Congridae: Heterocongrinae) from West Papua, Indonesia

aqua International Journal of Ichthyology

Vol. 15 (3), 20 July 2009

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Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia

Gerald R. Allen 1 and Mark V. Erdmann 2

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected] 2) Conservation International Indonesia Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235 Indonesia

Received: 27 April 2009 – Accepted: 03 July 2009

Abstract präorbitalen und suborbitalen Randes, wodurch diese Two new species in the pomacentrid genus Chromis are Bereiche dunkler wirken. Weitere Unterscheidungsmerk - described from Indonesian seas. Chromis albicauda is male sind ein schwarzer Fleck über der Analöffnung, die described from nine specimens, 79.6-133.7 mm SL, col - größere Körperlänge und die längeren Rückenstacheln. lected at Nusa Penida, off the east coast of Bali. It closely Die zweite Art, Chromis unipa, wird anhand von fünf resembles C. analis , which ranges widely in the western Exemplaren mit 42,8-50,4 mm SL beschrieben, die in Pacific. Both species have a similar yellow colouration, but 45-60 m Tiefe in der Cenderawasih-Bucht, West-Papua, C. albicauda has an abruptly white caudal fin compared to Indonesien, gefangen wurden. Zu den Bestimmungsmerk - the yellow caudal of C. analis. It also differs in having a malen zählen: XIV,11 Rückenflossenstrahlen; II,11 After - more densely scaled preorbital and a suborbital margin that flossenstrahlen, 18 Brustflossenstrahlen; drei stachelför - is obscured by scales. Additional differences include a black mige Schwanzflossenstrahlen; 14-17 röhrenförmige blotch covering the anal opening, a larger size, and taller Schuppen auf der Seitenlinie; eine Körpertiefe von 2,0-2,2 dorsal spines. The other new species, Chromis unipa , is mm SL; sowie ein gut unterscheidbares Farbmuster mit described from five specimens, 42.8-50.4 mm SL collected abwechselnd blaugrauen und messingartig gelben Streifen in 45-60 m in Cenderawasih Bay, Western Papua, Indone - an der Rumpfseite sowie blaugrauer Tönung bauchwärts sia. Diagnostic features include XIV,11 dorsal rays; II,11 an Kopf und Rumpf sowie bläulichen Flossen, abgesehen anal rays; 18 pectoral rays; 3 spiniform caudal rays; 14-17 von einem braunen basalen Anteil der Schwanzflosse und tubed lateral-line scales; body depth 2.0-2.2 in SL, and a einem deutlichen schwarzen Fleck auf der Afterflosse. C. distinctive colouration of alternating blue-grey and brassy unipa zählt zu einer Gruppe tief lebender Chromis- Arten yellow stripes on the side of the body, blue-grey ventrally on mit 14 Rückenstacheln und ähnelt am meisten C. degruyi, the head and body, bluish fins except for brown basal por - der aus 85 bis 120 m Tiefe von den Carolineninseln tion of the caudal fin, and a prominent black spot on the bekannt ist. Obwohl beide Arten eine ähnliche Zahl von anal fin. It belongs to a complex of deep-dwelling Chromis Brustflossenstrahlen, röhrenförmigen Seitenlinien-Schup - possessing 14 dorsal spines and it appears most similar to C. pen und Kiemenblättchen aufweisen, unterscheiden sie degruyi , known from 85-120 m depths in the Caroline sich im Hinblick auf die Modalzahlen der weichen Rü- Islands. Although both species have similar counts for pec - ckenflossen- und Analflossenstrahlen, die Körpertiefe und toral-fin rays, tubed lateral-line scales, and gill rakers, they die Schnauzenlänge. Außerdem sind die Farbmuster deut - differ with respect to modal numbers of soft dorsal and lich verschieden. anal-fin rays as well as body depth and snout length. They also exhibit significant colour pattern differences. Résumé Sont décrites deux nouvelles espèces du genre pomacentridé Zusammenfassung Chromi s des mers indonésiennes. Chromis albicauda est Zwei neue Riffbarscharten der Gattung Chromis aus dem décrit sur base de neuf spécimens, 79,6-133,7 mm de LS, Meer um Indonesien werden beschrieben. Chromis albi - collectés à Nusa Penida, au large de la côte est de Bali. cauda wird auf der Grundlage von neun Exemplaren mit L’espèce ressemble de près à C. analis, largement répandu 79,6-133,7 mm SL gekennzeichnet, die bei Nusa Penida dans le Pacifique occidental. Les deux espèces ont une colo- an der Ostküste Balis gefangen wurden. Die neue Art ration jaune similaire, mais C. albicauda possède une caudale ähnelt C. analis, der im Westpazifik weit verbreitet ist. abruptement blanche différente de la caudale jaune de C. Beide Arten zeigen eine ähnliche gelbe Farbgebung, doch analis. Elle se distingue aussi par un préorbital plus densé - unterscheidet sich C. albicauda durch seine deutlich abge - ment couvert d’écailles et par une marge suborbitale assom - setzte weiße Schwanzflosse, während bei C. analis auch die brie par des écailles. D’autres différences concernent une Schwanzflosse gelb gefärbt ist. Die neue Art unterscheidet tache noire couvrant l’anus, une taille plus grande et de plus sich außerdem durch eine dichtere Beschuppung des grandes épines dorsales. L’autre espèce nouvelle, Chromis

121 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia unipa, est décrite sur base de cinq spécimens, 42,8-50,4 mm INTRODUCTION de LS, collectés à 45-60 m dans la Cenderawasih Bay, The pomacentrid genus Chromis Cuvier, 1814 is Papouasie occidentale, Indonésie. Les données diagnostiques abundant throughout tropical and warm temperate comprennent XIV,11 rayons dorsaux; II,11 rayons à l’anale; seas (Allen 1991). The group is particularly con - 18 rayons pectoraux; 3 rayons spiniformes à la caudale; 14-17 écailles canaliculées sur la ligne latérale; hauteur du spicuous on coral reefs of the Indo-west and central corps 2,0-2,2 de LS, et une coloration voyante de lignes Pacific region, typically found in large mid-water alternées bleu-gris et jaune cuivre sur les flancs, bleu-gris ven - feeding aggregations. Similar to most pomacen - tralement sur la tête et le corps, des nageoires bleutées à l’ex - trids, the majority of Chromis are restricted to ception d’une portion basale brune à la caudale et une tache depths of less than 30 m, with maximum species noire voyante sur l’anale. Cette espèce appartient à un groupe diversity occurring in considerably shallower de Chromis vivant en profondeur, ayant 14 épines dorsales, et depths (in about 5-18 m). Sun-drenched shallow elle semble la plus proche de C. degruyi qui vit à une pro - reefs provide optimum conditions for benthic algal fondeur de 85 à 120 m dans les îles Caroline. Bien que les deux espèces aient un nombre similaire de rayons pectoraux, growth as well as supporting a diverse community d’écailles canalicues sur la ligne latérale et de branchiospines, of small invertebrates, both crucial components in elles se distinguent par le nombre de rayons dorsaux mous et the diet of many pomacentrids. The general lack of de rayons à l’anale ainsi que par la hauteur du corps et la filamentous algae below 40-50 m depth in particu - longueur du rostre. Elles présentent aussi des différences si- lar seems to be a limiting factor in the depth dis - gnificatives sur le plan de la coloration. tribution of most damselfish species, although the structural matrix of the reef that provides shelter Sommario and nest sites, also decreases rapidly with increasing Due nuove specie di pomacentridi del genere Chromis depth. Chromis are primarily zooplankton feeders, sono descritte dai mari dell’Indonesia. Chromis albicauda è descritta sulla base di nove esemplari di 79.6-133.7 mm a trait that has allowed penetration well beyond SL, raccolti a Nusa Penida, al largo della costa orientale di depths frequented by most pomacentrids. Deep Bali. È molto simile a C. analis, ampiamente diffusa nel trawling, submarine observations (Colin, personal Pacifico occidentale. Entrambe le specie hanno una colo- communication), the use of mixed-gas SCUBA for razione simile gialla ma C. albicauda ha una ben marcata deep diving (Pyle, personal communication), as pinna caudale bianca mentre quella di C. analis è gialla. well as standard SCUBA diving by the authors to Differisce anche per avere un preorbitale più densamente 50-70 m depths reveal that some Chromis are ricoperto di scaglie e il margine suborbitale nascosto dalle restricted to depths below 50 m and may routinely scaglie. Altre differenze includono una chiazza nera che ri- copre l’apertura anale, una taglia maggiore e spine dorsali penetrate to at least 175 m. Consequently a large più lunghe. L’altra nuova specie, Chromis unipa, è descritta proportion of recent pomacentrid discoveries (e.g. sulla base di cinque esemplari di 42.8-50.4 mm SL raccolti Pyle et al. 2008, Allen & Erdmann 2008), includ - a 45-60 m presso la baia Cenderawasih, Papua Occiden - ing one of the new species described herein, tale, Indonesia. Caratteristiche diagnostiche includono involve deep-dwelling Chromis , and many more XIV,11 raggi dorsali; II,11 raggi anali; 18 raggi pettorali; 3 can be expected as deep reef exploration continues raggi caudali spiniformi; 14-17 scaglie con tubicino lungo to expand. la linea laterale; altezza del corpo 2.0-2.2 in SL, e una di- The genus Chromis , which is the largest in the stinta colorazione di strie blu-grigio e giallo ottone alter - nantesi sul lato del corpo, una colorazione blu-grigio ven - family, now contains 94 species (including the new tralmente sul capo e sul corpo, pinne bluastre tranne per la taxa described herein). Allen (1991) provided pho - porzione basale marrone della caudale, e una cospicua tos and a brief diagnosis for 75 of the 76 described macchia nera sulla pinna anale. Appartiene al complesso di species ( C. dispilus Griffin, 1923 from New specie di Chromis di profondità che possiedono 14 spine Zealand unintentionally omitted). An additional dorsali e sembra più vicina a C. degruyi, diffusa a profon - 17 species were described in recent years (Randall dità di 85-120 m intorno alle Isole Caroline. Sebbene & McCosker 1992, Randall 1994, Moura 1995, entrambe le specie abbiano un numero simile di raggi pet - Randall 2001, Allen & Randall 2004, Lecchini & torali, di scaglie con tubicino in linea laterale e di rastrelli branchiali, esse differiscono per il valore modale dei raggi Williams 2004; Senou & Kudo 2007, Pyle et al. dorsali molli e di quelli anali come pure per l’altezza del 2008, Allen & Erdmann 2008). In addition, Allen corpo e la lunghezza del muso. Inoltre, esse mostrano si- (1993) showed that C. megalopsis Allen, 1976 from gnificative differenze nella colorazione. Western Australia is a junior synonym of C. mira - tionis Tanaka, 1917 of southern Japan. The present paper describes two new species that were collected with spear and rotenone during aqua vol. 15 no. 3 - 20 July 2009 122 Gerald R. Allen and Mark V. Erdmann recent surveys at Nusa Penida (near Bali) and Cen - measured from the anterior end of the upper lip to derawasih Bay, West Papua during November the anterior edge of the eye; orbit diameter is the 2009. The Nusa Penida fish was first mistaken for horizontal fleshy diameter, and interorbital width Chromis analis (Cuvier, 1830), a similar co-occur - the least fleshy width; upper jaw length is taken ring species, but we noted it differed by having a from the front of the upper lip to the posterior end larger size and different caudal-fin colouration. It of the maxilla; caudal peduncle depth is the least was found in areas of cool upwelling at Nusa depth, and caudal peduncle length is the horizon - Penida and based on photographic evidence is also tal distance between verticals at the rear base of the present at Japan, providing additional evidence for anal fin and the caudal fin base; lengths of fin spines the occurrence of warm-temperate Japanese species and rays are measured to their extreme bases (i.e. not on the reefs exposed to frequent cold-water from the point where the ray or spine emerges from upwelling in the deep straits that surround Bali. the basal scaly sheath); caudal fin length is the hori - The discovery at Cenderawasih Bay was made in zontal length from the posterior edge of the hypural connection with our on-going studies of the Bird’s plate to a vertical at the tip of the longest ray; caudal Head Region of West Papua. This large (approxi - concavity is the horizontal distance between verticals mately 59,000 km 2) bay situated on New Guinea’s at the tips of the shortest and longest rays; pectoral north coast separates the Bird’s Head or Vogelkop fin length is the length of the longest ray; pelvic fin Peninsula from the main body of the island. length is measured from the base of the pelvic spine Results of two (2006 and 2008) field trips co-spon - to the filamentous tip of the longest soft ray; pec - sored by the Cenderawasih Bay National Park toral ray counts include the small splint-like, upper - Authority, the State University of Papua and Con - most rudimentary ray; only the tube-bearing ante - servation International revealed a rich reef commu - rior lateral-line scales are counted; a separate count nity with several unique peculiarities indicative of is given for the deeply pitted scales occurring in a isolation from neighbouring areas in the geological continuous series midlaterally on the caudal pedun - past. The most obvious of these included at least cle; the decimal figure “0.5” appearing in the scale seven new fishes and 18 new coral species that row count above and below the lateral line refers to appear to be endemic to the bay, additional fishes a small truncated scale at the respective bases of the that occur widely in neighbouring regions, but dorsal and anal fins; gill raker counts include all exhibit unusual colour variation in the bay, and rudiments and are presented as separate counts for several deep-reef fishes that occur in unusually the upper and lower limbs as well as a combined shallow water within the bay. Moreover, a genetic count; the last fin ray element of the dorsal and anal study of connectivity patterns among marine fins is usually branched near the base and is counted organisms by Paul Barber of the University of Cal - as a single ray. ifornia at Los Angeles and Hamid Toha of the State Counts and proportions appearing in parentheses University of Papua (Barber, personal communica - apply to the range for the paratypes if different tion) has noted the presence of distinctive clades in from the holotype. Proportional measurements Cenderawasih Bay for several molluscs and crus - expressed as percentage of the standard length are taceans. Based on this evidence, we hypothesize provided in Tables I-II. Type specimens are that the inner portion of the bay was essentially deposited at Pusat Penelitian dan Pengembangan isolated for a substantial period over the past five Oseanologi, Jakarta, Indonesia (NCIP), United million years (Allen & Erdmann 2008). States National Museum of Natural History, Wash - ington, D.C. (USNM), and the Western Aus - MATERIALS AND METHODS tralian Museum, Perth (WAM). Lengths of specimens are given as standard length (SL) measured from the anterior end of the upper lip to the base of the caudal fin (posterior edge of Chromis albicauda n. sp. hypural plate); head length is measured from the (Figs 1-2, 4-5; Table I) same anterior point to the posterior edge of the opercle flap; body depth is the maximum depth Holotype: NCIP 6350, 133.7 mm SL, Crystal taken vertically between the belly and base of the Bay, 8 °42.887’S 115 °27.331’E, Nusa Penida, dorsal spines; body width is the maximum width Indonesia; 30 m depth, spear, M. Erdmann, 28 just posterior to the gill opening; snout length is November 2008.

123 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia

Paratypes (collected with holotype) : NCIP 6351, line to origin of dorsal fin 2.5; scales above lateral 131.8 mm SL; USNM 391796 , 2 specimens, 81.5- line to base of middle dorsal spine 2.5; scales below 120.5 mm SL; WAM P.33101-005, 5 specimens, lateral line to origin of anal fin 8.5; gill rakers 7-8 79.6-127.0 mm SL. + 16-19, total rakers 23-26; branchiostegal rays 6; Diagnosis: Dorsal rays XIII,12 (rarely 11); anal supraneural (predorsal) bones 3; vertebrae 11 + 15. rays II,12 (rarely 11); pectoral rays 18-20; spini - Body moderately deep, depth 1.9 (1.7-1.9) in SL, form caudal rays 3; tubed lateral-line scales 17-19; and compressed, the width 2.5 (2.5-2.9) in body gill rakers 7-8 + 24-26, total rakers 24-26; body depth; head length 3.1 (3.1-3.3) in SL; dorsal pro - depth 1.8-1.9 in SL; edge of suborbital hidden by file of head evenly rounded; snout shorter than scales; 4-5 scale rows on anterior portion of preo - orbit diameter, its length 3.7 (3.4-4.5) in head bital; generally yellow in life, grading to brown on length; orbit diameter 3.3 (2.8-3.2) in head; upper back with abruptly white caudal fin; black - interorbital space convex, its width 3.3 (2.8-3.3) in ish anal opening (also evident in preservative). head; caudal-peduncle depth 2.1 (1.9-2.1) in head; Description: Dorsal rays XIII,12 (one paratype caudal-peduncle length 3.0 (2.5-3.1) in head. with XIII,11); anal rays II,12 (one paratype with Mouth terminal, small, and oblique, forming an II,11); all dorsal and anal soft rays branched angle of about 30°-32° to horizontal axis of head (except first dorsal ray unbranched in one and body; posterior edge of maxilla reaching a ver - paratype), the last dorsal and anal rays branched to tical slightly behind anterior edge of pupil, the base; pectoral rays 19 (18-20), the upper and low - upper-jaw length 3.2 (2.8-3.3) in head; teeth mul - ermost pair unbranched; pelvic rays I,5; principal tiserial, an outer row of conical teeth in each jaw, caudal rays 15, the upper and lowermost largest anteriorly; 22 (17-23) upper and 20 (17-20) unbranched; spiniform caudal rays 3, followed by lower teeth on each side of jaw; a narrow band of 2 accessory segmented rays; scales in longitudinal villiform teeth lingual to outer row, in 2-3 irregu - series 28; tubed lateral-line scales 19/18 (17-19); lar rows anteriorly, narrowing to a single row on posterior midlateral scales with a pore or deep pit side of jaws; tongue triangular with rounded tip; (in continuous series) 8 (6-8); scales above lateral gill rakers long and slender, the longest on lower

Fig. 1. Chromis albicauda , holotype, 133.7 mm SL, Nusa Penida, Indonesia. Photo by G. R. Allen. aqua vol. 15 no. 3 - 20 July 2009 124 Gerald R. Allen and Mark V. Erdmann

Table I. Proportional measurements of selected type specimens of Chromis albicauda as percentage of the standard length.

Holotype Paratype Paratype Paratype Paratype Paratype NCIP NCIP WAM USNM USNM USNM 6350 6351 P.33101 391796 391796 P.33101

Standard length (mm) 133.7 131.8 127.0 120.5 81.5 79.6 Body depth 53.3 55.1 54.0 55.5 55.2 51.8 Body width 21.7 21.5 21.5 19.4 19.4 18.7 Head length 32.2 32.0 32.4 30.6 32.0 31.0 Snout length 8.6 8.4 9.0 9.1 7.2 6.9 Orbit diameter 9.7 10.0 10.3 9.6 11.3 11.1 Interorbital width 9.7 10.4 10.9 10.8 9.8 9.4 Depth of caudal peduncle 15.4 15.6 17.2 15.8 16.6 16.3 Length of caudal peduncle 10.8 10.6 10.3 11.2 12.0 12.2 Upper jaw length 10.2 10.1 10.4 11.0 9.8 10.6 Predorsal distance 43.4 41.3 42.8 43.3 42.8 42.0 Preanal distance 72.6 72.4 71.1 69.4 71.0 74.4 Prepelvic distance 44.6 44.7 43.1 42.1 42.0 47.1 Length of dorsal fin base 62.7 62.0 63.4 63.7 62.3 58.5 Length of anal fin base 23.6 24.7 25.6 23.8 24.9 24.6 Pectoral fin length 36.2 34.6 34.4 34.5 35.7 33.4 Pelvc fin length 32.8 30.3 29.1 32.3 34.6 33.3 Pelvic fin spine length 14.9 15.0 17.4 16.9 18.5 17.8 1st dorsal spine 7.1 7.1 6.9 7.2 8.5 8.4 2nd dorsal spine 12.7 12.1 11.5 11.4 12.6 13.4 7th dorsal spine 18.1 18.4 18.6 18.2 18.8 18.7 Last dorsal spine 14.8 12.9 14.3 13.4 15.0 14.1 Longest soft dorsal ray 20.8 20.9 20.9 22.0 22.7 21.4 1st anal spine 8.3 6.4 7.2 8.7 9.0 7.8 2nd anal spine 20.9 18.7 19.1 18.9 21.7 19.5 Longest soft anal ray 20.4 18.8 19.1 20.4 22.1 23.1 Caudal fin length 26.2 30.0 24.5 29.8 30.8 33.7 Caudal concavity 10.8 12.3 7.2 8.1 12.4 damaged

Fig. 2. Underwater photograph of Chromis albicauda , approximately 130 mm SL, 25 m depth, Nusa Penida Indonesia. Photo by G. R. Allen.

