Paracheilinus rennyae, a new species of flasherwrasse (Perciformes: Labridae) from southern Indonesia.
The Indo-Pacific labrid fish Paracheilinus rennyae is described from four male specimens, 52.2-60.4 mm SL, collected in 15-21 m depth off southwestern Flores Island in the Lesser Sunda island chain of Indonesia. It is distinguished from most congeners by the lack of filamentous extensions of the dorsal fin rays in males and a rounded caudal fin margin, a combination of features shared only by P. octotaenia (Red Sea). It differs from the Red Sea species in having 13-14 rakers (versus 16-18) on the first gill arch and several colour pattern differences. Genetic analysis (C01) indicates it is closely related to P. angulatus from the Philippines and northern Borneo (Brunei, Sabah, and Kalimantan), but the two species exhibit marked differences in the shape of the median fins.
Beschrieben wird der indo-pazifische Lippfisch Paracheilinus rennyae auf der Grundlage von vier mannlichen Exemplaren, 52,2-60,4 mm SL, die in 15 bis 21 m Tiefe vor dem Sudwesten der Insel Flores in der Kerte der Lesser-Sunda-Inseln, Indonesien, gefangen wurden. Von den meisten Angehorigen der Gattung lassen sich die Exemplare der neuen Art durch das Fehlen fadiger Anhange an den Ruckenflossenstrahlen bei den Mannchen unterscheiden sowie durch einen rundlichen Schwanzflossenrand--eine Merkmalskombination, die sonst nur bei P. octotaenia (Rotes Meer) vorkommt. Von den Arten des Roten Meeres unterscheiden sic sich durch 13-14 Kiemenreusen (im Vergleich zu 16-18) auf dem ersten Kiemenbogen und durch einige Farbmerkmale. Die Genanalyse (CO1) spricht fur eine nahe Verwandtschaft zu P. angulatus von den Philippinen und Nordborneo (Brunei, Sabah und Kalimantan), doch zeigen die beiden neuen Arten einen deutlichen Unterschied in der Fom der mittleren Flossen.
Le labride de l'Indo-Pacifique, Paracheilinus rennyae, est decrit sur base de quatre specimens males, 52,2-60,4 mm de LS, collectes a une profondeur de 15-21 m au large du sud-ouest de l'ile de Flores, dans les Petites Iles de la Sonde, en Indonesie. L'espece se distingue de la plupart de ses congeneres par l'absence de filaments sur les rayons de la dorsale pour les males et an bord arrondi de la caudale, an ensemble de caracteristiques que ne partage que P. octotaenia (Mer Rouge). Elle differe de cette derniere par 13-14 branchiospines (contre 16-18) sur le premier arc branchial et par plusieurs details du patron de coloration. L'analyse genetique (CO1) montre qu'elle est proche de P. angulatus des Philippines et du nord de Borneo (Brunei, Sabah et Kalimantan), mais les deux especes montrent de nettes differences dans la forme des nageoires medianes.
II labride dell'Indo-Pacifico Paracheilinus rennyaee descritto sulla base di quattro esemplari maschi di 52,260,4 mm SL raccolti a 15-21 m di profondita al largo della costa sud-ovest dell'Isola di Flores, arcipelago della Sonda Minore, Indonesia. Si distingue dalla maggior parte dei congeneri per la mancanza di estensioni filamentose dei raggi dorsali nei maschi e un margine della caudale arrotondato, una combinazione di caratteristiche condivise solo da P. octotaenia (Mar Rosso). Si differenzia dalla specie del Mar Rosso per avere 13-14 rastrelli (contro 16-18) sul primo arco branchiale e molte differenze di colorazione. L'analisi genetica (CO1) indica the e molto vicino a P. angulatus diffuso nelle Filippine e nel Borneo settentrionale (Brunei, Sabah e Kalimantan), ma le due specie presentano marcate differenze nella forma delle pinne mediane.
