melanotaenia flavipinnis, a new species of rainbowfish ...1)-melanotaenia-flavipinnis.pdf ·...

AbstractA new species of melanotaeniid rainbowfish, Melanotae-

nia flavipinnis, is described from Misool Island in the BirdsHead region of western New Guinea (West PapuaProvince, Indonesia). The new taxon is described from 20specimens, 38.5-77.2 mm SL, collected from two creekson southeastern Misool. It is the second rainbowfishknown from this island, which lies about 67 km south ofthe New Guinea mainland. It is distinguished from con-geners, and especially from M. misoolensis, by a combina-tion of features that include a distinctive colour patternwith yellow dorsal, anal, and pelvic fins, usually 18-19 rak-ers on the first gill arch, 11-16 cheek scales, 12-13 circum-peduncular scales, first dorsal fin situated well behind analfin origin, presence of both vomerine and palatine teeth,conspicuously rounded anal fin with relatively elongaterays on the anterior half, and relatively slender body inboth sexes (greatest depth usually less than 35% SL).

ZusammenfassungBeschrieben wird eine neue Art der Regenbogenfische,

die den Melanotaeniiden zuzurechnen ist, von der InselMisool in der Region Birds Head, West-Neuguinea (Prov-inz West-Papua, Indonesien). Grundlage für die Erst-beschreibung sind 20 Exemplare von 38,5-77,2 mm SL,die in zwei Fließgewässern im Südosten Misools gefangenwurden. Es ist der zweite Nachweis einer Regenbogenfis-chart auf dieser Insel, die etwa 67 km südlich vom FestlandNeuguineas liegt. Die neue Art unterscheidet sich von denanderen Angehörigen der Gattung und insbesondere vonM. misoolensis durch folgende Kombination von Merk-malen: auffällige Farbgebung mit gelben Rücken-, After-und Bauchflossen; gewöhnlich 18-19 Kiemenreusen aufdem ersten Kiemenbogen; 11-16 Schuppen im Wangen-bereich, 12-13 Schuppen um den Schwanzstiel; ersteRückenflosse deutlich hinter dem Afterflossenansatz;

Vorhandensein von Schlund- und Gaumenzähnen; deut-lich rundliche Afterflosse mit relativ langen Flossen-strahlen in der vorderen Hälfte; relativ schlanker Körperbei beiden Geschlechtern (größte Tiefe normalerweiseweniger als 35% der Körperlänge SL).

RésuméUne nouvelle espèce de Mélanotaeniidé, Melanotaenia

flavipinnis, est décrit provenant de l’île Misool, dans la régionde la Birds Head, à l’ouest de la Nouvelle-Guinée (provincede West Papua, Indonésie). Elle est décrite sur base de 20spécimens, 38,5-77,2 mm de LS, collectés dans deux petitscours d’eau, au sud-est de Misool. C’est le deuxième poissonarc-en-ciel connu dans cette île qui se trouve à environ 67km au sud de la Nouvelle-Guinée proprement dite. Cetteespèce se distingue de ses congénères, et notamment de M.mi soolensis, par une combinaison de caractéristiques com-prenant un patron de coloration particulier avec dorsale,anale et pelviennes jaunes, 18-19 branchiospines enmoyenne sur le premier arc branchial, 11-16 écailles sur lesjoues, 12-13 écailles circumpédonculaires, la première dor-sale située bien après la naissance de l’anale, la présence dedents vomérines aussi bien que palatines,, une anale claire-ment arrondie avec des rayons relativement allongés sur lamoitié antérieure et un corps plutôt élancé pour les deuxsexes (la plus grande hauteur habituellement moins de 35%de la LS).

SommarioUna nuova specie di pesci arcobaleno, Melanotaenia

flavipinnis, è descritta dall’isola di Misool nella regione diBirds Head, Nuova Guinea occidentale, Indonesia. Ilnuovo taxon è descritto sulla base di 20 esemplari di 38,5-77,2 mm SL, raccolti in due torrenti della regione sudori-entale di Misool. È la seconda specie di pesci arcobalenonoti in questa isola, che si trova a circa 67 km a sud della

aqua vol. 20 no. 1 - 15 January 201435

aqua, International Journal of Ichthyology

Melanotaenia flavipinnis, a new species of Rainbowfish (Melanotaeniidae) from Misool Island, West Papua Province, Indonesia

Gerald R. Allen1, Renny K. Hadiaty2 and Peter J. Unmack3, 4

1) Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected]

2) Museum Zoologicum Bogoriense (MZB), Division of Zoology , Research Centre for Biology, Indonesian Institute of Sciences (LIPI), Jalan Raya Bogor, Km 46, Cibinong 16911, Indonesia.

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

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

Received: 13 December 2013 – Accepted: 12 January 2014

Nuova Guinea. Si distingue dalle congeneri, e soprattuttoda M. misoolensis, per una combinazione di tratti morfo-logici che includono una caratteristica colorazione gialladelle pinne dorsale, anale e pelviche, 18-19 rastrelli sulprimo arco branchiale, 11-16 scaglie sulla guancia, 12-13scaglie circumpeduncolari, prima pinna dorsale inseritaben dietro l’origine della pinna anale, presenza di dentivomerini e palatini, pinna anale vistosamente arrotondatacon i raggi della metà anteriore relativamente allungati ecorpo relativamente snello in entrambi i sessi (altezza mas-sima di solito meno del 35% SL).