125 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia limb near angle about two-thirds to three-fourths one-fourth of pectoral fins; series of progressively length of longest gill filaments; nostril with a fleshy tapering scales forming median scaly process poste - rim, more elevated on posterior edge and located at riorly from between base of pelvic fins, its length level of middle of pupil, slightly closer to orbit than slightly more than one-third that of pelvic spine; edge of upper lip. axillary scale above base of pelvic spine about one- Opercle ending posteriorly in a flat spine, the tip half length of spine. relatively obtuse and obscured by a large scale; Origin of dorsal fin above third lateral-line scale, the margin of preopercle smooth, the posterior margin predorsal distance 2.3 (2.3-2.4) in SL; base of soft extending dorsally to level of upper edge of pupil, portion of dorsal fin contained 2.7 (2.6-2.7) times in the anterior margin extending to level of middle of base of spinous portion; first dorsal spine 4.5 (3.7- pupil; preorbital relatively broad, its width tapering 4.7) in head; second dorsal spine 2.5 (2.3-2.9) in posteriorly, with 4-5 irregular scale rows at widest head; fourth to ninth dorsal spines subequal, the point; suborbital covered with 1-2 rows of small longest 1.8 (1.6-1.7) in head; last dorsal spine 2.2 scales that obscure its lower margin. (2.0-2.5) in head; membranes of spinous portion of Scales finely ctenoid; anterior lateral line ending dorsal fin moderately incised; third to fifth dorsal soft beneath base of last dorsal spine; head scaled except rays longest, 1.5 (1.3-1.6) in head; first anal spine 3.9 lips, chin, tip of snout, and a narrow zone from (3.1-5.0) in head; second anal spine 1.5 (1.4-1.7) in orbit to edge of snout containing nostrils; a scaly head; first three anal soft rays subequal, the longest sheath at base of dorsal and anal fins, about one- 1.6 (1.3-1.7) in head; caudal fin forked, its length 3.8 half pupil diameter at base of middle of spinous (2.7-4.1) in SL, the caudal concavity 3.0 (2.6-4.5) in portion of dorsal fin, progressively narrower on soft head; fourth pectoral ray longest, 2.8 (2.7-3.0) in SL; portion; a column of scales on each membrane of pelvic spine 2.2 (1.6-2.1) in head; first soft ray of dorsal and anal fins, narrowing distally, those on pelvic fin filamentous, 3.1 (2.8-3.4) in SL. spinous portion of dorsal progressively longer, Colour of holotype in alcohol (Fig. 1): reaching about two-thirds distance to spine tips on Overall yellowish tan grading to brown on upper posterior membranes, then progressively shorter on third of head and body; scaly sheath on basal half soft portion; small scales on caudal fin extending of spinous dorsal fin brown, the brown colouration nearly to posterior margin; small scales on basal also extending to base of soft dorsal fin, which is

Fig. 3. Underwater photograph of Chromis analis , approximately 90 mm SL, 30 m depth, Nusa Penida, Indonesia. Photo by G. R. Allen. aqua vol. 15 no. 3 - 20 July 2009 126 Gerald R. Allen and Mark V. Erdmann otherwise pale tan; caudal fin pale tan; anal fin Colour in life (Fig. 2): Overall yellow except mainly pale tan, except light brown on basal por - caudal fin abruptly white; upper third of body tion; pectoral and pelvic fins yellowish tan, pec - grading to brown and scales on lower two-thirds of toral-fin base with small brown spot on upper side with narrow brown margins; dorsal fin brown edge; small blackish blotch covering anal opening. on basal half, yellow on outer portion, except dis -

Fig. 4. Chromis albicauda , paratype, 127.0 mm SL (upper) and C. analis , 89.0 mm SL, Watubela Islands, Maluku Province, Indonesia (WAM P.33085-006). Note size discrepancy of these adult specimens, black anal patch of C. albicauda , and more conspicuous dark marking on upper pectoral base of C. analis .

127 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia

Fig. 5. Comparison of preorbital/suborbital scale patterns of Chromis albicauda (upper) and C. analis . aqua vol. 15 no. 3 - 20 July 2009 128 Gerald R. Allen and Mark V. Erdmann tal half of soft dorsal fin translucent; anal and pec - two species (Fig. 4) with the largest known adults toral fins yellow; pectoral fins translucent yellow - reaching 133.7 mm SL in C. albicauda and about ish, the upper edge of its base brown, small black - 100 mm SL for C. analis , although the latter fish is ish blotch covering anal opening. Chromis albi - often reported reaching a larger size in the literature cauda was observed to instantaneously “switch” its due to confusion between the two species. Compar - white tail on and off underwater. When the normal isons are based on five specimens of C. analis , 82.0- white colour is temporarily turned off the tail is 92.4 mm SL, from the Watubela Islands, Maluku yellow and the fish is difficult to distinguish from Province, Indonesia (WAM P.33085-006). the similar C. analis (Fig. 3). Chromis albicauda is also similar to C . pembae Remarks: Chromis albicauda is very similar in Smith, 1960 from the western Indian Ocean, Red general appearance to C. analis, which co-occurs at Sea, and southern Arabian Peninsula. Both species Bali and is widely distributed in the Western have a similar shape and exhibit a white caudal fin, Pacific from the Great Barrier Reef of Australia to although the overall colour of C. pembae is mainly Japan, ranging east to Micronesia (except Marshall brown or yellowish brown rather than yellow. Islands) and Fiji. Both species have an overall yel - There are also differences in number of soft anal low colouration, brownish back and brown dorsal- rays (11 for C. pembae versus a usual count of 12) fin base (compare Figs 1 and 3). However, C. albi - and total gill rakers on the first gill arch (27-31 rak - cauda differs most noticeably in possessing an ers, usually 28-30, for C. pembae versus 24-26). abruptly white caudal fin, compared to that of C. Moreover the preorbital/suborbital scalation of C. analis , which is yellow (same as adjacent body pembae is similar to that of C. analis as illustrated colour) with a translucent margin. In preservative in Fig. 5. There is also a substantial size difference (Fig. 4) the two species are further distinguished by for the two species with C. pembae attaining a max - the blackish blotch that covers the anal opening of imum standard length of approximately 95.0 mm C. albicauda , a feature lacking in C. analis . Also, compared to at least 133. 3 mm for C. albicauda. the brown spot on the upper pectoral-fin base of C. Distribution and habitat: The new species was albicauda is small and inconspicuous, being mainly commonly observed in areas of cool (18°-24 °C) confined to the dorsal edge of the fin base com - upwelling along the northern and western shores of pared to the much larger, wedge-shaped spot of C. Nusa Penida at depths between about 25-50 m, analis that covers the uppermost portion of the fin although we observed a pair in only 10 m on one base. There is also an important difference related occasion. The species typically forms aggregations, to the pattern of scales on the preorbital and sub - which feed high in the water column on zooplank - orbital series (Fig. 5). The preorbital of C. albi - ton when currents are strong. In addition to its cauda is densely covered with 4-5 irregular rows of occurrence at Nusa Penida (a small island lying small scales that taper posteriorly to 1-2 rows on approximately 15 km east of Bali), C. albicauda the suborbital. In comparison, the preorbital of C. apparently also occurs in southern Japanese seas analis has 1-2 rows of larger scales on the preorbital based on photographic evidence (Masuda & and a single row on the suborbital. Furthermore, Kobayashi 1994, Kuiter & Debelius 2006). There - the lower margin of the suborbital is obscured by fore, it likely has a disjunct distribution associated scales in C. albicauda , but is exposed in C. analis . with cool temperature regimes. At least two other Morphometric differences include consistently species found at Nusa Penida also have Japanese taller dorsal spines in C. albicauda . For example the distributions including Apogon schlegeli Bleeker, length of the first spine ranges from 3.7 to 4.7 in 1854, and labrid Bodianus masudai Araga & the HL for C. albicauda compared to 5.1-6.0 in C. Yoshino, 1975. Nusa Penida is characterized by analis and that of the longest spine ranges from cold upwelling and populations of typically cool 1.6-1.8 (usually 1.6-1.7) compared to 1.8-2.0. water species are therefore able to thrive. The pres - Meristic differences include lateral line scales (17- ence of the acanthurid Prionurus chrysurus Randall, 19 for C. albicauda versus 15-16 for C. analis) and 2001 at Nusa Penida (also Bali, Lombok and a tendency towards lower gill raker counts in C. Komodo) provides an additional example of this albicaudata (average total rakers on the first phenomenon. The genus Prionurus Lacépède, branchial arch 25.0 versus 27.0, and rakers on 1804 typically inhabits lower latitudes with repre - lower limb 17.5 versus 19.8). sentatives known from Japan, temperate Australia, Finally, there is a significant size difference in the and the Eastern Pacific Ocean.

129 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia

Etymology: This species is named albicauda segmented rays; scales in longitudinal series 27; (Latin: “white tail”) with reference to the diagnostic tubed lateral-line scales 15 (13-15); posterior mid - feature that separates it from the similar C. analis . lateral scales with a pore or deep pit (in continuous series) 7/6 (7); scales above lateral line to origin of Chromis unipa n. sp. dorsal fin 2.5; scales above lateral line to base of (Figs 6-7; Table II) middle dorsal spine 1.5; scales below lateral line to origin of anal fin 8.5; gill rakers 8 + 19 = 27 (7-8 + Holotype: NCIP 6349, 50.4 mm SL, Tridacna 19-20 = 27-28; branchiostegal rays 6; supraneural Reef, 2 °29.672’S 134 °58.990’E, eastern side of (predorsal) bones 3; vertebrae 11 + 15. Cenderawasih Bay, Papua Barat Province, Indone - Body relatively elongate, depth 2.2 (2.0-2.2) in sia; 45-60 m depth, rotenone, M. Erdmann and G. SL, and compressed, the width 2.7 (2.6-2.8) in Allen, 13 November 2008. body depth; head length 3.1 (3.0-3.2) in SL; dor - Paratypes (collected with holotype) : WAM sal profile of head with a slight convexity above P.33043-002, 4 specimens, 43.2-47.5 mm SL. eye; snout shorter than orbit diameter, its length Diagnosis: Dorsal rays XIV,11; anal rays II,11; 5.7 (4.7-6.0) in head length; orbit diameter 2.5 pectoral rays 18; spiniform caudal rays 3; tubed lat - (2.4-2.7) in head; interorbital space convex, its eral-line scales 14-17; gill rakers 7-8 + 19-20 = 27- width 3.7 (3.7-4.2) in head; caudal-peduncle 28; body depth 2.0-2.2 in SL; alternating blue grey depth 2.4 (2.2-2.4) in head; caudal-peduncle and brassy yellow stripes on side of body; abdomen length 2.7 (2.3-2.9) in head. and lower part of head blue grey; fins bluish except Mouth terminal, small, and oblique, forming an base of caudal fin brown; a prominent black spot at angle of about 44° to horizontal axis of head and apex of anal fin. body; posterior edge of maxilla reaching a vertical Description: Dorsal rays XIV,11; anal rays II,11; slightly behind anterior edge of pupil, the upper-jaw all dorsal and anal soft rays branched except first length 3.5 (3.2-3.4) in head; teeth multiserial, an dorsal ray, the last dorsal and anal rays branched to outer row of conical teeth in each jaw, largest anteri - base; pectoral rays 18, the upper- and lowermost orly; 22 upper and 20 lower teeth on each side of pair unbranched; pelvic rays I,5; principal caudal jaw; a narrow band of villiform teeth lingual to outer rays 15, the upper and lowermost unbranched; row, in 2-3 irregular rows anteriorly, narrowing to a spiniform caudal rays 3, followed by 2 accessory single row on side of jaws; tongue triangular with

Fig. 6. Chromis unipa , holotype, 50.4 mm SL, Cenderawasih Bay, West Papua, Indonesia. Photo by G. R. Allen. aqua vol. 15 no. 3 - 20 July 2009 130 Gerald R. Allen and Mark V. Erdmann

Table II. Proportional measurements of type specimens of Chromis unipa as percentage of the standard length.

Holotype Paratype Paratype Paratype Paratype NCIP WAM WAM WAM WAM 6349 P.33043 P.33043 P.33043 P.33043 Standard length (mm) 50.4 47.5 44.1 43.2 42.8 Body depth 46.0 49.3 47.6 46.8 46.5 Body width 16.9 18.9 17.0 18.1 17.3 Head length 32.7 32.8 31.7 32.4 32.2 Snout length 5.8 6.9 6.6 6.7 5.4 Orbit diameter 13.3 12.2 13.4 13.0 12.4 Interorbital width 8.9 8.8 8.2 7.6 7.7 Depth of caudal peduncle 13.5 13.7 14.5 14.1 14.3 Length of caudal peduncle 12.3 11.4 11.3 12.0 13.8 Upper jaw length 9.3 9.7 10.0 9.7 9.6 Predorsal distance 42.7 41.5 40.8 40.0 42.1 Preanal distance 71.6 68.4 70.7 70.6 70.8 Prepelvic distance 41.3 38.7 41.3 42.1 41.1 Length of dorsal fin base 60.7 63.2 58.7 61.1 59.1 Length of anal fin base 21.4 21.9 22.0 22.2 19.9 Pectoral fin length 35.7 30.1 33.1 31.9 35.3 Pelvic fin length 34.5 26.3 32.9 32.4 32.7 Pelvic fin spine length 19.0 17.9 16.8 17.1 18.5 1st dorsal spine 6.0 5.7 5.7 6.3 6.5 2nd dorsal spine 10.3 9.5 10.7 10.9 11.9 7th dorsal spine 15.3 16.0 14.5 14.4 15.2 Last dorsal spine 10.1 11.6 11.1 11.6 11.2 Longest soft dorsal ray 17.1 18.9 17.9 19.2 17.3 1st anal spine 5.6 6.3 6.3 6.5 6.5 2nd anal spine 23.4 23.4 22.0 21.3 22.0 Longest soft anal ray 21.0 21.5 20.9 19.9 21.5 Caudal fin length 39.9 35.8 37.9 36.6 35.0 Caudal concavity 21.6 19.4 20.6 19.7 20.6 rounded tip; gill rakers long and slender, the longest a column of scales on each membrane of dorsal and on lower limb near angle about three-fourths length anal fins, narrowing distally, those on spinous por - of longest gill filaments; nostril with a fleshy rim, tion of dorsal progressively longer, reaching about more elevated on posterior edge and located at level two-thirds distance to spine tips on posterior mem - of middle of pupil, slightly less than one-half dis - branes, then progressively shorter on soft portion; tance from front of orbit to edge of upper lip. small scales on caudal fin extending slightly more Opercle ending posteriorly in a flat spine, the tip than two-thirds distance to posterior margin; small relatively obtuse and obscured by a large scale; scales on basal one-fifth of pectoral fins; a median margin of preopercle smooth, the posterior margin scaly process extending posteriorly from between extending dorsally to level of upper edge of pupil, base of pelvic fins, its length slightly more than one- the anterior margin extending to level of middle of third that of pelvic spine; axillary scale above base of pupil; narrow suborbital covered with single row of pelvic spine about one-half length of spine. small scales, with free lower margin extending Origin of dorsal fin above fourth lateral-line scale, nearly to a vertical at posterior edge of pupil. the predorsal distance 2.3 (2.4-2.5) in SL; base of Scales finely ctenoid; anterior lateral line ending soft portion of dorsal fin contained 3.6 (3.1-4.1) beneath base of twelfth dorsal spine; head scaled times in base of spinous portion; first dorsal spine except lips, tip of snout, and a narrow zone from 5.5 (5.9-5.8) in head; second dorsal spine 3.2 (2.7- orbit to edge of snout containing nostrils; a scaly 3.5) in head; fourth to ninth dorsal spines sube - sheath at base of dorsal and anal fins, about one-half qual, the longest 2.1 (2.1-2.3) in head; last dorsal pupil diameter at base of middle of spinous portion spine 3.2 (2.8-2.9) in head; membranes of spinous of dorsal fin, progressively narrower on soft portion; portion of dorsal fin moderately incised; third to

131 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia fifth dorsal soft rays longest, 1.9 (1.7-1.9) in head; dorsal fin grey-brown with bronzy yellowish basal first anal spine 5.9 (4.9-5.2) in head; second anal portion and narrow whitish margin on spinous spine 1.4 (1.4-1.5) in head; first three anal soft rays portion, posteriormost portion of fin translucent subequal, the longest 1.6 (1.5-1.6) in head; caudal with dusky grey rays; anal fin mainly bluish with fin forked, its length 2.5 (2.6-2.9) in SL, the cau - narrow white anterior margin, large black spot dis - dal concavity 1.5 (1.5-1.7) in head; fourth pectoral tally on middle portion, and translucent on poste - ray longest, 2.8 (2.8-3.3) in SL; pelvic spine 1.7 rior margin; caudal fin yellowish brown at base of (1.7-1.9) in head; first soft ray of pelvic fin fila - each lobe, the posterior margin broadly translucent mentous, 2.9 (3.0-3.8) in SL. with bluish grey rays; pelvic fins bluish white with Colour of holotype in alcohol (Fig. 6): white anterior margin; pectoral rays translucent Generally brown, paler where scales have been dis - with large black spot on upper half of fin base, lodged; median fins also brown except outer half of extending onto outer face of axil . soft dorsal fin and caudal fin translucent with Remarks: The new species is distinguished from dusky grey rays; pelvic fins light dusky brown with the majority of Chromis by having 14 dorsal-fin paler anterior margin; pectoral rays translucent spines instead of the usual counts of 12 or 13, with large black spot on upper half of fin base, rarely 15 (one species). The five species from the extending onto outer face of axil. central and eastern Atlantic generally have 14 Colour of holotype when freshly spines, but within the vast Indo-Pacific region only captured (Fig. 7): Head brownish dorsally and 14 currently valid species (including the new pale bluish below level of eye; about 6-7 blue spots taxon) possess 14 spines. This group, which on operculum; series of alternating blue and exhibits diverse morphological characters and is bronzy yellowish stripes on side, each blue stripe apparently polyphyletic, primarily inhabits either about half of pupil width and covering centre of subtropical areas such as Hawaii, and Japan or is each scale row; bronzy yellowish stripes slightly mainly confined to deep reefs in about 50-183 m wider and overlapping lower and upper portions of depth. Pyle et al. (2008) discussed the members of adjacent scale rows; belly and lower portion of side the deep-dwelling complex with 14 dorsal spines, (below level of lower pectoral-fin base) bluish grey; which includes C. abyssus Pyle, Earle & Greene,

Fig. 7. Chromis unipa , freshly collected (anesthetised) holotype, 50.4 mm SL, Cenderawasih Bay, West Papua, Indonesia. Photo by G. R. Allen. aqua vol. 15 no. 3 - 20 July 2009 132 Gerald R. Allen and Mark V. Erdmann

2008 (Palau), C. axillaris (Bennett, 1831) (East counts with C. degruyi having 12 (rarely 11) com - Africa and Mauritius), C. circumaurea Pyle, Earle pared to 11 rays in C. unipa . There are also differ - & Greene, 2008 (Yap, Caroline Islands), C. degruyi ences in body depth and snout length. The new Pyle, Earle & Greene, 2008 (Caroline Islands), C. species has a slightly more slender body (46.0- mirationis Tanaka, 1917 (Japan and Western Aus - 49.3% of SL, compared to 50.1-54.4%) and tralia), C. okamuri Yamakawa & Randall, 1989 shorter snout (5.4-6.9% of SL vs 7.4-8.7% of SL). (Okinawa Trough, China Sea), C. onumai Senou & The comparative data for the five type specimens Kudo, 2007 (southern Taiwan and Izu Islands) , C. of C. degruyi were taken from Pyle et al. (2008) planesi Lecchini & Williams, 2004 (Rapa Island, and although the three largest individuals (76.6- Polynesia) , C. struhsakeri Randall & Swerdloff, 81.0 mm SL) were considerably larger than our 1973 (Hawaiian Islands), and C. woodsi Bruner & specimens of C. unipa , the two smaller specimens Arnam, 1979 (East Africa) . The new species differs (38.3-38.7 mm SL) of C. degruyi exhibit a consis - from all of these in its distinctive colour pattern of tently deeper body (50.2-50.8% of SL) and longer alternating blue and bronzy yellowish stripes with snout (7.8-8.2% of SL). There are also significant a large black spot on the anal fin, and from most colour pattern differences, although both species species on the basis of its relatively low number possess a large black spot on the upper pectoral-fin (11) of soft dorsal and anal rays compared with the base. Chromis degruyi is basically dull brownish yel - usual range of 12-14 rays. Chromis woodsi possesses low with a thin, lavender-grey stripe on each scale 11-12 soft dorsal-fin rays, but differs in having a row, and lacks the prominent black spot on the lower gill-raker count (23-25 versus 27-28) and a anal fin that is characteristic of C. unipa . Further - very different colour pattern (overall whitish with more, individuals of C. degruyi under about broad, black bar on caudal peduncle). 50 mm SL have broad yellow dorsal and ventral Of the taxa mentioned above, C. unipa appears to margins on the caudal fin and yellow pigmentation be most similar to C. degruyi (Fig. 8). Both species on the posterior dorsal and anal fins. have 18 pectoral rays, relatively low lateral-line Distribution and habitat: The type locality and scale counts (14-17), and 27-28 gill rakers. They only known collection site of C. unipa consists of differ, however, in modal dorsal and anal ray rubble and low-profile coral reef substratum at the

Fig. 8. Chromis degruyi , holotype, 109.5 mm SL, Palau Islands. Photo by R. Pyle.