The species of the labrid genus Paracheilinus Fourmanoir Roux-Esteve and Fourmanoir, 1955, commonly known as flasherwrasses, are a colourful component of Indo-Pacific coral reef fish assemblages. The genus was almost unknown until the widespread use of scuba equipment by ichthyologists. Moreover, the predilection of these fishes for deeper reef environments, particularly rubble bottoms that are otherwise avoided by scuba divers, was also responsible for their lack of detection until relatively recent times. Fourmanoir in Roux-Esteve and Fourmanoir, 1955 described the first member of the genus, P. octotaenia Fourmanoir, a Red Sea endemic. Nineteen years later the second species, P. filamentosus (Allen, 1974) was described from northern Papua New Guinea. Subsequently, divers and especially underwater photographers became more aware of these fishes, resulting in the discovery of numerous species (Randall & Harmelin-Vivien, 1977; Randall & Lubbock, 1981; Comic, 1987; Randall, 1988; Kuiter & Allen, 1999; Randall, 1999; Randall & Allen, 2003, Allen & Erdmann, 2006 and 2008). The genus now contains 17 species including the new taxon described herein.
Flasherwrasses are commonly found in aggregations with an apparent haremic social structure consisting of relatively few large terminal phase males and numerous smaller females and juveniles. Typical habitat consists of low profile coral rubble or Halimeda algae at the base of sloping reefs, usually below 20 m, but occasionally as shallow as 3-5 m. They typically rise well above the bottom to feed on plankton, particularly when currents are prevalent. These fishes are relatively inconspicuous due to their small size (to about 75 mm SL) and usually drab brownish-red colouration (even less conspicuous at depths of 20 m or more due to the rapid attenuation of red wave lengths, rendering the fish grey to the human eye unless illuminated with artificial light). However, during courtship, which occurs daily with a peak about one hour before sunset, terminal males exhibit spectacular nuptial displays, characterized by rapid swimming bursts and enhanced colouration, consisting in part of electric blue, while erecting the dorsal and anal fins.
Although the members of the genus are remarkably similar with regards to most morphometric and meristic features as well as general appearance of initial phase individuals, adult males are readily differentiated, particularly on the basis of their courtship colour patterns. Additional diagnostic characters of male individuals include the shape of the caudal fin, which in most species is lunate or emarginate, and the shape of the soft dorsal rays. With regards to the latter feature, most species have one or more of the soft rays prolonged as thread-like filaments, which may extend for more than half of the standard length in some species.
The present paper describes a new species from southwestern Flores Island and the adjacent Komodo and Rinca islands collected by the authors in July 2013. It first came to our attention when a photo of the fish taken at Rinca Island in the Komodo National Park was sent to us in 2010 by diving colleague Zafer Kizilkaya. The adult male of the new species is highly unique among the genus in both lacking elongate filaments on the dorsal fin and having a rounded caudal fin, a combination of features shared only by P. octotaenia of the Red Sea.
MATERIALS AND METHODS
Lengths given for specimens are standard length (SL), the straight-line distance from the front of the upper lip to the base of the caudal fin (posterior end of the hypural plate). Head length is measured from the same median anterior point to the posteriormost edge of the opercular membrane, and snout length from the same point to the fleshy edge of the orbit. Body depth is the maximum depth, and body width the greatest width just posterior to the gill opening. Orbit diameter is the greatest fleshy diameter, and interorbital width the least bony width. Caudal peduncle depth is the least depth; caudal peduncle length is measured horizontally from the rear base of the anal fin to the caudal fin base. Spines and rays are measured to their extreme base. Pectoral ray counts include the uppermost rudimentary ray. Gill raker counts were made on the first gill arch and include rudiments.
Proportional measurements are presented in Table I as percentages of the standard length. Data in parentheses in the description refer to the range for paratypes if different from that of the holotype.
