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aqua International Journal of

Vol. 19 (1), 21 January 2013

Aquapress ISSN 0945-9871 AQUA19(1):AQUA 24/01/13 12:37 Pagina 102

aqua - International Journal of Ichthyology Managing Editor: Scope aqua is an international journal which publishes original Heiko Bleher scientific articles in the fields of systematics, , Via G. Falcone 11, bio geography, ethology, ecology, and general biology of 27010 Miradolo Terme (PV), Italy fishes. Papers on freshwater, brackish, and marine fishes Tel. & Fax: +39-0382-754129 will be considered. aqua is fully refereed and aims at pub- E-mail: [email protected] lishing manuscripts within 2-4 months of acceptance. In www.aqua-aquapress.com view of the importance of color patterns in identi - fication and ethology, authors are encouraged to submit color illustrations in addition to descriptions of Scientific Editor: coloration. It is our aim to provide the international sci- entific community with an efficiently published journal Frank Pezold meeting high scientific and technical standards. College of Science & Engineering Texas A&M University – Corpus Christi Call for papers 6300 Ocean Drive – Corpus Christi, TX 78412-5806 The editors welcome the submission of original manu- Tel. 361-825-2349 scripts which should be sent in digital format to the scien- E-mail: [email protected] tific editor. Full length research papers and short notes will be considered for publication. There are no page charges and color illustrations will be published free of charge. Authors will receive one free copy of the issue in which Editorial Board: their paper is published and an e-print in PDF format. Gerald R. Allen Department of Aquatic , Subscription Notice Western Australian Museum, Perth, Australia At least one volume (4 issues) of aqua is being published per year, each issue comprising 38-64 pages (incl. cover). Nina G. Bogutskaya The subscription rate (for one volume = 4 issues): Zoological Institute of the Russian Academy of Personal subscription: Euro 75,00 (online edition) - Euro Sciences, St. Petersburg, Russia 100,00 (online edition + print edition incl. priority mail); Institutional subscription: Euro 800,00 (online edition) - Friedhelm Krupp Euro 1000,00 (online edition + print edition incl. priority Curator of Fishes, Senckenberg Research Institute and mail). Subscription enquires should be sent to the Natural History Museum, Frankfurt am Main, Germany publisher at the address given below or by e-mail to: Flávio C. T. Lima [email protected] - [email protected] Museu de Zoologia da Universidade de São Paulo São Paulo, Brasil Special Publication Since 2003 Aquapress publishes a series of Special Publi- Axel Meyer cations, which are produced at irregular intervals. All Spe- Lehrstuhl für Zoologie und Evolutions biologie, cial Publications have about 100 or more pages and are Universität Konstanz, Germany available separately from regular issues of aqua. Enquiries about subscriptions and prices should be sent to the pub- Paolo Parenti lisher at the address given here above or by e-mail to: Department of Enviromental Sciences, [email protected] - [email protected] University of Milano-Bicocca, Milan, Italy Mário de Pinna Museu de Zoologia da USP, São Paulo, Brazil John E. Randall Bishop Museum, Honolulu, Hawaii, U.S.A. Richard Winterbottom ISSN 0945-9871 Centre of Biodiversity & Conservation Biology, Publisher: Aquapress, Redazione aqua, Royal Ontario Museum, Toronto, Canada I-27010 Miradolo Terme (Pavia), Italy www.aqua-aquapress.com Helen K. Larson Printer: Pronto Stampa Srl – Bergamo – Italy Curator Emeritus, Fishes - Museum and Art Gallery of Copyediting and layout: Rossella Bulla the Northern Territory. Darwin, Australia © 2013 aqua, International Journal of Ichthyology AQUA19(1):AQUA 24/01/13 12:37 Pagina 1

aqua, International Journal of Ichthyology

A new species of () from the Indian Ocean

John E. Randall1 and Joseph D. DiBattista2

1) Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA. E-mail: [email protected] 2) Red Sea Research Center, King Abdullah University of Science and Technology, Thuwal, 23955-6900, Saudi Arabia. E-mail: [email protected]

Received: 29 July 2012 – Accepted: 14 December 2012

Abstract morphologische Unterscheidung zwischen C. dimidiata am The pomacentrid fish dimidiata (Klunzinger, Beispiel von Exemplaren aus dem zentralen Roten Meer und 1871), locality Red Sea, formerly believed to be wide- C. fieldi anhand von Proben aus dem Indischen Ozean als ranging into the Indian Ocean, is restricted to the Red Sea. stammesgeschichtliche Trennung bestätigen (d = 0,019). The Indian Ocean population is described as a new Die beiden Arten unterscheiden sich von der nahe ver- species, Chromis fieldi. It differs in having modally 17 pec- wandten Art C. iomelas Jordan & Seale, 1906 vom west- toral rays and 17 lateral-line scales, compared to modally lichen und zentralen Pazifik durch 12 Weichflossenstrahlen 16 pectoral rays and 15 lateral-line scales for C. dimidiata, in der Rücken- und der Afterflosse (Modalwert) (bei C. and the demarcation of the dark brown anterior part of the iomelas sind es 13), Abgrenzung des dunkelbraunen und des body from the white posterior part is convex, compared to weißen Teils deutlich vor dem Ansatz der Afterflosse sowie nearly straight in C. dimidiata, and not as sharply defined den phylogenetischen Unterscheidungswert anhand der dorsally and ventrally. Phylogenetic comparisons based on Cyto chrom-b-Sequenz d = 0,085 im Vergleich zu d = 0,087. mitochondrial DNA (mtDNA) cytochrome b sequences support morphological differentiation with evolutionary Résumé separation of C. dimidiata sampled in the central Red Sea Le pomacentridé Chromis dimidiata (Klunzinger, 1871), and C. fieldi sampled in the Indian Ocean (d = 0.019). localité-type en Mer Rouge, que a eu une large distribution These two species differ from the related C. iomelas Jordan dans l'Océan Indien pensait-on jadis, se trouve confiné en & Seale, 1906 of the western and central Pacific in having Mer Rouge. La population de l’Océan Indien est décrite modally 12 dorsal and anal fin soft rays (vs. 13 for C. iome- com me nouvelle espèce, Chromis fieldi. Elle se caractérise par las), the demarcation of dark brown and white distinctly 17 rayons pectoraux modalement et 17 écailles sur la ligne anterior to the origin of the anal fin, and d = 0.085 to d = la térale, alors que C. dimidiata a 16 rayons pectoraux mo - 0.087 for cytochrome b. da lement et 17 écailles sur la ligne latérale, et par la démarca - tion convexe entre la partie antérieure brun foncé du corps Zusammenfassung et la partie postérieure blanche, alors qu’elle est presque Der Riffbarsch Chromis dimidiata (Klunzinger, 1871), droite chez C. dimidiata et moins nettement dessinée et ven - Typuslokalität Rotes Meer, von dem man bisher angenom- tra lement. Des comparaisons phylogénétiques basées sur des men hat, dass er ein weites Verbreitungsgebiet bis in den sé quences de cytochrome b de l’ADN mitochondrial Indischen Ozean hat, ist nach neueren Erkenntnissen auf (mtADN) rélèvent une différenciation morphologique avec das Rote Meer beschränkt. Der Bestand im Indischen di vergence évolutive de C. dimidiata collecté au centre de la Ozean wird als neue Art Chromis fieldi beschrieben. Ihre Mer Rouge et C. fieldi collecté dans l’Océan Indien (d = Vertreter unterscheiden sich durch 17 Brustflossenstrahlen 0.019). Ces deux espèces se distinguent de C. iomelas Jordan und 17 Seitenlinienschuppen (Modalwert) im Gegensatz zu & Seale, 1906 apparenté, originaire du Pacifique ouest et 16 Brustflossenstrahlen und 15 Seitenlinienschuppen bei C. central, par 12 rayons mous modalement à la dorsale et à dimidiata; außerdem ist die Grenzlinie zwischen dem l’anale (pour 13 chez C. iomelas), par la démarcation entre le dunkelbraunen vorderen Teil des Körpers und dem weißen brun foncé et le blanc situee clairement avant la naissance de hinteren Teil konvex, während sie bei C. dimidiata nahezu l’anale, et par un cytochrome b de d = 0,085 à d = 0,087. gerade verläuft, und die Abgrenzung ist rücken- und bauch- wärts weniger scharf gezeichnet. Beim phylogenetischen Sommario Vergleich auf der Grundlage der mitochondrialen DNA Il pomacentride Chromis dimidiata (Klunzinger, 1871), (mtDNA) in den Cytochrom-b-Sequenzen lässt sich die località tipo Mar Rosso, ritenuto ampiamente diffuso nel-

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l'Oceano Indiano, è in realtà limitato al Mar Rosso. La cauda and C. unipa, raising the total species of the popolazione dell’Oceano Indiano è descritta come una to 96. Quéro et al. (2010) described C. nuova specie, Chromis fieldi. Si differenzia per avere, come durvillei, the 97th species, from the island of Réu- valori modali, 17 raggi pettorali e 17 scaglie in linea late- nion. We provide here the description of the 98th rale anziché 16 raggi pettorali e 15 scaglie in linea laterale come in C. dimidiata e la demarcazione della parte anterio - species. re marrone scuro del corpo dalla parte bianca posteriore The species of Chromis feed primarily on zoo- convessa, rispetto a quasi dritta in C. dimidiata e non così plankton (Fig. 1); they are therefore not restricted nettamente definita dorsalmente e ventralmente. Con- to the photic zone of the sea, as are the species of fronti filogenetici basati su sequenze del DNA mitocon- the other large damselfish genera Abudefduf, Poma- driale (mtDNA) del citocromo b in C. dimidiata campi- centrus, and Stegastes that graze on benthic algae. onato nel Mar Rosso e C. fieldi campionato nell'Oceano The ability to colonize deeper habitats may have Indiano sono a favore di una differenziazione morfologica provided more opportunity for speciation in the e una separazione evolutiva delle due specie (d = 0.019). Esse differiscono dal relativo C. iomelas Jordan & Seale, genus Chromis. Most species of Chromis described 1906 del Pacifico occidentale e centrale per avere come va - in recent years have been collected from deeper lore modale 12 raggi molli nella pinna dorsale e anale (vs. than conventional scuba-diving depths. Pyle et al. 13 per C. iomelas), la delimitazione dell’area di colore mar- (2008), for example, named five new species of rone scuro da quella bianca distintamente anteriore all’ori- Chromis from the western Pacific, four of which gine della pinna anale e valori d tra 0.085 e 0.087 per il were collected from over 85 m, and the fifth from cito cromo b. 60 m. These authors used mixed-gas, closed-circuit rebreather gear to make their collections. Allen & INTRODUCTION Erdmann (2009) reported that species of Chromis The damselfish genus Chromis is the largest of the are known to penetrate depths as great as 175 m. family Pomacentridae. Allen & Erdmann (2009) Chromis durvillei, which was found at the surface described two new species from Indonesia, C. albi- after lava from an eruption on Réunion in 2007

Fig. 1. Chromis dimidiata feeding on zooplankton with C. ternatensis and Pseudanthias gibbosus, Strait of Tiran, Red Sea. Photo by J. L. Rose.

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flowed into the deep sea, may well have been from the species from Mauritius by name only, and a record depth. Harmelin-Vivien (1976) from Réunion at depths The species described here is a common inshore of 6-40 m. Allen in Smith & Heemstra (1986) also fish of the Indian Ocean that has usually been iden- used Smith’s painting of Chromis dimidiata and tified as Chromis dimidiata (Klunzinger, 1871), gave the distribution as “Tropical Indian Ocean type locality Red Sea, known by the English com- reefs down to at least 30 m; Red Sea to Durban, mon names Half & Half Chromis, Chocolate Dip common.” In a review of of the Chromis, and Two-tone Chromis. Klunzinger’s world, Allen (1991: 66) illustrated Chromis dimidi- name has been applied to more than one species by ata from Eilat, Red Sea and gave the distribution as authors that followed. Günther (1909) used the “Widespread in the Indian Ocean including name for a species from the Society Islands with a Kenya, Mauritius, Réunion, Chagos Archipelago, dark brown body, except for the white caudal Maldive Islands, Sri Lanka, Andaman Sea, and peduncle and adjacent median fins, now known to Christmas Island.” Allen also provided a color fig- be Chromis margaritifer Fowler, 1946, type locality ure of Chromis iomelas Jordan & Seale, 1906 (also Ryukyu Islands, first described as a subspecies of C. one in Randall et al. 1990) that has long been con- dimidiata. De Beaufort (1940) placed Barnard’s fused with C. dimidiata. It is easily seen why (com- (1927) C. xanthura, now a valid species from the pare Fig. 1 through Fig. 9 to Fig. 10 through Fig. Pacific to the eastern Indian Ocean, in the syn- 12). Fowler (1928), for example, placed C. iomelas onymy of C. dimidiata and listed localities from the in the synonymy of C. dimidiata and gave the dis- Red Sea and coast of Natal to the Hawaiian Islands tribution of the latter as Red Sea, East Indies, Poly- and Society Islands. In a report on fishes of the nesia, and Hawai‘i. Allen (1991) gave the correct Philippines and Indonesia collected by the United distribution for C. iomelas: Great Barrier Reef, States steamer Albatross, Fowler & Bean (1928) Coral Sea, New Guinea, Vanuatu, New Caledonia, listed Chromis bicolor (Macleay, 1882) from New Fiji, Samoa Islands, and Society Islands. Although Guinea, C. leucurus Gilbert, 1905 from the Hawai- Allen did not specifically differentiate C. iomelas ian Islands, and C. iomelas Jordan & Seale, 1906 from C. dimidiata, as by a key, the counts he gave from Samoa as synonyms of C. dimidiata. He com- for the dorsal and anal rays of the two species pro- mented on the extreme variation in the demarca- vide complete separation. tion of the dark brown anterior part of the body Khalaf & Disi (2007: 135) illustrated Chromis and the posterior white. dimidiata in color in their book on the fishes of the Smith (1949) illustrated Chromis dimidiata in Gulf of Aqaba, noting that the species occurs in the color, dark brown anteriorly, abruptly pale yellow Red Sea and the Indian Ocean. Richard and Mary and white posterior to a demarcation above the Field also figured Chromis dimidiata in color from base of the second anal spine, and gave the distrib- an underwater photograph in their guidebook Reef ution as central tropical Indo-Pacific to the east Fishes of the Red Sea (1998). Richard commented coast of Africa, south to Zululand. Smith & Smith to the first author that the demarcation of the dark (1963) used the same illustration in recording the brown anterior half of fish in the Red Sea is a little species from the Seychelles. Baissac (1976) listed posterior to that of individuals in the Indian

Fig. 2. Chromis dimidiata, Gulf of Aqaba, Red Sea. Photo Fig. 3. Chromis dimidiata, Strait of Tiran, Red Sea. Photo by J. E. Randall. by J. L. Rose.

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Ocean. This led to our taking routine counts of fin rounded to the nearest 0.05. Data in parentheses in rays, scales, and gill rakers. With modal differences the description refer to paratypes. apparent (Tables I-III), the second author made a A total of nine tissue samples were included for molecular study that conclusively determined that genetic analysis: four C. dimidiata collected from the Indian Ocean population of C. dimidiata is dis- the Red Sea (Thuwal, Kingdom of Saudi Arabia), tinct from that in the Red Sea and represents an two C. dimidiata (now C. fieldi) from the Indian undescribed species. We also provide meristic, Ocean (Republic of Seychelles), and three C. iome- genetic, and color separation from the similar las from Fiji in the Pacific Ocean for comparison. South Pacific C. iomelas Jordan & Seale. Eight previously published "C. dimidiata" (Réu- nion, N = 4: GenBank Accession Numbers MATERIAL AND METHODS JF458039, JF458040, JF458041, and JF458045 Type specimens have been deposited in the [Hubert et al. 2012]; Republic of Madagascar, Bishop Museum, Honolulu (BPBM); California N = 4: GenBank Accession Numbers JF458042, Academy of Sciences, San Francisco (CAS); JF458043, JF458044, and JF458046 [Hubert et Hebrew University, Jerusalem (HUJ); Royal al. 2012]) and two C. iomelas (Moorea, French Ontario Museum, Toronto (ROM); Senckenberg Polynesia, N = 1: GenBank Accession Number Museum, Frankfurt (SMF); South African Insti- JF458049 [Hubert et al. 2012]; New Caledonia, tute for Aquatic Biodiversity, Grahamstown N = 1: GenBank Accession Number AY208531 (SAIAB); and the U. S. National Museum of Nat- [Quenouille et al. 2004]) mitochondrial DNA ural History, Washington, D.C. (USNM). (mtDNA) cyto chro me b (cyt b) sequences further Lengths given for specimens are standard length supplemented gene tic analyses. Despite a relatively (SL), the straight-line distance from the median slower mutation rate than some other mtDNA anterior point of the upper lip or most anterior genes (i.e., control region; Lee et al. 1995), we here teeth (whichever is longest) to the base of the cau- use cyt b because of pre-existing vouchered speci- dal fin (posterior end of hypural plate). Head mens to increase our Indian Ocean sample, and an length is measured from the same anterior point to approximate molecular clock calibrated in other the posterior end of the opercular membrane, and reef fish (Bowen et al. 2001; Lessios 2008; Reece et snout length from the same point to the fleshy al. 2010). Indeed, the cyt b gene is one of the most edge of the orbit. Body depth is the greatest depth widely used markers in molecular systematics. to the base of the dorsal spines; body width is the Total genomic DNA was extracted from all tissue greatest width measured just posterior to the gill samples using the “HotSHOT” protocol (Meeker opening. Orbit diameter is the greatest fleshy et al. 2007). A 670 base pair (bp) segment of the diameter, and interorbital width the least fleshy mtDNA cyt b gene was amplified using a heavy- width. Caudal-peduncle depth is the least depth; strand (5’ - GTGACTTGAAAAACCACCGTTG caudal-peduncle length is measured horizontally - 3’; Song et al. 1998) and light-strand primer (5’ - from the rear base of the anal fin to the ventralmost AATAGGAAGTATCATTCGGGTTTGATG - 3’; principal caudal ray. Predorsal, preanal, and pre- Taberlet et al. 1992). Polymerase chain reaction pelvic lengths are from the front of the upper lip (PCR) mixes contained BioMix Red (Bioline Inc., (or upper teeth) to the origin of the respective fins. Springfield, NJ, USA), 0.26 mM of each primer, Lengths of fin spines and soft rays are measured and 5 to 50 ng template DNA in a 15 ml total vol- from the extreme base of these elements to the tips. ume. PCR cycling parameters were as follows: ini- Some soft rays of the dorsal, anal, and pelvic of tial 95 °C denaturation for 3 min., followed by 35 many species of Chromis are long and filamentous; cycles of 94 °C for 30 sec., 55 °C for 45 sec., and therefore, there is often great variation in recording 72 °C for 45 sec., with a final elongation step of 72 the maximum length of the rays of these fins. Pec- °C for 10 min. toral-ray counts include the short unbranched All PCR products were cleaned by incubating upper ray. Gill-raker counts were made on the first with exonuclease I and FastAP™ Thermosensitive gill arch; the raker at the angle is included in the Alkaline Phosphatase (ExoFAP; USB, Cleveland, lower-limb count. OH, USA) at 37 °C for 60 minutes, followed by Table I gives the proportional measurements of 85 °C for 15 minutes, sequenced in both the for- the new species as percentages of the standard ward and reverse directions with fluorescently length. Proportional measurements in the text are labeled dye terminators following manufacturer’s

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protocols (BigDye, Applied Biosystems Inc., Foster Sequence divergence (d) between identified groups City, CA, USA), and analyzed using an ABI was then estimated based on the corrected average 3130XL Genetic Analyzer (Applied Biosystems). pairwise nucleotide difference from Arlequin vers. The sequences were aligned, edited, and trimmed 3.5 software (Excoffier et al. 2005), divided by the to a common length using Geneious Pro vers. 4.8.4 total number of base pairs in the sequence. software (Drummond et al. 2009). Representative haplotypes were deposited in GenBank (accession numbers: KC311941 to KC311949). jModelTest Chromis fieldi, n. sp. vers. 1.0.1 (Posada 2008) was used to determine (Figs 4-8; Table I) the best nucleotide substitution model under Akaike information criterion (AIC), which selected Heliastes xanthurus (non Bleeker) Regan, 1917: 459 the HKY model (Hasegawa et al. 1985) and a (Natal). gamma parameter of 0.071. Chromis xanthurus (non Bleeker) Barnard, 1927: Maximum-likelihood (ML) and Bayesian (BA) 732 (Natal coast of South Africa). tree-building methods were applied using PAUP* Chromis dimidiatus (non Klunzinger) Smith, 1949: vers. 4.0 (Swofford 2000) and MRBAYES (Ronquist 507, pl. 51, fig. 746a (east coast of Africa to Zul- & Huelsenbeck 2003) Bayesian Markov Chain uland). Monte Carlo (MCMC) coalescent approach Chromis dimidiatus (non Klunzinger) Fourmanoir, implemented in Geneious Pro. The Bayesian 1954: 226 (Comoro Islands). MCMC search strategy consisted of the default Chromis dimidiatus (non Klunzinger) Smith, 1955: four heated, 1 million step chains with an initial 888 (Aldabra). burn-in of 150,000 steps. A previously published Chromis dimidiatus (non Klunzinger) Smith, Green Chromis (Chromis viridis) cyt b sequence 1960: 324, pl. 31, fig. I (Zululand and Inhaca (Genbank accession number: JF458062) was used Island, Mozambique, north over all of W Indian to root the tree. Support for the tree topology was Ocean). evaluated by bootstrapping over 10,000 replicates Chromis dimidiatus (non Klunzinger) Smith & for ML and posterior probabilities for BA. Smith, 1963: 34, pl. 71, fig. I (Seychelles).