133 aqua vol. 15 no. 3 - 20 July 2009 Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia base of a steep, nearly vertical dropoff at 42 m to at of C. unipa . We are also extremely grateful to least 65 m depth. The Tridacna Reef location con - Martha Davis and the Gordon and Betty Moore sists of a submerged atoll lying approximately Foundation for their financial support of this 38.5 km directly east of the Wandamen Peninsula. capacity building course. We thank Suharsono and The species was common and seen in the company Rianta Afriadi of the Indonesian Institute of Sci - of C. brevirostris Pyle, Earle & Greene, 2008, a ences for their support and assistance with type species previously known from Bali, Caroline specimens, and the owners, captain and crew of the Islands (Palau to Puluwat), Marshall Islands, Van - MV Temu Kira for providing an excellent diving uatu, and Fiji. The sighting of this species at Cen - platform for the Cendrawasih field survey. Dan derawasih Bay is noteworthy as the depth range Polhemus provided valuable inputs on the isolation (42-65 m) was much shallower than the usual of Cendrawasih Bay from a geological perspective. range of 90-120 m reported by Pyle et al. (2008). This observation is consistent with several other REFERENCES normally deep-dwelling reef species that are found ALLEN G. R. 1991. Damselfishes of the World . Aquarium Sys - in relatively shallow water at Cenderawasih Bay. tems, Mentor, Ohio, 271 pp. ALLEN , G. R . 1993. Two new species of damselfishes (Poma - Therefore, it is possible that C. unipa may be centrus), with comments on the validity of two additional found much deeper at locations outside the bay. pomacentrid fishes. Revue française d’Aquariologie 20 (1): Etymology: This species is named unipa with refer - 21-26. ence to the acronym for Universitas Negeri Papua ALLEN , G. R . & E RDMANN , M. V. 2008. Pterocaesio monikae , (State University of Papua, Manokwari), who invited a new species of fusilier (Caesionidae) from western New the authors to conduct an ichthyology field course Guinea (Papua and Papua Barat Provinces, Indonesia. that led to the discovery of this species. The Univer - aqua, International Journal of Ichthyology 13 (3-4): 163-170. sity has played a critical role in training young Pap- ALLEN , G. R. & R ANDALL , J. E. 2004. Two new species of damselfishes (Pomacentridae: Chromis) from Indonesian uans in the natural sciences, and it is our sincere hope seas. aqua, Journal of Ichthyology and Aquatic Biology 9 (1): that this work will inspire students and lecturers at 17-24. the University to actively explore the unique natural KUITER , R. H. & D EBELIUS , H. 2006. World Atlas of Marine laboratory of Cenderawasih Bay at their doorstep. Fishes . Ikan-Unterwasserarchiv, Frankfurt, 720 pp. The name is treated as a noun in apposition. LECCHINI , D. & W ILLIAMS , J.T. 2004. Description of a new species of damselfish (Pomacentridae: Chromis) from Rapa ACKNOWLEDGEMENTS Island, French Polynesia. aqua, Journal of Ichthyology and The authors would like to thank the following Aquatic Biology 8 (3): 97-102. MASUDA , H. & K OBAYASHI , Y. 1994. Grand Atlas of fish life agencies for supporting the rapid biodiversity modes. Tokai Universiy Press, Tokyo, 465 pp. assessment of Nusa Penida which led to the dis - MOURA , R. L. DE . 1995. A new species of Chromis (Perci - covery of C. albicauda: the Ministry of Research formes: Pomacentridae) from the southeastern coast of and Technology, the Indonesian Institute of Sci - Brazil, with comments on other species of the genus. Revue ences, the Ministry of Fisheries and Marine Affairs, française d’Aquariologie 21 (3-4): 91-96. the Bali Department of Nature Conservation PYLE , R. L., E ARLE , J. L. & G REENE , B. D . 2008. Five new (BKSDA), the Klungkung regional government, species of the damselfish genus Chromis (Perciformes: and Conservation International. Michael Corten - Labroidei: Pomacentridae) from deep coral reefs in the bach and Sven Fautz at Bali Diving Academy pro - tropical western Pacific. Zootaxa . 1671: 3-31. RANDALL , J. E . 1994. Two new damselfishes (Perciformes: vided excellent logistical support for the survey as Pomacentridae) from Arabian waters. Revue française well. Iwan Dewantama and Ketut Sarjana Putra of d’Aquariologie 21 (1-2): 39-48. Conservation International are owed special thanks RANDALL , J. E. 2001. Four new damselfishes (Perciformes: for their efforts to organize and oversee the Nusa Pomacentridae) from the Marquesas Islands. Copeia Penida assessment. Likewise, we gratefully 2001 (no. 1): 92-107. acknowledge the State University of Papua (partic - RANDALL , J. E. &MCCOSKER , J. M. 1992. Two new dam - ularly Paulus Boli and Emmanuel Manangkalangi), selfishes of the genus Chromis (Perciformes: Pomacentri - dae) from the South Pacific. Proceedings of the California the Cenderawasih Bay National Park Authority Academy of Sciences 47 (12): 329-337. (particularly Kemal Amas and John Sroyer), and SENOU , H. & K UDO , T . 2007. A new species of the genus Conservation International (especially Chris Chromis (Perciformes: Pomacentridae) from Taiwan and Rotinsulu) for arranging and supporting the Japan. Bulletin of the National Museum of Natural Science, ichthyology field course which led to the discovery Series A , Supplement 1, 51-57. aqua vol. 15 no. 3 - 20 July 2009 134 aqua, International Journal of Ichthyology

Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia

Gerald R. Allen 1 and Mark V. Erdmann 2

Received: 23 April 2009 – Accepted: 03 July 2009

Abstract 71 (65 en moyenne) de vertèbres préanales, 204 à 213 A new species of heterocongrine garden eel is described (207,4 en moyenne) de vertèbres au total et 63 à 69 (65,7 from West Papua, Indonesia based on 13 specimens, 106.3- en moyenne) de pores préanales sur la ligne latérale. Une clé 678.1 mm TL. It differs from other members of the genus est fournie pour les espèces d’ Heteroconger de l’Indo-Paci - by its distinct colour pattern, consisting of zebra-like black fique ouest. and white barring on the head grading to a complex black and white maze pattern on the body and a combination of Sommario features that include dorsal fin origin anterior to gill open - Una nuova specie di grongo di giardino (sottofamiglia He- ing, pterygoid teeth present, body depth at gill opening terocongrinae) è descritta sulla base di 13 esemplari di 106.3- 28.8-35.2 percent of head length, 62-71 (mean = 65) pre - 678.1 mm TL raccolti nelle acque di Papua Occidentale, anal vertebrae, 204-213 (mean =207.4) total vertebrae, and Indonesia. Differisce dagli altri membri del genere per la 63-69 (mean = 65.7) preanal lateral-line pores. A key is pro - caratteristica colorazione, costituita da una zebratura bianca e vided for the Indo-west Pacific species of Heteroconger . nera sul capo che degrada verso un complesso labirinto di motivi neri e bianchi sul corpo, e per una combinazione di Zusammenfassung caratteristiche che comprendono l’origine della pinna dorsale Beschrieben wird eine neue Art der Röhrenaale (Unterfam - anteriore all’apertura branchiale, presenza di denti pterigoi- ilie Heterocongrinae) von West-Papua, Indonesien, auf der dei, altezza del corpo a livello della fessura branchiale 28.8- Grundlage von 13 Exemplaren mit 106,3-678,1 mm TL. Sie 35.2 percento della lunghezza della testa, 62-71 (valore me- unterscheidet sich von anderen Angehörigen der Gattung dio = 65) vertebre preanali, 204-213 (valore medio =207.4) durch die unverkennbare Farbgebung: schwarz-weiße vertebre totali and 63-69 (valore medio = 65.7) pori preanali Zebrastreifen am Kopf, die übergehen in ein komplexes Net - della linea laterale. Si fornisce una chiave dicotomica per le zmuster auf dem Rumpf, sowie durch eine Kombination specie di Heteroconger dell’Indo-Pacifico occidentale. weiterer Merkmale: u.a. Ansatz der Rückenflosse vor der Kiemenöffnung, vorhandene Pterygoidzähne, Körpertiefe an INTRODUCTION der Kiemenöffnung 28,8-35,2 Prozent der Kopflänge, 62-71 Heterocongrine garden eels are common inhabi - (Mittel 65) präanale Wirbel, Gesamtzahl der Wirbel 204- 213 (Mittel 207,4), 63-69 (Mittel 65,7) präanale Seitenlin - tants of sand-rubble areas, frequently in the vicin - ien-Poren. Angefügt ist ein Bestimmungsschlüssel für die ity of coral reefs. The group contains 32 species indo-westpazifischen Arten der Gattung Heteroconger. worldwide in two genera (Eschmeyer & Fricke 2009), Gorgasia Meek & Hildebrand, 1923 and Résumé Heteroconger Bleeker, 1868. The majority, with the Une nouvelle espèce d’anguille jardinière hétérocongrine exception of eight species from the eastern Pacific est décrite originaire de Papouasie occidentale, Indonésie, and Atlantic, are restricted to the Indo-Pacific sur base de 13 spécimens, 106,3-678,1 mm de LT. Elle se region. They were reviewed by Castle & Randall distingue des autres membres du genre par son patron de (1999), who also provided descriptions of five new coloration qui se compose de zébrures blanches et noires sur species. An additional species, la tête passant à un patron complexe labyrinthique noir et Gorgasia thamani blanc sur le corps et par une combinaison de caractéri- Greenfield & Niesz, 2004 was later described from stiques qui comprennent une base de dorsale antérieure à Fiji. Earlier publications treating this group l’ouïe, la présence de dents ptérygoïdes, la hauteur du corps include those of Böhlke & Randall (1981) and à l’ouïe de 28,8 à 35,2 % de la longueur de la tête, de 68 à Castle & Randall (1995).

135 aqua vol. 15 no. 3 - 20 July 2009 Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia

Garden eels occur at depths between about 2-55 Peninsula of West Papua, Indonesia during March m. The eels construct permanent burrows in the 2009. This general region has proved to be partic - sand, which they never leave. Mating occurs ularly rich for garden eels during our investigations between members of the opposite sex that occupy over the past three years having previously col - adjacent burrows by intertwining their bodies lected and/or observed Heteroconger enigmaticus while the tail tips remain in their respective bur - Castle & Randall, 1999, H. hassi (Klausewitz & rows (personal observations). Zooplankton is the Eibl-Eibesfeldt, 1959), H. perissodon Böhlke & primary dietary item and the eels rise nearly to full Randall, 1981, H. taylori Castle & Randall, 1995, extent out of their burrows when feeding in the Gorgasia barnesi Robison & Lancraft, 1984, G. passing current. When disturbed the eels retreat maculata Klausewitz & Eibl-Eibesfeldt, 1959, G. backward down their burrows. When a diver naeocepaea (Böhlke, 1951), and G. preclara Böhlke swims through an extensive colony a “wave effect” & Randall, 1981. is often created – the eels directly ahead gradually disappear into the sand, while those in the divers MATERIALS AND METHODS wake slowly reappear. Randall & Castle (1999) Terminology and methods follow Castle and provided further details of the biology and system - Randall (1999). Head pore terminology is from atics of the group. McCosker et al. (1989) and the following abbrevi - The present paper describes a strikingly-marked ations are used: SO (supraorbital series), IO (infra - species of Heteroconger that was discovered at orbital series), POM (preopercular-mandibular Papisol Bay (Fig. 1), on the southern Bird’s Head series), ST (supratemporal series). Total length and

Fig. 1. Satellite map of Bird’s Head Peninsula region of West Papua, Indonesia, showing type locality (yellow star) of Heteroconger mercyae . aqua vol. 15 no. 3 - 20 July 2009 136 Gerald R. Allen and Mark V. Erdmann head length are abbreviated as TL and HL respec - mean = 21.3); gill opening 9.2 (6.6-10.1; mean = tively. Vertebral counts were made from radi - 8.6); branchial interspace 21.5 (18.9-24.7; mean = ographs and include the hypural. Type specimens 21.6); pectoral-fin length 10.1 (7.3-10.6; mean = are deposited at Pusat Penelitian dan Pengemban - 9.0); depth at gill opening 32.9 (28.8-35.2; mean gan Oseanologi, Jakarta, Indonesia (NCIP), = 32.0); depth at anus 24.1 (23.5-28.4; mean = National Museum of Natural History, Washing - 25.7). ton, D.C. (USNM), and the Western Australian Body relatively elongate, slightly compressed Museum, Perth (WAM). along head, progressively more so to tail tip; great - est depth at pectoral fins; anus at about 30% TL; head little depressed, flexed downward at pectoral Heteroconger mercyae n. sp. fins; lower jaw slightly protruding; snout short and (Figs 2-4) rounded; upper and lower lips well developed, the upper confluent medially, enclosing anterior nos - Holotype: NCIP 6352, male, 641.5 mm SL, trils, and turned back on anterolateral face of Papisol Bay, 4 °4.601’S 133 °2.576’E, Papua Barat snout; lower lip similarly turned back on lower jaw; Province, Indonesia, 5-6 m depth, clove oil and anterior nostril a minute tube close to midline of hand net, M. Paine, M. Erdmann & G. Allen, 20 March 2009. Paratypes (collected with holotype) : NCIP 6353, 3 specimens, 560.7-653.9 mm TL; USNM 391795 , 4 specimens, 385.1-678.1 mm TL; WAM 33088-001, 5 specimens, 106.3-677.4 mm TL. Diagnosis: A relatively slender species of Hetero - conger (Congridae) with dorsal-fin origin anterior to gill opening, pterygoid teeth present, body depth at gill opening 28.8-35.2 percent of HL, 62- 71 (mean = 65) preanal vertebrae, 204-213 (mean =207.4) total vertebrae, 63-69 preanal lateral-line pores, and body colouration consisting of zebra- like black and white barring on head grading to complex black and white maze pattern on body. Description: Counts and proportions of holo - type, followed by range and mean value for 9 paratypes (in parentheses). Vertebral formulae 6/67/205 (4-9/62-71/204-213); average total ver - tebrae for holotype and nine paratypes 207.4; lat - eral-line pores before pectoral fin 6 (6-7); lateral- line pores before anus 66 (63-69, mean = 65.7); total lateral-line pores 201 (198-210; mean = 205.8); pectoral-fin rays 10 (9-11; mean = 10.3); dorsal-fin rays before anus 190 (173-215; mean = 193.8); total dorsal rays 588 (578-629; mean = 591.6); anal rays 395 (390-432; mean = 412.5); head pores on left side, holotype (and 9 paratypes): SO 1+2, IO 2+2, POM 5+2, ST 1+1. As percent of total length: snout-anus length 29.7 (27.0-31.3; mean = 29.4); as percent of snout-anus length: HL 11.8 (10.9-14.7; mean = 11.9); predorsal length 11.0 (10.1-13.8; mean = 11.4); as percent of HL: snout 14.9 (13.5-15.7; mean = 14.7); eye 19.7 Fig. 2. Heteroconger mercyae , underwater photo of (20.0-23.9; mean = 21.2); fleshy interorbital 11.4 paratype, 641.4 mm TL, Papisol Bay, West Papua, Indone - (10.2-14.1; mean = 10.9); mouth 21.9 (18.1-23.6; sia. Photo by M. Erdmann.

137 aqua vol. 15 no. 3 - 20 July 2009 Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia snout; posterior nostril a simple opening near rows on maxilla, tapering to two rows posteriorly; anterodorsal corner of eye; mouth oblique, reach - teeth in about five irregular rows on dentary, taper - ing to below anterior rim of eye; eye relatively ing to a single row posteriorly; intermaxillary- large, oval, scarcely intruding into dorsal profile; vomerine patch with 3-4 rows of teeth anteriorly, throat with moderately developed folds extending the tooth patch tapering posteriorly and nearly as to branchial aperture. long as maxillary row; pterygoid teeth in single Gill opening before pectoral fins, slightly oblique row. with its ventral end more posterior than dorsal end; Colour in life (Figs 2, 3): conspicuous black pectoral fins small rounded flaps; dorsal-fin origin and white zebra-like bars, spots, and oblique bands about one-half eye diameter anterior to gill open - on head, grading to bold maze of black and white ing and low in preserved specimens; anal fin simi - vermiculations on body; dorsal fin translucent with larly low; caudal fin much reduced, tip of tail occasional rays marked with alternating black and pointed. white bands, and alternating black and white spots Head pores inconspicuous, with very slightly along outer margin; a yellow-orange iris ring on raised rims; lateral-line pores minute and difficult eye; lips white. to distinguish, especially on posterior one-fourth of Colour in alcohol (Fig. 4): Generally dark body, approximately matching number of verte - brown, but close inspection reveals dark markings brae. as described in live colouration above; about poste - Teeth small, although those of pterygoid and rear rior one-fifth of total length light brown with portion of vomer slightly enlarged; teeth in three dense pattern of melanophores.

Fig. 3. Heteroconger mercyae, underwater photo of paratype, 641.4 mm TL, Papisol Bay, West Papua, Indonesia. Photo by M. Erdmann. aqua vol. 15 no. 3 - 20 July 2009 138 Gerald R. Allen and Mark V. Erdmann

Remarks: The new species has one of the highest numerous, relatively large black spots with white average number of vertebrae (207.4) for the genus. interspaces (Fig. 5), some individuals (Fig. 6) Other species with relatively high average counts exhibit a pattern that approaches that of H. mer - include H. cobra Böhlke & Randall 1981 from the cyae . However, the head is marked with irregular Solomon Islands (199.8), H. tomberua Castle & spots and bands rather than zebra-like bars and the Randall, 1999 from Fiji (195.1), and H. tricia Cas - vermicular pattern of the body is less complex. tle & Randall, 1999 from Flores, Indonesia (210, Moreover, H. taylori has relatively large black spots based on a single specimen). However, these three on the dorsal fin and lacks the pattern of alternat - species have very different colour patterns consist - ing black and white spots along the dorsal-fin mar - ing of either numerous small round spots and trio gin. The two species also show significant differ - of large u-shaped black saddles (with white cental ences in vertebral counts (204-213 for H. mercyae portion) on head and anterior body (H. cobra), 2- and 169-172 for H. taylori). The new species also 3 irregular rows of small dark spots above and appears to attain a larger size than most members below lateral-line (H. tomberua), or widely scat - of the genus, at least based on museum specimens. tered large dark tan (possibly orange in life) spots, Castle & Randall (1999) reported maximum total becoming more numerous on posterior half of lengths ranging from 331-537 mm TL, with most body (H. tricia). The only species that has a colour species under 450 mm. pattern similar to H. mercyae is H. taylori , which Etymology: The new species is named mercyae in ranges from Papua New Guinea westward to Bali, honour of Mercy Paine, who discovered the eel Indonesia. Although the usual pattern consists of colony and helped us collect the type specimens.