INTRODUCTIONRainbowfishes of the family Melanotaeniidae are

exceptionally abundant throughout their distribu-tional range, which encompasses the northern two-thirds of Australia, the island of New Guinea, andthe nearby Aru Archipelago. Clear, runningstreams and lakes are the preferred habitats,although they also occur in turbid waters, ponds,swamps, and isolated rocky pools in otherwise drystreambeds. Most species form loose aggregations,which swim either in midwater or just below thesurface. The main dietary items include insectswhich fall onto the surface and micro-crustaceans.Spawning occurs year round in most species, butreproductive activity often peaks at the onset ofrainy periods. The family contains seven genera (Allen 1980) of

which Melanotaenia Gill 1862 is by far the largestwith 65 valid, described species and numerous oth-ers still waiting description. Allen (1995) provideda popular account of the family, including colourillustrations, and a summary of biological informa-tion for the 53 species known at that time. Thecurrently recognized genera include: CairnsichthysAllen 1980 (1 species), Chilatherina Regan 1914(11 species), Glossolepis Weber 1907 (9 species),Iriatherina Meinken 1974 (1 species), Melanotae-nia (65 species), Pelangia Allen 1998 (1 species),and Rhadinocentrus Regan 1914 (1 species). Recent genetic evidence (Unmack et al. 2013)

indicates the genus contains three main lineages,western, northern and southern, correlated respec-tively with the following major biogeographicregions: the Birds Head Peninsula of western NewGuinea, northern New Guinea, and the combinedsouthern New Guinea/Australia region. These dataalso reveal that Chilatherina and Glossolepis ofnorthern New Guinea as presently understood arepolyphyletic and most likely should be integratedinto the genus Melanotaenia, which is the oldestname for the group.

The Birds Head region and adjacent Birds Neck ofWest Papua (Fig. 1) has proved a particularly fertilearea for recent discoveries and will certainly yieldadditional new species with further exploration.Allen (1990) provided a summary of the sevenspecies known to inhabit the Birds Head mainlandat that time. A total of 21 species are currentlyknown from the region, including the offshoreislands of the Raja Ampat Group (Table I). The pre-sent paper describes a new Melanotaenia from theisland of Misool, which lies about 67 km south ofthe Birds Head mainland. It was collected by thefirst author in 1999 during a survey of freshwaterfishes of the Raja Ampat Islands. The fish was ini-tially identified as M. misoolensis, the only memberof the family known to occur at Misool. However,the colour pattern did not match specimens fromthe type locality and a visit to a nearby stream inJanuary 2013 afforded the opportunity to obtainmore specimens and photographs, facilitating moredetailed comparisons with M. misoo lensis.

MATERIALS AND METHODSCounts and measurements that appear in paren-

theses refer to the range for paratypes if differentfrom the holotype. Type specimens are deposited atMuseum Zoologicum Bogoriense, Cibinong, Java,Indonesia (MZB), National Museum of NaturalHistory, Washington, D.C. (USNM), and WesternAustralian Museum, Perth (WAM). Comparativematerial of Melanotaenia misoolensis is deposited atthe Naturalis Biodiversity Center, Leiden, Nether-lands (formerly at Zoologisch Museum, Amster-dam and therefore bearing the acronym ZMA).Comparative material examined: Melanotaenia

misoolensis (all from Wai Tama, tributary of KasimRiver, 1°51.635’S, 129°55.401’E, northwesternMisool, Raja Ampat Islands, Papua Barat Province,Indonesia ) – ZMA 116.456, male, 58.5 mm SL(holotype); MZB 21472, 9 specimens, 33.0-62.3mm SL; USNM 227492, 3 specimens, 30.6-39.5mm SL (paratypes); WAM P. 27279-001, 2 speci-mens, 52.9-58.5 mm SL (paratypes); WAMP.31550-001, 17 specimens, 31.3-65.5 mm SL;WAM P.32240-001, 55 specimens, 24.1-62.1 mmSL; ZMA 116.457, 17 specimens, 21.9-52.7 mmSL (paratypes). M. salawati – WAM P.31469-001,15 specimens, 33.0-93.6 mm SL, Kali Doktur,1°01.036’S, 130°39.943’E, Salawati Island, RajaAmpat Islands, Papua Barat Province, Indonesia.The methods of counting and measuring are as

follows: dorsal and anal-rays – the last ray of the

aqua vol. 20 no. 1 - 15 January 2014 36

Melanotaenia flavipinnis, a New Species of Rainbowfish (Melanotaeniidae) from Misool Island, West Papua Province, Indonesia

anal and second dorsal fins is divided at the baseand counted as a single ray; lateral scales – numberof scales in horizontal row from upper edge of pec-toral-fin base to caudal-fin base, excluding thesmall 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 ofnape in front of first dorsal-fin; cheek scales – totalnumber of scales covering suborbital and preoper-culum; standard length (SL) – measured from tip ofupper lip to caudal-fin base; head length (HL) –

measured from tip of upper lip to upper rear edgeof gill opening; caudal peduncle depth is least depthand caudal peduncle length is measured betweentwo vertical lines, one passing through base of lastanal ray and the other through caudal-fin base;caudal concavity – horizontal distance between ver-ticals at tips of shortest and longest rays.Rainbowfish species used to generate sequence

data are shown in Table IV. Methods for obtainingDNA sequence data and their analyses followsAllen & Unmack (2012) except where noted as fol-lows. Phylogenetic analyses were performed withmaximum likelihood (ML) using GARLI 2.0