Fig. 4. Holotype of Chromis fieldi, BPBM 20241, 57 mm, Mauritius. Photo by J. E. Randall.

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Chromis dimidiatus (non Klunzinger) Baissac, Chromis dimidiatus (non Klunzinger) Satapoomin, 1976: 211 (Mauritius). 2007: 104, fig. 13 (Andaman Sea). Chromis dimidiatus (non Klunzinger) Harmelin- Chromis dimidiatus (non Klunzinger) Shibukawa in Vivien, 1976: 76 (Réunion). Kimura et al., 2009: 205, middle fig. (Andaman Chromis dimidiata (non Klunzinger) Allen in Smith Sea). & Heemstra, 1986: 674, pl. 86, fig. 219.14 (trop- ical Indian Ocean; Red Sea to Durban). Holotype. BPBM 20241, 57 mm, Mauritius, west Chromis dimidiata (non Klunzinger) Allen & coast off Medine (north of Flic en Flac), reef in 30 Steene, 1987: 92, fig, 257 (Christmas Island). m, quinaldine (anaesthetic), J. E. Randall, 19 Nov Chromis dimidiata (non Klunzinger) Winterbottom 1973. et al., 1989: 45, fig. 255 (Chagos Archipelago). Paratypes: USNM 275619, 2: 16.5-17 mm, Sri Chromis dimidiata (non Klunzinger) Randall, Lanka, Trincomalee, 300 yds S of entrance to Ft. 1992: 102, fig. 213 (Maldive Islands). Frederick, 6 m, C. C. Koenig, 6 Apr 1970; BPBM Chromis dimidiata (non Klunzinger) Allen & Adrim, 20066, 2: 31-45 mm, Réunion, off Cap Houssaye, 2003: 45 (Indian Ocean to Sumatra and west Java). 12 m, rotenone, J. E. Randall, 23 Oct 1973; Chromis dimidiata (non Klunzinger) Heemstra & BPBM 41005, 52.5 mm, same data as holotype, Heemstra, 2004: 332, fig. (KwaZulu-Natal, juve- except taken by multiprong spear; BPBM 16391, niles occasionally to Algoa Bay). 17 mm, Tanzania, Mafia Island, Chole Bay, reef, 0- 2 m, rotenone, J. E. Randall, L. Westfield, and A. Klosser, 11 Dec 1973; USNM 364137, 55 mm, Cargados Carajos (St. Brandon’s Shoals), 16°25’S, 59°36’E, 6-10.5m, rotenone, V.G. Springer et al., 6 Apr 1976; BPBM 21723, 4: 30-36 mm, South Africa, KwaZulu-Natal, Sodwana Bay, reef off north end of bay, 10-12 m, rotenone, J. E. Randall and M. Christensen, 21 Jun 1977; BPBM 28436, 11: 23-49 mm; Chagos Archipelago, Salomon Atoll, 5°22.07’S, 72°13.11’E, offshore reef at S end of Isle Boddam, reef top, some sand, hard and soft Fig. 5. Chromis fieldi, Mauritius. Photo by R. F. Myers. coral, 8 m, rotenone, R. Winterbottom, A. R.

Fig. 6. Chromis fieldi, ROM 59476, 42.5 mm, Recif Bambo, Comoro Islands. Photo by R. Winterbottom.

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Emery, et al., 15 Mar 1979; CAS 234256, 2: 41- in SL; head length 3.15-3.35 in SL; snout short 48.5 mm; HUJ 20132, 2: 23.5-41 mm; SAIAB and rounded, 3.1-3.3 in head length; color in 186323, 2: 36-44 mm; SMF 34723, 2: 40.5-44.0 preservative dark purplish brown anteriorly to a mm; USNM 405570, 2: 38-48 mm, all with same near-vertical demarcation usually at or slightly pos- data as BPBM 28436; ROM 37174, 2: 34-37 mm, terior to origin of anal fin (but rarely distinctly Peros Banhos Atoll, Isle du Coin, 5°25’55”S, anterior to anal-fin origin), ending on average at 71°45’46”E, lagoon reef slope of Acropora, 10 m, base of ninth dorsal spine, pale beige posteriorly, rotenone, R. Winterbottom, A. R. Emery, 1 Apr the median fins colored like adjacent body; a black 1979; BPBM 27275, 8: 35-50 mm, Kenya, Shi- spot over base and axil of pectoral fins; color in life moni, Kitangamwe, reef, 12-14 m, rotenone, J. E. very dark gray-brown anteriorly, abruptly white Randall, 22 Mar 1979; USNM 274833, 10: 15-57 posteriorly, usually with wash of pale yellow over mm, Seychelles, Aldabra Atoll, Picard Island, 1-10 the anterior white, especially dorsally. m, B. Kensley, et al., 23-26 Mar 1985; BPBM Description: Dorsal rays XII, 12 (12 or 13, 35567, 3: 15-29 mm, Seychelles, Amirantes, modally 12); anal soft rays II, 13 (12-14); all soft Poivre Atoll, north side, reef, 13 m, rotenone, J. E. dorsal and anal rays branched, the last to base; Randall, D. P. Boullé, and E. Grandcourt, 31 Dec principal caudal rays 15, the median 13 branched; 1992; BPBM 37663, 38 mm, Indonesia, second and third, and fourteenth and fifteenth Mentawai Islands, Siberut Island, off entrance to principal caudal rays with filamentous branched Sarabua Bay, 1°30’S, 99°10’E, reef and coral rub- tips, especially long in third and fourteenth rays; ble, 8 m, quinaldine, J. E. Randall, 24 Apr 1997. upper and lower procurrent caudal rays 4, the ante- Diagnosis: Dorsal rays XII,12 or 13 (modally rior 2 spiniform; pectoral rays 17 (16 or 17, rarely 12); anal soft rays II,12-14; pectoral rays 16 or 17 16), the uppermost splint-like, the second (rarely 16); caudal fin strongly forked; spiniform unbranched; pelvic rays I,5, all branched, the first upper and lower caudal rays 2; second and third, soft ray filamentous. and fourteenth and fifteenth principal caudal rays Longitudinal scale series 27; scales above lateral with filamentous branched tips, especially long in line to base of first dorsal spine 3; scales below lateral third and fourteenth rays; first soft ray of pelvic to base of first anal spine 8; circumpeduncular scales fins filamentous; longitudinal scale series 27; lat- 16; lateral line interrupted, the dorsoanterior part as eral-line scales 16 or 17 (rarely 16); gill rakers 7-9 tubed scales, 17 (16 or 17, rarely 16), ending (rarely 9) + 19-21 (rarely 21); body depth 2.2-2.35 beneath midbase of soft portion of ; poste-

Fig. 7. Chromis fieldi, Mahé, Seychelles. Photo by J. E. Randall.

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rior midlateral pored scales in continuous series 6 (6- and body to above eye, then continuing on nape at 13); median predorsal scales about 17 (difficult to progressively less angle; orbit diameter 2.8 (2.5- count due to interrupting overlapping scales). 2.75) in head length; no fleshy papillae on margin Gill rakers long, the longest equal to length of of orbit; interorbital space convex, the least width longest gill filaments, nearly half orbit diameter in 2.75 (2.65-2.8) in head length; caudal-peduncle holotype, (8 + 20 (7-9, rarely 9) + 19-21, rarely depth 1.9 (1.9-1.95) in head length; caudal- 21); branchiostegal rays 6; supraneural (predorsal) peduncle length 2.5 (2.3-3.5) in head length. bones 3; vertebrae 26. Body moderately deep, the Mouth terminal to slightly inferior and strongly depth 2.2 (2.05-2.15) in SL, and compressed, the oblique, forming an angle of about 45° to horizon- width in body depth about 2.6; head length 3.3 tal axis of body; mouth small, the upper-jaw length (3.15-3.35) in SL; snout length 3.2 (3.1-3.3) in 2.9 (2.8-3.1) in head length, the maxilla extending head length; dorsal profile of snout steep, forming to a vertical slightly posterior to anterior edge of an angle of about 60° to horizontal axis of head orbit; teeth in jaws conical and recurved, in three

Fig. 8. Paratype of Chromis fieldi, ROM 37174, 34 mm, Peros Banhos Atoll, Chagos Archipelago. Photo by R. Winterbottom.

Fig. 9. Chromis fieldi, Sawda, southern Oman. Photo by J. P. Hoover.

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irregular rows anteriorly, narrowing to a single row margin of orbit to front of snout, nearly round, posteriorly; teeth of outer row much the largest, with a slender rim that is low anteriorly and about the longest median teeth twice the diameter of four times longer posteriorly, forming a somewhat aperture of anterior nostril; 38 teeth in outer row pointed flap; short zone before nostril scaleless. of upper jaw, and 42 in lower jaw of holotype; The most prominent pores of the cephalic lateralis tongue triangular, the tip slightly rounded. system are ones directly posterior to nostril, closer to Nostril on side of snout about half distance from nostril than edge of orbit (another directly dorsal to

Fig. 10. Chromis iomelas, Lady Musgrave Is., southern Great Barrier Reef. Photo by J. E. Randall.

Fig. 11. Chromis iomelas, BPBM 22761, 43 mm, Tutuila, American Samoa. Photo by J. E. Randall.

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nostril much smaller), three along margin of snout fins with a patch of small scales basally; a triangular just above upper jaw, followed by six lesser pores scaly process of three scales midventrally between along ventral edge of subopercle; also prominent are bases of pelvic fins; a slender axillary scale above four pairs of pores anteriorly on mandible; remain- each pelvic fins, about half length of pelvic spine. ing pores, such as those closely encircling orbit and Membranes of spinous portion of dorsal and anal those on margin of preopercle, small to minute. fins deeply incised, the first membrane of dorsal fin No spine on opercle; margin of preopercle irregu- nearly to base, the last one-third distance to base; lar, but not serrate; edges of scales slightly overlap- membrane of first anal spine incised one-half dis- ping margins of opercle, preopercle, and subopercle. tance to base. Origin of dorsal fin above base of Scales finely ctenoid; tubed lateral-line scales end- fourth lateral-line scale, the predorsal length 2.6 ing below anterior soft portion of dorsal fin, with (2.45-2.55) in SL; dorsal spines progressively pored scales continuing to anterior caudal peduncle; longer, the first 2.75 (2.7-3.05) in head length; midlateral pored scales commencing below end of twelfth dorsal spine 1.4 (1.45) in head length; tubed lateral-line scales and ending on base of cau- fourth dorsal soft ray longest, 3.15 (1.75-3.4) in dal fin, but not continuously (always one or more SL; origin of anal fin below base of ninth dorsal scales in series without pores); about two-thirds of spine, the preanal length 1.6 (1.6-1.7) in SL; first opercle covered by six large scales; three curving anal spine 2.45 (2.3-2.6) in head length; second rows of large scales across widest part of preopercle; anal spine 1.15 (1.1-1.35) in head length; sixth or no very small scales on head; scales in an oblique seventh anal soft rays longest, 3.35 (3.3-4.45) in row across interorbital of holotype from above mid- SL; caudal-fin length (includes filament of longest dle of orbit to behind posterior nostril of other side ray) 1.85 (1.65-1.9) in SL; caudal concavity 3.2 6; scales in a zigzag row between nostrils 6; narrow (2.9-4.5); pectoral-fin length 3.3 (3.1-3.4) in SL; zone of snout anterior to nostrils largely naked; a pelvic-fin length 2.6 (2.2-3.0) in SL. broad basal sheath of scales at base of dorsal and anal Color of holotype in alcohol dark pur- fins, with a narrowing band of progressively small plish brown anterior to a demarcation that begins scales extending out to incised end of each inter- ventrally at origin of anal fin and ends at base of spinous membrane; scales continuing on soft por- about ninth dorsal spine but extends in middle of tion of fin progressively shorter; caudal fin with body posteriorly to a vertical from base of first anal small scales continuing, progressively smaller, nearly soft ray to base of eleventh dorsal spine; body pos- to posterior margin (disregarding filaments); paired terior to demarcation pale beige; median fins col-

Fig. 12. Chromis iomelas, Rangiroa, Tuamotu Archipelago. Photo by J. E. Randall.

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ored basally like adjacent body, the rays soon Fig. 4. Note the nebulous margin dorsally and ven- becoming pale gray-brown and the membranes trally that separates the dark brown anterior part of translucent; pectoral fins translucent with pale the body from the pale yellowish posterior part; also brown rays and a large black spot over base and the pale yellowish posterior body grading to pale axil; pelvic fins dark brown. bluish white on caudal peduncle. Figures 6 and 8 Color of holotype when fresh shown in provide the color of other specimens when fresh.

Fig. 13. Phylogeny showing relationships among mtDNA cytochrome b haplotypes (670 bp) for Chromis dimidiata speci- mens collected in the Red Sea (N = 4) and Chromis fieldi specimens collected in the Indian Ocean (N = 2). Chromis iomelas (N = 3) specimens collected in Fiji were included for comparison. Genetic analyses were further supplemented by the addi- tion of previously published DNA sequences of "C. dimidiata" from the Republic of Madagascar (N = 4) and Réunion (N = 4), as well as C. iomelas from Moorea, French Polynesia (N = 1) and New Caledonia (N = 1). Trees were constructed using maximum-likelihood (ML) and Bayesian (BA) approaches; maximum bootstrap (N = 10,000 reps) values and posterior prob- abilities for each method are shown above the nodes (ML/BA). Both tree analyses resulted in nearly identical topologies and so only the Bayesian tree is presented here. The sample size and collection location of those samples are in parentheses, and branch lengths are according to the indicated scale. Site abbreviations are as follows: Fiji, FIJ; Kingdom of Saudi Arabia, KSA; Republic of Madagascar, MAD; Moorea, French Polynesia, MOR; New Caledonia, NCL; Réunion, REU; Republic of Sey- chelles, SEY.

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Table I. Proportional measurements of type specimens of Chromis fieldi as percentages of the standard length.

Holotype Paratypes BPBM BPBM BPBM USNM BPBM BPBM 20241 37663 28436 405570 27275 41105 Standard length (mm) 57.0 38.0 40.5 46.5 50.0 52.5 Body depth 45.6 45.8 49.2 48.9 47.1 46.6 Body width1 14.01 18.0 19.1 18.6 17.9 15.51 Head length 30.2 30.5 31.6 31.9 30.6 29.8 Snout length 9.5 9.4 9.6 10.3 9.8 9.7 Orbit diameter 10.7 13.2 12.7 12.2 11.1 10.8 Interorbital width 11.0 11.1 11.8 11.5 11.2 11.2 Upper-jaw length 10.3 10.5 10.2 10.8 10.2 10.7 Caudal-peduncle depth 15.9 15.6 16.0 16.6 16.2 15.3 Caudal-peduncle length 12.2 13.4 12.6 13.7 13.1 12.4 Predorsal length 38.7 38.9 40.2 40.5 39.8 39.0 Preanal length 62.5 59.7 60.3 59.5 63.1 61.8 Prepelvic length 36.8 36.9 37.0 38.0 37.7 37.8 Base of dorsal fin 56.2 54.8 57.0 56.5 51.5 53.8 First dorsal spine 10.9 10.3 10.7 10.6 10.2 11.0 Twelfth dorsal spine 21.4 21.2 21.5 21.7 21.3 20.8 Longest dorsal ray 31.7 29.7 31.9 31.3 broken 32.3 Base of anal fin 26.5 26.1 26.3 26.2 25.8 26.7 First anal spine 12.3 13.3 12.2 12.9 13.0 11.5 Second anal spine 26.4 23.5 24.7 23.9 23.9 27.7 Longest anal ray 29.8 26.5 30.1 28.2 25.6 22.5 Caudal-fin length 53.5 52.5 58.7 61.5 53.9 61.0 Shortest caudal ray 22.1 23.4 23.9 24.8 23.2 23.1 Pectoral-fin length 30.0 30.6 32.3 32.0 30.4 29.5 Pelvic-spine length 19.6 18.0 20.8 19.3 19.7 18.5 Pelvic-fin length 38.1 33.2 45.0 44.8 36.4 36.2

1Holotype and largest paratype once dessicated, now abnormally thin.

The life color of Chromis fieldi is shown in the Ocean fish, and d = 0.087 or d = 0.085 when com- underwater photographs of Figs 5, 7 and 9. Note paring each of these groups to the more distantly the light blue on the leading edge of the dorsal and related C. iomelas in the Pacific Ocean. This is in anal spines and along the margin of the anterior contrast to d = 0.002 or d = 0.004 within the Red soft portion of the median fins, the bluish white Sea or Indian Ocean groups. Comparable genetic posterior margin of the black axillary spot, and the differentiation was obtained from the cytochrome c faint dark longitudinal banding on the rows of oxidase subunit I (COI) barcoding gene (data not scales of the postorbital head; there is often a faint shown), but we here focus our discussion on the bluish iridescence on the upper lip and suborbital. above outlined cyt b gene and samples. Etymology: This species is named in honour of GENETIC RESULTS Richard Field, who first suspected that the Indian The genetic analysis, based on 19 mtDNA cyt b Ocean population of Chromis dimidiata might rep- sequences within the Chromis complex, support the resent a different species. He also provided 18 outlined morphological differentiation among new underwater photographs of C. dimidiata taken in and existing species. Both phylogenetic methods the Red Sea in the vicinity of Jeddah that gave us show evolutionary separation between C. dimidiata the opportunity to note color variation, and col- sampled in the central Red Sea and that sampled in lected four specimens of this species from north of the Indian Ocean (including Madagascar, Réunion, Jeddah, Saudi Arabia. and the Seychelles; Fig. 13). We found that d = Distribution: Specimens of Chromis fieldi have 0.019 when comparing the Red Sea to Indian been examined or records verified from Kenya,

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Table II. Fin-ray counts of species of Chromis.