Fig. 4. Heteroconger mercyae , preserved holotype, 641.4 mm TL, Papisol Bay, West Papua, Indonesia. Photo by G. Allen.

139 aqua vol. 15 no. 3 - 20 July 2009 Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia

Key to the Indo-west Pacific species of 7a. Body markings as evenly distributed, rounded, Heteroconger small, discrete dark spots on a light back - (adapted from Castle & Randall 1999) ground, but with no other markings; total vertebrae 186-208 (Fiji) ...... 1a. Body white with numerous narrow black bars ...... H. tomberua Castle & Randall, 1999 on head and trunk, some on trunk meeting 7b. Body markings more intense, either consist - opposite neighbour ventrally; no pectoral ing of closely packed dark spots or spots more fins; total vertebrae 153-159 (central Indone - closely packed and/or joining to form con - sia to Japan and Vanuatu) ...... spicuous mottling, or maze-like network, ...... H. polyzona Bleeker, 1868 with or without additional, prominent and 1b. Colour not as in 1a; pectoral fins present .. 2 distinctive bars, bands or blotches; total verte - 2a. Body plain brown, with or without light head brae 164-213 ...... 8 markings ...... 3 8a. Dorsal-fin origin more or less over gill open - 2b. Body colour not as in 2a, variously spotted or ing or a little behind it ...... 11 mottled and/or with more prominent addi - 8b. Dorsal-fin origin clearly anterior of gill open - tional black blotches or marks ...... 5 ing ...... 9 3a. Body plain medium brown with a white-yel - 9a. Body colouration consisting of zebra-like low patch on postorbital and white margins black and white bars on head grading to com - on median fins; pterygoid teeth present; lat - plex black and white maze pattern on body; eral-line pores before anus 54-59; total verte - total vertebrae 204-213 ...... H. mercyae n. sp. brae 173-176 (Philippines and Indonesia) .... 9b. Body colouration not as in 9a, usually spot - ...... H. perissodon ted, although sometimes consisting of maze- 3b. Body dark brown without light head mark - like pattern on body without zebra-like bars ings or white margins on median fins; ptery - on head; total vertebrae 169-176 ...... 10 goid teeth present or absent; lateral-line pores 10a. Body evenly spotted to mottled or occasion - before anus 47-55 ...... 4 ally with maze-like pattern, without other 4a. Pterygoid teeth present on most specimens; markings; total lateral-line pores 157-160; total vertebrae 154-163 (eastern Indonesia) .. mouth 24-30% of HL; pterygoid teeth pre - ...... H. enigmaticus sent; total vertebrae 169-172 (Indonesia and 4b. No pterygoid teeth; total vertebrae 144 Papua New Guinea) ...... H. taylori (Nicobar Islands) ...... 10b. Body evenly and closely spotted, with a black - H. obscurus (Klausewitz & Eibl-Eibesfeldt, 1959) ish anterior head and light throat; total lat - 5a. Body with relatively large, evenly spaced, round eral-line pores 161-168; mouth 18.4-21.3% dark tan spots (orange in life in H. balteatus ), of HL; no pterygoid teeth; total vertebrae with or without other markings ...... 6 173-176 (Marshall Islands and French Poly - 5b. Body markings as small closely packed dark nesia) ...... spots or freckles on a pale background; the ...... H. lentiginosus Böhlke & Randall, 1981 spots may link together to form irregular 11a. Three prominent black patches, one encom - mottling or a maze-like network, with or passing gill opening and pectoral fin, a second without several other prominent dark and/or halfway along midlateral portion of trunk, light bars or patches ...... 7 and the third encompassing anus; total verte - 6a. A wide oblique white bar on trunk and a less brae 164-175 (Indo-Pacific) ...... H. hassi prominent white patch on postorbital head; 11b. Three prominent dark and light (combined), lateral-line pores before anus 52-56; HL 15.0- u-shaped saddles, one on postorbital and two 18.0% of snout-anus length; total vertebrae spaced along anterior trunk; vertebrae 198- 158-161 (Red Sea) ...... 204 (Solomon Islands) ...... H. balteatus Castle & Randall, 1999 ...... H. cobra Böhlke & Randall, 1981 6b. No white bar on trunk and no white patch on head; lateral-line pores before anus 64-68; HL Zoogeography and habitat: The new species is 13.5-14.5% of snout-anus length; total verte - currently known for certain only from the type brae 210 (central Indonesia) ...... locality (Fig. 1), which is situated about 95 km ...... H. tricia Castle & Randall, 1999 southwest of the town of Kaimana, West Papua, aqua vol. 15 no. 3 - 20 July 2009 140 Gerald R. Allen and Mark V. Erdmann

Indonesia. However, it likely ranges at least as far east as Rabaul, New Britain, based on an underwa - ter photograph by R. Steene, showing a close-up view of the head of a garden eel that appears to be H. mercyae . The type specimens were obtained from a single colony, occupying about 100 m 2 and containing approximately 50 eels in 5-6 m depth. The bottom consisted of fine, silty sand and was no doubt influenced by a freshwater stream along the adja - cent shoreline, about 100 m distance. The nearest reef formation was about 50 m from the site.

ACKNOWLEDGEMENTS We are especially grateful to Dexter and Susan Paine and their three children, Mercy, Sam, and Honor, for inviting us to participate on the Silolona cruise to Banda and West Papua during March 2009. Special thanks are also due Mercy Paine for her diving and collecting assistance, including her discovery and capture of the first specimen of H. mercyae . We anticipate a bright future for her in ichthyology. We also express sin - cere thanks to Patti Seery, owner of the live-aboard Fig. 5. Heteroconger taylori , underwater photo, approxi - dive vessel Silolona, and her wonderfully efficient mately 450 mm TL, Bali, Indonesia. Photo by T. and hospitable crew. We thank the local govern - Tonozuka. ments of Kaimana and Raja Ampat for continued

Fig. 6. Heteroconger taylori , underwater photo, approximately 400 mm TL, Bali, Indonesia. Photo by T. Tonozuka.

141 aqua vol. 15 no. 3 - 20 July 2009 Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia support of Conservation International’s marine (Congridae, Heterocongrinae) from Papua New Guinea conservation initiatives in the Bird’s Head, and Dr. and Indonesia. Revue française d’Aquariologie 22 (1-2): 3-6. Suharsono and Rianta Pratiwi of the Indonesian CASTLE , P. H. J & R ANDALL , J. E. 1999. Revision of the Institute of Sciences for support and assistance Indo-Pacific garden eels (Congridae: Heterocongrinae), with descriptions of five new species. Indo-Pacific Fishes 30: with curation of specimens. Sue Morrison of 1-52. WAM prepared radiographs and offered curatorial ESCHMEYER , W. N. & F RICKE , R. (eds.) Catalog of Fishes assistance. electronic version (updated 13 Mar. 2009). http://research.calacademy.org/ichthyology/catalog/fishc atsearch.html REFERENCES MCCOSKER , J. E., B ÖHLKE , E. B. & B ÖHLKE , J. E. 1989. BÖHLKE , J. E. & R ANDALL , J. E. 1981. Four new garden eels Family Ophichthidae. In: Fishes of the Western North (Congridae, Heterocongrinae) from the Pacific and Indian Atlantic . Part Nine. Volume 1: Orders Anguilliformes Oceans. Bulletin of Marine Science 31 (2): 366-382. and Saccopharyngiformes. (Ed. E. B. Böhlke): 254-412. CASTLE , P. H. J & R ANDALL , J. E. 1995. A new garden eel Sears Foundation for Marine Research, New Haven.

aqua vol. 15 no. 3 - 20 July 2009 142 aqua, International Journal of Ichthyology

Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

Stefano Valdesalici 1, Marc Bellemans 2, Kiril Kardashev 3, Alexander Golubtsov 4

1) Via Cà Bertacchi 5, 42030 Viano (RE), Italy. E-mail: [email protected] and [email protected] 2) Mortselsesteenweg 138, 2540 Hove, Belgium 3) Stramna 4, 2600 Dupnitsa, Bulgaria 4) Laboratory of Lower Vertebrate Ecology, A.N. Severtsov Institute of Ecology & Evolution, Russian Academy of Sciences, Leninskii Pr. 33, Moscow119071, Russia

Received: 19 March 2009 – Accepted: 04 July 2009

Abstract Résumé A new species of annual killifish, Nothobranchius nubaensis , Une nouvelle espèce de killy annuel, Nothobranchius is described based on specimens collected from ephemeral nubaensis, est décrite sur base de spécimens collectés dans water bodies in central Sudan and south -west Ethiopia. The des poches d’eau temporaires du centre du Soudan et du new species is distinguished from the other members of the sud-ouest de l’Ethiopie. La nouvelle espèce se distingue des Nothobranchius ugandensis species group by the following autres membres du groupe Nothobranchius ugandensis par combination of characters : 17-19 dorsal fin rays; 17-19 anal la combinaison des caractéristiques suivantes: 17 à 19 fin rays; 29-30 scales in median lateral series; dorsal and anal rayons dorsaux; 17 à 19 rayons à l’anale, 29 à 30 écailles fins in males with short filamentous rays; pelvic fins short, dans les rangées latérales médianes; dorsale et anale tips reaching the anus; orange red head with three distinct des mâles avec de courts rayons filamenteux; pelviennes oblique bars on the operculum ; dorsal fin light blue with a courtes, les extrémités touchant l’anus; tête rouge orange pattern of irregular orange -red spots and elongated yellow avec trois barres obliques nettes sur l’opercule; dorsale spots in distal areas; anal fin yellow, light blue at base, with a bleu clair avec un patron irrégulier de taches rouge-oran- pattern of orange -red spots; pelvic fins yellow with a pattern ge; pelviennes jaunes avec un patron de taches rouge of orange -red spots; pectoral fins orange -red with a light blue orange; pectorales rouge orange avec un liseré bleu clair; margin; caudal fin orange -red with short pale or dark red caudale rouge orange avec de petites lignes rouges pâles ou lines , extending from fin base onto fin rays. foncées s’étendant de la base de la nageoire jusqu’aux rayons. Zusammenfassung Eine neue Art der einjährigen Killifische: Nothobranchius Sommario nubaensi, wird auf der Basis von Exemplaren beschrieben, Una nuova specie di killifish annuale, Nothobranchius die in kurzzeitigen Wasserlöchern im Zentralsudan und in nubaensis, è descritta sulla base di esemplari raccolti in corpi SW-Äthiopien gesammelt wurden. Sie unterscheidet sich idrici temporanei del Sudan centrale e dell’Etiopia sudocci - von anderen Angehörigen der Nothobranchius-ugandensis- dentale. La nuova specie si distingue dagli altri membri del Artengruppe durch die folgende Merkmale: 17-19 Rück - complesso di specie Nothobranchius ugandensis per la se- enflossenstrahlen; 17-19 Afterflossenstrahlen; 29-30 guente combinazione di caratteri: 17-19 raggi dorsali; 17-19 Schuppen in median-lateralen Reihen; die dorsalen und raggi anali; 29-30 scaglie lungo la serie mediana laterale; analen Flossen bei Männchen mit kurzen fadenförmigen pinna dorsale e anale nei maschi con brevi raggi filamentosi; Flossenstrahlen; Bauchflossen kurz, die Spitzen reichen bis pinne pelviche brevi, con le punte che arrivano fino all’ano; zum After; orangeroter Kopf mit drei deutlichen schrägen capo rosso-arancio con tre distinte barre oblique sull’oper - Streifen auf dem Kiemendeckel; Rückenflosse hellblau mit colo; pinna dorsale blu chiaro con un motivo di macchioline einem unregelmäßigen Muster orangeroter Flecken und irregolari rosso-arancio e macchie allungate gialle nelle aree länglichen gelben Flecken im distalen Bereich; Afterflosse distali; pinna anale gialla, blu chiaro alla base, con un gelb, hellblau am Grund, mit einem Muster orangeroter motivo di macchioline rosso-arancio; pinne pelviche gialle Flecken; Brustflossen gelb mit einem Muster orangeroter con un motivo di macchioline rosso-arancio; pinne pettorali Flecken; Brustflossen orangerot mit hellblauem Rand; rosso-arancio con un margine blu chiaro; pinna caudale Schwanzflosse orangerot mit kurzen blassen oder dunkel - rosso-arancio con brevi linee pallide o rosso scuro che si roten Linien, die sich von der Flossenbasis bis zu den estendono dalla base della pinna fino ai raggi. Flossenstrahlen erstrecken.

143 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

INTRODUCTION peduncle length , which is measured from the poste - The freshwater fish fauna of Sudan is poorly rior end of the anal fin base to the posterior end of known. It was studied by Sandon (1950a, 1950b, the hypural plate ; eye diameter , which is measured 1957) and more recently by Bailey & Hickley between anterior and posterior orbital walls ; and (1986 ), Hickley & Bailey (1987) , Lévêque et al. snout length , which is measured from the anterior (1991), and Bailey (1994). The freshwater fish fauna tip of the lower jaw to the anterior edge of the of Ethiopia is better known (Tedla 1973, Tedla & orbital wall. Measurements were taken with digital Meskel 1981, Dgebuadze et al. 1994, Golubtsov et callipers to the nearest 0.1 mm, partly under a dis - al. 1995, de Graaf et al. 2000). secting microscope. All measurements and counts Nothobranchius virgatus Chambers, 1984 is the were taken from the left side. All visible rays of the only species of Nothobranchius , which has been dorsal and anal fins were counted. The counts of described from Sudan. It was collected in October scales on the median longitudinal series is the num - 2008. From Ethiopia it is known from Gambella, ber of scales between the superior junction of the Obela Stream , near the road from Gambela to Gog opercular membrane and the hypural plate . The (J. Friel , pers. comm.). scales on the base of the caudal fin were counted sep - On 5 December 2002, live specimens of a red arately. All measurements are presented as percent - species of Nothobranchius were collected by M. Belle - ages of standard length (SL); except of eye diameter mans in Khor Angarko (which is part of the Khor and snout length, which are given as percentage of Abu Habl system ), South Kordofan , about 21 km head length (HL). Terminology for the cephalic south of Dilling and 100 km north of Kadugli (Belle - neuromast series follows Scheel (1968) , the frontal mans 2003). This species was provisionally described squamation following Hoedeman (1958). as N. nubaensis (Bellemans 2003), but this name is Type specimens and additional material is not available, for the following reasons: (1) the men - deposited in the following institutions:, Musée Royal tion of a “provisional description ” may questionably de l’Afrique Centrale , Tervuren, Belgium (MRAC ); be considered to violate ICZN Article 16.1; (2) no Natural History Museum, London, UK (BMNH ); type has been designated in the paper, the author Muséum National d’Histoire Naturelle, Paris, France rather states “types will be designated …”. Article (MNHN ); Zoological Museum, Moscow State Uni - 16.4.1 of the ICZN clearly requires an “explicit fixa - versity, Moscow, Russia (ZMM ); South African tion of a holotype, or syntypes ”, even if not immedi - Institute for Aquatic Biodiversity, Grahamstown, ately deposited. The paper mentions several speci - South Africa (SAIAB ); and, Museo Civico di Storia mens without giving numbers. The name N. nubaen - Naturale “G. Doria ”, Genova, Italy (MSNG ). sis is not a “nomen nudum ” as a description is given, Comparative material is listed in Blache & Miton but an “anoplonym ” according to Dubois (2000). (1960), Chambers (1984, Rahad el Fula population), In November 2005 and November 2006, K. Kar - and Wildekamp (1995) . Further collections exami - dashev and others collected specimens of red nated include: Stefano Valdesalici private collection: coloured Nothobranchius in Khor Maada Mulaha , Nothobranchius robustus , 4 males, 31,7-36,3 mm SL, belonging to the Wadi Al Ghallah system (Karda - Uganda, Kalisizo 0°33’S 31°36’E ; N. rubroreticula - shev 2007 a , b). In November 1986, November tus , 2 males, 34,3-36,9 mm SL, Tchad, Zakouma 1994 and December 2006, A. Golubtsov collected National Park 10°50’N 19°38’E ; N. ugandensis , 1 specimens of red coloured Nothobranchius near male, 47,3 mm SL, Uganda, Butiaba 01 °53’N Abobo , Ethiopia. They occured in seasonal ponds of 31°24’E; 1 male, 43,7 mm SL , Uganda, Busesa the Alvero River drainage , White Nile basin. Speci - 00°30’N 33°31 E. mens of these three populations from Sudan and Ethiopia are considered to belong to the same species on the basis of the coloration of males and Nothobranchius nubaensis n. sp. morphological characters . Here below this annual (Figs 1-6, Table I) fish is described as a new species , Nothobranchius nubaensis . Nothobranchius nubaensis Bellemans, 2003 (anoplonym ) MATERIAL AND METHODS Measurements and counts were taken according to Holotype: MRAC 2008-05-P-1, male, 42,9 mm Amiet (1987) ; except for the addition of the caudal SL; Sudan: Angarko, 21 km south of Dilling and aqua vol. 15 no. 3 - 20 July 2009 144 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov

100 km north of Kadugli, 11°53’ 05.23”N base, with a pattern of orange -red spots; pelvic fins 29°42’31.82”E, Marc Bellemans, 5 December yellow with a pattern of orange -red spots; pectoral 2002. Collected with hand net and preserved after fins orange -red with a light blue margin; caudal fin living for about six months in aquaria . orange -red , with short pale or dark red lines extend - Paratypes (collected with holotype ): MRAC 2008- ing from fin base onto fin rays. 05-P-2, 1 male, 43,9 mm SL ; MRAC 2008-05-P-3- Description: Morphometric measurements are 4, 2 females, 28,1-33,2 mm SL ; MSNG 54286, 1 given in Table I. Robust, deep bodied Notho - male, 40,8 mm SL ; MNHN 2008-1140–1141- branchius with rounded snout ; terminal mouth 1179, 3 males, 38,4-45,8 mm SL ; SAIAB 80410, 1 directed upward; dorsal fin rays 17-19; anal fin rays male, 36,5 mm SL ; BMNH 2008.4.2.11-12, 1 17-19; scales in median lateral series 29-30 + 2-3 on male, 45,1 mm SL, 1 female, 33,1 mm SL . caudal fin base; cephalic squamation pattern vari - None-type material: SAIAB 80411, 1 male, able; anterior neuromast series of ‘open’ type; central 49 mm SL, Sudan: Maada Mulaha, 11°09.405’N series in shallow groove with low lobes and with 3 40° 19.441’ E, Kiril Kardashev and Plamen Ivanov, neuromasts; posterior cephalic neuromast series 22 November 2005 , collected with large hand nets curved with 3 neuromasts; preopercular neuromast and preserved in the field ; SAIAB 80412, 1 female, system in open groove, distal ridge overlaping oper - 40,8 mm SL , BMNH 2008.4.2.13-14, 1 male, cle slightly; one neuromast on each scale of median 47,1 mm SL, 1 female, 39,8 mm SL, same data as longitudinal series. SAIAB 80411 ; ZMMGU P22009, 3 males, 38,4- Maximum observed length 49.9 mm SL in males; 49,9 mm SL , Ethiopia: Alvero River drainage, 7 km body deep and moderately compressed ; dorsal pro - S of the southern end of the Alvero Dam, 7° 47’N file slightly concave to nearly straight on head, con - 34° 25’E, altitude about 500 m, Alexander Gol - vex from nape to end of dorsal fin base ; ventral pro - ubtsov, 4 December 2006 , collected with large nets file convex, slightly concave on caudal peduncle pos - and preserved in the field. terior to dorsal and anal fins; maximum body height Diagnosis: Nothobranchius nubaensis can be distin - occurring at the base of the pelvic fins; dorsal and guished from all other species of the genus by the anal fins shallow and rounded, tips with short fila - following combination of characters: 17-19 dorsal mentous rays; dorsal and anal fins with papillate fin rays; 17-19 anal fin rays; 29-30 scales in median contact organs , distributed over fin rays; pelvic fins lateral series; dorsal and anal fin with short filamen - shorts, tips reaching the anus; caudal fin subtrun - tous rays in males; pelvic fins short, tips reaching the cate ; branchiostegal membrane projecting from anus; orange -red head with 3 distinct oblique bars opercle, distal edge slightly wrinkled. on the operculum ; dorsal fin light blue with a pat - Female smaller than male, maximum observed size tern of irregular orange -red spots and elongated yel - 40.8 mm SL; body less compressed and less deep low spots in distal areas; anal fin yellow, light blue at than in males; dorsal fin rounded; anal fin triangu -

Fig. 1. Adult male of Nothobranchius nubaensis (not preserved) from Angarko. Photo by S. Valdesalici.