Gerald R. Allen, Renny K. Hadiaty and Peter J. Unmack

Fig. 1. Map of West Papua Province, Indonesia showing main areas inhabited by Melanotaenia including the Birds HeadPeninsula, Bomberai Peninsula, Birds Neck region and islands of Waigeo, Batanta, Salawati, and Misool. Black dots and starson Misool represent the collecting sites for M. misoolensis (Wai Tama) and M. flavipinnis (Gam River and Ifaupan Creek)respectively. The thick black line through the Birds Head represents the geographic limits of the “northern” and “southernBirds Head” groups. The yellow region represents the southern lineage of New Guinea, while the red region represents apoorly explored region between the southern and western lineages. The dotted line represents the -135 bathymetric contourwhich is the approximate position of the coastline during the last glacial phase.

(Zwickl 2006) with bootstrapping for 1,000pseudoreplicates using the model of sequence evo-lution TrN+G+I, which was the best one identifiedby ModelTest 3.7 (Posada & Crandall 1998). Toget the ML topology we ran GARLI with 10 searchreplicates with the following default setting valueschanged: streefname = random; attachmentsper-taxon = 34; genthreshfortopoterm = 100,000;scorethreshforterm = 0.05; significanttopochange =0.00001. For bootstrapping we ran 1000 replicateswith the previous settings with the followingchanges: genthreshfortopoterm = 10,000; signifi-canttopochange = 0.001; treerejectionthreshold =20 as suggested in the GARLI manual to speed upbootstrapping. Trees were mid-point rooted.

Melanotaenia flavipinnis, n. sp.Misool Yellowfin Rainbowfish(Figs 2-5; Tables II-III)

Holotype. MZB 21470, male, 71.5 mm SL, Ifau-pan Creek, 2°00.653’S, 130°14.124’E, Misool,Raja Ampat Islands, Papua Barat Province, Indone-sia, 0-.1.5 m, seine net, G. Allen, M. Erdmann &H. Bleher, 20 January 2013.

Paratypes (collected with holotype unless statedotherwise): MZB 21471, 4 specimens, 55.3-61.5mm SL; USNM 427072, 4 specimens, 51.4-60.0mm SL; WAM P. 33830-001, 5 specimens: 47.5-77.2 mm SL; WAM P.31552-001, 6 specimens, 38.5-71.4 mm SL, small tributary creek at junction withGam River, approximately 1°57’S, 130°11’E, Mis-ool, Raja Ampat Islands, Papua Barat Province, Indo -nesia, 0-1 m, seine, G. & M. Allen, 24 April 1999.Diagnosis: A species of melanotaeniid rainbow-

fish distinguished by the following combination ofcharacters: dorsal rays usually V or VI-I, 11 to 14(usually 13); anal-rays I,22-26 (usually 23 to 25);pectoral-rays 12 to 14 (usually 14); lateral scales34-36 (usually 35 or 36), predorsal scales 16 to 19(usually 17 or 18); circumpeduncular scales 12 or13; total gill rakers on first arch 16 to 19 (usually18 or 19); total scales covering preoperculum 11-16 (average 13.2); origin of first dorsal-fin clearlyanterior to anal-fin origin; longest anal-fin ray con-spicuously greater than caudal peduncle depth;greatest body depth of adult male relatively slender,2.8-3.3 in SL; colour in life generally greenish dor-sally with faint yellow stripe between each horizon-tal scale row; prominent midlateral blue stripe onposterior half of body, initially one scale row in



Table I. List of valid Melanotaenia species from the western lineage currently known from the Birds Head and Birds Neckregion of Papua Barat Province, Indonesia.

Species and author(s) Approximate distribution

M. ajamaruensis Allen & Cross, 1980 Ajamaru Lakes vicinity, Birds HeadM. ammeri Allen et al, 2008 Arguni Bay vicinity, Birds NeckM. angfa Allen, 1990 Yakati River district, Birds NeckM. arfakensis Allen, 1990 Near Manokwari, Birds HeadM. arguni Kadarusman et al, 2012 Arguni Bay vicinity, Birds NeckM. batanta Allen & Renyaan, 1998 Batanta, Raja Ampat IslandsM. boesemani Allen & Cross, 1980 Ajamaru Lakes vicinity, Birds HeadM. catherinae de Beaufort, 1910 Waigeo, Raja Ampat IslandsM. fasiensis Kadarusman et al, 2010 Lake Ajamaru vicinity, Birds HeadM. flavipinnis Allen et al, new species Misool, Raja Ampat IslandsM. fredericki (Fowler, 1939) Western Birds HeadM. irianjaya Allen, 1985 Bomberai Peninsula and Birds HeadM. kokasensis Allen et al, 2008 Bomberai PeninsulaM. mairasi Allen & Hadiaty, 2011 Kaimana district, Birds NeckM. misoolensis Allen, 1982 Misool, Raja Ampat IslandsM. parva Allen, 1990 Yakati River district, Birds NeckM. salawati Kadarusman et al, 2011 Salawati, Raja Ampat IslandsM. sneideri Allen & Hadiaty, 2013 Bomberai Peninsula, Birds HeadM. synergos Allen & Unmack, 2008 Batanta, Raja Ampat IslandsM. urisa Kadarusman et al, 2012 Arguni Bay vicinity, Birds NeckM. veoliae Kadarusman et al, 2012 Arguni Bay vicinity, Birds Neck

width, but expanding to two rows (midlateral androw just below) on caudal peduncle; median finsgenerally yellow, except caudal fin greyish brown.Description: Dorsal-rays V-I,12 (IV to VI,11 to