Dorsal Soft Rays Anal Soft Rays Pectoral Rays 11 12 13 14 11 12 13 14 16 17 18 C. dimidiata 2 15 1 1 9 6 2 16 2 C. fieldi 11 6 4 12 1 1 16 C. iomelas 15 3 1 7 7 3 2 14 2

Table III. Lateral-line scale counts of species of Chromis. Table IV. Gill-raker counts of species of Chromis. Lateral line Scales Upper Limb Lower Limb 14 15 16 17 7891018192021 C. dimidiata 2106 C. dimidiata 89 1 2 8 6 2 C. fieldi 116 C. fieldi 4121 4121 C. iomelas 1134 C. iomelas 15 3 1 7 7 3

Tanzania, Mozambique, KwaZulu-Natal, Sey- species differ also in the dark brown-white demar- chelles (including Amirantes), Comoro Islands, cation on the body. It is straighter in C. dimidiata Réunion, Mauritius, Cargados Carajos Shoals, and more sharply defined dorsally and ventrally. Chagos Archipelago, Maldive Islands, southern The demarcation may be slightly before or poste- Oman, Socotra in the Gulf of Aden, Sri Lanka, rior to the origin of the anal fin in both species, but Mentawai Islands (Sumatra), and Christmas because of the convexity of the demarcation in C. Island. In addition, Gerald R. Allen (pers. comm.) fieldi, it is more posterior. The tip of the pectoral provided records for Pulu Weh (off western Suma- fin reaches or extends posterior to the demarcation tra) and western Java, and Tilman J. Alpermann in 13 of the 14 Bishop Museum specimens of C. identified uncataloged specimens in the Sencken- dimidiata. Except for juveniles, the pectoral fin berg Museum, Frankfurt that were collected by rarely reaches the demarcation in C. fieldi. In addi- him and Uwe Zajonz at Socotra Island in the Gulf tion, the demarcation in C. fieldi remains distinct of Aden, using a copy of this manuscript as a guide. dorsally, though it may be irregular as it passes Remarks: Chromis fieldi is found on coral reefs, through the dorsal fin, whereas it becomes diffuse both lagoon and outer reefs, but not in areas and curves strongly anteriorly in C. dimidiata. exposed to heavy wave action, from the depth The genetic analysis indicates evolutionary sepa- range of 1 to at least 40 m. It feeds on zooplank- ration between specimens sampled in the central ton, but rarely more than a meter above the shelter Red Sea and those sampled in the Indian Ocean. of the reef. The larger species of Chromis, Lepidozy- Previous research in Chromis viridis, a genetic out- gus tapeinosoma, and the species of Pseudanthias group we use here, shows a similar distinction feed much higher in the water column. Winterbot- between the Red Sea and Indonesian Archipelago tom et al. (1989) collected 491 specimens of C. using mtDNA control region (Froukh & Kochzius fieldi, 14-51 mm SL, in 37 lots from 0.5-36 m, 2008). Such lineages may be explained by isolation from reef tops and drop-offs in the Chagos Archi- between each body of water (Red Sea, Indian pelago. Thirty-one percent of their specimens were Ocean, and Pacific Ocean) and limited exchange from 6-15 m, and 66% in 16-25 m. They added, during glacial periods when sea levels were much “It was ubiquitous along the rim of the drop-off, lower (up to 120 m; Voris, 2000; Siddall et al. where it formed large associations feeding on 2003), though the discovery of cryptic species plankton in the water column.” Richard Winter- within ocean basins is not uncommon for Poma- bottom also collected 23 lots of this species in the centrids (Chrysiptera arnazae, Allen et al. 2010; Comoro Islands. Chrysiptera rex, Drew et al. 2010; Pomacentrus This species is differentiated from the closely maafu, Allen & Drew 2012). related Chromis dimidiata in having strongly Although no exact benchmark is available for modal 17 pectoral rays and 17 tubed lateral-line species-level divergence among reef fish taxa based scales vs. modally 15 for C. dimidiata. The two on cyt b (but see Johns & Avise 1998), our findings

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are consistent with the 2% cyt b divergence among Aqaba, Ras Mukabeila; BPBM 18232, 8: 40-52 recognized Chaetodon species (McMillan & Palumbi mm, Red Sea, Gulf of Aqaba, Sinai east coast north 1995) and approached the level of divergence of Coral Island, 15 m; BPBM 27457, 4: 32–39 observed among some closely related species of Das- mm, Sudan, Towartit Reef, 10-13 m; BPBM cyllus (Bernardi & Crane 1999). Genetic differenti- 38779, 4: 24-41 m, Red Sea, reef north of Jeddah ation in this study (d = 0.019) is also comparable to (21°42’29”N, 39°5’, 11”E). some (d = 0.017; Abudefduf concolor versus A. tau- Chromis iomelas: BPBM 14718, 48 mm, Great rus), but not other (d = 0.099; Chromis atrilobata Barrier Reef, Pixie Reef (NE of Cairns), 9 m. versus C. multilineata), recognized geminate species BPBM 33806, 13: 17-50 mm, Coral Sea, Chester- of damselfish separated more than three million field Bank, Ile Longue, 15 m. BPBM 28676, 3: 42- years ago by the rise of the isthmus of Panama 43 mm, New Caledonia, 55 m. BPBM 32734, 8: (Lessios 2008). In line with recent studies of Poma- 16-45 mm, Rotuma, 11-17 m. BPBM 14579, 5: centridae in the Indo-Pacific (Allen et al. 2010; 18-45 mm, Fiji, Viti Levu, 15-22 m. BPBM Allen & Drew 2012), we found that intra-specific 38160, 7: 26-52 mm, Tonga, Vava’u, 16 m. genetic differentiation was nearly an order of mag- BPBM 22781, 2: 38-47 mm, American Samoa, nitude lower than inter-specific differentiation, Tutuila. BPBM 13690, 31 mm, Austral Islands, despite the slower relative mutation rate of cyt b. Rurutu, 27 m. BPBM 5880, 17: 43-56 mm, Soci- Thus, based on reciprocally monophyletic groups ety Islands, Tahiti, 21-27 m; BPBM 8373, 3L 35- and concordant meristic and color differences 47 mm. BPBM 39256, 54 mm, Marquesas Islands, between the Red Sea and Indian Ocean form, we Fatu Hiva, 15 m. feel C. fieldi warrants species-level designation. The similar Chromis iomelas Jordan & Seale (Figs. ACKNOWLEDGEMENTS 10-12), type locality Samoa Islands, is readily sep- This research was supported by the National Sci- arated from both C. fieldi and C. dimidiata by hav- ence Foundation grants OCE-0453167 and OCE- ing 13 or 14 dorsal soft rays compared to modally 0929031 to Dr. Brian W. Bowen, as well as NOAA 12 dorsal rays, the demarcation of dark brown National Marine Sanctuaries Program MOA No. anterior and white posterior colors anterior to the 2005-008/66882. It was also funded in part by a origin of the anal fin (the pectoral-fin tips passing Natural Sciences and Engineering Research Coun- distinctly posterior to the demarcation), and the cil of Canada (NSERC) postgraduate fellowship genetic differentiation is distinct at mtDNA (see and a National Geographic Society Grant (9024- Fig. 13). It is distributed from the Great Barrier 11) to the second author. We thank foremost Reef and Coral Sea north to Papua New Guinea, Richard Winterbottom, who contributed his pho- and east to New Caledonia, Vanuatu, Fiji, Samoa tographs of the paratypes of Chromis fieldi of Fig- Islands, Society Islands, and Tuamotu Archipelago ures 6 and 8, and whose large collections of this (Randall et al. 1990; Allen 1991). There are no species from the Chagos Archipelago at the Royal valid records of C. iomelas in the western Pacific Ontario Museum provided paratypes for five other north of New Guinea, and none for the islands of institutions. Thanks are also due Dr. Tilman J. Micronesia. Alpermann of the Senckenberg Museum, David The common names Half-and-Half Chromis, Catania of the California Academy of Science, and Two-tone Chromis, and Chocolate-dip Chromis Shirleen Smith of the U. S. National Museum of have been variously used for this complex of Natural History for loans and/or information on species. We propose that Chocolate-dip Chromis specimens in their care; Anthony Nahacky and be reserved for the Hawaiian endemic Chromis Kish Patel of Aquarium Fish Ltd. for genetic mate- hanui Randall & Swerdloff, as first used by Randall rial from Fiji; Wouter Holleman for assistance with (1980); Half-and-Half Chromis for Chromis literature; Drs. Luiz A. Rocha and Matthew T. dimidiata, as by Randall (1983) for C. dimidiata in Craig for genetic material from the Republic of the Red Sea (wrongly illustrated from a photo of C. Seychelles; Dr. Michael Berumen at the King fieldi from Mauritius); and Two-Tone Chromis for Abdullah University of Science and Technology, Chromis fieldi, as by Kuiter (1998) and Kimura et associated staff from the Coastal and Marine al. (2009). Resources Core Lab at the Red Sea Research Cen- Comparative material: Chromis dimidiata: ter, and Drs. Brian Bowen and Michelle Gaither BPBM 14680, 2: 36-44.5 mm, Red Sea, Gulf of for facilitating collections in the Kingdom of Saudi

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Arabia; the Center for Genomics, Proteomics, and EXCOFFIER, R., L., LAVAL, G. & SCHNEIDER, S. 2005. Bioinformatics at the University of Hawai‘i Arlequin ver. 3.0: An integrated software package for pop- (Manoa Campus) for assistance with DNA ulation genetics data analysis. Evolutionary Biology Online sequencing; and Dr. Gerald R. Allen, Helen A. 1: 47-50. FIELD, R. & FIELD, M. 1998. Reef Fishes of the Red Sea. Randall, and Jean Michel Rose for their review of Regan Paul International, London, 192 pp. the manuscript, with valuable suggestions for FOURMANOIR, P. 1954. Ichthyologie et Pêche aux improvement. This is contribution no. 8806 from Comores. Mémoires de l’Institut Scientifique de Madagas- the Hawai‘i Institute of Marine Biology and no. car, sér. A, 9: 189-238. 1533 from the School of Ocean and Earth Science FOWLER, H. W. 1928. The Fishes of Oceania. Memoirs of and Technology of the University of Hawai‘i. the Bernice. P. Bishop Museum. 10: 1-540. FOWLER, H. W. 1946. A collection of fishes obtained in the Riu Kiu Islands by Captain Ernest R. Tinkham, REFERENCES A.U.S. Proceedings of the Academy of Natural Sciences of ALLEN, G. R. 1991. Damselfishes of the World. Mergus, Philadelphia 98: 123-218. Melle, Germany, 271 pp. FOWLER, H. W. & BEAN, B. A. 1928. Contributions to the ALLEN, G. R. & ADRIM, M. 2003. Coral reef fishes of biology of the Philippine Archipelago and adjacent Indonesia. Zoological Studies 42 (1): 1-72. regions. The fishes of the families Pomacentridae, Labri- ALLEN, G. R. & ERDMANN, M. V. 2009. Two new species dae, and Callyodontidae, collected by the United States of damselfishes (Pomacentridae: Chromis) from Indone- Bureau of Fisheries steamer “Albatross,” chiefly in Philip- sia. aqua, International Journal of Ichthyology 15 (3): pine seas and adjacent waters. Bulletin of the United States 121.134. National Museum 100, vol. 7: i-viii + 1-525. ALLEN, G. R. & STEENE, R. C. 1987. Reef Fishes of the FROUKH, T. & KOCHZIUS, M. 2008. 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Poissons de mer des eaux de l’Ile HARMELIN-VIVIEN, M. 1976. Ichthyofaune de quelques Maurice. Proceedings of the Royal Society of Arts and Sci- recifs corallines des Iles Maurice et La Réunion. The ences of Mauritius 3 (2): 191-226. Mauritius Institute Bulletin 8 (2): 69-104. BARNARD, K. H. 1927. A monograph of the marine fishes HASEGAWA, M., KISHINO, K. & YANO, T. 1985. Dating the of South Africa. Annals of the South African Museum 21 human-ape splitting by a molecular clock of mitochon- (2): vii + 419-1065. drial DNA. Journal of Molecular Evolution 22: 160-174. BERNARDI, G. & CRANE, N. L. 1999. Molecular phy- HEEMSTRA, P. & HEMMSTRA, E. 2004. Coastal Fishes of logeny of the humbug damselfishes inferred from Southern Africa. South African Institute for Aquatic Bio- mtDNA sequences. Journal of Fish Biology 54: 1210- diversity and National Inquiry Service Centre, Graham- 1217. stown, xxiv + 488 pp. BOWEN, B. W., BASS, A. L., ROCHA, L. A., GRANT, W. S. HUBERT, N., MEYER, C. P., BRUGGEMANN, H. J., GUÉRIN, & ROBERTSON, D. R. 2001. Phylogeography of the F., KOMENO, R. J. L., ESPIAU, B., CAUSSE, R., WILLIAMS, trumpetfishes (Aulostomus): Ring species complex on a J. T. & PLANES, S. 2012. Cryptic diversity in Indo-Pacific global scale. Evolution 55: 1029-1039. coral-reef fishes revealed by DNA-barcoding provides DE BEAUFORT, L. F. 1940. The Fishes of the Indo-Australian new support to the centre-of-overlap hypothesis. PLoS Archipelago, vol. 8, E. J. Brill, Leiden, xv + 508 pp. ONE 7 (3): e28987. DREW, J. A., ALLEN, G. R. & ERDMANN, M. V. 2010. JOHNS, G. C. & AVISE, J. C. 1998. A comparative sum- Congruence between mitochondrial genes and color mary of genetic distances in the vertebrates from the morphs in a coral reef fish: population variability in the mitochondrial cytochrome b gene. Molecular Biology and Indo-Pacific damselfish Chrysiptera rex (Snyder, 1909). Evolution 15: 1481-1490. Coral Reefs 29: 439-444. JORDAN, D. S. & SEALE, A. 1906. The fishes of Samoa. DRUMMOND, A. J., ASHTON, B., CHEUNG, M., HELED, J., Description of the species found in the archipelago, with KEARSE, M., MOIR, R., STONES-HAVAS, S., THIERER, T. a provisional check-list of the fishes of Oceania. Bulletin & WILSON, A. 2009. Geneious v4.8. Available from of the Bureau of Fisheries 25: 173-488. http://www.geneious.com. KIMURA, S., SATAPOOMIN, U. & MATSUURA, K. (eds.).

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A new species of damselfish (Pomacentridae) from the Indian Ocean

2009. Fishes of Andaman Sea. National Museum of REECE, J. S., BOWEN, B. W., SMITH, D. G. & LARSON, A. Nature and Science, Tokyo, vi + 346 pp. 2010. Molecular phylogenetics of moray eels KHALAF, M. A. & DISI, A. D. 2007. Fishes of the Gulf of (Muraenidae) demonstrates multiple origins of a shell- Aqaba. Marine Science Station, Jordan, Aqaba, 252 pp. crushing jaw (Gymnomuraena, Echidna) and multiple KLUNZINGER, C. B. 1871. Synopsis der Fische des Rothen colonizations of the Atlantic Ocean. Molecular Phyloge- Meeres. II. Theil. Verhandlungen der K.-K. zoologisch- netics and Evolution 57: 829-835. botanischen Gesellschaft in Wien. 21: 441-688. REGAN, C. T. 1917. Additions to the fish fauna of Natal. KUITER, R. H. 1998. Photo Guide to Fishes of the Maldives. Annals of the Durban Museum 1 (5): 458-459. Atoll Editions, Apollo Bay, Victoria, 257 pp. RONQUIST, F. & HUELSENBECK, J. P. 2003. MrBayes 3: LEE, W., CONROY, J., HOWELL, W. H. & KOCHER, T. D. Bayesian phylogenetic inference under mixed models. 1995. Structure and evolution of teleost mitochondrial Bioinformatics 19: 1572-1574. control regions. Journal of Molecular Evolution 4: 1, 54- SATAPOOMIN, K. 2007. A Guide to the Reef Fishes of the 66. Andaman Sea, Thailand. Phuket Marine Biological Cen- LESSIOS, H. A. 2008. The Great American Schism: Diver- ter, vi + 231 pp. gence of marine organisms after the rise of the Central SIDDALL, M., ROHLING, E. J., ALMOGI-LABIN, A., HEM- American Isthmus. Annual Review of Ecology, Evolution, LEBEN, C., MEISCHNER, D., SCHMELZER, I. & SMEED,D. and Systematics 39: 63-91. A. 2003. Sea-level fluctuations during the last glacial MACLEAY, W. 1882. Contribution to a knowledge of the cycle. Nature 423: 853-858. fishes of New Guinea. – No. II. Proceedings of the Linnean SMITH, J. L. B. 1949. The Sea Fishes of Southern Africa. Society of New South Wales. 7 (3): 351-366. Central News Agency, Cape Town, 550 pp. MCMILLAN, W. O. & PALUMBI, S. R. 1995. Concordant SMITH, J. L. B. 1955. The fishes of Aldabra. – Part III. evolutionary patterns among Indo-West Pacific butterfly- Annals and Magazine of Natural History, ser, 1, 8: 886- fishes. Proceedings of the Royal Society of London. Series B, 896. Biological Sciences 260: 229-236. SMITH, J. L. B. 1960. Coral fishes of the family Pomacen- MEEKER, N. D., HUTCHINSON, S. A., HO, L. & TREDE, tridae from the Western Indian Ocean and the Red Sea. N. S. 2007. Method for isolation of PCR-ready genomic Ichthyological Bulletin No. 19: 317–349. DNA from zebrafish tissues. Biotechniques 43: 610-614. SMITH, J. L. B. & SMITH, M. M. 1963. The Fishes of Sey- POSADA, D. 2008. jModeltest: Phylogenetic model averag- chelles. 215 pp. Department of Ichthyology, Rhodes ing. Molecular Biology and Evolution 25: 1253-1256. University, Grahamstown. PYLE, R. L., EARLE, J. L. & GREENE, B. D. 2008. SMITH, M. M. & HEEMSTRA, P. C. (eds.). 1986. Smiths’ Sea Five new species of the damselfish genus Chromis (Perci- Fishes. Johannesburg: Macmillan South Africa, xx + 1047 formes: Pomacentridae) from deep coral reefs in the trop- pp. ical western Pacific. Zootaxa 1671: 3-31. SONG, C. B., NEAR, T. J. & PAGE, J. M. 1998. Phyloge- QUENOUILLE, B., BERMINGHAM, E. & PLANES, S. 2004. netic relations among percid fishes are inferred from Molecular systematic of the damselfishes (Teleostei: mitochondrial cytochrome b DNA sequence data. Mole- Pomacentridae): Bayesian phylogenetic analyses mito- cular Phylogenetics and Evolution 10: 343-353. chondrial and nuclear DNA sequences. Molecular Phylo- SWOFFORD, D. L. 2000. PAUP*: phylogenetic analysis by genetics and Evolution 31: 66-88. parsimony, version 4. Sinauer Associates. Massachusetts: QUÉRO, J.-C., SPITZ, J. & VAYNE, J.-J. 2010. Chromis Sunderland. durvillei: une nouvelle espèce de Pomacentridae TABERLET, P., MEYER, A. & BOUVERT, T, J. 1992. Unusu- (Acinopterygii: ) de l’île de la Réunion ally large mitochondrial variation in populations of the (France, Océan Indien) et 1er signalement four l’ile de blue tit, Parus caeruleus. Molecular Ecology 1: 27-36. Chromis axillaris. Cybium 33(4): 321-326. VORIS, H. K. 2000. Maps of Pleistocene sea levels in RANDALL, J. E. 1980. Guide to Hawaiian Reef Fishes. Har- Southeast Asia: shorelines, river systems and time dura- rowood Books, Newtown Square, PA., 74 pp. tions. Journal of Biogeography 27: 1153-1167. RANDALL, J. E. 1983. Red Sea Reef Fishes. Immel Publish- WINTERBOTTOM, R., EMERY, A. R. & HOLM, E. 1989. An ing, London, 192 pp. annotated checklist of the fishes of the Chagos Archipel- RANDALL, J. E. 1992. Diver’s Guide to Fishes of Maldives. ago, Central Indian Ocean. Royal Ontario Museum Life Immel Publishing, London, 193 pp. Sciences Contributions 145: 1-226. RANDALL, J. E., ALLEN, G. R. & STEENE, R. C. 1990. Fishes of the Great Barrier Reef and Coral Sea. University of Hawai‘i Press, Honolulu, xx + 557 pp.