Table I. Proportional measurements of type specimens of Paracheilinus rennyae expressed as percentage of the standard length. Holotype Paratype Paratype Paratype MZB 21463 USNM WAM WAM 410701 P.33900 P.33900 Sex male male male male Standard length 58.4 60.4 59.2 52.2 (mm) Body depth 30.7 32.3 35.0 29.5 Body width 13.5 13.7 15.2 15.1 Head length 28.1 31.0 29.1 31.4 Snout length 7.0 8.4 7.3 8.2 Eye diameter 7.4 6.6 7.3 8.0 Interorbital 7.4 8.3 7.8 7.9 width Upper jaw 6.7 6.8 6.9 7.3 Depth of caudal 14.9 15.7 14.4 15.3 peduncle Length of 18.2 18.2 18.2 19.2 caudal peduncle Predorsal 30.0 29.1 30.6 31.8 distance Preanal 52.4 54.3 56.4 56.3 distance Prepelvic 32.5 35.3 27.4 34.9 distance Length of 57.0 57.5 56.1 55.6 dorsal fin base 1st dorsal 4.5 4.5 3.9 4.0 spine Last dorsal 17.1 16.6 15.2 14.4 spine Longest soft 27.7 23.7 23.1 21.1 dorsal ray Length of anal 31.5 35.3 29.4 29.3 fin base 1st anal spine 8.7 9.3 8.4 7.3 2nd anal spine 9.8 9.6 9.6 8.4 3rd anal spine 11.0 10.4 11.3 9.4 Longest soft 28.6 25.0 26.2 24.1 anal ray Caudal fin 22.3 24.3 23.5 24.5 length Pectoral fin 20.5 23.7 20.9 21.8 length Pelvic fin 10.4 9.9 9.5 10.3 spine length Pelvic fin 17.1 17.1 16.7 17.6 length
Type specimens are deposited at the Museum Zoologicum Bogoriense, Cibinong, Java, Indonesia (MZB), United States National Museum of Natural History, Washington, D.C. (USNM), and Western Australian Museum, Perth (WAM). In addition, comparative material of P. octotaenia and I? angulatus was examined at Museum national d'Histoire naturelle, Paris (MNHN), the Bishop Museum, Honolulu (BPBM), WAM, and USNM.
Comparative specimens of Paracheilinus angulatus examined include: BPBM 22526 (paratype), 59.9 mm SL, Batangas, Luzon, Philippines; WAM P.33128-004, 40.2 mm SL, Brunei.
Comparative specimens of Paracheilinus octotaenia examined include: BPBM 13856, 8 specimens, 25-69 mm SL, Gulf of Aqaba, Red Sea; MNHN 52-296, 64 mm SL (holotype), Abulat Island, Saudi Arabia, Red Sea; USNM 277592, 86 specimens, 33.9-90.6 mm SL, El Himeira, Gulf of Aqaba, Red Sea.
Tissue samples were obtained from two of the male paratypes of the new species and from 12 individuals of other congeners from the East Indian region as follows: two specimens of P. mccoskeri from the Mergui Archipelago in the Andaman Sea, one specimen of P. carpenteri from Brunei, two specimens of P. filamentosus from New Britain in Papua New Guinea, two specimens of P. cyaneus from Raja Ampat in West Papua, one specimen of 1? walton from Manokwari in West Papua, two specimens of P. angulatus (one from Palawan in the Philippines and one from Brunei), and two specimens of P. flavianalis (one from Nusa Penida in SE Bali and one from SW Flores). All specimens were fixed in 95% EtOH and stored in the Indonesian Biodiversity Research Center (IBRC) Laboratory in Denpasar, Bali (see Table 2 for specimen numbers). Mitochondrial DNA was extracted using a 10% Chelex solution (Walsh et al., 1991). A portion of the cytochrome coxidase subunit I (COI) region was amplified via PCR using the primers: Fish BCL: 5'-TCAACYAATCAYAAAGATATYGGCAC-3' and Fish BCH: 5'-TAAACTTCAGGGTGACCAAAAAATCA-3' and jgLCO: 5'-TITCIACIAAYCAYAARGAYATTGG-3' and jgHCO: 5'-TAIACYTCIGGRTGICCRAARAAYCA-3' (Matt Craig, pers. comm.). The PCR reaction was carried out in 25 [micro]L volumes, using 1 [micro]L of template. Each reaction included 2.5 [micro]L (10x PCR buffer Gold), 2.5 [micro]L (8 mMdNTPs), 2 [micro]L (25 mM Mg[Cl.sup.2] solution), 1.25 [micro]L of each primer at 10 [micro]M, 0.125 [micro]L Amply-TaqGold[TM] (Applied Biosystems) and PE Amplitaq (5 units/[micro]L) (red taq) and 14.5 [micro]L dd [H.sup.2]O.