145 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

Table I. Morphometric measurements of Nothobranchius nubaensis (including type series and additional specimens).

Holotype Males (n=12) Females (n=5) SL (mm) 42.9 36.5-49.9 33.1-40.8 Depth at the pelvic fins 31.4 25.9-34.0 21.4-30.4 Preanal length 60.3 56.5-65.7 58.7-68.8 Predorsal length In percents of standard length (SL) 56.8 54.4-61.8 57.8-61.4 Prepelvic length 47.5 44.0-54.9 49.7-56.0 Caudal peduncle lenght 20.2 18.0-22.0 14.9-22.3 Caudal peduncle depth 12.8 11.7-15.4 9.3-12.9 Head length 29.3 26.3-33.6 25.6-33.9 Snouth length 23.8 21.6-32.3 20.7-26.8 In percents of head length (HL) Eye diameter 23.8 20.7-29.8 23.7-29.1 lar with rounded tip; anal fin positioned more pos - Female (Fig. 4): body pale olive-brown, lighter terior ; short snout and narrower caudal peduncle brown to silvery ventrally; unpaired and paired fins than in males; opercular membrane not projecting hyaline; iris golden, with faint black vertical bar from opercle. through center of eye. Color in life : Male (Figs 1-3): body and head Color in alcohol: Male (Fig. 5): body scales scales light blue with broad orange -red margin, cre - light brown to whitish, almost all scales with distinct ating a reticulated pattern on body ; almost com - red margin; dorsal and anal fins light brown to pletely orange -red on head; lips, snout, throat, whitish with a pattern red spots; caudal fin of a light frontal and superior portion of head all orange -red; pale red; pelvic fins light brown to whitish with red opercular region with three distinct oblique bars on spots near base; pectoral fins of a light pale red; iris the operculum, extending from the eye to the upper bluish. part of the head; branchiostegal membrane orange - Female (Fig. 6): body light brown to whitish; oper - red with light blue rim; dorsal fin light blue with cular and ventral area yellowish; unpaired and paired broad dark red reticulation, forming approximately fins whitish; iris bluish. horizontal arch-like stripes, broad border with inter - Distribution (Fig. 7) : Populations have been found rupted red pattern; some elongated yellow spots in scattered over a large area. Two groups of localities distal areas; anal fin yellow, light blue at base, with can be distinguished: (1) Khor Angarko (in South pattern of large orange -red spots ; spots in the mid - Kordofan around the foothills of the Nuba Moun - dle of the fin dark red, distal spots orange -red; cau - tains) and the area near Khor Maada Mulaha (in the dal fin orange -red with pale or dark red lines extend - plains of West Kordofan) in Sudan ; (2) near Abobo ing from base onto fin rays , some specimens having (Gambella Region) in Ethiopia. a faint dark red marginal band; pelvic fins yellow Habitat (Figs 8-10) : The specimens of Notho - with orange -red spots, spots near base dark red; pec - branchius nubaensis described here were found in toral fins orange -red with light blue margin, some residual pools of seasonal rivers (called “Khor”) and specimens with yellow spots and short dark or pale rainwater filled depressions (Bellemans 2003, Kar - red lines extending onto fin rays; iris golden, with dashev 2007 a , b). All populations of Notho - faint black vertical bar through center of eye. branchius in central Sudan and south -west Ethiopia undergo important annual fluctuations in abun - dance and extent of distribution area. In Khor Angarko , N. nubaensis was the only fish species pre - sent in 2002. During successive surveys at the same collection site during the post-rain period in 2005 and 2006 no specimens of N. nubaensis were found, not even in the mud collected from the residual pools (M. B.). Fig. 2. Adult male of Nothobranchius nubaensis (not pre - In Khor Maada Mulaha Nothobranchius virgatus served) from Khor Maada Mulaha. Photo by K. Kardashev. and N. aff. rubroreticulatus were found in in the aqua vol. 15 no. 3 - 20 July 2009 146 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov same locality in 1998 and 1999 (Bellemans 2000) there are two seasonal ponds near Abobo where and in 2004 (J. Bulterman pers. comm.). In 2005 Nothobranchius nubaensis is known to occur, both and 2006 N. virgatus , N. nubaensis and Protopterus belonging to the Alvero river drainage (White Nile sp. (probably Protopterus annectens Owen, 1939) basin). were found in the same locality, but not a single In late November 1994 , two males and three specimen of N. aff . rubroreticulatus . In this locality females of Nothobranchius were collected from a all males Nothobranchius nubaensis were collected at swamp-like pond at the land divide between the the surface near rocks. Almost all females were col - Alvero and Gilo rivers (the latter river seems to be a lected in about 2 to 2.5 m deep water. No aquatic tributary of the Pibor River, affluent of Sobat, afflu - vegetation was present, with only dry grass along the ent of White Nile). The pond was muddy and cov - shores. The water was brown, slightly turbid, with a ered with dense macrophyte vegetation. The Pro - pH of 7.8- 8.2. At 12:30 h the temperature was topterus sp. larvae were sampled simultaneously with 22°C, at 1 m below the water surface. The air tem - the Nothobranchius specimens. Based on the anec - perature was 42°C. The bottom substrate of these dotal information from the local Anuak people, the habitats was brown-black clay (K.K.). In Ethiopia , pond is never connected to any river system; it is

Fig. 3. Freshly collected adult male of Nothobranchius nubaensis, ZMMGU P22009, from Akobo. Photo by A. Golubtsov.

Fig. 4. Adult female of Nothobranchius nubaensis (not preserved) from Angarko. Photo by S. Valdesalici.

147 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia filled with rain water. The same pond was re-sam - elephants (judging from foot prints). The pond was pled on 4 December 2006. No Protopterus were never re-sampled until present (A. G.). found. New villages and cultivated lands appeared in Etymology: The new species is named after type the vicinity of the pond, which became a source of locality, which lies in the Nuba mountains. drinking water for the nearest settlement located Discussion: Based on the known distribution, on about 2.5 km to the north . the coloration of males and on morphology, Notho - In the other pond, in late November 1986, about branchius nubaensis can be grouped in the Notho - 25 km W of Abobo (now 15 km W of a dam) branchius ugandensis species group, which is defined 7° 54’ N 34° 20’ E, altitude about 430 m, only one here . This clade includes N. ugandensis Wildekamp, female of Nothobranchius was sampled from the 1995, an undescribed species of Nothobranchius pan-like pond in the flood plain of the Alvero River ”Lake Victoria” (see comments in Wildekamp 1990: (a tributary of the Baro River, affluent of Sobat, 199, 1996: 147, and pictures in Seegers 1997: 82- affluent of White Nile). The pond had a light clay 83) and N. nubaensis , which is described here. bed and no dense macrophyte vegetation. Judging Synapomorphies to define this group are: male col - from the fish species composition (occurring oration consisting of light blue scales with a large together with the only Nothobranchius specimen), irregularly reticulated pattern on body , forming i.e. small Barbus sp ., Ctenopoma sp ., Neolebias sp ., oblique bars in vivid red color, vivid red coloration Aplocheilichthys (? , possibly Micropanchax ), the pond on the head and in dorsal areas, yellow to yellowish was connected to the river during the rainy season. anal fin, large vivid red spot pattern on dorsal and When we tried to re-sample the pond in early Janu - anal fins, red caudal fin of subtruncated shape; rela - ary 1987, no fish was found there; the water tem - tively deep body (up to 38% of SL) ; moderately perature was about 43°C; it was a watering place of compressed, and rounded head ; dorsal profile of

Fig. 5. Nothobranchius nubaensis , MRAC 2008-05-P-1, holotype, 42 .9 mm SL, male. Photo by S. Valdesalici.

Fig. 6. Nothobranchius nubaensis , MRAC 2008-05-P-3, paratype, 33.2 mm SL, female. Photo by S. Valdesalici. aqua vol. 15 no. 3 - 20 July 2009 148 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov head slightly concave to nearly straight, convex from spots pattern (broad and arranged in horizontal row nape to end of dorsal fin base ; variable cephalic vs. small spots , sometime arranged in thin lines squamation. extending onto fin rays) ; anal fin spot pattern (broad Male N. nubaensis differ from a similarly red male vs. absent or if present, few and arranged in lines morph of N. ugandensis by coloration of the whole extending to fin rays ) ; pectoral fin color (red vs. hya - body (orange -red broad scales margin vs. narrow red line). Males and females with relatively a reduced scale margins) and head (almost completely orange - number of scales in the longitudinal series (29-30 vs. red vs. red on lips and superior part only ); dorsal fin 28-33), modally less deep body in males (25.9-34. 0

Fig. 7. Geographical distribution of Nothobranchius nubaensis (asterisk; type locality: open asterisk ), Nothobranchius virgatus (circle) and N. aff . rubroreticulatus (orange squares) (black squares are cities with their names). Drawing by S. Valdesalici.

149 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia vs. 25.0-37.3% of SL) and females (21.4-30. 4 vs. line) ; caudal fin coloration (red vs . light blue with a 22.4-34. 0% SL) ; dorsal fin positioned more anterior black margin followed by a submarginal white band in males (54 .4-61.8 vs. 54.5-65.2% of SL) and and a faint black vertical bar in the inner region ); females (57 .8-61.4 vs. 57.0-66.5% of SL) ; origin of and females color pattern (hyaline without particu - pelvic fins relatively more anterior in males (44 .0- lar pattern vs. with faint oblique bars) ; more dorsal - 54.9 vs. 47.2-57.0% of SL) ; shorter head in males fin rays in males and in females (17-19 vs. 13-16); (26 .3-33.6 vs. 26.6-36.8% of SL) and females (25 .6- more anal -fin rays in males and females (17-19 vs. 33.9 vs. 27.6-37.8% of SL) ; body scales (without 14-16) ; modally less deep caudal peduncle in males ctenii vs. with ctenii), ctenii structures on operculum (11 .7-15.4 vs. 14.3-16. 4% SL) ; dorsal fin positioned (absent vs. present). more anterior in males (54 .4-61.8 vs. 59.0-62.8% of Among other species of Nothobranchius occurring SL) ; shorter head in males (26 .3-33.6 vs. 31.4- in the area, male N. nubaensis differ from male N. 35.7% of SL) ; pelvic fin shorter (tips reaching the virgatus (found in the same habitat in Khor Maada anus vs. tips reaching anal fin). Mulaha) by the coloration pattern of the whole Male N. nubaensis differ from male N. robustus by body (broad orange -red scale margins vs. thin red coloration of the whole body (light blue with broad scale margins , forming about 10-16 oblique bars) orange -red scale margins vs. completely red to red - and head (almost completely orange red vs. light brown) ; dorsal and anal fin pattern (spotted vs. plain blue with narrow red scale margins) ; anal fin spot red) ; pectoral fin coloration (red vs. hyaline) ; males pattern and coloration (yellow, light blue at base, and females with more dorsal -fin rays (17-19 vs. 14- with pattern of large orange -red spots, spots in the 16); shorter head in males (26 .3-33.,6 vs. 29.2- middle of fin dark red, distal spots orange -red vs. 38.8% of SL); different shape of head and snout light blue with red spots forming 6-8 curved lines (rounded vs. pointed). and a broad margin) ; pectoral fin color (red vs. hya - Male N nubaensis differ from male N. rubroreticu - latus by coloration of the whole body (broad orange - red scale margins vs. red broad scale margins form - ing a reticulated irregular pattern) and head (almost completely orange -red vs. light blue with narrow red scale margins); dorsal fin spot pattern (broad and arranged in horizontal rows vs. spots near the fin base , arranged in an irregular horizontal line, spots in the median part arranged in broad lines extend - ing onto fin rays, light blue submarginal band and broad black margin); anal fin coloration and spot pattern (yellow, light blue at base, with pattern of large orange -red spots, spots in the median part dark red, distal spots orange red vs. light blue with a series of red spots near the base forming a broad irregular horizontal line, spots in the median part arranged in broad red lines extending onto fin rays, light blue submarginal band and broad black margin) ; pectoral fin color (red vs. hyaline) ; caudal fin coloration (red vs. light blue with a pattern of broad red lines extending onto fin rays, light blue submarginal band and broad black margin ; deeper body in males (26 .3-34.0 vs. 25.4-29.7% of SL) ; anal fin posi - tioned more posteriorly in males (56 .5-65.7 vs. 55.1-59.3% of SL) ; shorter head in female (25 .6- 33.9 vs. 32.4-37.2% of SL) ; different shape of head and snout (rounded vs. pointed). The present disjunct distribution pattern of N. Fig. 8. Sudan: Angarko; type locality of Nothobranchius nubaensis , similar to that of N. virgatus can be nubaensis . Photo by M. Bellemans. explained by the dynamics of the changing rain pat - aqua vol. 15 no. 3 - 20 July 2009 150 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov tern and climatic conditions over the vast plains of in Chambers (1984), Hoelzmann et al. (2000) and central and southern Sudan and SW Ethiopia (see Leblanc et al. (2006). also Lévêque 1990). The higher water levels of the A comparison of the samples did not reveal any Nile in the past and the presence of a fossil embank - apparent differences in morphometry, color pattern ment surrounding a pre-historic lake south of Khar - or meristics among populations. Determining the toum, suggest a more continuous and permanent full extent of the distribution of N. nubaensis will connection between the presently known popula - require additional collections. tions of N. nubaensis (and N. virgatus), as reported

Fig. 9. Sudan: Khor Maada Mulaha. Photo by K. Kardashev.

Fig. 10. Ethiopia: Alvero River drainage, 7 km S of the southern end of the Alvero Dam (pond after intensive sampling). Photo by A. Golubtsov.

151 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

ACKNOWLEDGEMENTS HOEDEMAN , J. 1958. The frontal scalation pattern in some The first author wishes to thank Francesca Fontana, groups of tooth carps (Pisces: Cyprinodontiformes). Daniele Cavazzoni, and Ruud Wildekamp for their Bulletin of Aquatic Biology 1: 23-28. assistance with the first draft on the manuscript; HOELZMANN , P., KRUSE , H.-J. & ROTTINGER , F. 2000. Pre - cipitation estimates of the E-Saharan Palaeomonsoon: sed - Alain Dubois for suggestions regarding nomencla - imentary geochemistry; satellite image remote sensing and ture; Maurice Kottelat for suggesting the etymology; a regional water balance model of the West-Nubia-Lake. Dirk Neumann for help with references; C. E. Global and Planetary Change 26 : 105-120. Cherenkov for the “clipping path” of Ethiopian fish GOLUBTSOV , A. S., DARKOV , A. A., DGEBUADZE , Y U. Y U. & picture; and John Friel for information on N. virga - MINA , M. V. 1995. An artificial key to fish species of the tus in Ethiopia . We thank two anonymous reviewers Gambella region (the White Nile basin in the limits of for their their critical comments on the manuscript. Ethiopia). Joint Ethio-Russian Biological Expedition. We would also like to express our gratitude to Miguël Addis Ababa, Ethiopia, 84 pages. Parrent (MRAC); James Maclaine (BMNH); Patrice KARDASHEV , K. 2007 a. Sudan 2005-2006. AIK Notizie Killi 3: 5-9. Pruvost and Romain Causse (MNHN); Roger Bills, KARDASHEV , K. 2007 b. Catturare Notho nella Terra Dei Paul Skelton and Bernard MacKenzie (SAIAB) and Neri - Sudan 2005. AIK Notizie Killi 5: 6-14. Giuliano Doria (MSNG) for access to collections LEBLANC , M. J., LEDUC , C., STAGNITTI , F., VAN OEVELEN , under their care. P. J., JONES , C., MOFOR , L. A., RAZACK , M. & FAVREAU , G. 2006. Evidence for Megalake Chad, north-central REFERENCES Africa,during the late Quaternary from satellite data. AMIET , J. 1987. Faune du Cameroun. Fauna of Cameroon. 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A new species of Hemigrammus from the upper rio Negro basin, Brazil, with comments on the presence and arrangement of anal-fin hooks in Hemigrammus and related genera (Ostariophysi: Characiformes: Characidae)

Flávio C. T. Lima and Leandro M. Sousa

Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04299-970, São Paulo, SP, Brazil. E-mail: [email protected]; [email protected]

Received: 20 June 2009 – Accepted: 15 July 2009

Abstract H. pulcher , pela presença de duas machas umerais. Ela pode A new species of Hemigrammus is described from the rio ser distinguida de todas estas espécies por não possuir uma Tiquié, a tributary of the rio Uaupés, upper rio Negro basin, mancha escura no pedúnculo caudal (vs. mancha no pedún - Amazonas, Brazil. The new species can be distinguished culo caudal presente). É proposto um grupo monofilético from all congeners by the presence (in life specimens) of an dentro de Hemigrammus , englobando a espécie nova, junto orange blotch, situated between the two dark humeral a Hemigrammus ocellifer , H. neptunus , H. guyanensis , H. blotches, immediately ahead and slightly above the poste - luelingi , H. pulcher e H. haraldi , baseado na presença com - rior, darker second humeral blotch. Additionally, the new partilhada de derivado arranjo de ganchos na nadadeira anal species can be distinguished from all congeners, except H. de machos maduros e em algumas características de pig - haraldi, H. luelingi, H. neptunus, H. ocellifer, H. pretoensis, mentação. Uma discussão sobre a presença e arranjo de gan - and H. pulcher, by the possession of two humeral blotches. chos na nadadeira anal no gênero Hemigrammus e nos It can be distinguished from these species by lacking a blotch gêneros Parapristella e Petitella é apresentada, baseado num on the caudal peduncle (vs. caudal peduncle blotch present). extenso exame de espécimes pertencentes a esses gêneros. É A monophyletic group within Hemigrammus, the Hemi - hipotetizado que a associação entre ganchos na nadadeira grammus ocellifer species-group, encompassing the new anal e tecido denso presumivelmente secretório, encontrado species, plus Hemigrammus ocellifer, H. neptunus, H. guya - em todas as espécies examinadas de Hemigrammus , Parapri- nensis, H. luelingi, H. pulcher, and H. haraldi, is proposed, stella , Petitella e Hyphessobrycon , é um mecanismo para faci- based on the shared possession of a derived hook arrange - litar a ruptura das paredes celulares e provocar, em conse - ment in the anal-fin of mature males and some pigmentary qüência, a liberação das secreções celulares. features. A discussion of the hooks’ presence and arrange - ment within the genus Hemigrammus and in the related gen - Zusammenfassung era Parapristella and Petitella is presented, based on an exten - Beschrieben wird eine neue Hemigrammus- Art vom Tiquié- sive examination of specimens belonging to these genera. It Fluss, einem Nebenfluss des Uaupés im oberen Rio-Negro- is hypothesized that the close association between anal-fin Einzugsgebiet, Amazonas, Brasilien. Die neue Art lässt sich hooks and dense, presumably secretory tissue, found in all leicht von allen anderen Angehörigen der Gattung unter - Hemigrammus, Parapristella, Petitella, and Hyphessobrycon scheiden: Bei lebenden Exemplaren zeigt sich ein orangefar - species examined, is a mechanism to facilitate cell rupture, bener Fleck zwischen den beiden dunklen Schulterflecken, enhancing the setting free of cell secretions. direkt vor und etwas über dem hinteren, dunkleren zweiten Humeralfleck. Außerdem unterscheidet sich die neue Art Resumo von allen anderen in der Gattung – außer H. haraldi, H. Uma nova espécie de Hemigrammus é descrita da bacia do luelingi, H. neptunus, H. ocellifer, H. pretoensis und H. pulcher rio Tiquié, um afluente do rio Uaupés, sistema do alto rio – durch das Vorhandensein von zwei Schulterflecken. Ein Negro, Amazonas, Brasil. A nova espécie pode ser facilmente Unterscheidungsmerkmal von all den Arten ist das Fehlen diagnosticada de todas as congêneres pela presença, em vida, eines Schwanzflecks (bei den anderen befindet sich ein Fleck de uma mancha alaranjada, situada entre as duas manchas auf dem Schwanzstiel). Vorgeschlagen wird die Annahme umerais, imediatamente à frente e ligeiramente acima da einer monophyletischen Gruppe innerhalb der Gattung segunda mancha umeral. Além disso, a nova espécie pode ser Hemigrammus, wozu außer der neuen Art Hemigrammus diferenciada de todas suas congêneres, com exceção de H. ocellifer, H. neptunus, H. guyanensis, H. luelingi, H. pulcher haraldi , H. luelingi , H. neptunus , H. ocellifer , H. pretoensis e und H. haraldi zu zählen wären; Merkmale für die Einord -