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aqua, International Journal of Ichthyology

Social structure and reproductive behavior of the callionymid fish bairdi (Gobiesociformes: Callionymidae): with notes on male alternative reproductive tactics

Matthew L. Wittenrich1* and Adeljean L. F. C. Ho2

1) University of Florida, Tropical Aquaculture Laboratory, School of Forest Resources and Conservation, 1408 24th Street SE, Ruskin, FL 33570, USA 2) Florida Institute of Technology, Department of Biological Sciences, 150 West University Blvd., Melbourne, FL 32901, USA

Received: 22 August 2012 – Accepted: 14 December 2012

Abstract zen leben, korreliert deutlich mit der Größe des Reviers. Despite the abundance and diversity of callionymids in Diskutiert werden alternative männliche Taktiken beim marine environments, information on the social structure Fortpflanzungsverhalten, darunter Weibchenmimikry und and mating systems of the group are rare. Most research Sabotageverhalten. has suggested that callionymids are non-territorial, polyg- ynous spawners where male mating opportunities are con- Résumé trolled by dominance hierarchies, which in turn, are gov- Malgré l’abondance et la diversité des Callionymidés en erned by body size. Here, we investigate the social struc- milieu marin, nous manquons d’information sur la struc- ture and reproductive behavior of Callionymus bairdi ture sociale et les système de reproduction de ce groupe. La through observations of wild and captive fish. Males plupart des recherches suggèrent que les Callionymidés ne greater than 8 cm SL held territories ranging in size from sont pas territoriaux, s’accouplent avec plusieurs femelles 6.81-10.9 m2, within which, females maintained territo- quand les occasions des mâles reproducteurs sont con- ries of 0.182-1.41 m2. Subordinate males were found to trôlées par la dominance hiérarchique, qui, de son côté, est move freely through the territories of larger males and induite par la taille du corps. Ici, nous étudions la structure compete for mating opportunities with females. The num- sociale et le comportement reproducteur de Callionymus ber of subordinate males and females living within territo- bairdi par des observations sur des poissons sauvages et rial boundaries correlated strongly with the size of the ter- captifs. Les mâles de plus de 8 cm de LS occupent des ter- ritory. Male alternative reproductive tactics, including ritoires qui vont de 6,81 à 10,9 m2 où les femelles ont des female mimicry and saboteur males, are discussed. territoires de 0,182 à 1,41 m2. Des mâles dominés ont été observés traversant librement les territoires de mâles plus Zusammenfassung grands et entrant en concurrance pour d’éventuels accou- Trotz der Häufigkeit und Vielfalt der Callionymiden im plements avec les femelles. Le nombre de mâles dominés et Lebensraum Meer gibt es bisher wenige Informationen de femelles vivant dans les limites territoriales est en rela- über Sozialstruktur und Paarungsverhalten der Gruppe. tion étroite avec la taille du territoire. Les tactiques de Nach der bisherigen Forschung bilden männliche Leier- reproduction des mâles, comprenant le mimétisme femelle fische keine Reviere und verhalten sich polygyn, wobei die et les saboteurs mâles, sont discutées. Gelegenheiten der Männchen zur Paarung durch Domi- nanzhierarchien gesteuert werden, die wiederum von der Sommario Körpergröße abhängen. Die Ergebnisse der hier vor ge- Nonostante l'abbondanza e la diversità dei callionimidi legten Arbeit gründen sich auf Beobachtungen an freile- negli ambienti marini, ci sono pochi dati sulla struttura benden und gefangen gehaltenen Fischen. Die Männchen sociale e sui sistemi di accoppiamento del gruppo. Gran parte mit einer Größe von über 8 cm SL bildeten Reviere von delle ricerche suggerisce che i callionimidi siano non territo- 6,81-10,9 m2 Fläche, innerhalb deren die Weibchen riali e poligami, dove le opportunità di accoppiamento del Reviere von 0,182-1,41 m2 bilden. Rangniedere Män- sesso maschile sono controllate da gerarchie di dominanza, nchen bewegen sich frei durch die Reviere der großen che a loro volta, sono regolate in base alle dimensioni dell’in- Männchen und bemühen sich um Gelegenheiten zur dividuo. In questo lavoro, si è indagata la struttura sociale e Paarung mit den Weibchen. Die Zahl der rangniederen il comportamento riproduttivo di Callionymus bairdi attra- Männchen und Weibchen, die innerhalb der Reviergren- verso osservazioni su esemplari selvatici e in cattività. I

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

maschi più grandi di 8 cm SL mantengono territori che va- son 1978). Most courtship and spawning behavior riano da 6.81 a 10.9 m2, all'interno del quale le femmine observations within the Callionymidae stem from 2 hanno un territorio di 0.182-1.41 m . Sono stati trovati captive observations (Holt 1898; Wilson 1978; maschi subordinati che si muovevano liberamente nei terri- tori di grandi maschi entrando in competizione per l’accop- Takita & Okamoto 1979; Takita et al. 1983; Zhu piamento con le femmine. Il numero di maschi e femmine et al. 1991; Gonzales et al. 1996; Harrington subordinate presente entro i confini territoriali ha una forte 1997), which may have impeded the ability of correlazione con la dimensione del territorio. Infine, sono researchers to observe such strategies within the discusse tattiche riproduttive alternative adottate dai maschi, family since natural behaviors may be constrained tra cui il mimetismo femminile e i maschi saboteur. in captivity. Here, we investigate the social structure, repro- INTRODUCTION ductive behavior, male alternative reproductive tac- More than 180 species of callionymids (Gobieso- tics, and reproductive success of the lancer drag- ciformes: Callionymidae) are recognized world- onet, Callionymus bairdi, in the wild and in the wide (Fricke 2002). They occupy a diversity of laboratory. Callionymus bairdi (Jordan 1888) is one habitats from shallow sand and mud flats to coastal of two shallow water callionymids from the West rubble fields, seagrass beds, and coral reefs. Their Atlantic, distributed from , southern larvae are often the most abundant representatives Florida and the Gulf of Mexico, and the Bahamas of inshore ichthyoplankton surveys; highlighting to northern South America (Scott & Scott 1988). the ecological importance of this family in marine Virtually nothing is known about the life history of systems (Houde & Lovda 1984). Despite their C. bairdi. prevalence, information relating to the social struc- ture and mating systems of callionymids are rare. MATERIALS AND METHODS Most research has suggested that callionymids are Field Observations non-territorial, polygynous spawners where male Demographic Parameters: The present study was mating opportunities are controlled by dominance conducted in Lake Worth Lagoon, Riviera Beach, hierarchies, which in turn, are governed by body Florida (26°46 59.658’N, 80°2 31.813’W) in size (Awata et al. 2010; Rasotto et al. 2010). depths 2-5 m (Fig. 1). During preliminary obser- Observations, however, suggest that the reproduc- vations we located a study area (20 m × 10 m) tive strategies of callionymids may be more com- where C. bairdi appeared abundant. The habitat plex than previously thought. Takita et al. (1983) consisted of open sand, crushed shell, and rubble observed interruption of spawning ascents by sub- with very little vertical relief. The sea urchin, ordinate males of Callionymus and Thresher (1984) Lytechinus variegatus was often the only form of references potential sneaker males in Diplogram- relief in the rubble fields. Using SCUBA, we con- mus. Interruption of courtship and spawning ducted 14 separate underwater observations from behavior by subordinate males was recently 12 June to 23 September 2011 yielding 37 hours observed in the lancer , Callionymus bairdi of total observations. Observations were timed (Jordan 1888) from the West Atlantic (pers. obs.), with tide changes. Water temperature during warranting further investigation into the social observations ranged from 25-28 °C. structure and mating system of the species. Initial observations were conducted by locating Male alternative reproductive tactics of fishes and following large males, which were the most have been well documented in recent years with a conspicuous in the habitat. These observations broad diversity of strategies being observed across allowed us to map the home range of males, and multiple families (e.g. Kodric-Brown 1986, locate neighboring males, as well as the females liv- Oliveira et al. 2001). Evolutionary studies are rec- ing within the range of respective males. Calliony- ognizing the importance of alternative mating mus bairdi exhibits extreme sexual dimorphism; strategies to understand how and why social inter- males have an enlarged first dorsal fin, and an over- actions give rise to phenotypic alternatives all larger body size. We followed four territory- (Dominey 1984, Gross 1984, 1996). The ability to holding males (identified as T-HM1-T-HM4) detect the presence and diversity of alternative tac- independently for up to two hours per observation. tics, however, may be hindered by captive observa- Once the females and subordinate males living tions of specimens maintained under unnatural within a male-dominated territory were identified, environmental conditions or social densities (Wil- we followed each member of the group for a mini-

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Matthew L. Wittenrich and Adeljean L. F. C. Ho

mum of 30 minutes on two separate observation lengths (SL) of territory-holding males or subordi- days to determine their respective ranges within nate males, and females were determined by cali- the larger territory. Subsequent observations were brated photography (Nikon D200). To test for dif- used to confirm ranges of individuals. Standard ferences in mean standard length of territory-hold-

Fig. 1. Map illustrating the study site for Callionymus bairdi located in Riviera Beach, Florida, USA. The star represents the study site. A) Lake Worth inlet, B) Peanut Island Park, and C) Phil Foster Park.

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

ing males, subordinate males, and females, a one- determined by the number of successful matings way Brown-Forsythe corrected ANOVA was defined as spawning ascents ending in gamete employed with and α-level of P < 0.05 and a Tukey release. HSD post-hoc test after 1/x transformation to Laboratory Observations: On 13 June 2011 six meet normality requirements. To test for the asso- adults were collected from outside the study area ciations between territory size, SL, and abundance, and acclimated to a 30 × 30 × 30 cm glass aquar- a series of both Spearman Rank and Pearson Cor- ium to observe the reproductive success of subor- relation tests were conducted for the appropriate dinate males in detail. To facilitate these observa- datasets. tions, a social group consisting of one territory- Reproductive success: Four separate field observa- holding male (9.4 cm SL), one subordinate male tions (12 hours total) were made around sunset to (7.2 cm SL), and four females ranging in size from quantify reproductive success of territory-holding 4.2-5.8 cm SL were collected to mimic wild repro- and subordinate males. Reproductive success was ductive groups. Fish were fed two times daily with

Fig. 2. Size frequency distribution of Callionymus bairdi specimens (N = 54) surveyed in this study (bottom pane) and back- transformed mean (± SE) standard length for each group (top pane). Different lower case letters denote significantly differ- ent groups after a Brown–Forsythe corrected ANOVA: F = 133.1, d.f. = 2,28.4, P < 0.001. SM = subordinate males, T-HM = territory-holding males.

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Matthew L. Wittenrich and Adeljean L. F. C. Ho

Table I. Territory size, number of subordinate males and females of Callionymus bairdi associated with territory-holding males.

Male SL (cm) Territory size (m2) Number of Subordinate Males Number of Females T-HM1 9.6 10.9 4 14 T-HM2 9.2 8.56 3 11 T-HM3 8.4 7.81 2 9 T-HM4 8.1 6.81 2 5

frozen Mysis shrimp (Piscine Energetics, Canada) Two distinct color patterns were observed in ter- and maintained on a constant photo/thermal cycle ritory-holding males that were identified as (14L/10D at 26 °C) to initiate reproduction. A 30 courtship and agonistic patterns. Courting males minute dusk and dawn period was provided by use exhibited a conspicuous dark mottled body color, a of two 24 W fluorescent bulbs (one 560 nm yellow face with iridescent blue markings, yellow actinic, and one 10,000 K) with independent light and orange dorsal fin with contrasting brown timers. The actinic light was set to go on 30 min- bands, and blue markings along the anal fin. When utes before, and turn off 30 minutes after, the two neighboring territory-holding males encoun- 10,000 K light. Two blue LED lights (moon lights) tered one another the body color in both fish were set at 24L/0D to offer subdued lighting dur- turned pale with a near total loss of yellow and ing nocturnal observations. Substratum consisted orange color from the body and dorsal fin. The of fine sand to facilitate burying behavior. No face turned dark with distinct rows of dots under other substratum was provided. Daily observations the eye, above the mouth and below the preoper- were made from 20:00-22:00 from 8 July to 21 cular spine. During four territory-holding male September 2011. encounters, both males also developed blackish bands under the eye, forward to the mouth and RESULTS rearward to the insertion of the of the preopercular Field Observations spine. Males would approach one another from up Territory-Holding Males: The four large territory- to one meter away with outstretched fins and holding males identified in this study were greater mouths in a slow, hovering manner. Most territor- than 8 cm SL (Fig. 2). These territory-holding ial disputes consisted of two males circling one males commanded territory sizes ranging from another with outstretched fins before aligning side 6.81-10. 9 m2 (Table I, Fig. 3), and the territory by side and performing undulating body waves in size correlated strongly and significantly with the the direction of the opponent. When males were SL of the territory-holding male (Pearson’s r = substantially different in size (>20 mm), the 0.943, P = 0.029, Fig. 4). During most of the smaller male fled without further behavior. For observation periods these territory-holding males males of nearly equal size (<20 mm), territorial were observed to wander haphazardly within their behaviors escalated to include biting the preoper- territorial boundaries consuming prey. Each time a cular spine of their opponent, jaw locking, and bit- territory-holding male encountered a female, he ing fins and body. Territorial disputes rarely lasted would approach her rapidly with fins and mouth more than 5 minutes and resulted in both males outstretched, forming a “T” in front of the female retreating to their respective territories. (“T-configuration”). The male would wave his Subordinate & Saboteur Males: Four subordinate body in undulating motions while moving closer males were found in the territory of T-HM1, three to the female. During daylight hours, females in T-HM2, and both T-HM3 and T-HM4 had two rejected such advances, slowly raising and lowering subordinate males living within their respective ter- their dorsal fin several times before fleeing from the ritories (Table I). The number of subordinate male. This behavior persisted until all of the males in the territory-holding males territories had females within the territory had been greeted. Soon a strong, significant correlation with the size of the thereafter, males tended to bury in the substratum territory that the territory-holding male com- for up to one hour, before reemerging and wander- manded (Pearson’s r = 0.956, P = 0.022, Fig. 5). ing their territory once more. The subordinate males had significantly smaller

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

mean SL than territory-holding males (Brown- observed throughout the day in all parts of the ter- Forsythe corrected ANOVA: F = 133.1, d.f. = ritory and did not seem confined to any particular 2,28.4, P < 0.001; Fig. 2). These males, in all cases, area. The term saboteur male is introduced here to seemed to wander the entire territory of larger, ter- define the role of these males in the social unit. ritory-holding males, often traveling outside the Females: Females had significantly smaller mean territory-holding male’s range by as much as 3 m SL than subordinate males with some overlap in (Fig. 3). Subordinate males were inconspicuously size (Brown–Forsythe corrected ANOVA: F = colored and, with the exception of the dorsal fin 133.1, d.f. = 2,28.4, P < 0.001; Fig. 2) and main- length, resembled females. During times when the tained territory sizes of 0.18-1.41 m2 located larger, territory-holding male was actively courting within the boundaries of the territory-holding or greeting females some of these subordinate male’s territory. The size of female territories corre- males attempted to chase the female away by biting lated significantly, albeit moderately, with the and ramming the female. This behavior was female SL (Spearman’s = 0.645, P < 0.001, Fig. 6).

Fig. 3. Map illustrating the territory of the territory-holding males of Callionymus bairdi in the black outlines with the ter- ritory of each female within the territory-holding male’s territory shaded in grey. The dotted lines represent an observed hap- hazard wandering path of one of the subordinate males within each territory.

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Matthew L. Wittenrich and Adeljean L. F. C. Ho

Thirteen females were observed in the territory of configuration”; Fig. 7A). Undulating body behav- T-HM1, 11 females in T-HM2, 9 females in T- ior, as observed during daylight hours, was not HM3, and 5 females in T-HM4 (Table I). The observed during dusk spawning periods. Instead, number of females in the territory-holding male’s the male quickly positioned himself to the side of territory had a strong, significant correlation with the female (Fig. 7B), inserted his pectoral fins the size of the territory held by the territory-hold- below the female and attempted to lift her off the ing male (Pearson’s r = 0.960, P = 0.020, Fig. 5). substratum. Prior to sunset, females rejected the Females spent near equal time wandering their advances of the male and either buried into the home ranges in search of prey and exhibiting bury- substratum or fled. After an unsuccessful attempt, ing behavior. When two females came within 15 territory-holding male’s rapidly swam in search of cm of each other they flashed their dorsal fins in an other females. Successful spawning occurred when up and down and slightly rolling motion before the male aligned himself parallel to the female with moving on. Agonistic behavior was not observed. his pectoral fin below her body (Fig. 7B). The pair Spawning behavior: Spawning occurred within 30 swam rapidly across the substratum for roughly 2 minutes of sunset in all observations. During obser- m before initiating a spawning assent (Fig. 7C). vations where spawning was observed, the male This horizontal swimming behavior was never in a exhibited heightened courtship activity one hour straight line and always within ~7.5 cm of the sub- before sunset. Ovulation of females was readily stratum. Once the spawning assent was initiated observed at this time as the abdomen became well (Fig. 7D) the rapid swimming stopped and the pair rounded and appeared pink from the stretched skin. slowly ascended to 1.0-1.5 m above the substratum In the absence of saboteur males, territory-hold- (Fig. 7E). At the peak of the spawning rise the male ing males approached females rapidly with out- assumed a curved body position, while the female stretched fins and mouth from a perpendicular remained straight immediately prior to and during position to form a “T” in front of the female (“T- initial gamete release (Fig. 7F). Chains of pelagic

Fig. 4. Pearson’s correlation between standard length squared of territory holding males of Callionymus bairdi with their ter- ritory size (r = 0.943, P = 0.029).

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

Fig. 5. Pearson’s correlation between territory size of territory-holding males of Callionymus bairdi with number of females (♀ r = 0.960, P = 0.020) and subordinate males (♂ r = 0.956, P = 0.022) within their territory.

Fig. 6. Spearman’s rank correlation between standard length squared of Callionymus bairdi females with their territory size (r = 0.645, P < 0.001).

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eggs were released for approximately 3 s. The pair attempting to maintain his position beside the then parted and swam rapidly to the bottom (Fig. female. Of 17 successful spawning rises observed 7G) where the male then began to search for during sunset observations, saboteur males another female and repeat the process. attempted to interrupt the courtship of the terri- Spawning behavior and reproductive success of tory-holding male 10 times. Of these 10 attempts, saboteur males: Whenever a territory-holding saboteur males were able to interrupt the territory- male was present, saboteur males exhibited a very holding male’s courtship on four occasions. In each low profile, often with bodies and mouth pressed case, the saboteur male successfully mated with the firmly to the substratum while displaying an courted female. Interruptions from saboteur males inconspicuous color pattern. As the territory-hold- led to the territory-holding male initiating a rapid ing male courted a target female, the saboteur male horizontal swim before prior to upward spawning would rush in, bite or ram the female in an attempt assents. Four of the 10 interruptions mentioned to make her flee. During these encounters, sabo- above led to the territory-holding male eventually teur males would always swim with darting moving on to court other females. When the sabo- motions while maintaining a low body profile. In teur male was successful in interrupting the - response, the territory-holding male would posi- ing behavior of the territory-holding male it would tion himself between the female and the saboteur then display somewhat subdued yellow and orange male. These events often became quite chaotic with colors on the face and dorsal fin as it courted the the territory-holding male circling the female, female and initiated a spawning assent. Spawning

Fig. 7. Callionymus bairdi spawning sequence showing A) Male approaching female in “T-configuration” B) male displaying to female perpendicularly, C) Male position his pectoral fin under the female and starting to swim horizontally, just above the substratum, D) Initiation of spawning rise, E) mid-water rise, F) Male assumes a curved body while female remains straight in immediately prior to, and during initial gamete release, G) Immediately post final gamete release both male and female make a sharp turn and quickly dart back for the substratum in opposite directions. ♂ = male, ♀ = female.