The thermocyding profile for COI included an initial denaturation of 94[degrees]C for 3 min, 35 cycles of 94[degrees]C for 30s, 53[degrees]C for 30s, and 72[degrees]C for 60s, with a final extension of 72[degrees]C for 2 min. PCR reactions were checked on 1% agarose gels stained with ethidium bromide. PCR product was sequenced at the University of California Berkeley sequencing facility, with all sequences accessioned in GenBank (Table II).
Table II. Data on tissue samples utilized in genetic analysis of southeast Asian Paracheilinus species, including specimen voucher numbers from Indonesian Biodiversity Research Center (IBRC), sample locality name and GenBank accession numbers. Species IBRC Sample Locality of Sample Gen Bank Voucher Number Accession No. P. walton MB040101 Indonesia: KF709097 Manokwari P. mccoskeri MB040301 Myanmar: Mergui KF709098 Archipelago P. mccoskeri MB040302 Myanmar: Mergui KF709099 Archipelago P. cyaneus MB040402 Indonesia: Raja KF709100 Ampar P. carpenteri MB040501 Brunei: Colombo KF709101 Reef P. angulatus MB040601 Philippines: El KF709102 Nido, Palawan P. MB040801 Indonesia: SE Bali KF709103 flavianalis P. angulatus MB040901 Brunei: Colombo KF709104 Reef P. MB041101 Indonesia: SW KF709105 fiavianalis Flores P. rennyae MB041401 Indonesia: SW KF709106 Flores P. rennyae MB041501 Indonesia: SW KF709107 Flores P. cyaneus MB041601 Indonesia: Raja KF709108 Ampat P. MB042002 Papua New Guinea: KF709109 filamentosus New Britain P. MB042003 Papua New Guinea: KF709110 filamentosus New Britain
Forward and reverse sequences were proofread in MEGA5 then aligned using ClustalW (Tamura et al. 2011). Two methods were used to generate phylogenetic reconstructions: maximum likelihood using MEGA5 (Tamura et al., 2011) and MRBAYES 3.2 (Ronquist & Hulsenbeck, 2003). Maximum likelihood analysis was used to assess the model of best fit for the nucleotide substitution. Bayesian Information Criterion (BIC) ranked the Hasegawa Kishino Yano (HKY) model with a discrete Gamma distribution (HKY+G) as having the best fit to the data. This model assumes different rates of transitions and transversions as well as different nucleotide frequencies, and was chosen as the appropriate model of evolution as determined by JModelTest 0.1.1 (Posada, 2008; Guindon & Gascuel, 2003). Bootstrap support was determined using 1000 bootstrap replicates in MEGA5. For the Bayesian analysis we used a Markov Chain Monte Carlo approach with four chains. Analyses were run for 10,000,000 generations with an initial burn-in of 250,000 generations. After 10,000,000 generations, the resulting phylogeny was checked for convergence using the TRACER program (Rambaut & Drummond, 2007).
Paracheilinus rennyae n. sp.