153 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) nung wären die Gestaltung der Haken an der Afterflosse bei caudale (vs. presenza di chiazza su peduncolo caudale). Sul- den geschlechtsreifen Männchen sowie einige Besonder - la base di una comune disposizione degli uncini sulla pinna heiten im Farbmuster. Diskutiert werden abschließend anale dei maschi maturi e di alcune caratteristiche della co- Vorhandensein und Gestaltung der Flossenhaken bei Hemi - lorazione, si propone l’esistenza di gruppo monofiletico al- grammus und den verwandten Gattungen Parapristella und l’interno di Hemigrammus comprendente la nuova specie, Petitella auf der Grundlage einer genauen Untersuchung von Hemigrammus ocellifer, H. neptunus, H. guyanensis, H. Exemplaren dieser Gattungen. Der enge Zusammenhang luelingi, H. pulcher e H. haraldi. La presenza e la disposi- zwischen Analflossenhaken und dem dichten Gewebe mit zione degli uncini entro il genere Hemigrammus e nei ge- wahrscheinlich sekretorischen Aufgaben, wie er bei allen neri affini Parapristella e Petitella è discussa sulla base del - Arten der Gattungen Hemigrammus, Parapristella, Petitella l’esame di un ampio numero di esemplari appartenenti a und Hyphessobrycon anzutreffen ist, wird in einer Arbeitshy - questi generi. Si ipotizza che la stretta associazione tra gli pothese als Mechanismus erklärt, der das Aufreißen von uncini anali e il denso tessuto, presumibilmente a funzione Zellen erleichtert, wodurch Sekretionen der Zelle ermöglicht secretoria, presente in tutte le specie esaminate di Hemi - werden. grammus, Parapristella, Petitella e Hyphessobrycon, rappre - senti un meccanismo per facilitare la lisi cellulare, Résumé favorendo la liberazione dei secreti citosolici. Une nouvelle espèce d’ Hemigrammus est décrite, originaire du rio Tiquié, un tributaire du rio Uaupés, dans le bassin supérieur du rio Negro, Amazonie, Brésil. La nouvelle espèce INTRODUCTION est facile à distinguer de tous ses congénères par la présence, The genus Hemigrammus Gill, 1858 is one of the in vivo, d’une tache orange située entre les deux taches most speciose genera within the family Characidae, humérales sombres juste avant et légèrement au-dessus de la with 50 species currently recognized as valid (Lima seconde tache humérale postérieure et plus sombre. En outre, la nouvelle espèce peut être distinguée de tous ses congénères, et al. 2003, Lima et al. 2009). Similar to most gen - sauf H. haraldi, H. luelingi, H. neptunus, H.ocellifer, H. pre - era of Characidae previously assigned to the sub - toensis, et H. pulcher, par la présence de deux taches hu- family Tetragonopterinae (sensu Géry 1977), Hem - mérales. Elle peut être distinguée de toutes ces espèces par igrammus almost certainly does not constitute a l’absence de marque sur la tache caudale (contre la présence monophyletic assemblage. In fact, doubts regard - d’une marque sur le pédoncule caudal). Un groupe mono - ing its monophyletic nature, and specially its dis - phylétique à l’intérieur du genre Hemigrammus est proposé, tinction from Hyphessobrycon Durbin, 1908 were englobant la nouvelle espèce en plus d’ Hemigrammus ocellifer, first raised by Ellis (in Eigenmann 1918: 135) and H. neptunus, H. guyanensis, H. luelingi, H. pulcher et H. haraldi, sur base de la possession commune d’une forme reiterated by several subsequent authors, particu - dérivée de crochet sur l’anale des mâles adultes et de quelques larly Böhlke (1955: 233-234) and Weitzman & données pigmentaires. Une discussion sur la présence et la Fink (1983: 342). The alpha taxonomy of the configuration de crochets dans le genre Hemigrammus et les group is also still poorly known, and much revi - genres proches Parapristella et Petitella est présentée, basée sur sionary and descriptive work will be necessary until un examen exhaustif de spécimens appartenant à ces genres. a proper assessment of its diversity can be achieved. L’hypothèse proposée est que la relation étroite entre les cro - The rio Tiquié is the main tributary of the rio chets de l’anale et le tissu dense, probablement sécrété, trouvé Uaupés, which is one of the main tributaries of the chez toutes les espèces d’ Hemigrammus, Parapristella, Petitella et Hyphessobrycon examinées, est expliquée comme un méca- rio Negro, running 374 km from its sources at the nisme pour faciliter la rupture cellulaire, augmentant la Departamento Vaupés in Colombia to its mouth at libération de sécrétions cellulaires. the lower rio Uaupés in Brazil, 321 km being on Brazilian territory. From the year 2000 until pre - Sommario sent, the first author has been conducting an exten - Una nuova specie di Hemigrammus è descritta dal rio sive fish survey in this river basin, which resulted in Tiquié, un affluente del rio Uaupés, bacino superiore del the discovery of several fish species (Lima & rio Negro, Amazzonia, Brasile. La nuova specie può essere Toledo-Piza 2001, Vari & Lima 2003, Britto & facilmente diagnosticata da tutti i suoi congeneri per la Lima 2003, Zanata & Lima 2005, Ferreira & Lima presenza, negli esemplari vivi, di una chiazza arancione, si- 2006, Marinho & Lima 2009) and an inventory of tuata tra due macchie omerali scure, immediatamente da- the fish species occurring in the upper portion of vanti e leggermente sopra la seconda macchia omerale po- steriore più scura. Inoltre, la nuova specie può essere dia- the basin (Cabalzar et al. 2005). The new Hemi - gnosticata da tutti i congeneri, ad eccezione di H. haraldi, grammus species presented herein was collected in H. luelingi, H. neptunus, H. ocellifer, H. pretoensis e H. pul - the course of an ongoing survey of the fish diver - cher, per avere due macchie omerali. Si distingue da tutte sity in the middle and lower portions of the rio queste specie per l’assenza di una chiazza sulla macchia Tiquié basin. It was immediately recognized as new aqua vol. 15 no. 3 - 20 July 2009 154 due to its unique life color pattern. In addition, inae; Malabarba 1998: 211-212, and Malabarba & mature males of the new species possess an unusual Weitman 2000: 277, 279 for Cheirodontinae). arrangement of anal-fin hooks, characterized by The conclusion that anal-fin hook arrangements the presence of a single mid-sized hook per ray, and morphology in Hemigrammus constituted a extending from the last unbranched to the third to hardly exploited source of morphological informa - sixth branched anal-fin rays, surrounded by a dense tion, generally neglected in standard phylogeneti - pad of whitish tissue, which was discernible even in cal studies of characids due to the relative scarcity life specimens kept in aquaria. of mature material bearing these structures, Characid fishes generally possess bony projections prompted us to conduct a preliminary survey of in their fin rays, which are generally spiny and are the variability of this character. For practical rea - most commonly present in the anal, but also often sons, we restricted our study primarily to species of in the pelvic, and, more rarely, in the dorsal, pec - Hemigrammus , but we also examined species toral, and caudal fins (Malabarba & Weitzman belonging to a few other related genera (Petitella 2003). These projections develop on the surface of Géry & Boutière, 1964, Parapristella Géry, 1964, the lepidotrichia and are a dimorphic feature, pre - and a single species of Hyphessobrycon , H. dian - sent in mature males, with a few records of their cistrus). In fact, as it stands now, Hemigrammus is presence in females (Malabarba & Weitzman almost certainly a paraphyletic assemblage and 2003). Since apparently the majority of the there is no point in limiting the analysis merely to characid fishes possess the so-called fin hooks, the the species currently assigned to this genus. It is presence of these structures was hypothesized to our hope that this effort will stimulate other represent a synapomorphy of the family Characi - researchers to look further into the variation of this dae (Malabarba & Weitzman 2003). There are, structure, which, we believe, has a great potential however, records of similar structures on fin rays of in helping to understand the systematics of this the genus Nannocharax Günther, 1867, which complex and poorly known group. The aims of the belongs to the family Distichodontidae (Vari & present paper are thus twofold, to formally describe Ferraris Jr. 2004), and in some species of the genus a new species of Hemigrammus discovered in the Characidium Reinhardt, 1867, family Crenuchidae rio Tiquié, and to document the variation in anal- (Graça Jr. et al. 2008). fin hook arrangements in Hemigrammus and Examination of the literature revealed that a sim - related genera. ilar anal-fin hooks arrangement to the one found in the new species was previously reported for some MATERIAL AND METHODS of its congeners. Géry (1965: 19) noticed the pres - Counts and measurements were taken according ence of “1-2 fort crochets sur les premiers rayons to Fink & Weitzman (1974) and Menezes & ramifiés chez le mâle” in Hemigrammus ocellifer Weitzman (1990), except for counts of the hori - (Steindachner, 1882). The same species was zontal scale rows below the lateral line, which are reported by Weitzman (1977: 356-357) as possess - counted to the pelvic-fin insertion. Horizontal ing “one fairly large hook on each side of the ante - scale row counts between dorsal-fin origin and the rior five branched anal-fin rays”. A similar, though lateral line do not include scales of the median pre - not identical, condition was depicted by Géry dorsal series situated immediately anterior to first (1961: 50, figs. 5-6) for Hemigrammus pulcher pul - dorsal-fin ray. In the descriptions, the frequency of cher Ladiges, 1937 and H. pulcher haraldi Géry, each count is given in parentheses after the respec - 1961 (= Hemigrammus haraldi; Lima et al. 2003). tive count. An asterisk indicates counts of the holo - A single hook in the last unbranched and the ante - type. Counts of supraneurals, procurrent caudal- riormost branched anal-fin rays was also reported fin rays, branchiostegal rays, gill rakers, for H. guyanensis Géry, 1959 (Planquette et al. unbranched anal-fin rays, and dentary teeth and 1996: 276) and H. neptunus Zarske & Géry, 2002 their cusps were taken from cleared and stained (Zarske & Géry 2002: 28). paratypes (CS), prepared according to Taylor & Relatively few researchers have previously Van Dyke (1985). Vertebrae of the Weberian appa - explored the potential of anal-fin hook arrange - ratus were counted as four elements and included ments as a source of information on the systemat - in the vertebral counts, and the fused PU1+U1 of ics of any given characid group (for exceptions see the caudal region as a single element. Patterns of Weitzman & Fink 1985: 30-31 for Glandulocaud - scale circuli and radii were described for scales sam -

155 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) pled from the area between the lateral line and the ANSP, Academy of Natural Science of Philadelp - anterior origin of the dorsal fin. In the list of spec - hia, Philadelphia, USA; and MCP, Museu de Ciên - imens examined, the number of whole specimens cias e Tecnologia da Pontifícia Universidade Cató - in the lot is given first, followed by the number of lica do Rio Grande do Sul, Porto Alegre. those cleared and stained (if any). In the compara - tive material section, the number following the number of specimens in a lot refers to specimens Hemigrammus yinyang n. sp. bearing hooks on the anal fin, the size range refer - (Figs 1-3) ring only to those specimens. Institutional abbre - viations are: INPA, Instituto Nacional de Pesquisas Holotype: MZUSP 99300, 29.2 mm SL, sex unk - da Amazônia, Manaus; MZUSP, Museu de Zoolo - nown (presumably female), Brazil, Amazonas, iga - gia da Universidade de São Paulo, São Paulo; rapé Castanha, near Sítio São Pedro, 0°11’N

Fig. 1. Hemigrammus yinyang, holotype, MZUSP 99300, 29.2 mm SL, sex unknown (presumably female), immediately before preservation (above); after two years of preservation in alcohol (below); Brazil, Amazonas, igarapé Castanha, trib utary of rio Tiquié, near Sítio São Pedro. Photos by L. M. Sousa. aqua vol. 15 no. 3 - 20 July 2009 156 Table I. Morphometric characters of Hemigrammus yinyang. Data of the holotype (MZUSP 99300) and paratypes (MZUSP 93037, MZUSP 93053, MZUSP 92534, MZUSP 93250, ANSP 189317, INPA 31879, and MCP 43843).

n mean range SD holotype

Standard Length (mm) 36 - 16.8 - 29.2 - 29.2 Porcentages of SL Depth at dorsal-fin origin 36 29.8 26.1 - 36.4 1.8 36.4 Snout to dorsal-fin origin 36 53.5 49.5 - 55.7 1.2 55.5 Snout to pectoral-fin origin 36 26.9 25.3 - 28.6 0.8 26.3 Snout to pelvic-fin origin 36 45.1 42.9 - 47.9 1.1 47.9 Snout to anal-fin origin 36 59.1 56.3 - 62.9 1.5 61.2 Caudal-peduncle depth 36 8.8 7.6 - 10.2 0.6 10.2 Caudal peduncle length 36 13.1 11.7 - 14.9 0.9 13.2 Pectoral-fin length 36 21.5 19.3 - 24.4 1.3 19.3 Pelvic-fin length 36 16.5 14.7 - 18.8 0.9 15.5 Dorsal-fin base length 36 12.9 11.6 - 14.7 0.8 12.6 Dorsal-fin length 33 26.5 21.9 - 29.5 1.4 25.5 Anal-fin base length 36 31.0 26.8 - 33.7 1.5 31.3 Anal-fin lobe length 35 21.2 18.3 - 24.5 1.6 21.2 Eye to dorsal-fin origin 36 40.2 36.8 - 43.6 1.4 42.2 Dorsal-fin origin to caudal-fin base 36 50.0 47.3 - 54.3 1.4 50.6 Bony head length 36 25.9 23.8 - 28.6 0.9 24.8 Percentages of head length Horizontal eye diameter 36 39.4 35.2 - 43.0 1.7 39.0 Snout length 36 23.3 20.1 - 28.5 1.7 22.5 Least interorbital distance 36 38.3 34.5 - 41.2 1.6 38.5 Upper jaw length 36 44.5 41.0 - 47.4 1.3 44.8

69°35’W; F.C.T. Lima and others, 14-30 Novem - 15.4 mm SL, Igarapé Castanha, beach downstream ber 2006. from Santa Rosa village, 0°5’N 69°39’W, F. C. T. Paratypes: All from Brazil, Amazonas, rio Tiquié Lima and others, 3 September 2006. MZUSP basin. MZUSP 93053, 9, 16.8-18.8 mm SL, same 92572, 9, 10.0-12.9 mm SL, igarapé Castanha, data as holotype. MZUSP 101238, 1, 26.6 mm igapó (flooded forest) lake, near Santa Rosa village, SL, same data as holotype (reared in an aquarium 0°5’41”N 69°39’0”W, F. C. T. Lima and others, 3 for c. 6 months). MZUSP 93250, 28, 14.7-25.1 September 2006. mm SL, 3 CS, 16.6-23.3 mm SL; ANSP 189317, Diagnosis: Hemigrammus yinyang can be distin - 2, 16.0-24.2 mm SL; INPA 31879, 2, 17.5- guished from all congeners by the presence (in life 24.4 mm SL; igarapé Cunuri (or Macucu), near specimens) of an orange blotch, situated between São José II village, 0°13’N 69°36’W; F. C. T. Lima two dark humeral blotches, immediately ahead and and others, 16 November 2006. MZUSP 93037, slightly above the posterior, darker second humeral 7, 18.2-21.2 mm SL, stream at São José village, blotch. Additionally, Hemigrammus yinyang can be 0°13’N 69°36’W, F. C. T. Lima, 25 November distinguished from all congeners, except H. 2006. MZUSP 92292, 1, 15.8 mm SL, rio Tiquié, haraldi , H. luelingi , H. neptunus , H. ocellifer , H. between São José and Floresta villages, 0°13’N pretoensis , and H. pulcher , by the possession of two 69°36’W, F. C. T. Lima and others, 28 August – 1 humeral blotches. It can be distinguished from all September 2006. MZUSP 92534, 27, 12.7- these species by lacking a blotch on the caudal 25.3 mm SL; MCP 43843 , 2, 19.4-22.0 mm SL, peduncle (vs. caudal peduncle blotch present). igarapé Castanha, near the mouth, 0°12’ N Description: Morphometric data of holotype and 69°35’W, F. C. T. Lima, M. C. Lopes and others, paratypes presented in Table I. Body compressed, 30-31 August 2006. MZUSP 92927, 3, 17.1-21.2 moderately elongate; greatest body depth slightly mm SL , 1 CS, 17.8 mm SL, igarapé Castanha, anterior to dorsal fin origin; dorsal profile of head Sítio Belém, downstream from Santa Rosa village, convex from upper lip to vertical through anterior 0°5’23”N 69°39’57”W, F. C. T. Lima and others, nostril; straight to slightly concave from latter 28 November 2006. MZUSP 92591, 3, 13.1- point to tip of supraoccipital spine; predorsal pro -

