowledge the support of the Ministry of Research and Technology (RIS-TEK) and the Indonesian Institute of Sciences (LIPI), who provided collection and research permits. We are indebted to Hans-George Evers and Jeffrey Christian, who collected the type specimens of M laticlavia and also provided critical tissue samples of this and other species. Heiko Bleher and Wolfgang Tins aided Gerald R. Allen with the collection of M ericrobertsi in 1982. Gary Lange and Natasha Khardina generously provided important photographs of live specimens of M ericrobertsi and M. irianjaya respectively. We are also grateful for the assistance provided by Sopian (MZB), Sue Morrison (WAM), and Jeff Clayton (USNM) in assisting with fin ray counts and providing museum registration numbers. Most of the DNA sequencing was conducted in the laboratory of Jerald Johnson (Brigham Young University) whose help is gratefully acknowledged.

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1. Melanotaenia ajamaruensis, males, photo by H.-G. Evers. 2. Melanotaenia ajamaruensis, female, photo by EL-G. Evers. 3. Melanotaenia boesemani, male, Lake Ayamaru, Irian Jaya, Indonesia, photo by H. Bleher. 4. Melanotaenia boesemanni, Aytinjo Lake, bred by M. Dielen, photo by N. Khardina. 5. Melanotaenia fasinensis, female, photo by H.-G. Evers. 6. Melanotaenia fasinemis, male, photo by H.-G. Evers.

Gerald R. Allen(1), Peter J. Unmack(2, 3) and Renny K. Hadiaty(4)

1) Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986.

E-mail: tropical_reef@bigpond.com

2) Institute for Applied Ecology and Collaborative Research Network for Murray-Darling Basin Futures, University of Canberra, ACT 2601, Australia.

3) Department of Biology, 401 WIDB, Brigham Young University, Provo UT 84602, USA.

4) Museum Zoologicum Bogoriense (MZB), Division of Zoology, Research Centre for Biology, Indonesian Institute of Sciences (LIPI), Jalan Raya Bogor, Km 46, Cibinong 16911, Indonesia.
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Author:Allen, Gerald R.; Unmack, Peter J.; Hadiaty, Renny K.
Publication:aqua: International Journal of Ichthyology
Date:Jul 29, 2014
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