(Figs 1-4; Table I)
Holotype: MZB 21463, male, 58.4 mm SL, southwestern Flores, 08[degrees]48.573'S, 119[degrees]50.200'E, Lesser Sunda Islands, Indonesia, 15-20 m, clove oil and hand net, M. Erdmann, 28 July 2013.
Paratypes (collected with holotype unless stated otherwise): USNM 410701, male, 60.4 mm SL, same data as holotype except 15-21 m, collected with spear, 8 September 2010; WAM P.33900001, 2 male specimens, 52.2-59.2 mm SL.
Diagnosis: Profile of dorsal, anal and caudal fins prominently rounded; tubed scales in posterior section of lateral line 6-9; gill rakers on first arch 13-14; none of dorsal soft rays of males prolonged as filaments; colour of adult males overall orange-red with about 10 broken or complete greyish to blue stripes on side of body, several blue stripes on head (most prominent on snout, below eye, and behind eye), wine-red median and pelvic fins, with broken blue stripes on basal portion and blue outer or anterior margins on pelvic, dorsal and anal fins, frequently yellowish to orange zone at base of last 2-3 dorsal and anal fin rays, and two broken blue stripes and a broad yellow outer margin on caudal fin.
Description: Dorsal fin rays IX, 11; anal fin rays 111,9; pectoral fin rays (including upper rudimentary ray) 14; pored lateral line scales 16 + 9 (one paratype with 15 + 6, and another with 16 + 7); scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 6; median predorsal scales 5 (4-5); median preventral scales 6 (one paratype with 5); rows of scales on cheek 2; circumpeduncular scales 15 (15-16); gill rakers 13 (two paratypes with 14); pseudobranchs 13 (12-13).
Body depth 3.3 (2.9-3.4) in SL; body width 2.3 (1.9-2.3) in body depth; head length 3.6 (3.2-3.4) in SL; snout short, 4.0 (3.7-4.0) in head length; orbit diameter 3.8 (3.9-4.7) in head length; interorbital width 3.8 (3.7-4.0) in head length; least depth of caudal peduncle 1.9 (1.9-2.1) in head length; caudal peduncle length 1.5 (1.6-1.7) in head len Mouth small, oblique, maxilla not reaching vertical at front edge of orbit; three pairs of curved canine teeth anteriorly in upper jaw, progressively more laterally projecting, the third (posteriormost) pair largest; single pair of canine teeth anteriorly in lower jaw, very strongly curved laterally; no canine tooth at corner of mouth; side of jaws with single row of small close-set conical teeth; no teeth on palate; fleshy flap on side of lower lip; gill rakers short, longest about one-third length of longest gill filaments on first gill arch; posterior nostril an oval opening about twice size of cephalic sensory pores, anterior to fleshy upper edge of orbit and on vertical at anterior bony edge of orbit; anterior nostril small with a short fleshy rim anterior and slightly ventral to posterior; internarial space about 3.7-3.8 in orbit diameter; cephalic sensory system consisting of nine infraorbital pores, two supraocular pores, five occipital pores, single mid-interorbital pore, four small pores around nostril area, four mandibular pores, and five preopercular pores.
Head covered with scales except for interorbital space, snout, and chin; row of pointed scales on base of dorsal and anal fins; basal half of caudal fin with large scales; axillary scale of pelvic fin about equal to pelvic spine length; midventral scaly process of pelvic fins slightly longer than pelvic spine.
Free ventral margin of preopercle extending forward to vertical at centre of eye, vertical posterior margin dorsally to level of lower edge of pupil; exposed bony edge of preopercle smooth without serrations.