14); anal-rays I,23 (I,22 to 26); pectoral-rays 14(12 to 14); pelvic-rays I,5; branched caudal-rays15; lateral scales 35 (34 or 36); transverse scales 10;predorsal scales 18 (16 to18); cheek scales 14 (11-16); gill rakers on first branchial arch 3 + 16 (2-3 +13-17), total gill rakers on first arch 19 (16-19).Body depth 2.9 (2.8-3.6) in SL, head length 3.7

(3.5-3.9) in SL; greatest width of body 2.7 (2.2-2.8) in greatest body depth; snout length 2.9 (2.7-3.2) in HL; eye diameter 3.5 (2.9-3.6) in HL;interorbital width 2.8 (2.6-3.1) in HL; depth ofcaudal peduncle 2.1 (2.2-2.6) in HL; length ofcaudal peduncle 1.8 (1.4-1.8) in HL.Jaws about equal, oblique, premaxilla with an

abrupt bend between the anterior horizontal portionand lateral part; maxilla ends below about anterioredge of pupil or just anterior to this point; maxillarylength 2.8 (2.7-3.2) in HL; lips thin; teeth conical

with slightly curved tips, extending on to outer sur-face of lips; teeth of upper jaw in 4-5 irregular rowsanteriorly, reduced to a single row posteriorly, wherethey are exposed when mouth is closed; teeth inlower jaw in about 7-10 irregular rows anteriorly,reduced to 1 or 2 rows posteriorly; narrow row con-taining small, conical teeth on vomer and palatines(usually embedded in congealed mucous). Scales of body cycloid, relatively large, and arranged

in regular horizontal rows; scale margins weaklycrenulate; predorsal scales extending forward toabout middle of interorbital space; preopercle with 2-3 scale rows between its posterior angle and eye.Predorsal length 1.9 (1.9-2.1) in SL; preanal length

2.1 (2.0-2.2) in SL; prepelvic length 2.7 (2.6-2.8) inSL; length of second-dorsal fin base 4.2 (4.2-4.8) inSL; length of anal-fin base 2.2 (2.3-2.6).First dorsal-fin origin about level with base of

fifth or sixth soft anal-fin ray; longest spines (usu-ally third or fourth) of first dorsal-fin 1.7 (1.7-2.5)in HL, its depressed tip reaching spine or first softray of second dorsal-fin in females and reaching to



Fig. 2. Aquarium photograph of freshly collected holotype of Melanotaenia flavipinnis, male (lower), 71.5 mm SL and female,58.5 mm SL (MZB 21470). Photo by G. R. Allen

about base of second or third soft ray in maturemales; longest (generally anteriormost in femalesand middle rays in males) rays of second dorsal-fin1.8 (1.9-2.4) in HL, depressed posterior raysextending about one-half length of caudal pedun-cle in females and full length of caudal peduncle inmature males; longest (sixth to ninth rays in malesand females) anal-rays 1.6 (1.6-2.0) in HL; pelvicfin tips when depressed reaching to base of thirdor fourth soft anal-fin ray in mature adults; lengthof pelvic fins 1.4 (1.3-1.7); length of pectoral fins1.3 (1.2-1.4) in HL; length of caudal-fin 1.3 (1.1-

1.4) in HL; caudal-fin moderately forked, caudalconcavity 3.2 (2.8-4.8) in head length.C o l o u r o f m a l e h o l o t y p e i n l i f e (Fig.

2, lower fish): head brownish dorsally and onsnout; preoperculum yellowish to white; opeculumpale blue with dark blue stripe crossing uppermostportion with silvery white patch immediatelybelow; green on dorsal half of body; pale yellowstripe between each scale row of body; dark bluemidlateral stripe on posterior half of body, initiallyone scale row in width, but expanding to two rows(midlateral and row just below) on caudal pedun-



Fig. 3. Aquarium photograph of freshly collected paratype of Melanotaenia flavipinnis, male, 71.4 mm SL (WAM P.31552-001), Gam River, Misool. Photo by G. R. Allen.

Fig. 4. Melanotaenia flavipinnis, freshly collected male holotype, 71.5 mm SL. Photo by G. R. Allen.



Fig. 5. Melanotaenia flavipinnis, preserved male holotype, 71.5 mm SL and female paratype (WAM P. 33830-001), 77.2 mmSL. Photo by G. R. Allen.

Table II. Rainbowfish species from the Birds Head region of Papua Barat Province used in the phylogenetic analysis includ-ing locality data, the number of individuals examined and their GenBank accession number. In the locality field the code ASindicates fish were sourced from rainbowfish aquarium hobbyists.