157 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) file of body slightly convex, posteroventrally scales, with few slightly divergent radii extending inclined along dorsal-fin base; body profile straight to posterior margin of scales; lateral line slightly from dorsal-fin base terminus to adipose fin, decurved ventrally, incompletely pored, with 4 (4), slightly concave between latter point to origin of 5 (21), or 6* (10) perforated scales; lateral series anteriormost procurrent caudal-fin ray; ventral scales including perforated scales 29 (2), 30 (2), 31 profile of head and body convex to straight from (4), 32 (6), 33 (10), 34* (4), or 35 (7); scales from lower lip to anal-fin origin; body profile along four uppermost longitudinal scale rows consider - anal-fin base straight and posterodorsally inclined; ably larger than scales from three lowermost scale ventral profile of caudal peduncle slightly concave. rows, including lateral line scale row; horizontal Jaws equal, mouth terminal; posterior terminus of scale rows between dorsal-fin origin and lateral line maxilla reaching vertical through anterior third of 5; horizontal scale rows between lateral line and orbit; premaxillary teeth in two rows; outer row pelvic-fin insertion 3; scales along middorsal line with 2 (3), relatively compressed, unicuspid teeth; between distal tip of supraoccipital process and ori - inner row with 5 (1), 6 (1), or 7(1) relatively bulky, gin of dorsal fin 10 (8), 11 (21), or 12* (6); hori - tricuspid teeth; maxilla with 3 (1) or 4(3) tri- to zontal scale rows around caudal peduncle 9 (1), 10 unicuspid teeth; anteriormost tooth being the (10), 11* (16), or 12 (8); single row of 4-6 scales largest; dentary with 7 (1), 11 (1), 13(1), 14 (1) covering base of anteriormost anal-fin rays; caudal teeth; 4 anteriormost teeth larger, bulky, tricuspid, fin with few, small scales present on basal third of posterior ones unicuspid (Fig. 4). upper and lower caudal lobes. Scales cycloid, circuli absent in exposed areas of Dorsal-fin rays II,9; small ossification anterior to first unbranched ray, absent in the 4 CS specimens examined; distal margin of dorsal fin straight; dor - sal-fin origin at middle of standard length; base of posteriormost dorsal-fin ray slightly behind vertical through anal-fin origin; first dorsal-fin pterygio - phore inserting posterior to neural spine of tenth (4) vertebra; adipose fin present; anal-fin rays III (1) or IV (3), 19 (1), 21* (16), 22 (14), or 23 (3); last unbranched and 4 anterior anal-fin rays longest, remaining rays progressively shorter towards anal- fin end; anteriormost anal-fin pterygiophore insert - ing behind haemal spine of fifteenth (4) vertebra; pectoral-fin rays I,10* (2), 11 (12), 12 (19), or 13 (3); tip of pectoral fin reaching slightly beyond ver - tical through pelvic-fin insertion; pelvic-fin rays I,7; caudal fin forked, lobes slightly rounded, similar to each other in size; principal caudal-fin rays 10+9 (3); 8 (3) or 9 (1) dorsal procurrent caudal-fin rays, and 6 (1) or 7 (3) ventral procurrent caudal-fin rays; first gill arch with 0 (2), 1 (1), or 2 (1) hypo - branchial, 7 (3) or 8 (1) ceratobranchial, and 5 (1) or 6 (3) epibranchial gill-rakers; vertebrae 33 (1) or 34 (3); supraneurals 5 (4); branchiostegal rays 4 (4): 3 originating from anterior ceratohyal and one from posterior ceratohyal. Color in alcohol: Ground color clear; snout, tip of dentary, anterior portion of maxilla and top of head densely covered by small dark Fig. 2. Hemigrammus yinyang, male (above) and female chromatophores, resulting in overall dark pigmen - (below); specimens in aquarium, about 11 months after tation; infraorbitals with scattered dark chro - capture, not preserved; both collected with holotype. matophores; opercle with a dense concentration of Photos by L. M. Sousa. relatively large dark chromatophores, forming a aqua vol. 15 no. 3 - 20 July 2009 158 dark blotch; opercle translucent due to the loss of Sexual dimorphism: Bony hooks were detected guanine pigmentation. on anal fins of 14 specimens, ranging from 18.4 to Scales on middorsal area with dense concentra - 24.9 mm SL, collected during the months of tion of dark chromatophores; 2 humeral blotches August and November. One dissected specimen present; anterior humeral blotch large, relatively (MZUSP 92534, 23.9 mm SL) proved to be a faint, formed by relatively large dark chro - male. Another large dissected specimen lacking matophores, transversed at its center by the lateral hooks (MZUSP 92534, 25.3 mm SL) was a female line, about 4 scales long and 2 scales high; second with well-developed oocytes. Hooks are present as humeral blotch formed by relatively large, densely a single bilateral pair per anal-fin ray, from the last concentrated dark chromatophores, resulting in an unbranched a ray to the third to sixth branched overall intensely dark pigmented blotch, at the rays. The hooks are arranged in a row, positioned level of the seventh to eight longitudinal scales; lat - immediately above the branching point in eral surfaces of body above midlateral line with branched fin rays (Figs 5-6). A dense, apparently dark chromatophores mostly concentrated on glandular tissue is surrounds each hook (Fig. 5). scales margins, outlining scales; dark chro - Females also apparently grow larger than males, the matophores below midlateral line arranged along largest female recorded reaching 29.2 mm SL margins of epaxial muscles bundles from area (MZUSP 99300, holotype), whereas the largest above anal fin to caudal peduncle; a relatively male reached 24.9 mm SL (MZUSP 93250). broad, faint longitudinal stripe formed by scattered Geographical distribution: Hemigrammus yiny- large dark chromatophores, along the midlateral ang is only known from tributaries of the middle line, extending into the caudal peduncle; all fins rio Tiquié, a tributary of the rio Uaupés, upper rio with scattered, dark chromatophores, more numer - Negro basin, Amazonas state, Brazil (Fig. 7). ous on dorsal, caudal, and anal-fins. Ecological notes: Hemigrammus yinyang was col - Color in life: Based on the examination of lected in moderately large (2-6 m wide) streams, several freshly caught specimens, some of which tributaries of rio Tiquié and including igarapé Cas - were reared in aquaria for additional observations tanha, the largest tributary of rio Tiquié at its cen - and photos: ground color clear, mostly translucent, tral course. These streams, like igarapé Cunuri (Fig. except facial bones, abdominal area, and area along 8) are clear-water, carrying little suspended sedi - midlateral line, which are silvery, with a bright ment, in contrast with the dark-colored waters of greenish tinge; dorsal portion of eye red; orange the rio Tiquié or the muddy, silt-laden waters of blotch located between 2 dark humeral blotches, the igarapé Castanha, where only few specimens of immediately ahead and slightly above the posterior, Hemigrammus yinyang were collected. A few juve - darker second humeral blotch; dorsal, anal, and nile specimens were collected in a small igapó caudal fins yellowish; anterior portion of dorsal, (flooded forest) lake marginal to igarapé Castanha. pelvic, and anal fins whitish (Figs 1-2). Etymology: The dual concept of yin and yang

Fig. 3. Hemigrammus yinyang, paratype, MZUSP 93037, 21.2 mm SL, sex unknown (presumably female); Brazil, Amazonas, igarapé Cunuri, tributary to the rio Tiquié, near São José II village. Photo by L. M. Sousa.

159 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) derives from the ancient Taoistic Chinese philoso - Zarske & Géry, 2007 (Zarske & Géry 2007: 8), H. phy and religion, describing two primal opposing guyanensis (Planquette et al. 1996: 276), H. haraldi but complementary principles said to be found in (as H. pulcher haraldi; Géry 1961: 50, fig. 6), H. all non-static objects and processes in the universe. mahnerti Uj & Géry, 1989 (Uj & Géry 1989: Yin , in Chinese, originally meant “sunless”, as the 156), H. mimus Böhlke, 1955 (Taphorn 1992: northern side of a mountain, and as a concept 218, Zarske & Géry 2007: 13), H. neptunus evolved to embody the dark, passive, feminine ele - (Zarske & Géry 2002: 28), H. ocellifer (Géry 1965: ment, corresponding to the night, the winter, the 19, Weitzman 1977: 356), H. pulcher (as H. pul - water, and the earth. Yang originally meant cher pulcher ; Géry 1961: 50, fig. 5), H. rhodostomus “sunny”, as the southern side of a mountain, and it Ahl, 1924 (Géry & Boutière 1964: 478, Géry & came to embody the bright, active, masculine ele - Mahnert 1986: 45), H. schmardae (Steindachner, ment, corresponding to the day, the summer, the 1882) (as H. proneki Géry, 1963: Géry, 1963: 14, air and the fire. All forces in nature are expressions Géry 1965: 19), H. taphorni Benine & Lopes, of yin and yang states. It is in their complementar - 2007 (Benine & Lopes 2007: 56), H. unilineatus ity that balance is given to the universe. The new (Gill, 1858) (Géry 1966: 112), and H. vorderwin - species is named in allusion to its complementary kleri Géry, 1963 (Géry 1963: 11). In addition, orange and black humeral blotches, which are rem - Eigenmann (1918: 152) mentions that males of H. iniscent of the Taiji diagram, the pictorial repre - hyanuary possess an “anal armature”, which is pre - sentation of the state of undifferentiated absolute, sumably a reference of the occurrence of anal-fin encompassing both the yin and yang qualities hooks in this species. However, we were unable to [from Wikipedia and other sources]. confirm the occurrence of fin hooks in specimens Discussion: Several species of Hemigrammus are of this species examined in this study. known to possess anal-fin hooks (Malabarba & On other hand, some recently described species of Weitzman 2003). More specifically, fin hooks were Hemigrammus were reported as lacking fin hooks, reported for Hemigrammus bleheri Géry & Mahn - e.g. H. skolioplatos Bertaco & Carvalho (2005: ert, 1986 (Géry & Mahnert 1986: 45), H. geisleri 145), H. ora Zarske et al. (2006: 21), H. parana Marinho et al. (2008: 56), and H. silimoni Britski & Lima (2008: 568). However, H. ora , H. skolio - platus , and H. silimoni were described based on few specimens, the latter two species collected during the meridional dry season (July and September, respectively). Hence it is uncertain whether these structures are truly absent in those species. As for H. parana , a sizable portion of the type-series was collected during the meridional rainy season (Jan - uary) and it seems plausible that the lack of hooks

Fig. 4. Hemigrammus yinyang, paratype, MZUSP 93250, Fig. 5. Hemigrammus yinyang , paratype, MZUSP 93250, 23.3 mm SL. Upper and lower jaws, lateral view, left side. 23.3 mm SL. Close-up of anal fin showing fin hooks and Scale bar 1 mm. surrounding dense whitish tissue. Photo by E. G. Baena. aqua vol. 15 no. 3 - 20 July 2009 160 noticed may indeed indicate the absence of those in the rio Amazonas/Solimões near Manaus. structures in that species. Though it has been suggested that fin hooks, once We conducted an extensive survey of the Hemi - developed, are permanent fin structures grammus material deposited in the MZUSP collec - (Gonçalves et al. 2005), we think that the question tion for specimens bearing hooks on fins. During is still not yet entirely settled. For example, the this search, we focused on examining samples col - large-sized characids Salminus and Brycon clearly lected during the meridional rainy season, i.e. from only develop fin hooks during the breeding period November to March, since in a preliminary search (F.C.T. Lima, unpublished data). Our observations for specimens bearing hooks, we noticed that sam - on fin hook presence in Hemigrammus and related ples collected during the dry season (April to Sep - genera support a view earlier expressed by Ferreira tember) generally did not yield specimens with & Lima (2006), who stated that either the presence hooks. This even holds true for samples collected or absence of hooks in characids can only be in near-equatorial latitudes as, for example, the assessed with some confidence when, for a given Central Amazon basin, i.e. the lower rio Negro and species, numerically adequate samples are available,

Fig. 6. Anal-fin and hooks in selected Hemigrammus species and related genera. Hemigrammus schmardae, MZUSP 85693, 24.0 mm SL; Parapristella georgiae, MZUSP 95250, 30.9 mm SL; Hemigrammus analis, MZUSP 74763, 41.0 mm SL; Hemigrammus stictus, MZUSP 95269, 31.7 mm SL; Hemigrammus ocellifer, MZUSP 85608, 31.2 mm SL; Hemigrammus yinyang, MZUSP 93250, 23.3 mm SL; Hemigrammus haraldi, MZUSP 18734, 22.5 mm SL.

161 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) collected during various seasons of the year. Anal erature search, we noticed a considerable variation and pelvic fin hooks were found in specimens of in the number, size and arrangement of anal-fin the following Hemigrammus species: H. analis hooks in Hemigrammus and some related genera. Durbin, 1909, H. barrigonae Eigenmann & Henn, Most species possess numerous hooks in the first 1914 , H. bellottii (Steindachner, 1882), H. bleheri , branched and anteriormost branched anal-fin rays. H. boesemani Géry, 1959, H. haraldi , H. levis This condition is found among species examined Durbin, 1908, H. luelingi Géry, 1964, H. lunatus of Hemigrammus analis (Fig. 6), H. barrigonae, H. Durbin, 1918, H. ocellifer , H. pretoensis Géry, bellottii , H. levis , H. lunatus , H. pretoensis , H. 1965, H. rhodostomus , H. schmardae , H. stictus schmardae (Fig. 6), H. vorderwinkleri and Parapris - (Durbin, 1909), and H. vorderwinkleri . On the tella georgiae (Fig. 6), being apparently also the other hand, no fin hooks were found in any speci - condition found in H. geisleri (Zarske & Géry men examined of H. coeruleus Durbin, 1908, H. 2007: 8), H. mahnerti (Uj & Géry 1989: 156), H. marginatus Ellis, 1911, and H. ulreyi (Boulenger, mimus (Taphorn 1992: 218, Zarske & Géry 2008: 1895), even though for each of these species hun - 13), H. taphorni (Benine & Lopes 2007: 56), and dreds of large-sized, apparently mature specimens H. unilineatus (Géry 1966: 112). There is, how - collected during all seasons were checked for those ever, a considerable variation in the number, size structures (see Comparative material examined). and relative position of hooks in the species pos - Apparently, mature males belonging to these sessing several anal-fin hooks. In these species species never develop fin hooks. hooks range from tiny (Hemigrammus bellottii) to Based on the examination of specimens and a lit - large-sized (H. analis , H. levis , and Parapristella

Fig. 7. Map of northern South America showing collecting sites of Hemigrammus yinyang. The open circle represents the type-locality. aqua vol. 15 no. 3 - 20 July 2009 162 georgiae Géry, 1964). Some species possess both to generally possess more than one hook per anal- small- and medium-sized hooks (e.g., Hemigram - fin ray). The second condition, found in H. stictus mus lunatus ). We have not attempted to identify (Figs 6, 9), is the presence of a single mid-sized the various potential distinct conditions found hook per ray, situated in the central portion of the within species having numerous hooks in the anal anteriormost 5 to 8 branched anal-fin rays, fol - fin because we feel that a broader analysis of hook lowed by several tiny hooks along the posterior presence and arrangement in Hemigrammus and branch of the anal-fin rays. The third condition, related genera is necessary to provide a classifica - displayed by H. boesemani and apparently identical tion. to the condition found in Hyphessobrycon otrynus It is, however, possible to distinguish five distinct Benine & Lopes (2008, fig. 4), is the presence of conditions of hook arrangement among species of two large-sized hooks, in the central portion of the Hemigrammus and some belonging to related gen - last unbranched and the first branched anal-fin era possessing a small number of anal-fin hooks. rays, followed by tiny hooks along the posterior The first condition, found in Hemigrammus ble - margin of the unbranched and subsequent heri , H. rhodostomus , and in the related Petitella branched anal-fin rays. A fourth condition, dis - georgiae Géry & Boutière, 1964, is the presence of played by Hemigrammus yinyang , H. ocellifer (Fig. relatively few (1-3 per ray) medium-sized hooks 6), H. neptunus (Zarske & Géry 2002: 28), and H. per ray located in the central portion of the anteri - guyanensis (Planquette et al. 1996: 276), is the ormost 6 to 8 branched anal-fin rays (contrary to presence of a single medium-sized hook per anal- Géry & Mahnert 1986, we found H. rhodostomus fin ray, extending from the last unbranched to the sixth (H. yinyang) or seventh ( H. ocellifer ) branched anal-fin rays, generally located at the ray’s branch - ing point or immediately above it. Contrary to Géry (1965: 19) and Weitzman (1977: 356), who mention the presence of hooks only in the branched fin rays of H. ocellifer , all specimens of this species examined in the present study possess a well-developed hook in the last unbranched anal- fin ray. Lastly, a fifth condition, found in Hemi - grammus luelingi , H. pulcher and H. haraldi (Fig. 8) is the presence of a single, large-sized hook per ray extending from the last unbranched to the second (H. haraldi) or third (H. luelingi and H. pulcher) branched anal-fin rays. The presence of a single, medium-sized hook per anal-fin ray, arranged in a row along the anterior - most anal-fin rays, as displayed in H. yinyang , H. ocellifer , H. neptunus , and H. guyanensis is clearly a derived feature within Hemigrammus . Apparently, a similar condition among characids is found only in the distantly related genus Mimagoniates , family Glandulocaudinae (Menezes & Weitzman 1990, Menezes & Weitzman in press). Hemigrammus luelingi , H. pulcher , and H. haraldi possess a condi - tion very similar to the one shared by H. yinyang , H. ocellifer , H. neptunus , and H. guyanensis , differ - ing only by the fact that in the former species the hooks are further reduced in number (i.e. restricted to the last unbranched up to the third branched Fig. 8. Igarapé Cunuri, a tributary of rio Tiquié, where part anal-fin ray) and are considerably larger than those of the type series of Hemigrammus yinyang was collected, at present in the latter species. This condition is very low water. Photo by F. C. T. Lima. likely a further specialization in the hook develop -

163 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) ment found in H. yinyang , H. ocellifer , H. nep - hypothesize that H. guyanensis , H. haraldi , H. nep - tunus , and H. guyanensis , and is therefore inter - tunus , H. ocellifer , H. luelingi , H. pulcher , and H. preted herein as being a homologous character yinyang constitute a monophyletic entity within shared by these species. All species in question the genus Hemigrammus , for which we suggest the share a color pattern consisting in two well-devel - informal name “ Hemigrammus ocellifer species- oped humeral blotches and a conspicuous caudal group”, referring to the best-known and most peduncle blotch (the latter being absent in H. widespread species within the group. This putative yinyang), a condition, which within Hemigrammus monophyletic group is obviously more restricted is only shared with H. pretoensis , a species which than the homonymous group proposed by Géry possesses numerous relatively small hooks on the (1977: 502-507) as an “artificial group” including anal-fin rays (see above). In addition, H. guyanen - Hemigrammus species possessing one or two sis , H. haraldi , H. neptunus , H. ocellifer , H. luelingi , humeral blotches and a caudal peduncle blotch. H. pulcher , and H. yinyang all share another pig - The conditions displayed by H. bleheri , H. rho - mentary feature, the upper margin of the eye being dostomus , and Petitella georgiae , on one hand, and red in life. A red upper eye margin is, however, also H. stictus , on the other hand, are similar to the one present in at least a few more congeners (e.g. H. present in species belonging to the H. ocellifer analis). Though not unique, these pigmentary fea - species-group, but the former species generally lack tures, combined with the derived anal-fin hook hooks in the last unbranched anal-fin ray (a few arrangement found in these species, allow us to specimens of H. stictus , however, bear a small to medium-sized hook in that ray). The reduction in the number and size of the anal-fin hooks dis - played by these species may indicate a close rela - tionship with the species belonging to the H. ocel - lifer species-group. Further investigations on the systematics of the group are necessary to properly address this issue. The similarity in anal-fin hook morphology and arrangement in Hemigrammus rhodostomus , H. bleheri and Petitella georgiae , in addition to the fact that these species share a color pattern unique among characids, which is com - posed of barred anal- and caudal-fins and a red head in life specimens, suggests that these species form a monophyletic assemblage. Petitella georgiae , however, possesses a teeth, dentary and premaxil - lary morphology, which is very distinct from H. rhodostomus and H. bleheri , the reason for its assignment to a distinct genus (Géry & Boutière 1964, Géry & Mahnert 1986). Although these dif - ferences may simply constitute autapomorphic fea - tures of Petitella georgiae , given the very likely para - phyletic nature of Hemigrammus , a synonymiza - tion of Petitella with Hemigrammus or, conversely, the transferal of H. rhodostomus and H. bleheri into Petitella is considered premature. All characid specimens examined in the present study, which have anal-fin hooks (belonging to the genera Hemigrammus , Parapristella , Petitella , and Hyphessobrycon) possess a variably developed dense, Fig. 9. Close-up of anal fin of Hemigrammus haraldi, gelatinous whitish tissue on their fin membranes, MZUSP 18734, 22.5 mm SL, and Hemigrammus stictus, bordering or in some cases surrounding the hooks. MZUSP 95269, 31.7 mm SL, showing fin hooks and sur - The close association between hooks and this rounding dense whitish tissue. Photos by E. G. Baena. whitish tissue is more obvious in species possessing aqua vol. 15 no. 3 - 20 July 2009 164 few, medium-sized to large hooks, as for example release of cell secretions is an assumption that mer - H. haraldi and H. stictus (Fig. 9), or Hyphessobrycon its further investigation. diancistrus Weitzman, 1977 (Weitzman 1977, fig. Comparative material examined: Hemigrammus 2). The feature can even be discerned in as whitish analis: MZUSP 74463, 112, 34, 23.8-38.8 mm spots situated at the mid-length of the anterior - SL, Brazil, Amazonas, rio Negro drainage. MZUSP most anal-fin rays in life specimens (e.g., H. 100150, 1, 23.3 mm SL, Brazil, Pará, rio Amazo - yinyang , Fig. 2, upper photograph, and H. ocellifer , nas drainage. MZUSP 81373, 36, 13, 27.2- Géry 1977: 506, specimen to the right). In these 32.7 mm SL, Brazil, Amazonas, rio Tiquié. species there is a thick mass of tissue surrounding MZUSP 95272, 216, 45, 22.2-34.8 mm SL, Bra - the hooks, which is more developed in species hav - zil, Amazonas, rio Negro drainage. MZUSP ing larger hooks. Weitzman et al. (2005: 350) 85667, 139, 29, 31.6-34.6 mm SL, Brazil, Amazo - analysed histologically the dense tissue surround - nas, Rio Preto da Eva. MZUSP 74761, 135, 21, ing the large anal-fin hooks in Hyphessobrycon dian - 28.6-39.6 mm SL; 2, 34.3-36.6 mm SL, CS, Bra - cistrus and found it to possess a great number of zil, Amazonas, rio Negro drainage. MZUSP club cells. In ostariophysans club cells are linked to 92302, 395, 5, 32.7-37.3 mm SL, Brazil, Amazo - the production and release of alarm substances nas, rio Tiquié. Hemigrammus barrigonae: MZUSP (Pfeiffer 1977), and, in some cases, possibly 85106, 17, 13, 31.7-40.4 mm SL; 1, 34.3 mm SL, pheromones (Weitzman et al. 2005). We have not CS, Brazil, Amazonas, rio Tiquié. MZUSP 85186, performed a histological analysis of the tissues sur - 1, 33.4 mm SL, Brazil, Amazonas, rio Tiquié. rounding or bordering anal-fin hooks in the species MZUSP 85018, 25, 1, 29.6 mm SL, Colombia, of Hemigrammus , Parapristella , and Petitella exam - Depto. Vaupés, rio Tiquié. Hemigrammus bellottii : ined in this study, but it seems very likely that this MZUSP 85729, 303, 3, 22.7-25.0 mm SL, Brazil, tissue is homologous with the one found in Amazonas, rio Urubu. MZUSP 84987, 138, 62, Hyphessobrycon diancistrus , and, as such, is formed 16.1-25.9 mm SL; 5, 22.1-25.4 mm SL, CS, Bra - by club cells. It is interesting to remark that a sim - sil, Amazonas, rio Tiquié. MZUSP 85056, 109, ilar concentration of tissue surrounding anal-fin 42, 20.2-27.3 mm SL, Brazil, Amazonas, rio hooks, found in a member of the family Stervardi - Tiquié. Hemigrammus bleheri: MZUSP 29435, inae, Diapoma speculiferum Cope, 1894, is com - 6701, 6, 24.8-27.0 mm SL, CS, Brazil, Amazonas, posed exclusively of club cells (K. M. Ferreira, pers. rio Negro. Hemigrammus boesemani: MZUSP comm.). The close association between a presum - 76579, 61, 2, 27.2-29.6 mm SL, Surinam, Suri - ably secretory tissue and the anal-fin hooks, as well nam River drainage. Hemigrammus coeruleus: as the correlation between hook size and tissue MZUSP 7459, 101 of 414, 28.1-52.4 mm SL, development, calls for a causal explanation. Club Brazil, Amazonas, rio Solimões drainage. MZUSP cells can only release their contents when the skin 7333, 54 of 280, 27.1-39.2 mm SL, Brazil, Ama - is damaged (Pfeiffer 1977, Kristensen & Closs zonas, rio Solimões drainage. MZUSP 75473, 91 2004). We suggest that the close association of of 550, 32.4-45.9 mm SL, Brazil, Amazonas, rio secretory tissues and hooks in the anal fin of Negro drainage. Hemigrammus haraldi: MZUSP mature characid males is a mechanism that causes 17394, 1, 23.4 mm SL, Brazil, Amazonas, rio Soli - friction and cell rupture and consequently mões. MZUSP 26326, 2, 1, 21.2 mm SL, Peru, enhances the release of the cell secretions. This Loreto, Río Amazonas drainage. MZUSP 26432, hypothesis, however, does not account for the pres - 3, 1, 21.2 mm SL, Peru, Loreto, Río Ucayali basin. ence of hooks in pelvic fins and other fins of MZUSP 88700, 17, 1, 22.0 mm SL, Brazil, Ama - characids where these structures also appear, nor zonas, rio Solimões basin. MZUSP 18734, 25, 5, for species that possess well-developed pads of tis - 21.2-25.6 mm SL; 1, 23.3 mm SL, CS, Brazil, sue formed mainly by club cells over the anal fin Amazonas, rio Solimões drainage. Hemigrammus but which do not have associated hooks, as is the hyanuary: MZUSP 63215, 8 of 13, 20.2-26.8 mm case in Bryconadenos tanaothoros Weitzman, SL; MZUSP 63191, 3, 19.2-25.6 mm SL; Menezes, Evers & Burns, 2005 (Weitzman et al. MZUSP 63218, 1, 24.6 mm SL, Brazil, Amazo - 2005: 340, 354-355, figs. 5, 7). Fin hooks have nas, rio Japurá basin. MZUSP 92114, 109 of 129, been traditionally interpreted as serving as contact 22.4-31.7 mm SL, Brazil, Amazonas, rio Tiquié. organs used during spawning in characids (e.g. Hemigrammus levis: MZUSP 100612, 24, 7, 29.8- Wiley & Collette 1970). Their function in the 33.9 mm SL; 1, 31.8 mm SL, CS, Brazil, Amazo -