Origin of dorsal fin above second or third lateral line scale, predorsal length 3.3 (3.1-3.4) in SL; dorsal fin spines progressively longer, first 6.3 (6.9-7.8) in head length, and ninth 1.6 (1.9-2.2) in head length; longest (ninth) soft dorsal fin ray 3.6 (4.2-4.7) in SL; origin of anal fin below base of first soft dorsal fin ray, preanal length 1.9 (1.8) in SL; first anal fin spine 3.2 (3.3-4.3) in head length; second anal fin spine 2.9 (3.0-3.7) in head length; third anal fin spine 2.6 (2.6-3.3) in head length; longest (penultimate) soft anal fin ray 3.5 (3.8-4.1) in SL; caudal fin rounded, 1.3 (1.2-1.3) in head length; pectoral fin length 1.4 (1.3-1.4) in head length; pelvic fin length 1.6 (1.7-1.8) in head length.
Colour of live holotype (Fig. 1): head and body generally orange-red, grading to yellowish orange on breast and belly region; 10-12 narrow, mainly irregular, stripes on side, most reddish to pink, except ventralmost and those of caudal peduncle often electric blue; electric blue stripes on head including pair on side of snout, lowermost passing from posterior margin of eye, another immediately below on lower cheek, and pair radiating from posterior margin of eye; dorsal, anal, and pelvic fins burgundy red with purple rays and narrow electric blue outer or anterior margin, except orange at base of last 2-3 dorsal and anal rays, and prominent pair of broken, electric blue stripes along basal third of dorsal and anal fins; caudal fin burgundy red with pair of blue stripes on basal two-thirds and broad yellow outer margin; pectoral fins translucent.
Live colour of courting male (Fig. 2): similar to live colour of holotype described above except electric blue areas on side more extensive, involving zones both dorsally and ventrally, separated by a broad orange red area that tapers in width posteriorly.
Colour of holotype when fresh (Fig. 3): body generally brownish red, grading to red-orange on breast and belly region; 10-12 narrow, grey to reddish stripes on side, those on upper half irregular, sometimes broken, and less vivid compared to those of lower half (tend to wider and clearly reddish); head maroon with bluish stripes including pair on side of snout, lowermost passing from posterior margin of eye, and pair radiating from posterior margin of eye more centrally; dorsal, anal, and pelvic fins burgundy red with purple rays, except orange at base of last 2-3 dorsal and anal rays of holotype and two largest paratypes; caudal fin bluish on basal two-thirds and yellow on outer third; pectoral fins translucent.
Live colour of female: overall orange red with series of about six diffuse, narrow bluish grey stripes on side and translucent to reddish fins.
Colour after two months in alcohol (Fig. 4): overall pale greyish; scattered grey-brown spots/broken stripes on dorsal third of body; similar, but bolder stripes on head including pair on side of snout, one below eye, and pair radiating from posterior margin of eye; median and pelvic fins translucent with purple rays; pectoral fins translucent.
Colour of largest paratype after three years in alcohol: similar to specimens described above but paler and markings faded.
DNA Analysis: We resolved relationships between I? rennyae and other Paracheilinus species using a 655 base pair segment of the mtDNA COI gene from 14 Paracheilinus individuals. All Paracheilinus samples had different haplotypes except for the two P. filamentosus samples, which were identical. The two P. rennyae samples had an intraspecific genetic distance of 0.002, while the other species for which two samples were available showed intraspecific genetic distances ranging from 0.000 for P. cyaneus to 0.005 for P. flavianalis (whereby the two individuals were sampled from populations approximately 500 km apart). In the total Paracheilinus alignment there were 111 parsimony-informative characters. Nucleotide frequencies of P. rennyae were similar to the other species, and average frequencies for the combined Paracheilinus samples were as follows: A = 21.9, C = 26, G = 19.5, and T = 32.7. Pairwise distances between the 8 species of East Indian Paracheilinus ranged from 0.012-0.119, with the least distance between P. filamentosus and P. cyaneus, and the greatest distance recorded between P. rennyae and P. carpenteri (Table III).