Species Locality N GenBank #

Melanotaenia ammeri trib to Arguni Bay, Birds Neck 4 KC133615.1Melanotaenia angfa AS, Yakati R, Birds Neck 3 KC133616.1Melanotaenia arfakensis AS, Prafi R, Birds Head 2 KC133617.1Melanotaenia batanta Warmon Ck, Batanta 3 JQ282008.1Melanotaenia boesemani AS, Birds Head 3 KC133618.1Melanotaenia fredericki I AS, Birds Head 2 KC133621.1Melanotaenia fredericki II AS, Sth of Sorong, Birds Head 2 KC133619.1Melanotaenia flavipinnis Ifaupan Ck, Misool 2 KF954094-5Melanotaenia kokasensis Kokas village, Bomberai 7 KC133623.1Melanotaenia misoolensis I Wai Tama, Misool 5 KC133624.1Melanotaenia misoolensis II Wai Tama, Misool 2 KC133625.1Melanotaenia parva AS, L Kurumoi, Birds Neck 4 KC133626.1Melanotaenia salawati Kali Doktur, Salawati Is 9 KC133620.1Melanotaenia sp. Rawarra Sebyar R, Birds Head 2 KC133627.1Melanotaenia sp. Suswa Karabara R, Birds Head 2 KC133628.1

cle; dark blue stripe on upper operculum contin-ued just above and behind pectoral-fin base, incor-porating 2-3 midlateral scales and connected topreviously mentioned stripe by row of pale bluescales with dark blue margins; three lowermosthorizontal scale rows of side with mauve streak oncentre of each scale; side of breast bluish, some-times with pink blush; dorsal, anal and pelvic-finsyellow; caudal-fin translucent grey to brownish;pectoral fins translucent. C o l o u r p a t t e r n o f f e m a l e i n l i f e (Fig.

2, upper fish): similar to that of male except some-times lacking prominent dark blue midlateralstripe, in which case this feature mainly light blue.Live male paratypes (Fig. 3) from the nearby GamRiver were generally similar in colour to the holo-type, although they exhibit a slightly more yellow-ish hue on the dorsal half of the body with less yel-low on the ventral half. In addition, the black mid-lateral stripe on the posterior body of the pho-tographed specimen was uniformly two scaleswide, rather than initially covering a single scalerow. Additional photographs of the new specieswere provided by Tuskin (2010).

C o l o u r o f h o l o t y p e i n a l c o h o l (Fig.5): head brownish dorsally, grading to tan ven-trally; opeculum with blackish stripe crossinguppermost portion with silvery patch immediatelybelow; light brown on back, grading to whitishventrally; each scale of back with dark brown pos-terior margin; midlateral stripe similar to livedescription above, but generally blackish; blackishstripe on upper operculum continued just aboveand behind pectoral-fin base, incorporating 2-3midlateral scales and connected to previously men-tioned stripe by row of tan scales with blackishmargins; fins generally translucent, except caudaldusky grey. Paratypes collected in 1999 (WAMP.31552-001) exhibit slightly less vivid dark mark-ings, but are generally similar, although overall yel-lowish tan.Sexual dimorphism: less pronounced compared

with many Melanotaenia, which are frequently char-acterised by males having a much deeper body (upto about 50 percent of SL) and longer dorsal andanal fin-rays. In M. flavipinnis the adult male is onlyslightly deeper bodied, and has slightly longer fin-ray elements than in females. The longest soft dor-



Table III. Proportional measurements of selected type specimens of Melanotaenia flavipinnis expressed as percentage of thestandard length.

Holotype Paratype Paratype Paratype Paratype Paratype Paratype ParatypeMZB WAM WAM WAM MZB USNM MZB WAM21470 P.33830 P.31552 P.33830 21471 427072 21471 P.33830

Sex male female female male male male female femaleStandard length (mm) 71.5 77.2 66.8 66.6 60.7 59.6 58.5 47.5Body depth 34.8 31.3 28.7 32.4 31.6 32.0 28.7 27.8Body width 13.0 13.7 12.6 12.8 12.4 13.8 12.5 12.0Head length 27.3 26.8 25.6 25.8 26.4 26.3 26.3 26.5Snout length 9.4 9.3 9.3 9.6 9.6 9.4 9.1 8.4Maxillary length 9.7 9.7 9.1 9.5 9.4 9.1 8.9 8.6Eye diameter 7.7 7.5 7.6 7.8 8.6 8.6 8.5 8.4Bony interorbital width 9.7 9.6 10.0 9.8 9.2 9.7 9.1 8.6Depth of caudal peduncle 13.0 11.1 11.4 11.7 11.4 11.6 10.6 10.9Length of caudal peduncle 15.4 15.0 17.1 16.8 17.8 17.3 18.3 17.1Predorsal distance 52.9 52.3 49.0 49.1 52.4 49.5 48.5 49.9Preanal distance 47.6 50.1 46.3 49.2 49.8 50.2 47.0 48.6Prepelvic distance 37.5 37.0 35.3 36.0 37.1 37.8 36.8 37.32nd dorsal-fin base 24.1 24.1 21.4 21.3 23.4 24.0 23.1 22.3Anal-fin base 45.2 41.2 39.8 40.5 40.9 42.3 42.9 41.3Pectoral-fin length 21.0 19.0 20.1 19.8 19.4 20.5 19.8 18.3Pelvic-fin length 18.9 16.2 19.6 15.8 17.6 17.8 18.1 15.6Longest ray 1st dorsal-fin 16.4 12.4 12.6 14.4 13.7 15.9 13.7 13.5Longest ray 2nd dorsal-fin 15.1 11.0 13.6 12.6 13.5 14.1 12.3 14.3Longest anal-ray 16.9 14.2 13.5 14.6 14.3 14.1 13.7 14.3Caudal-fin length 20.4 19.6 21.0 20.7 2.8 21.0 22.4 21.9Caudal concavity 8.5 6.6 7.5 7.8 7.4 7.4 5.5 8.0