165 aqua vol. 15 no. 3 - 20 July 2009 A new species of Hemigrammus from the upper rio Negro basin, Brazil, (Ostariophysi: Characiformes: Characidae) nas, Rio Solimões. MZUSP 96411, 18, 8, 30.3- Solimões drainage. MZUSP 5450, 1, 31.1 mm SL, 35.0 mm SL, Venezuela, Estado Bolívar, Río Ori - Brazil, Pará, rio Trombetas drainage. MZUSP noco drainage. Hemigrammus luelingi: MZUSP 75471, 42, 6, 30.9-36.2 mm SL, Brazil, Amazonas, 85587, 2, 1, 28.6 mm SL, Peru, Depto. Loreto, rio Negro drainage. MZUSP 75470, 16, 6, 32.6- Río Ucayali basin. Hemigrammus lunatus: MZUSP 37.6 mm SL, Brazil, Amazonas, rio Negro drai - 54028, 55, 17, 26.8-33.1 mm SL; 1, 30.8 mm SL, nage. MZUSP 74762, 31, 19, 31.2-34.6 mm SL, CS, Paraguay, Depto. Concepcion, Río Apa. Brazil, Amazonas, rio Negro drainage. MZUSP MZUSP 54027, 21, 7, 30.9-33.5 mm SL; Para - 95269, 83, 42, 29.5-33.9 mm SL; 2, 30.8- guay, Depto. Concepcion, Río Apa. Hemigrammus 31.6 mm SL, CS, Brazil, Amazonas, rio Negro marginatus: MZUSP 17088, 37 of 77, 32.2- drainage. MZUSP 96535, 1, 30.1 mm SL, Vene - 41.5 mm SL, Brazil, Minas Gerais, rio São Fran - zuela, Estado Bolívar, Río Orinoco drainage. cisco. MZUSP 57523, 212 of 1073, 19.6- Hemigrammus ulreyi: MZUSP 96663, 35, 18.1- 33.4 mm SL, Brazil, Bahia, rio de Contas. MZUSP 31.8 mm SL, Brazil, Mato Grosso, rio Cuiabá 38039, 107 of 552, 21.2-30.8 mm SL; Brazil, basin. MZUSP 19082, 2, 26.8-30.8 mm SL, Bra - Minas Gerais, rio São Francisco. Hemigrammus zil, Mato Grosso, rio Paraguai drainage. MZUSP ocellifer: MZUSP 85608, 26; 12, 29.1-33.4 mm 44389, 12, 28.1-33.0 mm SL, Brazil, Mato SL; Peru, Depto. Loreto, Río Ucayali basin. Grosso, rio Paraguai basin. MZUSP 96696, 23, MZUSP 17409, 84; 10, 26.3-31.5; 2, 29.4- 21.1-32.8 mm SL; Brazil, Mato Grosso, rio 29.7 mm SL, Brazil, Amazonas, rio Solimões basin. Piquiri. MZUSP 59538, 60 of 206, 23.5-35.8 mm MZUSP 77891, 26; 8, 29.9-36.5 mm SL, Brazil, SL, Brazil, Mato Grosso do Sul, rio Negro. Hemi - Amazonas, rio Japurá basin. MZUSP 85399, 3, grammus vorderwinkleri: MZUSP 18684, 60, 10, 28.0-29.5 mm SL, Brazil, Rondônia, rio Machado. 20.8-24.3 mm SL; 1, 23.1 mm SL, CS, Brazil, Hemigrammus pretoensis: MZUSP 17627, 6, 2, Mato Grosso, rio Guaporé. MZUSP 16950, 3, 2, 45.7-50.0 mm SL, Brazil, Amazonas, rio Solimões. 26.9-29.3 mm SL, Brazil, Amazonas, rio Negro. Hemigrammus pulcher : MZUSP uncat., 3, 1, MZUSP 25547, 1, 22.2 mm SL, Brazil, Pará, rio 30.5 mm SL, Peru, Depto. Loreto, Río Amazonas Tapajós. MZUSP 96551, 15, 8, 26.8-28.0 mm SL, basin. MZUSP uncat., 2, 1, 26.6 mm SL, Peru, Venezuela, Depto. Bolívar, Río Orinoco. Hyphesso - Depto. Loreto, Río Amazonas basin. Hemigram - brycon diancistrus: MZUSP 13179, paratypes, 2, 1, mus rhodostomus: MZUSP 17960, 21, 2, 26.2-28.7 25.9 mm SL, Colombia, Vichada, Río Vichada. mm SL; Brazil, Pará, rio Capim. MZUSP 17997, MZUSP 17682, 3, 1, 33.5 mm SL, Brazil, Amazo - 152, 5, 26.4-26.8 mm SL; 1, 26.7 mm SL, CS, nas, rio Negro basin. MZUSP 96516, 16, 7, 25.4- Brazil, Pará, rio Capim. MZUSP 18632, 36, 3, 26.1 mm SL, Venezuela, Bolívar, Río Parguaza; 26.3-26.7 mm SL, Brazil, Pará. Hemigrammus MZUSP 96401, 11, 7, 26.5-29.1 mm SL, Vene - schmardae: MZUSP 52654, 5, 3, 22.2-23.2 mm zuela, Bolívar, Río Orinoco drainage. MZUSP SL, Brasil, Pará, rio Trombetas. MZUSP 85743, 1, 15744, 1, 29.9 mm SL, Brazil, Pará, rio Trombetas 23.6 mm SL, Brazil, Amazonas, rio Preto da Eva. drainage. MZUSP 29843, 147, 5, 31.0-33.3 mm MZUSP 85692, 158, 18, 22.1-27.6 mm SL; 1, SL, Brazil, Amazonas, rio Negro drainage. MZUSP 23.9 mm SL, CS, Brazil, Amazonas, rio Preto da 29846, 80, 26, 27.3-32.4 mm SL; 2, 28.1- Eva. MZUSP 77885, 364, 12, 21.2-24.1 mm SL, 28.9 mm SL, CS, Brazil, Amazonas, rio Negro Brazil, Amazonas, rio Negro. MZUSP 95271, 50, drainage. MZUSP 29847, 4, 2, 30.9-31.8 mm SL; 21, 18.4-28.0 mm SL, 1, 24.0 mm SL, CS; Brazil, 1, 30.9 mm SL, CS, Brazil, Amazonas, rio Negro Amazonas, rio Cuieiras. MZUSP 58621, 126, 8, drainage. Parapristella georgiae: MZUSP 95250, 21.3-24.2 mm SL, Brazil, Amazonas, rio Negro. 49, 18, 28.3-33.8 mm SL; 2, 29.8-31.6 mm SL, MZUSP 85693, 35, 14, 21.-26.1 mm SL, Brazil, CS, Brazil, Amazonas, rio Negro drainage. Amazonas, rio Preto da Eva. MZUSP 64699, 143, MZUSP 95247, 43, 20, 33.8-41.2 mm SL, Brazil, 8, 25.8-26.3 mm SL, Brazil, Amazonas, rio Tiquié. Amazonas, rio Negro drainage. Petitella georgiae: MZUSP 81169, 3, 20.6-22.9 mm SL; Brazil, MZUSP 77269, 398, 32, 28.7-31.4 mm SL; 5, Amazonas, rio Tiquié. MZUSP 85695, 5, 4, 20.5- 28.1-31.4 mm SL, CS, Brazil, Amazonas, rio Soli - 24.8 mm SL, Brazil, Amazonas, rio Preto da Eva. mões. MZUSP 85696, 3, 1, 23.3 mm SL, Brazil, Amazo - nas, rio Urubu. Hemigrammus stictus: MZUSP 7430, 2, 30.0-30.4 mm SL, Brazil, Amazonas, rio aqua vol. 15 no. 3 - 20 July 2009 166 ACKNOWLEDGEMENTS FERREIRA , K. M. & L IMA , F. C. T. 2006. A new species of All specimens of Hemigrammus yinyang were col - Knodus (Characiformes: Characidae) from the Rio lected during expeditions of the project “Peixes e Tiquié, Upper Rio Negro system, Brazil. Copeia 2006 pesca no rio Tiquié”, a joint programme of Insti - (4): 630-639. FINK , W. L. & W EITZMAN , S. H. 1974. The so-called tuto Sociambiental (ISA) and Federação das Orga - cheirodontin fishes of Central America with description nizações Indígenas do Alto Rio Negro (FOIRN). of two new species (Pisces, Characidae). Smithsonian We are grateful to A. Cabalzar and M. C. Lopes for Contributions to Zoology 172 : 1-46. their assistance during fieldwork. Special thanks GÉRY , J. 1961. Hemigrammus pulcher haraldi a new sub - are due to Tukano fishermen, and particularly to species of a well-known aquarium tetra. Tropical Fish Antenor, Roberval Pedrosa, and José Maria, for Hobbyist 10 (2): 42, 43-47, 50-51. their enthusiastic collaboration during collecting GÉRY , J. 1963. Three new tetras from the upper Rio Negro activities in the rio Tiquié basin. Eduardo G. Baena near Tapurucuara. Tropical Fish Hobbyist 12 (3): 9, 11, 13-15, 57-59, 62-63. prepared Figs 4, 5, 8 and 9. We are grateful to C. GÉRY , J. 1965 . Poissons characoides sud-américains du R. Moreira, K. M. Ferreira, and N. A. Menezes for Senckenberg Muséum, II. Characidae et Crenuchidae de enlightening discussions on fin hooks presence and l’Igarapé Préto (Haute Amazonie). Senckenbergiana Bio - function in characids. Luiz R. Malabarba and an logica 46 (1): 11-45, 4 pls. anonymous reviewer provided useful suggestions GÉRY , J. 1966. Notes on characoid fishes collected in Suri - which improved the present paper. The authors nam by Mr. H.P. 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Biollania, Edición Espe - Characidae) aus dem Einzugsgebiet des Rio Manuripi in cial 4: 537 pp. Bolivien (Departamento Pando). Zoologische Abhandlun - TAYLOR , W. R. & V AN DYKE , G. C. 1985. Revised proce - gen 52 (1): 23-34. dures for staining and clearing small fishes and other ver - ZARSKE , A. & G ÉRY , J. 2007. Hemigrammus geisleri sp.n. – tebrates for bone and cartilage study. Cybium 9 (2): 107- ein neuer Glassalmler aus dem zentralen Amazonasge - 109. biet, mit einer ergänzenden Beschreibung von Hemi - UJ, A. & G ÉRY , J. 1989. Deux nouvelles espèces de Tetras grammus mimus Böhlke, 1955 (Teleostei: Characiformes: (Poissons characoides, Characidae auct., Tetragonopteri - Characidae). Vertebrate Zoology 57 (1): 5-14.

aqua vol. 15 no. 3 - 20 July 2009 168 Guidelines for Authors 1. Manuscript preparation : manuscripts must be sub - BLABER , S. J. M. 1980. Fish of the Trinity inlet system mitted in English. In exceptional cases aqua may pro - of North Queensland, with notes on the ecology vide translations of manuscripts written in French, Ger - of fish faunas of tropical Indo-Pacific estuaries. man, Italian, or Spanish. Australian Journal of Marine and Freshwater Research 31 :137-46. Manuscripts must be word-processed in Microsoft WORD and submitted in an electronic form. Generic, DAY , J. H., B LABER , S. J. M., & W ALLACE , J. H. 1981. specific, and sub-specific names must be italicised. All Estuarine fishes. In: Estuarine Ecology with Parti - papers must conform to the International Code of Zoo - cular Reference to Southern Africa. (Ed. J.H. Day.): logical Nomenclature. Authors are strongly advised to 197-221. A. A. Balkema, Rotterdam. follow the format set out in previous publications of aqua . DIMMICH , W. W. 1988. Ultrastructure of North Ame - rican cyprinid maxillary barbels. Copeia 1988 (1) : 2. Title : the title must be short and should precisely 72-79. identify the main topic of the article. Names of genera TREWAVAS , E. 1983. Tilapiine Fishes of the Genera or species are followed by the systematic group to which Sarotherodon , Oreochromis and Danakilia. they belong. Author name(s) are given in full beneath British Museum (Natural History), London, 583 pp. the title, followed by the complete mailing and e-mail address(es). 6. Submission of manuscript and illustrations: The manuscript and illustrations must be submitted digi - 3. Abstract : the abstract should not exceed 250 words tally to the Scientific Editor: and draw attention to the principal conclusions. It should not contain any uncommon abbreviations or lit - Dr. Friedhelm Krupp erature citations. The inclusion of abstracts in other lan - Senckenberg Research Institute guages is optional. Senckenberganlage 25 60325 Frankfurt am Main, Germany 4. Subject matter : the text of the manuscript is usually E-mail: [email protected] arranged in four main sections: Introduction, Materials Tel: +49-69-7542.1255, Fax: +49-69-7542.1253 and methods (including a key to abbreviations), Results, and Discussion. Other subdivisions may be to whom all subsequent correspondence shall be chosen depending on the material presented. Acknowl - addressed. Texts, tables, and graphs should be in edgements should be placed between the text and refer - Microsoft-compatible electronic form. ences. After the paper has been accepted for publication, illus - Scientific names of genera, species, and subspecies trations as high-resolution TIF files or original pho - should be followed by the name(s) of author(s) and the tographs (ideally transparencies; otherwise glossy year of publication on first mention. A description of a prints, preferably in the size in which they will appear - new taxon must contain the following sections: Mater - the type area of aqua is 158 x 224 mm, one column is ial, Diagnosis, Description, and Affinities. Holotype 76 mm wide) must be sent to: and paratypes must be clearly identified, the institution Aquapress, The Managing Editor in which they have been deposited named, and the cat - Via G. Falcone 11, alogue numbers given. Private collections are not 27010 Miradolo Terme (Pavia), Italy acceptable as repositories for holotypes. E-mail: [email protected] Synonyms must be arranged in chronological order. Authors should retain copies of all materials for refer - Identification keys must be dichotomous. ence. The metric system and SI units must be used. Proofs of accepted papers will be sent as PDF files by e- Temperatures are given in °C. Fractions should not be mail attachment to the corresponding author. used. 7. Evaluation of manuscripts: manuscripts will be eval - 5. References to literature : the name-year system must uated by the editors and referees. Papers are accepted on be used. The list of references should be placed at the the understanding that they have not and will not be end of the paper, alphabetically arranged according to published elsewhere. author name. Only those publications cited in the paper may be included. Titles of journals must be given in 8. Reprints: Authors will receive one free copy of the full. issue in which their paper appears and an e-print in PDF format. Additional copies may be ordered from Examples of correct reference formats: Aquapress. aqua International Journal of Ichthyology Vol. 15 (3), 20 July 2009 Contents:

Gerald R. Allen and Mark V. Erdmann: Two new species of damselfishes (Pomacentridae: Chromis) from Indonesia ...... 121-134

Gerald R. Allen and Mark V. Erdmann: Heteroconger mercyae , a new species of garden eel (Congridae: Heterocongrinae) from West Papua, Indonesia ...... 135-142

Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov: Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia ...... 143-152

Flávio C. T. Lima & Leandro M. Sousa : A new species of Hemigrammus from the upper rio Negro basin, Brazil, with comments on the presence and arrangement of anal-fin hooks in Hemigrammus and related genera (Ostariophysi: Characiformes: Characidae) ...... 153-168

Papers appearing in this journal are indexed in: Zoological Record; BioLIS – Biologische Literatur Information Senckenberg; www.aqua-aquapress.com; www.aquapress-bleher.com; www.Joachim-Frische.com

Cover photo: Heteroconger mercyae, underwater photo of paratype, 641.4 mm TL, Papisol Bay, West Papua, Indonesia. Photo by M. Erdmann.

Pictichromis paccagnellae, Papua New Guinea. Underwater. Photo by J. E. Randall from an upcoming description of a new dottyback from Indonesia, in aqua 15 (4), October 2009.