Table III. Interspecific pairwise genetic distance matrix for mtDNA COI sequences among the 8 species included in this study. No. Species 1 2 3 4 5 6 7 1 P. walton 0.000 2 P. mccoskeri 0.098 3 P. cyaneus 0.035 0.105 4 P. 0.106 0.054 0.109 carpenteri 5 P. angulatus 0.084 0.105 0.090 0.115 6 P. 0.105 0.047 0.107 0.022 0.106 flavianalis 7 P. rennyae 0.089 0.107 0.095 0.119 0.013 0.108 8 P. 0.032 0.108 0.012 0.111 0.093 0.107 0.098 filamentosus No. Species 8 1 P. walton 2 P. mccoskeri 3 P. cyaneus 4 P. carpenteri 5 P. angulatus 6 P. flavianalis 7 P. rennyae 8 P. 0.000 filamentosus
Our results indicate that P. rennyae and P. angulatus are sister species, forming a clade with 99% bootstrap support using maximum likelihood analysis and 100% posterior probability using the Bayesian method (Fig. 5). The pairwise genetic distance between P. rennyae and P. angulatus is 0.013 (Table III), and both maximum likelihood (ML) and Bayesian trees clearly separate P. rennyae from P. angulatus. The phylogenetic analyses produced identical trees for most of the taxa, but could not resolve the relative positioning of the P. rennyae and P. angulatus clade with the clade consisting of P. filamentosus, P. cyaneus, and P. walton (with low bootstrap support of <50%). Maximum likelihood and Bayesian analyses resolved P. rennyae and P. angulatus as distinct lineages and together they were reciprocally monophyletic relative to the other Paracheilinus species (Fig. 5).
Distribution and habitat: The new species is currently known only from the type locality at southwestern Flores Island and has also been reported from nearby (about 18 km westward) southern Rinca and Nusa Kode islands in Komodo National Park on the basis of underwater photographs. Habitat at the type locality consisted primarily of a gently sloping rubble bottom with low coral outcrops situated along the base of a relatively steep slope at a depth of 15-20 m. The three collected male specimens were observed in company with approximately 30-40 much smaller females; no other males were observed in this group. A fourth male was observed and photographed at a nearby location, but was not collected. All males, except the smallest paratype, periodically exhibited courtship display, "flashing" bright electric blue colours a short distance above the bottom during the middle of the day.
Etymology: It is a pleasure to name this beautiful species rennyae in honour of Renny K. Hadiaty, Curator of Fishes at the Indonesian Institute of Sciences' Museum Zoologicum Bogoriense in Cibinong, for her valuable contributions to our knowledge of Indonesian freshwater and marine fishes and in recognition of many years of productive ichthyological collaborations with the authors.
Remarks: As mentioned in the introduction, the species of Paracheilinus are distinguished primarily on the basis of adult male characteristics including colour pattern (particularly during courtship) and shape of the median fins, especially the caudal fin (rounded, truncate, emarginated, lanceolate, or lunate), and number of elongated soft dorsal rays. The new species differs from all other members of the genus, except P. octotaenia of the Red Sea, in having rounded dorsal, anal, and caudal fins, and lacking prolonged dorsal fin rays. The two species differ in total number of gill rakers on the first branchial arch (13-14 in P. rennyae vs. 16-18 in P. octotaenia) and colour patterns of adult males. The pattern of P. octotaenia (Fig. 6) consists of a yellowish head and anterior body, grading to lavender bluish posteriorly with eight narrow bluish stripes on the side of the body, bright red median fins, and yellowish to orange pelvic fins. The primary differences of adult male P. rennyae include an overall red or reddish orange head and body, poorly defined stripes on the side, burgundy red versus bright red median fins, an orange area on the posterior dorsal and anal fins, burgundy red pelvic fins, broad yellow caudal-fin margin, and more prominent electric blue markings along the bases of the dorsal, anal, and caudal fins. Paracheilinus piscilineatus Comic, 1987 from Mauritius also lacks filamentous extensions of the dorsal fin rays, but has a truncate to slightly emarginate caudal fin and very different colouration (Fig. 7).