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

ascents initiated by saboteur males displayed descended slowly directly on top of the territory- reduced (typically less than a ¼ of the distance) holding male, closing his extended dorsal fin (Fig. horizontal swimming prior to the assent. 8C), as it positioned itself between the territory- Laboratory observations of reproductive success holding male and the female (Fig. 8D). From this and female mimicry of subordinate males: Under position, the subordinate male initiated a spawning captive conditions the subordinate male did not rise with the territory-holding male as the female spawn at all when the either fluorescent bulb was departed (Fig. 8D). Up to 15 male-male ascents on. During captive observations the territory-hold- (Fig. 8E-G) were observed following the initial ing male exhibited agonistic and display behavior interruption of the territory-holding male’s toward the subordinate male only near spawning courtship. Male-male ascents were mostly short in time. The subordinate male exhibited submissive duration and height. Following this behavior, behavior, fleeing at the territory-holding male’s which often lasted 10 minutes or more, the terri- approach. tory-holding male moved on to court other females Within minutes of the 560 nm fluorescent bulbs or buried in the substratum. The subordinate male going out the behavior of the subordinate male then courted and spawned with the target female. changed dramatically. As the territory-holding This behavior was observed each evening from 17 male courted a female with extended dorsal fin July to 21 September. The subordinate male suc- (Fig. 8A-B), the subordinate male approached cessfully spawned with a single female each from behind and above. The subordinate male evening. Although the subordinate male attempts

Fig. 8. Callionymus bairdi saboteur male false spawning sequence showing A) territory-holding male (T-HM) approaching and displaying to female in “T-configuration” B) T-HM positioning its pectoral fin under the female and initiating spawn- ing rise, D) Saboteur male (SbM) approaching the T-HM from opposite the female and depressing T-HM’s dorsal fin, F) SbM inserts itself between the T-HM and female, female departs, E) mid mid-water rise of T-HM and SbM, F) both males assume a curved body posture, G) Immediately post assent both males head back the substratum at a leisurely pace in oppo- ♂ ♂ ♀ site directions. T-H = territory-holding male, Sb = saboteur male, = female.

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Matthew L. Wittenrich and Adeljean L. F. C. Ho

to mimic the female, during the final phase the Most evaluations of callionymid behavior have ascent (Figs 7F and 8F), the female maintains a occurred in relatively small aquaria with unnatu- straight posture (Fig. 9A), while the subordinate rally high densities. This may have impeded evalu- male assumes a curved posture in the mirror of the ations of territoriality and alternative mating tac- territory-holding male (Fig. 9B1-B2). tics (Wilson 1978). Males of Callionymus lyra were regarded as territorial by Holt (1898) and Wilson DISCUSSION (1978) who both observed them in aquaria. Terri- Male C. bairdi use different reproductive tactics tory size, polygyny or female territories were not to gain mating opportunities. A minimum body mentioned. Takita et al. (1983) regarded C. enneac- size appears to be a prerequisite to becoming a ter- tis as territorial, although this study too was prob- ritory-holding male, where territory size and the ably impeded by the relatively small confines of the number of females and subordinate males are tank. Six males and 14 females were held in a related to body size. Subordinate and saboteur round tank where spawning behavior was observed males less than 8 cm body length are likely not at night using a “faint” light. They report only two capable of defending a territory and attracting of the six males actively spawned, and maintained mates. Living within the territorial boundaries of a territorial boundary in the middle of the tank. large, territory-holding males offers mating oppor- Takita et al. (1983) further mentions that the non- tunities without the energy expenditure of territo- spawning males would strike at actively spawning rial defense. This is similar to the behavior of pairs during ascents. This behavioral pattern of sneaker males, which live without the cost of terri- sabotage warrants further investigation into the torial defense, but differ in their approach to reproductive behavior of both C. enneactis and the acquiring mating opportunities. The behavior of family at large. It is likely that male alternative sneaker males, darting in to deposit sperm during mating tactics are more common than previously reproduction, might not be effective during pelagic thought. Results of this study suggest that ontoge- spawning rises where egg release occurs relatively netic state or size attributes govern the reproduc- quickly. Interfering with the courtship and mating tive pathway of males and that light levels influ- opportunities of the large male seems a successful ence the success of these alternative tactics. strategy since the territory-holding male was not This is the first study that documents territorial- observed to mate with more than three females in ity and polygyny in callionymids. Spawning behav- a single night. If all females in the territory were ior of C. bairdi observed in the wild and in captiv- receptive to mating, an overall decrease in fecun- ity was similar to that described for other cal- dity would be observed without successful mating lionymids, with the exception of horizontal pair by saboteur males. The diversity and evolutionary swimming prior to the vertical spawning ascent. significance of this behavior should be further This is likely an adaptation to deal with saboteur addressed within the family. males. Swimming rapidly across the substratum

♂ Fig. 9. Callionymus bairdi positions prior to gamete release. A ventral view (A) of territory-holding male ( T-H) assuming a ♀ curved body position, while female ( ) remains straight. A dorsal (B1) and ventral view (B2) of both the territory-holding ♂ ♂ male ( T-H) and the saboteur male ( Sb) assuming a curved body position.

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Social structure and reproductive behavior of the callionymid fish Callionymus bairdi: with notes on male alternative reproductive tactics

may offer the pair an opportunity to flee agonistic and tactics: diversity within sexes. Trends in Ecology & behavior of saboteur males prior to the vertical Evolution 11(2): 92-98. spawning ascent. Striking at the ascending pair as HARRINGTON, M. E. 1997. Behavior patterns and sexual observed by Takita et al. (1983) may be the out- dimorphism in the , Diplogrammus pau- ciradiatus (Pisces: Callionymidae). Bulletin of Marine Sci- come of immediate ascents in the vicinity of such ence 60 (3): 872-893. males. This horizontal swimming has not been HOLT, E. W. L. 1898. On the breeding of the dragonet noted elsewhere, but would likely be an indication (Callionymus lyra) in the Marine Biological Association’s of alternative mating tactics if observed. aquarium at Plymouth; with a preliminary account of the Of particular interest is the probable stochastic elements, and some remarks on the significance of sexual population flux of the species. Three months after dimorphism. Proceedings of the Zoological Society of Lon- the field study we visited the study location to don 66(2): 281-315. determine if the social structure had been main- HOUDE, E. D. & LOVDA, J. A. 1984. Seasonality of occur- rence, foods and food preferences of ichthyoplankton in tained long term. Only one of the original males Biscayne Bay, Florida. Estuarine, coastal and shelf science was present. During this observation we noticed 18 (4): 403-419. several scorpionfish in the vicinity of the study site JORDAN, D. S. 1888. Description of a new species of Cal- and observed predation of a small female C. bairdi lionymus (Callionymus bairdi) from the Gulf of Mexico. by one of these ambush predators. Not surprising, Proceedings of the United States National Museum 10: 501- predators such as the scorpionfish and recently 502. introduced lionfish, probably contribute to the KODRIC-BROWN, A. 1986. Satellites and sneakers: oppor- labile social structure patterns observed (Albins & tunistic male breeding tactics in pupfish (Cyprinodon pecosensis). Behavioral Ecology and Sociobiology 19: 425- Hixon 2008). 432. OLIVEIRA, R. F., CANARIO, A. V. M. & GROBER, M. S. REFERENCES 2001. Male sexual polymorphism, alternative reproduc- ALBINS, M. A. & HIXON, M. A. 2008. Invasive Indo- tive tactics, and androgens in combtooth blennies Pacific lionfish Pterois volitans reduce recruitment of (Pisces: Blenniidae). Hormones and Behavior 40 (2): 266- Atlantic coral-reef fishes. Marine Ecology Progress Series 275. 367: 233-238. RASOTTO, M. B., DE MITCHESON, Y. S. & MITCHESON, AWATA, S., KIMURA, M. R., SATO, N., SAKAI, K., ABE, T. & G. 2010. Male body size predicts sperm number in the MUNEHARA, H. 2010. Breeding season, spawning time, mandarinfish. Journal of Zoology 281 (3):161-167. and description of spawning behaviour in the Japanese SCOTT, W. B. & SCOTT, M. G. 1988. Atlantic fishes of ornate dragonet, Callionymus ornatipinnis: a preliminary Canada. University of Toronto Press, Toronto, 731 pp. field study at the northern limit of its range. Ichthyologi- TAKITA, T., IWAMOTO, T., KAI, S., & SOGABE, I. 1983. cal Research 57 (1): 16-23. Maturation and spawning of the dragonet, Callionymus DOMINEY, W. J. 1984. Alternative mating tactics and evo- enneactis, in an aquarium. Japanese Journal of Ichthyology lutionarily stable strategies. American Zoologist 24 (2): 30 (3): 221-226. 385-396. TAKITA, T. & OKAMOTO, E. 1979. Spawning behavior of FRICKE, R. 2002. Annotated checklist of the dragonet fam- the two , Callionymus flagris and C. richardsoni, ilies Callionymidae and Draconettidae (Teleostei: Cal- in the aquarium. Japanese Journal of Ichthyology 26 (3): lionymoidei), with comments on callionymid fish classi- 282-288. fication. Stuttgarter Beiträge zur Naturkunde Serie A THRESHER, R. E. 1984. Dragonets (Callionymidae). In: (Biologie) 645 (103s): 1-103. Thresher RE, editor. Reproduction in reef fishes. Nep- GONZALES, B., OKAMURA, O. & TANIGUCHI, N. 1996. tune City: TFH Publications, Inc Ltd. p. 334-335. Spawning behavior of laboratory-reared dragonet, Repo- WILSON, D. P. 1978. Territorial behaviour of male drag- mucenus huguenini, and development of its eggs and pro- onets (Callionymus lyra). Journal of the Marine Biological larvae. Suisanzoshoku 44 (1): 7-15. Association of the United Kingdom 58 (3): 731-734. GROSS, M.R. 1984. Sunfish, salmon, and the evolution of ZHU, Y., FURUKAWA, K., AIDA, K. & HANYU, I. 1991. alternative reproductive strategies and tactics in fishes. In: Daily spawning rhythm during spring and autumn Fish Reproduction. (Ed. R. Wooton & G. Potts) 55-75. spawning seasons in the Tobinumeri-dragonet Repomu- Academic Press, London. cenus beniteguri. Nippon Suisan Gakkaishi 57: 1865- GROSS, M. R. 1996. Alternative reproductive strategies 1870.

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aqua, International Journal of Ichthyology

Description of a new species of (Perciformes: ) from the Southern Gulf of Mexico

Jose Tavera1 and Arturo P. Acero2

1) Centro de Investigaciones Biologicas del Noroeste, CIBNOR, La Paz, México. E-mail: [email protected] 2) Universidad Nacional de Colombia sede Caribe, CECIMAR/INVEMAR, Santa Marta, Colombia. E-mail: [email protected]

Received: 22 September 2012 – Accepted: 15 December 2012

Abstract gitudes del rostro, mandíbula superior, aletas pectorales y Nine of the sixteen species of the western Atlantic genus pélvicas, además de la coloración. Es una especie café oscura, Hypoplectrus (Serranidae) are currently recognized to be sin antifaz ni mancha en el rostro, aleta o pedúnculo caudal, distributed in the Gulf of Mexico. Hypoplectrus atlahua n. y con las aletas pectorales transparentes. Presenta una serie sp. is only known from Tuxpan banks and Isla Lobos, de pecas y líneas violeta iridiscente en las mejillas y pecho así Veracruz. It differs from the only other similarly colored como un definido, pero pequeño, punto violeta sobre la species, H. nigricans (i.e black hamlet) in the number of espina plana en la parte posterior del opérculo. gill rakers on the first arch, snout length, upper jaw length, pectoral and pelvic fins lengths, and coloration. It is a dark Résumé brown species, lacking nose spots, mask, caudal fin dark Neuf des seize espèces du genre Hypoplectrus (Serranidae) de spots, peduncle saddle, and pectoral pigmentation, but has l’Atlantique ouest sont actuellement reconnues comme peu- iridescent violet speckles and lines on cheeks and chest, plant le Golfe du Mexique. Hypoplectrus atlahua n. sp. n’a été and also a well-defined but rather small violet dot over the repéré que près de Tuxpan banks et de l’Isla Lobos, Veracruz. dorsal flat spine on the rear edge of the opercle. Cette espèce se distingue de la seule autre espèce à co loration analogue, H. nigricans par le nombre de branchio spines sur le Zusammenfassung premier arc, la longueur du rostre, la lon gueur de la mâchoire Bei neun von sechzehn Arten der westatlantischen Gat- supérieure, la longueur des pectorales et des ventrales, et par tung Hypoplectrus (Serranidae) konnte bisher eine Verbrei- la couleur. C’est une espèce brun fon cé, sans taches nasales ni tung im Golf von Mexiko festgestellt werden. Hypoplectrus masque, ni taches sombres sur la caudale, ni selle pédoncu- atlahua n. sp. ist nur von den Tuxpan-Ufern und der laire, ni pigmentation pectorale, mais elle a une moucheture sla Lobos in Veracruz bekannt. Ihre Vertreter unterscheiden et des lignes d’un violet iridescent sur les joues et bien nette, sich von der einzigen ähnlich gefärbten Art H. nigricans ni masque, ni taches sombres sur la caudale, mais plutôt (Schwarzer Hamletbarsch) durch die Zahl der Kiemen- petite sur le bord arrière de l’opercule. reusen am ersten Kiemenbogen, Schnauzenlänge, Ober- kieferlänge, Länge der Brust- und der Bauchflosse und die Sommario Farbgebung. Die neue Art ist dunkelbraun, bei ihr fehlen Nove delle sedici specie del genere dell'Atlantico occiden- Nasenflecken, eine Maske, dunkle Flecken an der tale Hypoplectrus (Serranidae) sono distribuite nel Golfo del Schwanzflosse, ein Sattel am Schwanzstiel und Farbmuster Messico. Hypoplectrus atlahua n. sp è conosciuta solo dai im Brustbereich, zeigt aber schillernd violette Sprenkel und banchi del Tuxpan e dall’Isola Lobos, Veracruz. Si differen- Linien an Wangen und Brust sowie einen deutlichen, wenn zia dall’unica altra specie dello stesso colore, H. nigricans auch kleinen violetten Fleck über dem dorsalen flachen (volgarmente noto come hamlet nero), per il numero di ras- Stachel am rückwärtigen Ende des Kiemendeckels. trelli branchiali sul primo arco, la lunghezza del muso, la lunghezza della mascella superiore, la lunghezza delle pinne Resumen pet torali e pelviche e per la colorazione. Si tratta di una spe - Nueve de las dieciséis especies de Hypoplectrus (Serranidae) cie di colore marrone scuro, priva di macchie sul muso, di del Atlántico occidental se distribuyen en el Golfo de Méx- maschera, di macchie scure sulla pinna caudale, di sella sul ico. Hypoplectrus atlahua n. sp. solo se conoce de los bajos de peduncolo e di pigmentazione pettorale, ma ha macchioline Tuxpan e Isla Lobos, Veracruz. Difiere de su único con- e linee viola iridescenti sulle guance e sul petto, e anche una génere de coloración similar H. nigricans (i.e vaquita negra) macchia puntiforme viola ben definita sulla spina dorsale en el número de branquiespinas del primer arco, en las lon- piatta del bordo posteriore dell’opercolo.

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INTRODUCTION reverse) 0.16 µl of premixed dNTP’s [10 µM], 0.4 The serranid genus Hypoplectrus Gill, 1861, µl of MgCl2 [25 µM], 0.8 µl of Q solution, 1 µl of endemic to the Greater Caribbean, is one of the Taq, 1 µl of DNA and 5.35 µl of H20, for a 10 µl most fascinating examples of an ongoing marine total reaction. PCR program consisted on an initial evolutionary radiation, with at least 16 described 94°C denaturation of 2 min, 30 cycles at 94 °C for species, based mainly on coloration differences 30 s, followed by 30 s at an annealing temperature (Puebla et al., 2007; Holt et al., 2010; Lobel, of 54 °C, and 60 s at 72 °C with a final extension 2011). Eight species are known from the Gulf of 5 min at 72 °C. of Mexico (Hoese & Moore, 1998; McEachran & All available COI hamlet’s sequences were down- Fechhelm, 2005), including three recently loaded from genbank, accessions numbers can be described (Del Moral Flores et al., 2011; Victor et seen on Appendix Table I. Sequences were cleaned al., 2012). The limits of hamlets species have been and trimmed using Geneious 5.0 (Biomatters, extensively discussed; in the absence of morpho- NZ). A final 615 b.p fragment of the gene was logical and meristic characters, a general consent aligned using the MUSCLE algorithm. The soft- on using color patterns and geographic distribu- ware J model test (Posada, 2008) was used to cal- tion for defining them has proven to be reliable culate gene model of evolution to be used in the (Randall, 1968; Domeier, 1994; Fischer, 1980; maximum likelihood tree search. RAxML (Sta- Heemstra et al., 2002; Del Moral Flores et al., matakis, 2006) was used for this purpose including 2011; Lobel, 2011). Mate selection appears to be the 200 bootstraps. Tree was rooted using mid- the isolating mechanism restricting gene flow point option (Fig. 2). between the different color morphs, as hamlets dis- play strong mating preference for individuals of the same color pattern (Domeier, 1994). During a visit Hypoplectrus atlahua n. sp to a fish market in Tuxpan (Veracruz), we found (Fig. 1) two specimens of an undescribed species of the genus. The aim of this paper is to describe this new Holotype: CICIMAR-CI 8371, 104.1 mm SL, member of such an astonishing clade. Bajo de Tuxpan 21°8’N, 97°16’W, 7 Jun. 2011; hook and line, Tuxpan, Veracruz, México. MATERIAL AND METHODS Paratype: CICIMAR-CI 8372, 75.1 mm SL, Bajo The authors purchased fresh specimens directly de Tuxpan 21°4’N, 97°15’W, 8 Jun. 2011; hook from fishermen at Tuxpan, Veracruz, México. Dig- and line, Tuxpan, Veracruz, México. ital pictures of both fish were taken fresh and spec- Diagnosis: Hypoplectrus atlahua is distinguished imens were frozen after capture to preserve col- from its congeners by the combination of the fol- oration until examination and description. All lowing attributes: Total gill rakers (GR) on the first specimens were measured fresh to the nearest 0.1 arch, including rudiments, 14-16; snout length mm, lengths are given in mm and as percentage of rather small, 11.8-13 % SL; upper jaw large, 18.8- standard length (SL). Measurements follow those 23.2% SL; long pectoral fins, 35.3-38.8% SL, and described in detail by Lobel (2011). Rows of scales pelvic fins, 26.8-27.4 % SL; pelvic fins not reach- above lateral line were counted at the dorsal fin ori- ing anal fin; body basically dark brown with ante- gin; lateral line count includes pored scales over the rior violet speckles, lines, and a well-marked dot caudal fin. Fixed and preserved type specimens of over the rear edge of opercle, lacking distinct dark the new species have been deposited at the Centro spots (nose, pectoral-fin base, and caudal pedun- Interdisciplinario de Ciencias Marinas, Colección cle). Meristic and morphometric data of the type Ictiológica (CICIMAR-CI), La Paz, Baja Califor- series are summarized in Table I. Selected features nia Sur, México. distinguishing H. atlahua from the other Gulf of DNA extractions were performed with the Qia- Mexico species of hamlets can be seen on Table 2. gen extraction kit following manufacturer specifi- Description: Dorsal X, 14 or 15; anal III, 7; dor- cations. The mitochondrial gene COI was ampli- sal and anal soft rays marginally branched; pectoral fied using universal primers COI_VF2_t1 (Ward rays 13; pelvic I, 5; branched caudal rays 19 (18- et al., 2005) and COI_VR1d_t1 (Ivanova et al., 19); mouth terminal. The following morphometric 2007). PCR reactions contained 1 µl of Buffer data are given as percentages of the standard 10X, 0.1 µl of each primer [20 µM] (forward- length: head length (37.6-40.2); snout length

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Jose Tavera and Arturo P. Acero

(11.8-13.0); upper jaw length (18.8-23.2); interor- lateral line and dorsal fin; caudal fin truncate and bital width (5.9-6.8); eye diameter (9.6); maxi- moderately concave; pectoral fins elongated and mum body depth (44.2-49.4); maximum body rounded; pelvic fins do not reach anal fin; the first width (13.8-15.2); predorsal length (36.8-37.4); third of the basal area of the membranes of dorsal preanal length (60.9-63.6); prepelvic length (36.1- and anal fins covered with scales in the form of a 41.3); base of dorsal fin (52.5-55.7); base of anal right triangle, this coverage decreases past the sev- fin (17.8-22.7); caudal-peduncle depth (12.1- enth dorsal ray becoming at the end of the fin 12.6); longest dorsal spine (IV) (16.2-16.6); pec- about a fifth of the membrane total area; jaws teeth toral-fin length (35.3-38.8); pelvic-fin length distinct, small canines and a patch (several rows) of (26.8-27.4). 14 to 16 total GR on first arch, com- viliform teeth at the anterior tip; vomer with a well posed of 2-3 rudiments and 7 moderately long and developed chevron shaped patch of villiform teeth. slender GR on the lower limb, 1 median GR, slen- Life coloration: Body basically dark brown der but shorter than those of the lower limb, and with anterior violet speckles, lines, and a well- 4-5 blunt rudiments on the upper limb; lateral-line marked dot over the dorsal flat spine on rear edge scales 55 to 57; 10 rows of ctenoid scales between of opercle. A rather discontinuous violet line sur-

Fig. 1A-B. Fresh specimens of Hypoplectrus atlahua, n. sp. A) Holotype. B) Paratype.