sal-fin rays of males are located in the middle por-tion of the fin, in contrast to that of females, whichare situated anteriorly. In addition, the depressedfirst dorsal fin of adult males extends to the base ofthe second or third soft ray of the second dorsal fin,compared with the spine or first ray in females.Body depth generally increases, especially in

males, with increased growth and is therefore mostuseful as a distinguishing character when assessedon the basis of sex and size range as follows: males– 55-69 mm SL, 30.7-33.7% SL (x, = 32.3, N = 8);

males – 70+ mm SL, 34.8-36.1% SL (x, = 35.5, N = 2); females less than 45 mm SL – 29.1% SL (N = 1); females – 45-54 mm SL, 27.8-30.1% SL(x, = 28.6, N = 3); females – 55-69 mm SL, 28.7-32.1% SL (x, = 30.2, N = 5); females – 70+ mmSL, 31.3% SL (N = 1). All type specimens are sex-ually mature individuals. The smallest female withripe eggs is 38.5 mm SL (WAM P.31552-001).Remarks: Allen (1982) described M. misoolensis

from 23 specimens collected by a Dutch expedi-tion in 1948. The specimens were located during a



Fig. 7. Aquarium photograph of Melanotaenia salawati, male, 93.6 mm SL (WAM P.31469-001), Salawati, Raja AmpatIslands, West Papua Province, Indonesia. Photo by G. R. Allen.

Fig. 6. Aquarium photograph of Melanotaenia misoolensis, male, approximately 60 mm SL, Wai Tama, Misool. Photo by G. R. Allen.

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–

visit to the Zoological Museum in Amsterdam.Subsequent trips were made by the first author tothe type locality in 1999 and 2002 with the goal ofobtaining additional specimens as well as geneticsamples and live photographs. The type locality atthe Wai Tama (tributary of the Kasim River sys-tem, Fig. 1) is situated in the northwestern portionof the island. During the 1999 trip numerousstreams were sampled while circumnavigating theisland, but only one other location yielded Melano -taenia specimens, the Gam River in the southeast-ern portion, or approximately 40 km distance from

the Wai Tama site. Initially, the fishes from bothsites were identified as M. misoolensis. Although,they differed in colour pattern, most meristic andmorphometric features were very similar. Geneticanalysis by the third author indicated they weremost likely a single species. The Gam River fishwas also collected by Heiko Bleher and NatashaKhardina at nearby Ifaupan Creek in 2001. Thedistinctly different appearance of fish from the twolocations and the opportunity to collect additionalspecimens from Ifaupan Creek in 2013 by H. Ble-her, M. Erdmannn, and G. Allen, provided impe-



Table IV. Summary of dorsal, anal, and pectoral fin-ray, gill-raker, cheek-scale, and circumpeduncular-scale (CPS) countsfor Melanotaenia flavipinnis. Pectoral-ray counts were taken on both sides of each individual.

First Dorsal Fin Spines Soft Dorsal Rays Soft Anal Rays

IV V VI 11 12 13 14 22 23 24 25 26

1 12 7 11 4 11 4 3 4 7 5 1

Pectoral Rays Total Gill Rakers CPS Cheek

12 13 14 16 17 18 19 12 13 11 12 13 14 15 16

1 6 33 2 3 7 7 13 7 1 7 2 8 1 1

Fig. 8. Maximum likelihood tree for Birds Head region Melanotaenia species based on analysis of cytochrome b sequences(1,141 bp). Bootstrap values were obtained from 1,000 replicates.

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tus for a re-examination of their status. Althoughthe two main river systems containing Melanotae-nia, the Kasim and Gam, originate relatively close(within 4-5 km) to each other in central-southernMisool, the two systems comprise well separateddrainages, with the Kasim flowing towards thenorthwest and the Gam to the southeast (Fig. 1).The new species differs most notably from M.

misoolensis, with regards to colour pattern, relativepositions of the first dorsal and anal fins, andheight of the anal fin. Although both species havea prominent dark blue midlateral stripe (at least inadult males), that of M. misoolensis (Fig. 6) is moreirregular in appearance, consisting of a solid poste-rior section encompassing two scale rows (asopposed to 1-2 rows in M. flavipinnis), anteriorsection embracing about 4-5 scales and usuallyextending onto upper half of adjacent scale rowbelow (vs. 2-3 scales in M. flavipinnis), and 2-3 iso-lated dark patches (each covering 1-2 midlateralscales and part of adjacent scale row above)between the anterior and posterior sections. Otherconsistent differences found in M. misoolensisinclude the predominately whitish or translucent

dorsal and anal-fins that grade to pale yellow dis-tally (vs. entirely yellow), whitish pelvic-fins (vs.yellow), large red spot on the operculum (vs. whitespot), and lack of yellow colouration on the body. The first dorsal-fin of M. flavipinnis is situated

well behind that of anal-fin. The dorsal origin isopposite (or in vertical alignment) with the base ofthe fifth or sixth soft anal-ray. This position con-trasts notably with the condition in M. misoolensisin which the two fins are approximately oppositeeach other. Finally, there is a pronounced differ-ence in the height of the tallest anal-fin raysbetween the two species. The length of the tallestrays of M. flavipinnis is clearly much greater thanthe least depth of the caudal peduncle compared tothe condition for M. misoolensis in which the tallestrays are about equal to the peduncle depth.The new species is remarkably morphologically

similar to M. salawati (Fig. 7) from Salawati Island,Raja Ampat Islands, which is separated from theadjacent Birds Head Peninsula of far western NewGuinea by the narrow (1.5 km) Sele Strait. BothSalawati and Misool were formerly part of theBirds Head mainland during periods of lower sea