As noted above in Fig. 5 and Table III, our preliminary genetic analysis of the majority of Paracheilinus species found in the East Indian region reveals a close relationship between P. rennyae and P. angulatus Randall & Lubbock, 1981 originally described from Luzon, Philippines, but also occurring at Brunei, Malaysia (Sabah), and northeastern Indonesia (northeastern Kalimantan) (Allen & Erdmann, 2012). The pairwise genetic distance between these two taxa is only 0.013, at the lower end of distance values for COI generally considered to represent inter-specific differences in reef fishes (Steinke et al, 2009). We note however that these small genetic distances appear to be common within the genus Paracheilinus; Table III shows pairwise distances between P. ganeus and P. filamentosus of only 0.012 and between P. carpenteri and P. flavianalis of only 0.022. Preliminary data from a review of the genus Paracheilinus (including a more comprehensive genetic analysis) we are currently conducting suggests that divergences between many of these flasherwrasse species are relatively recent and have been driven by strong sexual selection (unpublished data). Such rapid and recent diversification under sexual selection could explain the overall low pairwise genetic differences seen in our analysis.
Genetic similarity notwithstanding, P. rennyae and P. angulatus are easily differentiated on the basis of median fin shape. In contrast to the evenly rounded dorsal and anal fin profile of P. rennyae, P. angulatus differs from all other adult male members of the genus in having a sharply angular profile posteriorly (Fig. 8). The caudal fin shape is also markedly different with P. rennyae having a rounded caudal fin compared to the emarginate to deeply lunate shape of P. angulatus.
We note that we deliberately avoided the collection of initial phase individuals in the type specimen series, primarily because of their similarity to small individuals of the other two sympatric species in the southwest Flores region, P. filamentosus (Fig. 9) and P. flavianalis Kuiter & Allen, 1999. Moreover, these three species readily form hybrids (e.g., Fig. 10), which appear to be locally abundant. It is easy to understand why this phenomenon is common given that all three species spawn daily in mixed aggregations and there appears to be fierce competition between males for spawning partners. Widespread hybridization of flasherwrasses was discussed by Kuiter and Allen (1999) and is a further indication of the relatively recent diversification within this genus.
We thank Zafer Kizilkaya for first calling our attention to this species with his photos from Rinca Island, and the captain and crew of the NusaTara liveaboard for their excellent dive support in collecting the type specimens in southwest Flores. Rudie Kuiter generously provided the photograph of P. octotaenia, and Ngurah Mahardika and his staff at the Indonesian Biodiversity Research Centre at Udayana University provided excellent support for the genetic analysis. As always, we thank Renny Hadiaty and her colleagues at the Indonesian Institute of Sciences (LIPI) for curation of the type specimens, and we thank the Paine Family Trust for their generous support of the field work involved in this study. Financial support for the genetic analysis was provided by the United States Agency for International Development's "Supporting Universities to Partner across the Pacific" program (Cooperative Agreement No. 497-A-00-10-00008-00). Finally, we thank Dita Cahyani, Paul Barber and two anonymous reviewers for their review and improvement of the manuscript.
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Gerald R. Allen (1), Mark V. Erdmann (2), (3) and Ni Luh Astria Yusmalinda (4)
1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986
2) Conservation International Indonesia Marine Program, J1. Dr. Muwardi No. 17, Renon, Denpasar 80235 Indonesia
3) California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 USA
4) Indonesian Biodiversity Research Centre, Faculty of Veterinary Medicine, Udayana University, Bali 80226 Indonesia
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|Author:||Allen, Gerald R.; Erdmann, Mark V.; Yusmalinda, Ni Luh Astria|
|Publication:||aqua: International Journal of Ichthyology|
|Date:||Oct 25, 2013|
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