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Description of a new species of Hypoplectrus (Perciformes: Serranidae) from the Southern Gulf of Mexico

rounding eye, the anterior portion extending as a Common names: We propose jarocho hamlet series of successive dots down to the preopercle; (English) and vaquita jarocha (Spanish) as the four violet lines starting on the inferior fringe of common names for the new species. the operculum and spreading into the chest, the Habits: Nothing is known of the ecology of the second one prolonging over the lateral edge of the new species, beyond that it occurs in shallow ; an additional fifth violet line crossing waters, less than 20 m, where it is hooked by fish- the body behind the pectoral fin from the free mar- ermen. gin of the operculum to the belly but not reaching Distribution: Only known in the Mexican part of the pelvic fin; soft portion of dorsal and caudal fins the Gulf of Mexico, referenced material collected at finely dotted; it lacks distinct dark spots (nose, pec- Bajo de Tuxpan (21°8’N, 97°16’W), and visual toral-fin base, and caudal peduncle spot). Pectoral records while snorkeling at Isla Lobos (21°28’N, fins transparent. 97°13’W), Veracruz. Etymology: Atlahua, used as a noun in apposi- Comparisons: Following the species = color tion; in Aztec mythology, Atlahua (“deity of the morphs rationale, we focus the comparisons on the waters”) was a water goddess, protector of fisher- only other similarly colored species, the black ham- men and archers. let. Hypoplectrus nigricans is one of three widely

Fig. 2. Hamlets COI maximum likelihood tree, including bootstrap support (only nodes over 70% are shown).

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Table I. Morphometrics of H. atlahua holotype and detailed meristic and morphometric comparisons paratype. GR: gill rakers; numbers within brackets corre- with other somehow similar and/or Gulf of Mexico spond to gill rakers rudiments. residing species, see Table II. In order to make data comparable minimum and maximum values of H. Hypoplectrus atlahua nigricans measurements published in Aguilar-Per- Holotype Paratype era (2004) were transformed to percentages of SL. Morphometrics Domeier (1994) described black hamlet body Total length (mm) 131.8 96.2 color as entire blackish, with variable intensity that Standard length (mm) 104.1 75.1 may vary from bluish to brownish with all fins pig- Head length (mm) 39.1 30.2 mented, including the pectorals. Variations in Percentage of SL morphology and coloration among black hamlets Head length 37.6 40.2 (Belize/Mexican-Caribbean and Puerto Rico) have Snout length 11.8 13.0 been detected (Aguilar-Perera, 2004). According to Upper jaw length 16.6 15.7 that author, H. nigricans (Mexican-Caribbean and Interorbital width 6.8 5.9 Belize) have darkish eyes, body and fins blackish to Eye diameter 9.6 9.6 intense dark, darkly pigmented pectoral fins (i.e., Max. Body depth 49.4 44.2 not transparent), pelvic fins short and rounded, Max. Body width 13.8 15.2 not reaching the third anal-fin spine, and a trun- Predorsal length 36.8 37.4 cated caudal fin. On the other hand, black hamlets Preanal length 60.9 63.6 from Puerto Rico have yellow eyes, exhibit a larger, Prepectoral length 37.2 39.0 brownish to grayish, body, transparent pectoral fins Prepelvic fin length 36.1 41.3 (no dark pigment), with anterior part of pelvic and Base of dorsal fin 55.7 52.5 anal fins intense blue and with long and pointed Base of anal fin 22.7 17.8 pelvic fins reaching at least to the third anal-fin Caudal peduncle depth 12.1 12.6 spine, and a lunate caudal fin with pointed lobes. Longest dorsal spine (IV) 16.6 (IV) 16.2 Hypoplectrus atlahua differs from above descrip- Pectoral fin length 38.8 35.3 tions in having dark brownish body, dark brown Pelvic fin length 27.4 26.8 eyes, pectoral fins transparent, pelvic fins short and Percentage of head length rounded, not reaching anal fin; when fresh, soft Snout length 31.5 32.5 dorsal and caudal fins have an orange finely dotted Upper jaw length 44.2 39.0 pattern. Violet speckles, dots, and lines in snout, Interorbital width 18.2 14.6 cheeks, and chest. Eye diameter 25.6 23.8 Moreover, H. atlahua not only varies in color but Meristics also in morphology and total GR count in first Dorsal fin X,15 X,14 arch. Jarocho hamlet has 14 to 16 total elements Anal fin III,7 III,7 including rudiments, compared to 18-21 in the Pectoral fin rays 13 13 black and yellowtail hamlets, 17-23 in the barred Lateral line scales 57 55 hamlet, 17-19 in H. ecosur, and 16-21 in H. flori- Total GR in the 1st arch 16 14 dae. The new species has a shorter snout, 11.8- GR; lower+upper limb (3) 7+1+ (5) (2) 7+1+ (4) 13% SL than those of the black, barred, and yellow Rows of scales above 10 10 tail hamlets. It also has longer pectoral fins, 35.3- lateral line 38.8% SL, and pelvic fins, 26.8-27.4% SL, than those of H. ecosur, H. floridae, H. nigricans,andH. distributed hamlets, although it is not as abundant puella, but comparable in the area to those of H. as the barred or butter hamlets (Domeier, 1994); chlorurus and H. castroaguirrei (Table II). these three species along with H. chlorurus and H. Almost every paper published on hamlets dis- gemma, formally presented for the Gulf of Mexico cusses the uncertainty of color morphs correspond- by Aguilar-Perera and Tuz-Sulub (2010), and the ing to valid species; the low amount of morpho- three recently described H. castroaguirrei, H. flori- logical differences and the low molecular differen- dae, and H. ecosur are the species distributed in the tiation, even with different type of molecular Gulf of Mexico (McEachran & Fechhelm, 2005; markers and/or genes (Graves & Rosenblatt, 1980; Del Moral Flores et al., 2011; Victor, 2012). For Ramon et al., 2003; García-Machado et al., 2004)

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Description of a new species of Hypoplectrus (Perciformes: Serranidae) from the Southern Gulf of Mexico

Table II. Comparisons of H.atlahua. GM: Gulf of Mexico; Bz: Belize; MC: Mexican Caribbean; PR: Puerto Rico. Super- script numbers match data taken from: 1Del Moral et al., 2012; 2McEachran and Fechhelm (2005); 3Victor, 2012; 4Aguilar- Perera and Tuz-Sulub, 2010. 5Aguilar-Perera, 2004; 6Lobel, 2011. Total GR in the first arch includes rudiments. McEachran and Fechhelm (2005) counts on pored scales for H. unicolor may be a typo. - - - - 6 Bz 7.6-11 16.1-17.8 42.5-46.4 16.8-26.9 41.8-44.9 49.8-57.7 22.4-28.6 31.9-32.4 16.1-17.3 5 PR -- -- 5 83.7-99.5 68.0-115.0 70.9-91.4 11.2-13.0 8.9-15.5 13.9-14.2 101.0-118.8 84.3-150.0 MC and Bz 2 GM 2 GM 2 GM ------4 GM 3 Hypoplectrus 12-15 5.3-6.4 GM 12 12 3 74-83 33.4-40.3 39-41 38-42 64-68 65-71 35-43 36-41 13-14 14-15 GM ------2 GM 12-13+6-8 12–13+5–6 11–14+5–7 12-13+6-7 11-15+6-8 12-15+6-7 13-14+7 12-13+1+6-7 11-12+1+6-7 ------1 (n=2) (n=2) (n=2) (n=3) (n=12) (n=20) (n=4) GM GM atlahua castroaguirrei chlorurus ecosur floridae gemma puella unicolor nigricans randallorum Total length (mm)Total 96.2-131.8 98.1-115 Standard length (mm)Standard 75.1-104.1 79.2-93.2 Predorsal lengthPredorsal 36.8-37.4 37.8-39.6 Eye diameterEye Body depth Max. Body widthMax. 44.2-49.4 44.3-48.5 13.8-15.2 9.6 11.9-14.9 9.5-10.5 45-47 39-42 9-10 10.1-10.9 38-42 11.2-11.9 41-43 8-9 36-46 8-9 41-51 44-46 8-9 39.0-48.2 28.8-55.9 8-9 44.2-47 8.0-10.1 6.6-9.5 9.6-10.4 Percentage of SL Percentage lengthHead lengthSnout jaw length Upper widthInterorbital 37.6-40.2 37.9-38.1 15.7-16.6 11.8-13 17.7-18.6 12.3-12.7 38-39 5.9-6.8 17-19 13-14 6.3-7.3 36-39 14.8-16.6 12.9-14.4 17.2-18.4 41-43 9.4-12.6 18-19 37-40 12-14 18-19 38-44 13-16 18-19 37-39 12-13 17-18 38-39 13-14 15.8-19.0 33.7-40.0 12.2-13.0 10.5-17.5 26.2-43.8 7.1-12.4 16.2-18.9 38.5-41.5 10.3-12.2 Preanal lengthPreanal 60.9-63.6 58.4-63.9 Prepectoral lengthPrepectoral fin lengthPrepelvic 37.2-39 36.1-41.3 Base of dorsal finBase of anal finBase Caudal peduncle depth 52.5-55.7 55.4-57.3 12.1-12.6 17.8-22.7 17.9-18.7 12.5 Longest dorsal Spine 16.2-16.6 Dorsal finDorsal Anal fin fin raysPectoral Lateral line scales X,14-15 GR in the 1st archTotal limbGR; lower+upper X,15 14-15 13 55-57 III,7 9-10+5-6 X,14-15 17-18 III,7 14 X,14-15 55 18-21 13-14 III,7 48-53 X,14 17-19 X,14-16 13–14 III,7 16-21 52 X,14-16 13-14 X,13-15 18-20 III,7 X,14-15 52 13-14 17-23 III,7 X,15-16 48-53 13-14 18-22 X,15-16 III,7 48-53 13-15 20-21 X,15 13-14 77-82 III,7 19-21 48-53 III,7 12-13 18-20 51-60 13 13-14 (14) 50-58 III,7 Pelvic fin lengthPelvic 26.8-27.4 24.4-26.7 24-27 17-21 15–16 24-26 23-26 24-25 25-26 Pectoral fin lengthPectoral 35.3-38.8 31.3-33.7 31-32 27-32 27–29 29-30 25-31 29-32 29-31 Rows of scales above LL of scales above Rows 10 12 Morphometrics Meristics Meristics

aqua vol. 19 no. 1 - 21 January 2013 34 AQUA19(1):AQUA 24/01/1312:37Pagina35 Bz: Belize; MC:Mexican Caribbean;PR:Puerto Rico. Superscript numbersmatchdatatakenfrom: 2 Table III 35 McEachran andFechhelm (2005);

atlahua castroaguirrei chlorurus ecosur floridae gemma puella unicolor nigricans randallorum GM GM1 GM2 GM3 GM3 GM4 GM2 GM2 GM2 MC and Bz5 PR5 Bz6 . ColorcomparisonsofallGulf ofMéxicohamletsincludingBelize endemic

Main body color Pale to dark yellow to dark Pale Dark brown Dark bluish, purple or Dark black brownish Dark bluish or bronish black bluish or bronish Dark yellow Pale yellow Pale blue to blach Iridescent Creamy white to pale yellow Creamy yellow Pale dark to intense Blackish to grayish Brownish to light brown Dark 3 Bar ✓✓ ✓ ✓ ✓ Victor, 2012;

Line or spots encircling eye ✓✓ ✓ ✓ ✓ ✓ ✓

Spots or speckles on snout ✓✓ ✓ ✓ ✓ ✓ ✓ Jose Tavera P. andArturo Acero 4 Blotch on snout ✓ ✓✓Aguilar-Perera and Tuz-Sulub, 2010.

Mask on eye ✓✓

Lines on operculum ✓✓ ✓ ✓ ✓ ✓ ✓

Lines on chest ✓✓ ✓ ✓ ✓ ✓

Dot over rear edge ✓✓✓✓v of opercle

Caudal saddle or blotch ✓✓✓ ✓ ✓

A pair of dark rounded ✓✓

spots atthe base of the H. randallorum

caudal fin 5 Aguilar-Perera, 2004;

Pectoral fin pigmented ✓✓✓ ✓ ✓ aqua vol. 19no. 1-21January 2013

Yellow caudal fin ✓✓ 1 Del Moral etal.,2012; . GM:Gulf ofMéxico; 6 Lobel, 2011. AQUA19(1):AQUA 24/01/13 12:37 Pagina 36

Description of a new species of Hypoplectrus (Perciformes: Serranidae) from the Southern Gulf of Mexico

Appendix Table I. Genbank hamlets collecting information and accession numbers for all species used for this paper; Species in alphabetical order. GM: Gulf of Mexico; C: Caribbean.

Species Collection site Genebanck accesion

Hypoplectrus aberrans C, Belize JX488035 Hypoplectrus atlahua (Holotype) GM, Mexico KC333451 Hypoplectrus ecosur GM, Mexico KC333450 Hypoplectrus ecosur GM, Mexico JN312341 Hypoplectrus ecosur GM, Mexico JN312343 Hypoplectrus ecosur GM, Mexico JN312342 Hypoplectrus floridae GM, USA JN444752 Hypoplectrus floridae GM, USA JN444756 Hypoplectrus floridae GM, USA JN444753 Hypoplectrus floridae GM, USA JN444755 Hypoplectrus floridae GM, USA JN444754 Hypoplectrus gemma FJ848375 Hypoplectrus indigo C, Belize JQ841607 Hypoplectrus indigo C, Belize JX488036 Hypoplectrus indigo C, Belize JQ840877 Hypoplectrus nigricans C, Belize JQ840879 Hypoplectrus nigricans C, Belize JQ840541 Hypoplectrus nigricans C, Belize JQ841608 Hypoplectrus nigricans C, Belize JQ840878 Hypoplectrus puella C, Trinidad and Tobago JQ842906 Hypoplectrus puella C, Belize JQ840542 Hypoplectrus puella C, Trinidad and Tobago JQ842907 Hypoplectrus puella C, FJ583582 Hypoplectrus puella C, Belize JQ840880 Hypoplectrus puella C, Cuba FJ583557 Hypoplectrus puella C, Cuba FJ583578 Hypoplectrus puella C, Belize JQ841609 Hypoplectrus puella C, Cuba FJ583579 Hypoplectrus puella C, Belize JX488038 Hypoplectrus puella C, Belize JQ840881 Hypoplectrus puella C, Mexico HQ573385 Hypoplectrus puella C, Trinidad and Tobago JQ842905 Hypoplectrus sp. C, Curacao JQ842169 Hypoplectrus sp. C, Curacao JQ842168 Hypoplectrus sp. C, Bahamas JQ839798 Hypoplectrus sp. C, Dominica JX516091 Hypoplectrus sp. C, Panama JX488042 Hypoplectrus sp. C, Mexico JN312340 Hypoplectrus sp. C, Panama JX488040 Hypoplectrus sp. C, Mexico JN312339 Hypoplectrus sp. C, Belize JQ840882 Hypoplectrus sp. C, Belize JQ840883

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Jose Tavera and Arturo P. Acero

have fueled this debate. One of the astonishing ACKNOWLEDGEMENTS results of the recent paper by Victor (2012), We thank fisherman Miguel Angel from Tuxpan, describing two species of hamlets from the Gulf of Veracuz, José De La Cruz-Agüero, Ana María Mil- Mexico, is the high level of mitochondrial differen- lán, Lucía Campos, Colección Ictiológica CICI- tiation of the two new species from the remaining MAR-IPN, and CECIMAR. JJT was scholarship ones. This amount of differentiation had not been holder from CONACYT. Insightful comments of detected before for any pair of Hypoplectrus species. three anonymus reviewers contributed in the The presence within hamlets of two divergent improvement of the final version of this manu- mtDNA clades largely corresponding to geograph- script. Contribution 389 of the Centro de Estudios ical areas suggests a longer history of restricted gene en Ciencias del Mar, CECIMAR, of the Universi- flow between Caribbean and the Gulf of Mexico dad Nacional de Colombia sede Caribe. species flocks and thus represents a fertile ground for testing speciation hypothesis. Our molecular results (Fig. 2) place H. atlahua REFERENCES deeply nested within H. ecosur. We strongly believe AGUILAR-PERERA, A. 2004. Variations in morphology and coloration in the black hamlet, Hypoplectrus nigricans color and morphological data (see Tables II and (Teleostei: Serranidae). Caribbean Journal of Science, 40: III), following rationale on species delimitation on 150-154. this clade of fishes, are sufficient evidence for spe- AGUILAR-PERERA, A. & GONZÁLEZ-SALAS, C. 2010. Dis- ciation despite the lack of genetic differentiation tribution of the genus Hypoplectrusv(Teleostei: Ser- for this single mitochondrial gene. A thorough ranidae) in the Greater Caribbean Region: support for a analysis of Amplified Fragment Length Polymor- color-based speciation. Marine Ecology, 31: 375-387. phisms (AFLPs) made by Holt et al. (2011) led the AGUILAR-PERERA, A. & TUZ-SULUB, A. N. 2010. Hypoplec- authors to conclude that even with low genetic dif- trus gemma (Teleostei: Serranidae) is not endemic to southern Florida waters. Pan-American Journal of Aquatic ferentiation in Hypoplectrus, color forms do repre- Sciences, 5 (1): 143-146. sent more than just color variants of a single DEL MORAL FLORES, L., TELLO-MUSI, J. & MARTÍINEZ- species. PÉREZ, J. 2011. Descripción de una nueva especie del Different hypotheses have been proposed for this género Hypoplectrus (Actinopterigi: Serranidae) del Sis- lack of genetic differentiation (Ramon et al., tema Arrecifal Veracruzano, suroeste del Golfo de Méx- 2003). One of them claims that hamlet species ico. Revista de Zoología, 22: 1-10. evolved so recently and rapidly that there has not DOMEIER, M. L. 1994. Speciation in the serranid fish been enough time for lineage sorting. The second Hypoplectrus. Bulletin of Marine Science, 54: 103-141. FISCHER, E. 1980. Speciation in the hamlets (Hypoplectrus: hypothesis is associated with ongoing hybridiza- Serranidae): A continuing enigma. Copeia, 1980: 649- tion, which prevents the mtDNA to reflect the his- 659. tory of speciation in the group, a possibility that GARCÍA-MACHADO E., CHEVALIER-MONTEAGUDO, P. P. & has been long recognized, but rarely given much SOLIGNAC, M. 2004. Lack of mtDNA differentiation consideration (Ramon et al., 2003). Hamlet among hamlets (Hypoplectrus, Serranidae). Marine Biolo - hybridization has been reported before both empir- gy, 144: 147-152. ically and experimentally (Fischer, 1980; Domeier GILL, T. N. 1861. Two new species of marine fishes. Pro- 1994). Both alternatives are possible, but discrimi- ceedings of the Academy of Natural Sciences of Philadelphia, nating between them is difficult and out of the 13: 98-99. GRAVES J. E. & ROSENBLATT R. H.1980. Genetic relation- scope of this paper. ships of the color morphs of the serranid fish Hypoplec- We speculate that the fact that H. atlahua have no trus unicolor. Evolution, 34 (2), 240-245. genetic differences with H. ecosur and H. gemma HEEMSTRA, P. C., ANDERSON, W. D. & LOBEL, P. S. 2002. (FJ848375) might be a local and similar case as Serranidae, (seabasses, creolefish, hinds, ham- that of the Caribbean flock where color morphs lets, anthiines, and soapfishes). In: K. E. Carpenter (Ed), can be distinguished even in the absence of com- FAO Species Identification Guide for Fishery Purposes. The plete genetic isolation for this particular gene. It living marine resources of the Western Central Atlantic. Vol. 2. Bony Fishes, Part 1 (Acipenseridae to Grammatidae). would be of great interest to see where in this tree Food and Agriculture Organization of the United the other two recently described species found in Nations, Rome, pp. 1308-1369. the Gulf of Mexico, H. maya and H. castroaguirrei, HOESE, H. D. & MOORE, R. H. 1998. Fishes of the Gulf of would be placed. Mexico: Texas, Louisiana, and adjacent waters (2nd ed.): Texas A&M University Press College Station.