Fig. 9. Habitat of Melanotaenia flavipinnis near the type locality, Ifaupan Creek, southeast Misool, West Papua Province.Photo by H. Bleher.

levels (Fig. 1), although the latter island is now sep-arated by 67 km of shallow sea (mainly less than 20-30 m depth). Judging from the close appearance ofthese species it would appear that either they aresimilar due to sharing the same common ancestorfrom which they evolved, or they independentlyconverged on a similar morphology. Both speciesshare most morphological and meristic features,including broadly overlapping scale and fin-raycounts. Moreover, the two species are characterisedby the posterior position of the first dorsal fin inrelation to the anal-fin and both exhibit relativelyelongate anal fin rays. They also have very similarcolour patterns, although the hues of M. salawati aregenerally less vivid than those of M. flavipinnis, par-ticularly with regards to the blue midlateral stripeand yellow colour of the dorsal, anal, and pelvic-fins.The only obvious difference between the twospecies, except for their genetic separation, is theusual possession of fewer preopercular (cheek) scalesin M. flavipinnis (11-16, average of 13.2 vs. 16-19,average of 17.5, n = 11 for M. salawati).Genetic results and discussion: A total of 52

individuals were sequenced from 14 species fromthe Birds Head region for the mtDNA cytochromeb (cytb) gene (Table II). Within species geneticdiversity was low, with only three species (M.flavipinnis, M. fredericki and M. misoolensis) eachcontaining two haplotypes. Sequence analysis ofthe 17 OTUs (Table II) yielded 898 invariant char-acters, 67 variable but parsimony uninformativecharacters, and 176 parsimony informative charac-ters. ML analysis recovered one tree with a likeli-hood score of -3488.148897 (Fig. 8). The relation-ships recovered were broadly congruent with thelarger sequence dataset in Unmack et al. (2013).Rainbowfishes in the Birds Head region form threedeeper lineages. The most divergent is the“Waigeo” group (which was not included in thecurrent analysis) which consists of the species M.catherinae and M. synergos from Waigeo andBatanta islands. The remaining species (includedhere) separate in two groups, “Northern BirdsHead” and the “Southern Birds Head” on thesouthern side of the Birds Head Peninsula and theBirds Neck region (Figs 1, 8). The northern groupis limited to drainages west of Berau Bay, as well asdrainages in the northern Birds Head Peninsulawhile the “Southern Birds Head” group occupiesdrainages that enter Berau Bay (and adjacent Bin-tuni Bay) and south through the Bomberai Penin-sula to drainages in the vicinity of Arguni Bay.

Genetically the relationship between M. flavipin-nis and M. misoolensis are confused based on thepresence of two mtDNA lineages within M. misoo -lensis (Fig. 8). It is unclear whether the OTU M.misoolensis II found in two individuals from WaiTama represents the original mtDNA lineage forthis species, or if it is an artifact of ancestral poly-morphism that has persisted for some unknownreason. Finding two distinct mtDNA lineageswithin a rainbowfish population is quite rare, andit usually only occurs in locations where activehybrid zones exist between rainbowfish species(Unmack, unpub. data). Alternatively, the OTUM. misoolensis I could be a result of past introgres-sion from M. flavipinnis, (which is commonbetween rainbowfishes, Unmack et al. 2013).Nuclear data from the S7 marker lacks sufficientvariation to be able to shed further light on thesealternative explanations. If M. misoolensis I doesrepresent the true mtDNA genome of that speciesthen it does not really differentiate the two speciesvery well. However, it is not uncommon for rain-bowfishes that can be distinguished by other traitsto be virtually identical genetically (Unmack et al.2013). For instance, the three rainbowfish speciesG. incisus, G. pseudoincisus and G. dorityi are allessentially identical genetically, yet have clear sub-stantial meristic, morphological, colour and behav-ioural differences (Unmack et al. 2013). Severalother pairs or trios of rainbowfish species are simi-lar genetically that have morphological differences(Unmack et al. 2013). It is important to recognisethat genetic markers such as cytb primarily differ-entiate species via neutral mutations which takeconsiderable time to accumulate, thus any recentspeciation events will not have had sufficient timefor many neutral differences to have evolvedbetween.Zoogeography and habitat: The new species is

known only from the southeastern portion of theisland of Misool in two drainage systems, GamRiver and Ifaupan Creek, which are separated by adistance of only 4 km. The type locality (Fig. 9)consists of a narrow (about 1-2 m wide), relativelyshallow (to about 0.5 m depth) stream with verygradual gradients flowing through second growthforest, about 3.5 km upstream from the sea. Thetype specimens were mainly collected over sandand gravel bottoms.Qualitative comparison of morphological data