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Description of a new species of Hypoplectrus (Perciformes: Serranidae) from the Southern Gulf of Mexico

HOLT, B. G., CÔTÉ, I. M. & EMERSON, B. C. 2010. Sig- MAN, E. 2007. Colour pattern as a single trait driving natures of speciation? Distribution and diversity of speciation in Hypoplectrus coral reef fishes? Proceedings. Hypoplectrus (Teleostei: Serranidae) colour morphotypes. Biological sciences / The Royal Society, 274: 1265-1271. Global Ecology and Biogeography, 19: 432-441. RAMON, M. L., LOBEL, P. S. & SORENSON, M. D.2003. HOLT, B. G., CÔTÉ, I. M. & EMERSON, B. C. 2011. Lack of mitochondrial genetic structure in hamlets Searching for speciation genes: Molecular evidence for (Hypoplectrus spp.): recent speciation or ongoing selection associated with colour morphotypes in the hybridization?. Molecular Ecology 12: 2975-2980. Caribbean reef fish genus Hypoplectrus. PLoS ONE, 6 (6): RANDALL, J. 1968.Caribbean Reef Fishes (First ed.). New e20394. York: T.F.H Publications, Inc. IVANOVA, N. V., ZEMLAK, T. S., HANNER, R. H & HEBERT, STAMATAKIS, A. 2006. RAxML–VI–HPC: Maximum like- P. D. N. 2007. BARCODING Universal primer cock- lihood–based phylogenetic analyses with thousands of tails for fish DNA barcoding. Molecular Ecology Notes, 7 taxa and mixed models. Bioinformatics, 22 (21), 2688- (4): 544-548. 2690. LOBEL, P. S. 2011. A review of the Caribbean hamlets (Ser- VICTOR, B. C. 2012. Hypoplectrus floridae n. sp. and ranidae, Hypoplectrus) with description of two new Hypoplectrus ecosur n. sp., two new Barred Hamlets from species. Zootaxa, 3096: 1-17. the Gulf of Mexico (Pisces: Serranidae): more than 3% MCEACHRAN, J. D. & FECHHELM, J. D. 2005. Fishes of the dif- ferent in COI mtDNA sequence from the Caribbean Gulf of Mexico: Scorpaeniformes to Tetraodontiformes (Vol. Hypoplectrus species flock. Journal of the Ocean Science 2). Austin: University of Texas Press. Foundation, 5: 1-19. POSADA, D. 2008. jModelTest: phylogenetic model aver- WARD, R. D., ZEMLAK, T. S., INNES, B. H., LAST, P. R. & aging. Molecular Biology and Evolution, 25 (7): 1253- HEBERT, P. D. N.2005. DNA barcoding Australia’s fish 1256. species. Philosophical Transactions of the Royal Society B, PUEBLA, O., BERMINGHAM, E., GUICHARD, F. & WHITE- 360: 1847-1857.

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aqua, International Journal of Ichthyology

Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

Jacques Géry

Received: 11 December 2012 – Accepted: 09 January 2013

Abstract Tetragonopterinae 43 species, 28 preserved in alcohol (including 19 spp. Knodus moenkhausii (Eigenmann & Kennedy, photographed) and 15 species not preserved (some still in 1903) (no photo). aquarium), all photographed. 8 ex., largest 31 mm (poor condition), Station 5. From the upper basins of the rios Tieté, Paraná, Paraguay, Alto Parana basin, km 78 on the road Tres Lagoas Tapajós, Xingú and Tocantins (the last three in the Serra do Roncador). to Campo Grande; Córrego da Lagoa, seemingly dry a few days ago, H. Bleher 13.12.2005. N. 134. Zusammenfassung Depth 3.70 and head 3.75 in SL; dorsal at mid- Präsentiert werden 43 Arten, von denen 28 als Alkohol- body; trunk scales 5/38/3´, 12 predorsal, 14 präparat vorliegen (darunter 19 auf Fotos), sowie 15 nicht around caudal peduncle; base of caudal covered präparierte Arten (einige leben noch in Aquarien), die alle with small scales extending onto the proximal 1/4 auf Fotos gezeigt werden. or 1/3 of the middle of the lobes, not along their Sie stammen aus den oberen Einzugsgebieten der Flüsse border; anal fin iii,18-19, no hooklets observed; Tieté, Paraná, Paraguay, Tapajós, Xingú und Tocantins (die drei letzteren befinden sich in der Serra do Roncador). SO3 entire, except a small “naked” triangle dorso- posteriorally; premaxilla with 4 tricuspid teeth in Résumé the outer row and 4 pentacuspid ones in the inner 43 espèces, 28 préservées dans l’alcool (comprenant 19 row, all teeth usually black, the cuspids long; max- espèces photographiées) et 15 espèces non préservées (cer- illa short, just reaching level of anterior margin of taines encore en aquarium), toutes photographiées. eye, with one broad black tooth; mandible 4 pen- En provenance des bassins supérieurs des rios Tieté, tacuspid followed by about 8 much smaller tricus- Paraná, Paraguay, Tapajós, Xingú et Tocantins (les trois pid teeth. A pale vertical humeral spot just poste- derniers dans la Serra do Roncador). rior to opercle, the chromatophores not dilated, a black longitudinal narrow band up to the end of Sommario 43 specie, 28 conservate in alcol (di cui 19 fotografate) e 15 the middle caudal fin rays, broadened onto the non conservate (alcune ancora in acquario), tutte fotografate. caudal peduncle. Known from Paraguay and also Dal bacino superiore dei rii Tieté, Paraná, Paraguay, Tapa - from Upper Amazon (same species??). jós, Xingú e Tocantins (gli ultimi tre nella Serra do Ronca - Knodus sp. 1 aff. moenkhausii (Eigenmann & dor). Kennedy, 1903) (Fig. 1 in situ). 5 ex., largest 42.7 mm SL, Station 15, Pantanal do Norte, Córrego do Bororo, into Rio Vermelho, near Poxoreu, on the road north of Rondonopolis Bryconinae (fast moving current), H. Bleher 18.12.2005. Brycon juvenile (photo in situ). Largest with depth 3.7 and head 4.12 in SL, ca. 8 cm total length (not preserved). Rio smallest with depth up to 4.25; dorsal fin anterior Paraguay basin, tributary of Rio Formosa near to midbody; no pelvic spines; trunk scales 5 or Pitangeiras, km 30 of the road Bonito to Miranda, 6/37-39/4, predorsal irregular but covering predor- Fazenda in a disturbed countryside H. Bleher sal line, about 12; 14 around caudal peduncle; only 15.12.2005. the base of caudal covered with scales (up to the

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

developed posteriorly, the teeth probably stronger, not black coloured, suggesting a different diet, the maxilla longer with one more tooth; the colour- pattern, through similar, is also different, but a photo of the fish from St. 4. was not available for comparison. Knodus sp. 2 aff. moenkhausii (Eigenmann & Kennedy, 1903) (possibly photo Fig. 2, “Moen - khausia cf. phaeonota with bacteria from Station Fig 1. Knodus sp. 1 aff. moenkhausii. Station 15, Pantanal 16”). do Norte, Córrego do Bororo, into Rio Vermelho, near 4 ex., largest 40.5 mm SL, Station 16, Rio das Poxoreu, on the road north of Rondonopolis (fast moving Mortes basin, km 52 after Primavera do Leste, near current). Photo by H. Bleher, 18.12.2005. Reserva Indigena Sangradouro, H. Bleher 18.12.2005 – (poor condition). proximal third or less); anal fin iii,19; head moder- Largest with depth 3.35 or 3.4 and head 3.8 in ate in length, probably shorter than in most SL; dorsal slightly anterior to midbody; no pelvic Knodus, but not very short as in K. breviceps; snout spines; pectoral and pelvic fins relatively short, not slightly anterior to mandible in profile; maxilla reaching anal fin; trunk scales 5 /38 + 2 on caudal/ reaching to vertical through pupil, with 2 teeth; 3; base of caudal fin covered with scales (up to the premaxilla with an outer, weavy row of 5 tricuspid proximal third or less, not on the margin of the teeth, the last one very close to the maxilla, and an fin); anal fin iii,18; head moderate in length; snout inner row of 4 pentacuspid ones, all the teeth quite slightly anterior to mandible in profile; maxilla heavy; mandible with 4 strong pentacuspid teeth in reaching vertical between eye and pupil, with 2 front, followed by smaller ones on sides; large sub- teeth; premaxilla with an outer, weavy row of 4 tri- orbital developed below, touching the preopercular cuspid teeth, the last one very close to the maxilla, canal, not quite complete posteriorly; prominent and an inner row of 4 pentacuspid ones with the black vertical humeral spot, an indistinct broad lat- concavity anterior; mandible with 4 pentacuspid eral band; no peduncular spot, but the middle cau- teeth in front followed by 7 smaller ones on sides; dal fin rays distinctly marked with black; eye gold; large suborbital as in the form from Station 15, a photo in situ shows the fish blue or bluish, not developed below, touching the preopercular canal, iridescent. not quite complete posteriorly; iridescent, the eye In alcohol, this bluish form differs from the pre- gold; an indistinct humeral spot, an indistinct ceeding one by several characters, some of them broad lateral band; a horizontally oval peduncular rather subjective; the head is shorter, the dorsal fin spot, the middle caudal fin rays distinctly marked more anterior, the caudal less scaled, the SO3 less with black; no live photo available.

Fig. 2. Knodus sp. 2 aff. moenkhausii. Station 16, Alto Rio das Mortes basin, road Cuiabá to Barra do Garça, km 52 after Pri- mavera do Leste, near Reserva Indigena Sangradouro (no more existing). Photo by H. Bleher, 18.12.2005.

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Jacques Géry

This form is quite similar to the two preceeding ing the subopercular canal; a distinct, vertical ones, but the single large specimen available is a lit- humeral spot; a lozenge-shaped peduncular spot tle deeper with a longer head. up to the tip of the middle caudal fin rays; back Inc. gen. sp. aff. Ceratobranchia (Hemi- dark; a grey lateral band from dorsal fin to pedun- bryconini?) (no photo). cular spot. A photo shot in situ shows a fish with- 1 ex., very small, Station 16, Rio das Mortes basin, out brillant colours, except for a shiny spot just km 52 after Primavera do Leste, near Reserva Indi- anterior to the humeral spot (rather indistinct in gena Sangradouro, H. Bleher 18.12.2005. life) and a shiny lateral band visible mostly from Elongate; LL complete, with 35 or more scales; dorsal fin to caudal peduncle. anal fin rather long; large suborbital elongate, not A species difficult to identify in a difficult group completely developed; mouth inferior, with the where the generic differences (between Astyanax, teeth in two rows rather numerous, narrow, close Jupiaba, Deuterodon etc.) are minimal; the geo- together, the external row covering the internal one graphically closest “Deuterodon” minor from Tapa- as in Ceratobranchia. The specimen is in poor con- jós basin has an indistinct humeral spot and a dition, unidentifiable. longer anal fin; “Deuterodon” pedri from Minas Astyanax (Astyanax) aff. pedri (formerly Gerais seems closer morphologically, with the head “Deuterodon” pedri (Fig. 3 “Characid sp. 1”, Sta- shorter, and a few more premaxillary, maxillary and tion 16)). dentary, teeth. 8 ex., largest 40.8 mm SL, Station 16, Rio das Astyanax (Jupiaba) acanthogaster (no photo; Mortes basin, km 52 after Primavera do Leste, near resembling more or less the preceeding, but with Reserva Indigena Sangradouro, H. Bleher pelvic spines). 18.12.2005 – (poor condition) N. 1073. 7 ex., largest ca 41.0 mm SL, (Station 9, Rio Depth 2.9 and head 3.78 in SL (largest); resem- Formosa near Rio Bonito, Alto Paraguay basin, bling the following species of the genus or sub- 15.12.2005 on label N. 812. genus Jupiaba with similar morphometrics, Depth 2.8 and head 3.66 in SL (two largest spec- Deuterodon-like teeth and flat breast, nevertheless imens); preventral area flattened, with pelvic spines without pelvic spines; dorsal fin slightly anterior to reaching to half pectoral fin length, under skin; midbody; trunk scales 6/37/4´, 10 in a regular pre- trunk scales 7/38/5; anal fin iii,20; anterior part of dorsal series, 14 around peduncle; anal fin iv,17; head Deuterodon-like, with multicuspid teeth, but anterior part of head with the facies of a the snout not very elongate, shorter than eye diam- Deuterodon, the maxilla short, reaching to vertical eter, equal to the maxilla in projection; maxilla through anterior margin of eye; premaxilla equally reaching to vertical through anterior margin of eye; short, not horizontal; teeth black, multicuspid, 2 premaxilla 2/5, broadest tooth with up to 9 cusps; pentacuspids on the anterior premaxillary series, 5 maxilla with 3 multicuspid teeth; mandible with 6- broad teeth with up to 9 cusps in the inner one; 7 teeth, anteriormost with 7-9 cusps; a humeral maxilla with 2 narrower teeth, with up to 6 cusps; spot, a longitudinal band from dorsal fin to end of mandible with 8-9 progressively diminuishing middle caudal fin rays, enlarged on peduncle. No teeth from anterior part to posterior part, the photo in situ. broadest with 9 cusps; large suborbital not touch- Astyanax (Poecilurichthys) bimaculatus (L. 1754), (1 ex. Fig. 4, photo marked "Astyanax sp. 1" taken on the spot at Station 22). 2 ex., largest 66 mm SL, Station 22 (not pre- served in situ but photographed, dead in aquar- ium): probably Rio Araguaia basin (near mouth of Rio das Mortes), second river on the road B242 from São Felix to Alto Boa Vista, H. Bleher 19.12.2005. N. 236. Depth 2.5 and depth 3.85 in SL; dorsal fin Fig. 3. Astyanax (Astyanax) aff. pedri. Station 16, Alto Rio das slightly anterior to midbody; predorsal series of Mortes basin, road Cuiabá to Barra do Garça, km 52 after scales regular for 3-4 scales just before dorsal, irreg- Primavera do Leste, near Reserva Indigena Sangradouro (no ular on the anterior two-thirds, not covering the more existing). Photo by H. Bleher, 18.12.2005. median line; trunk scales 7/39/5; anal fin iv,26;

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

Fig. 4. Astyanax (Poecilurichthys) bimaculatus. Station 22, probably Rio Araguaia basin (near mouth of Rio das Mortes into Araguaia), second córrego on the road B242 from São Felix to Alto Boa Vista. Photo by H. Bleher 20.12.2005.

large suborbital not developed, separated from the tributary of Rio Salobro (?) a few km from preopercular canal; premaxilla with 5 tricuspid Bodoquena, km 62 on the road Bonito to outer teeth and 5 pentacuspid inner ones, concave Miranda, H. Bleher 15.12.2005 (photo Astyanax anteriorly; maxilla reaching to vertical through sp. 1). pupil, without teeth; mandible with 5 strong, ante- Depth 2.6 in SL; dorsal fin slightly anterior to rior teeth followed by much smaller ones; humeral midbody; predorsal series of scales regular for 3-4 spot slightly horizontally oval with dorsal and ven- scales just before dorsal fin, irregular rows on the tral extension, followed by a faint, second spot; anterior two-thirds, not covering the median line; caudal fin pattern with a peduncular lozenge- trunk scales 7/(18)36/5´; anal fin iv,26; sexual shaped spot prolonged as a band onto the middle hooks numerous; large suborbital not developed, caudal fin rays up to their tips and extending ante- separated from the preopercular canal; premaxilla riorly to beneath dorsal fin; scales of the dorsum with 5-6 tricuspid outer teeth and 5 pentacuspid and flanks black bordered, forming a net; the 3 inner ones; maxilla relatively short, reaching to ver- photos shot in aquarium shows the anterior third of anal fin and the base of the caudal fin lobes rosy to dark red. See also A. argyrimarginatus Garutti 1999 “Ara- garças, Rio Araguaia” - id (group bimaculatus). Astyanax (Poecilurichthys) asuncionensis or n. sp.?? – (one photo in aquarium, fish coming from Station 11, road Bonito-Miranda km 62 (Fig. 5, fish from Station 11, on the road Bonito-Miranda km 62, Paraguay basin). 1 ex. male, 54.0 mm SL, Station 9, Rio Fig. 5. Astyanax (Poecilurichthys) asuncionensis or n. sp.?? For mosa near Rio Bonito, Alto Paraguay basin, Station 11, Alto Paraguay basin, tributary of Rio Salobro H. Bleher 15.12.2005 (one photo in aquarium (?) a few km from Bodoquena, km 62 on the road Bonito labelled A. cf. bimaculatus (with another species, to Miranda, same biotope; probably same fauna. Photo by not preserved, Station 11, Alto Paraguay basin, H. Bleher 15.12.2005.