suggests a closest relationship between M. flavipin-nis and M. salawati. Mitochondrial genetic data



provides poor resolution among four of the fiveclosely related species: M. fredericki, M. batanta,M. salawati, M. misoolensis II, and the lineage lead-ing to M. misoolensis I and M. flavipinnis; with thelatter two species well resolved as closely related sis-ter species. As noted by Ballard and Whitlock(2004), mtDNA can produce misleading resultsdue to hybridisation, introgression and gene tree -species tree issues. Unfortunately, the main nuclearmarker currently used in rainbowfishes, S7 pro-vides only minimal differences between thesespecies with no resolution. According to molecu-lar clock estimates (Unmack et al. 2013), based onmultiple mtDNA genes and the nuclear S7 marker,diversification within the group of the five closelyrelated rainbowfishes occurred between a meantime of 2.6-3.1 Ma with a 95% highest posteriordensity of 2.0-3.9 Ma. All five species are geo-graphically proximate to each other at the south-western end of Birds Head Peninsula, with all butM. fredericki occurring on the offshore islands ofBatanta, Salawati and Misool. All of these islandsare separated by shallow seas (20-30 m deep) andhave been linked by broader land connections inthe past during glacial low sea levels when the con-tinental shelf was exposed to the southwest of Mis-ool Island (Fig. 1).It is interesting to speculate on the origin of the

Misool rainbowfishes. The island lies in a complexpart of the earth’s crust, which has not been satisfac-torily explained by various geological interpretations(Froidevaux 1974). It is generally regarded as a por-tion of the Birds Head Plate, although there arearguments to make it a separate microplate that col-lided with the Birds Head in the Oligocene (Pigram& Davies 1987). Judging from rich layers of marinefossils, which overlie the basement rock, the islandwas submerged for much of its history (Simbolon etal. 1984). However, it underwent a series of upliftevents, shaping the modern landscape, which essen-tially consists of a mountainous (to about 400 m)southern coast, that either gradually descends north-ward (western half ) or is uniformly hilly, but lower(eastern half ). The likely ancestral Misool rainbow-fish was perhaps distributed throughout the interiorportion of the island, which may have been tra-versed by a single major river system lying to thenorth of the southern range. However, uplift eventscould have conceivably formed a barrier, resulting inthe current northwest and southeast flowing Kasimand Gam systems.Etymology: The new species is named flavipinnis

(Latin: yellow fin) with reference to the yellowcolour of the dorsal, anal, and pelvic fins.

ACKNOWLEDGEMENTSWe are grateful for the generous support of the

National Geographic Society for funding thatenabled the first author to survey fishes in the RajaAmpat Islands in 1999, including the first visit toMisool where the initial specimens of the newspecies were collected. The 1999 collections wereassisted by Mark Allen and Dan Polhemus. We alsothank Heiko Bleher and partner NatashaKhardina, who collected specimens of M. flavipin-nis at Ifaupan Creek in 2001 and were insistentthat these represented a new taxon. We are espe-cially grateful to William M. Brooks and wifePamela S. Rorke, who provided funding for the2013 visit to Misool. We also thank Ken andJosephine Wiedenhoft, the crew of MV Putiraja,Mark Erdmann, Heiko Bleher, and Chris Papo-rakis for their excellent assistance during this sametrip. Museum registration numbers and curatorialassistance were provided by Jeff Clayton (USNM)and Sue Morrison (WAM). Most of the DNAsequencing was conducted in the laboratory of Jer-ald Johnson (Brigham Young University) whosehelp is gratefully acknowledged.

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ALLEN, G. R. & HADIATY, R. K. 2011. A new species ofrainbowfish (Melanotaeniidae), from western NewGuinea (West Papua Province, Indonesia). Fishes of Sahul25 (1): 602-607.

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ALLEN, G. R., UNMACK, P. J. & HADIATY, R. K. 2008. Twonew species of rainbowfishes (Melanotaenia: Melano-taeniidae), from western New Guinea (Papua, BaratProvince, Indonesia). aqua, International Journal ofIchthyology 14 (4): 209-224.

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APPENDIX I – Aquarium photos taken at Ifaupan Creek, Misool in 2001.



Melanotaenia flavipinnis, (A+B = males, C = female), Ifaupan Creek, showing colour pattern variation. Photos by H. Bleher.

A

B

C



Melanotaenia misoolensis, Kasim river tributary, Misool Island, Irian Jaya, Indonesia. Aquarium specimen bred by MarcelDielen in Belgium. Photo by N.Khardina

Redigobius bikolanus, Ifaupan Creek, Misool.

Eleotris melanosoma, Ifaupan Creek, Misool.

APPENDIX II

The following photos were taken of fishes occurring with Melanotaenia flavipinnis at Ifaupan Creek, Misoolin 2013. Photos by H. Bleher.



Redigobius tambujon, female, Ifaupan Creek, Misool.

Redigobius chrysosoma, Ifaupan Creek, Misool.

Eleotris fusca, Ifaupan Creek, Misool.



Ophieleotris hoedti, Ifaupan Creek, Misool.

Butis amboinensis, Ifaupan Creek, Misool.

Glossogobius illimus, Ifaupan Creek, Misool.

melanotaenia flavipinnis, a new species of rainbowfish ...1)-melanotaenia-flavipinnis.pdf ·...

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