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tical through anterior margin of eye, with one scales below the lateral line is dichotomously small tooth; mandible with 5 strong, anterior teeth branched above the origin of the ventral, the main followed by much smaller ones; upper profile row being apparently deflected; other rows simi- rather pointed, lower lip developed into a flap larly branched; the rows of scales above the front of owing to probable lack of oxygen; humeral spot the anal are all oblique; from above the second almost round, not clearly horizontally oval as in third of the anal, there are two or three series of “typical” A. bimaculatus, followed by a faint second scales parallel with the lateral line”; anal fin long, spot; caudal pattern “typical” of the species; spots usually 40-46 rays in total (range 36-49), begin- on scales as in A. paraguayensis (renamed A. asun- ning under the first dorsal fin ray(s), its border cionensis), very indistinct on the specimen in alco- straight; mouth small with an almost vertical, short hol, but about 5 midlateral rows of spots in the maxilla with 0-2 teeth; colour-pattern quite com- photo (if it is the same species). mon in the Tetragonopterinae, with usually one or Discussion: the morphometrics of the single two humeral spots and a caudal spot at the end of specimen (one more in aquarium) agree with the the peduncle or astride the peduncle and base of common form; however, the slightly different caudal fin. They form putatively a clade, possibly humeral colour-pattern, the high number of outer sister group of Astyanax s.l., from La Plata and São premaxillary teeth (5-6 cf usually 4) and chiefly the Francisco basins to Venezuela. This group includes incompleteness of the lateral line making it techni- the following species (in chronological order): cally a Hyphessobrycon, suggests that we are dealing Tetragonopterus spilurus Valenciennes in Cuvier & with a distinct, possibly endemic, form. This will Valenciennes, 1849 (Guianas). remain putative until more collections. Tetragonopterus hauxwellianus Cope, 1870 (Ama- Ctenobrycon sp. (photo only) (Fig. 6), Station 13, zon basin). Posada Albuquerque near Albuquerque, on the road Tetragonopterus multiradiatus Steindachner, 1876 Campo Grande to Corumba, H. Bleher 16.12.05. (Tefé, Amazon basin), a junior of the pre- Scales and anal fin rays not counted. Habitus of a ceeding. Ctenobrycon (see comments); depth probably 2.35- Tetragonopterus correntinus Holmberg, 1891 2.45 in SL. Unpaired fins slightly rosy. (Corrientes, Argentina). Comments: a few Tetragonopterinae close to Tetragonopterus erythropterus Holmberg, 1891 Astyanax share some derived characters (at least in (Buenos Aires, Argentina). part): body relatively deep and compressed, rhom- Astyanax pellegrini Eigenmann & Kennedy, 1903 boid, with the original of dorsal and anal fins at (Tetragonopterus multiradiatus Eigenmann & mid-body; scales cyclo-ctenoid, at least in adults in Kennedy, non Steindachner, 1876 (Rio Paraguay). the preventral region, rather numerous, 9-13/40- Astyanax alleni Eigenmann & McAtee, 1907 (rio 55/7-11; caudal fin naked; predorsum irregularly Paraguay); redescribed by Britski & al 1999 as a scaled or even naked as in Poecilurichthys; some or Ctenobrycon (possibly a geographical form of T. many interpolated scales below lateral line, possi- hauxwellianus). bly in all species, their extent depending on the Psellogrammus kennedyi Eigenmann, 1908 (Rio species; this structure in C. alleni (the most derived Paraguay and Rio São Francisco, according to in this respect) is described by Eigenmann (The Eigenmann in Amer. Char. and Britski & al, 1984). American Characidae) in these terms: “the row of Redescribed by Britski & al 1999 as a Psellogrammus . Ctenobrycon rhabdops Fowler, 1913 (Rio Madeira), a synonym of T. hauxwellianus. The genus available for those species is Cteno- brycon Eigenmann, 1908 (type species Tetra- gonopterus hauxwellianus Cope), synonym Apoda - styanax Fowler, 1911 (type species Tetra gonopterus spilurus), possible synonym Psello grammus Eigen- mann, 1908 (type species Psellogrammus kennedyi Eigenmann). Psellogrammus differs from Ctenobry - con only in having an irregular lateral line, which Fig. 6. Ctenobrycon sp. Station 1, near Station 12 at Posada can be complete or incomplete, and more often Albuquerque. Photo by H. Bleher 16.12.05. irregular (“stuttering” sensu Eigenmann).

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

According to the descriptions, mostly by Eigen- head); unpaired fins rosy; interpolated rows of mann, the species differ in the following way: scales beginning above origin of anal fin ...... C. alleni (Rio Paraguay basin: Corumba, Asun- Artificial key to Ctenobrycon cion, Rio Otuquis; rare). 1a. Predorsum scaled with irregular rows 6a. Depth 2.5 in SL; anal fin 45; trunk scales 1b. Predorsum naked except near base of dorsal fin. 8/47/7-8; eye 1.7 in interorbital, 3.7 in head .. 2a. Lateral line complete (except in the Rio ...... C. correntinus (Corrientes, rare). Paranahyba); caudal spot usually vertical, not 6b. Depth 2.5-2.8 in SL; anal fin 41; trunk scales very apparent; largest...... size 96 mm SL). 9-11/45-50; eye 1.0 in interorbital (3.5-2.75 in 2b. Lateral line “stuttering” or incomplete (depth head); interpolated rows of scales beginning 2.16-2.60 and head 4.0-4.33 in SL; anal fin above anal fin only...... rays 40-46; trunk scales 10/40-46/8; a few C. pellegrini (Bahia Negra, Corumba, Asun- interpolated scales over anal fin; caudal spot cion, Villa Hays; not common; probably a larger than in C. hauxwellianus, roundish; no junior synonym of C. correntinus). red streak in front of anal fin; largest size ca 50 mm SL)...... When I studied the Paraguay collection in C. kennedyi (common in Rio Paraguay basin, Geneva, I identified some 400+ specimens as C. rare in Rio São Francisco basin). kennedyi and none as C. alleni (of which Eigen- 3a. Depth 2.25-3.0 and head 4.25-4.50 in SL; a mann mentioned 23 ex., usually sympatric). Thus red streak in front of anal fin ...... the two forms are likely to be sibling (then I over- C. spilurus (abundant in certain places in the looked this), or synonyms. Another difficulty: both Guianas). have a “hump” on the nape in Eigenmann’s figures, 3b. Depth 2.0 in average and head 3.7-4.0 in SL; more pronounced in C. alleni. The latter is also no red streak in front of anal...... much bigger: 104mm total versus 46mm total. A C. hauxwellianus (common in Amazon, rare in photo shot by H. Bleher (2005) in a spot not far Rio Parahyba). from that of C. alleni (retouched photo of speci- 4a. Lateral line 48-55. mens from Corumba) does not show the hump, 4b. Lateral line 42-50. and it seems that the lateral line is stuttering like 5a. Depth 2.65 in SL; anal 45; lateral line 55; eye that of C. kennedyi. The photographed specimens 0.66 in snout, 0.8-0.83 in interorbital; are still alive and may be studied in the future. In unpaired fins bright red...... tetras, important characters such as ctenoid scales, ...... C. erythropterus (Buenos Aires; rare). incomplete lateral line, naked predorsal line etc. are 5b. Depth 2.0-2.2 in SL; anal fin 40-45; lateral line not visible on photos. Finally there is the question 48-50; eye 1.0-1.1 in interorbital (2.75 in of the status of C. correntinus (Corrientes) and C.

Fig. 7. Moenkhausia intermedia. Station 9, Rio Formosa, Alto Paraguay basin, near Bonito. Photo by H. Bleher, 15.12.2005.

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erythropterus (Buenos Aires) succinctly described scales not very numerous, not on the border of the by Holmberg in 1891, cited as inc. sp. in CLOFF- lobes; maxilla long, reaching to pupil’s level, with 4 SCA. Photos from Argentina or Paraguay are teeth; colour-pattern characteristic (several exam- needed. The only sure fact is that C. erythropterus, ples are pale, having possibly been kept alive for a rare fish, has bright red fins (thus the name, and some time and dead in bag). Holmberg insisted on this). Some specimens Moenkhausia oligolepis (Günther, 1864) (Fig. 9 apparently belonging to this species have been in situ, from Station 20). studied in Geneva from Paraguayan Chaco, but the 2 ex., largest 37 mm, Station 20, on BR 242: dirt coloration in vivo was not recorded. road 118 km before São Felix do Araguaia, small Moenkhausia intermedia Eigenmann, 1908 (Fig. córrego not destroyed among fazendas, H. Bleher 7 from Station 9, shot in situ, labelled Characoid 19.12.2005 (probably Chapadinha, Alto Araguaia sp. 3 = Moenkhausia intermedia). basin (bad state) N. 168.1-2.2006. 9 ex., largest 36.4 mm SL, Station 4, km 54 on Lateral line complete, 29 pored scales. This po- the road Tres Lagoas to Campo Grande, H. Bleher pulation, according to a photo in life, has a pale 13.12.2005. N. 248.1-9.2006 – Alto Parana and yellow eye, not red. Alto Paraguay basins (same species? in Rio For- Small Tetras: 1) Extra-Amazonian basin. mosa, (marked M. cf. intermedia), and Station 7 Hyphessobrycon callistus (Boulenger, 1900) (just west of Campo Grande. near entrance to BR (photo in habitat, Alto Paraguay, plus two photos 262, "Pirapitinga", Altro Pardo basin, photo cf. le - in situ, (Fig. 10A-B) male and female from Station pi dura, not preserved). 1 (Rio Tiété basin). Depth 3.40 and head 4.15 in SL; dorsal fin at mid- 11 ex., largest 27.75 mm SL, Station 1, Rio Tiété body or slightly posterior; trunk scales 5/35/3, pre- dorsal 11: caudal fin scaled up to 2/3 of the lobes; anal fin iii,20; premaxilla 4-5 tricuspid/5 pentacus- pid; maxilla long, up to pupil’s level, apparently without tooth; a longitudinal line broadened at dor- sal level and forming a lozenge on peduncle, nar- rowed on caudal and ending on the tip of the mid- dle caudal rays; tips of lobes black without white or hyaline margin; a photo in aquarium of a specimen collected Station 7 (Pirapitinga) shows no striking colours; body pale green, caudal fin marks grey, not as well delimited as in the preserved specimens; a sil- very band. The photos from Paraguay Station 7 and Fig. 8. Moenkhausia phaenota. Station 7, Alto Pardo basin? (into Paraná), road Campo Grande to Corumba Vazante 9 (without specimens) are difficult to compare with do km 2 just west of Campo Grande, dirt road to Pirapi- the specimens from Parana (without photo). tanga, sign: Pousada Sol. Photo by H. Bleher, 14.12.2005. These specimens represent probably the nominal form. The extended distribution suggests a poly- typic species. Moenkhausia phaenota Fink, 1979 (Fig. 8. in situ and one in aquarium). 18 ex., largest 34.85 mm SL, Station 17 and 24, Rio das Mortes or Suia Missu basin, west flowing unnamed córrego in Fazenda Ipuá, just north of Agua Boa on the road Barra do Garça to São Felix do Araguaia, Bleher 19.12.2005. N. 943.1- 18.2006 (note: 6 ex. had been put in the bag labelled photo 1 (pale, probably kept alive and died in the field). Fig. 9. Moenkhausia oligolepis. Station 20, dirt road on BR Depth 3.45-3.50 in SL; scales about 38, 5´/3´, LL 242, 118 km before São Felix do Araguaia, small córrego “stuttering”, lacking on the middle of the body or not destroyed among fazendas, (probably Chapadinha, at the end of the peduncle on 5-6 scales; caudal Alto Araguaia basin). Photo by H. Bleher 19.12.2005.

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

A

A photo shot in the “Aquarium natural” (Station 9, Rio Sucuri) shows the same pattern; the whole body and the fins are cherry-red; the humeral spot and anal fin border are much less conspicuous than the deep black flag of the dorsal fin, which seems the prominent identification mark for members of the species. Hyphessobrycon luetkeni (Boulenger, 1894) (Fig. 11). B 1 ex., 29.7 mm SL, Station 11, tributary of Rio Formosa near Pitangeiras, km 30 of the road Fig. 10A-B. Hyphessobrycon callistus, male (A) and female Bonito to Miranda, Rio Paraguay basin, H. Bleher (B). Station 1, Rio Tiété basin, km 458 from São Paulo, after Araçatuba on the BR 300 "Marechal Rondon"; small 15.12.2005. N. 324.1.2006. creek flowing north, in a mostly destroyed farm. Photo by Depth 2.85 in SL; dorsal fin slightly anterior to H. Bleher 12.12.2005. midbody; trunk scales 5/(15)35-36/4; predorsal series complete but irregular; anal fin iii, 20 or 21; basin, km 458 from São Paulo after Araçatuba on large suborbital surrounded by a rather broad the BR 300 “Marechal Rondon”, H. Bleher “naked” zone (difference with H. santae); premax- 12.12.2005. N. 181. illa with 3 broad tricuspid teeth in the outer row Depth 2.65 in SL; scales 6/(6)33/4 (11 transverse and 5 broad flat pentacuspid teeth in the inner row total), 14 around caudal peduncle; premaxilla teeth (difference with H. anisitsi); maxilla of moderate 2/5, pentacuspid, regular; maxilla relatively short, length, reaching to pupil’s level, with 2 broad pen- with 2 teeth; humeral spot vertical, long, lozenge- tacuspid teeth; dentary with 5 broad pentacuspid shaped, extending astride the lateral line 3 scales teeth in front, followed by a series of minute teeth posterior to opercle onto ´4´ transverse scales; dor- laterally; a conspicuous triangular humeral spot sal fin almost entirely black in some specimens, its close to the opercle; a conspicuous spot almost the base red; anal fin border not prominent, but last entire depth of the caudal peduncle, with a black rays jet black, forming a blotch. This is the most line on the middle caudal rays. No photo in situ, eastern record of the species (first collected in the colour alive not known (?). upper Rio Paraguay). The photo of a male and a Hyphessobrycon herbertaxelrodi Géry (Fig. 12). female shot in situ shows the black anal fin margin Not preserved, Station 14, Córrego Gaucho north relatively narrow in this eastern population. of Coxím, Alto Paraguay basin. Coloration typical.

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Table I. The discriminant characters of four similar Hyphessobrycon species from the region.

Species Bleher – Station 17 H. loweae H. mutabilis H. heliacus

Nr of sp. 7 ex. 5 ex. 8 ex. 43 ex Max. size 29.6 mm 31.5 mm 26.9 mm 27.8 mm Base of caudal 1/3 scaled 10% scaled not scaled not scaled Depth 2.9-3.0 2.25-2.85 3.0-3.4 2.0-2.65 Head 3.65-3.70 3.3-3.6 3.45-3.65 3.30-3.75 Dorsal anterior posterior middle/ant. middle/ant. Longest ray not to to caudal not to to caudal adipose adipose Pectoral to pelvic to pelvic to pelvic to pelvic Pelvic overlapp. overlapp. not to anal overlapp. Squ.long. (9)37 (7)30-31 (6)31-32 (7-9)30-34 Squ.transv. 5/3 6-7/4 5/3´ 5/4-6/5 Anal br. 21 19-20 17-20 20-24 SO3 very large 1/2 cheek 1/2 cheek ? Maxilla long,thin ? long,thin ? Mx teeth 4 broad 2-3 broad 3-7 tri-con. 1-3 broad Pmx teeth 3/5 3-4/5 3/5 2-3/5 Broadest teeth up to up to up to up to 7 7 cusps 7 cusps 3 cusps 7 cusps Humeral spot none vertical none (humer. vertical (faint band) hiatus) Peduncle light zone no light no light no light Caudal middle, not not to end to end (?) vary large to end not to end Body yellow/blue golden green/golden golden golden band fins red/orange

Hyphessobrycon? n. sp. (Fig. 13, a red finned Depth about 3.4 in SL; dorsal about at midbody; species with lateral band). No N.G. (photo in sq. 5/(7)30-31/3, anal fin with ca, 23 branched rays; aquarium). maxilla long, up to pupil’s level, apparently without 3 ex., largest 19 mm SL, Station 4, Alto Parana teeth; premaxilla with 2 tricuspid and 5 broad tri- basin, km 54 on the road Tres Lagoas to Campo cuspid to pentacuspid teeth; large suborbital not Grande, H. Bleher 13.12.2005. developed, with a broad naked zone all around;

Fig. 11. Hyphessobrycon luetkeni. Station 11, Alto Paraguay Fig. 12. Hyphessobrycon herbertaxelrodi. Station 14, Cór- basin, tributary of Rio Salobro (?) a few km from Bodoquena, rego Gaucho north of Coxím, road Córrego Grande to km 62 on the road Bonito to Miranda, same biotope, proba- Cuiabá, into Soja plantation, Alto Paraguay basin. Photo bly same fauna. Photo by H. Bleher, 15.12.2005. by Bleher, 17.12.2005.

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

Fig. 13. Hyphessobrycon? n. sp. Station 4, Alto Parana basin, km 54 on the road Tres Lagoas to Campo Grande, tiny creek flowing south into cattle field. Photo by H. Bleher, 13.12.2005. Rather bad state, all fins rotten; a thin lateral band Rio Araguaia basin, Corrego Chega de Perto before from dorsal fin to end of middle caudal rays (?), per- São Felix do Araguaia, H. Bleher 19.12.2005. N. haps a little expanded onto the peduncle, preceeded 570.1-3.2006. by an aggregate of chromatophores from opercle to Depth 2.66 in SL; dorsal fin anterior to midbody; dorsal fin level; snout distinctly black. If a Hyphesso- pectoral and pelvic fins relatively developed, each brycon, differs from H. cachimbensis which has a reaching the next respective fin; trunk scales humeral spot, scales 6/4 and 14-17 branched anal 5/(5)33/3; anal fin iv,19; great suborbital devel- rays, and from H. vilmae which has no peduncular oped, not quite reaching the suborbital canal; teeth spot and bluish fins, and 19-20 branched anal fin relatively broad, broadest with 7 cusps in the front rays. However, the caudal fin rays are so damaged part of the mandible and in the second premaxil- that one cannot be sure that the apparent absence of lary series, 3 tri-to pentacuspid teeth in the first scales is “natural”: the fish might be a Hemigrammus. series, which is external, with the lips not devel- In the habitus and locality, it fits with the photo oped, black coloured between the teeth; maxilla in aquarium with the following comments: “Photo with 1 or 2 teeth, long, reaching pupil; mandible 4, collected only 3 specimens; I think not one pre- with 4 front teeth and about 8 much smaller ones served !” (the concordance of the photo and the on sides. A broad, dark, lateral band, not well dis- specimens is under debate with the collector). This tinct in alcohol, from opercle to beneath dorsal fin, photo shows the resemblance with a Hasemania continued up to the end of the middle caudal rays but with an adipose fin; a black lateral band, most by a black, much more distinct but narrower line, pronounced on caudal peduncle, where it is some- expanded onto the whole depth of the peduncle; what expanded in its posterior part and prolonged base of anal fin with a dark line; region above anal apparently up to end of middle caudal ray; pelvics fin, up to the lateral line, darkened. and unpaired fins, as well as anal base region, This small sample could well be classified within bright red; colour extending onto the body at dor- the small group of species with a distinct caudal spot sal and anal level; head structures not distinct. and without humeral spot; however, no species cor- responds, whereas in the group with longitudinal Small Tetras: 2) Hyphessobrycon and Hemigram- pat tern, it corresponds well to Hyphessobrycon ca - mus from Serra do Roncador. chim bensis, owing to morphometrics and locality; in Hyphessobrycon aff. cachimbensis Travassos, vivo, the fins are said to be yellowish (they are bluish 1964, differing in colour-pattern (no photo). in the near Hyphessobrycon vilmae). 31 ex. from Alto 3 ex., largest 21.45 mm SL, Station 19 and 20, Aripuana have about the same morphometrics, but

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the band is entire from opercle (included) to end of The collecting spot of Bleher being within the caudal fin, surmounted by a red band as in Hyphesso- limits of the collections of the Roncador Expedi- brycon heterorhabdus. Material for comparison, 4 ex. tion, it is necessary to check the toponyms to see to from Rio Batovi (near type locality) are not at hand which brook cited by Rosemary McConnell corre- (in Geneva). According to photos, examples from sponds with Station 17 of Bleher. According to the upper Tocantins in MTD show a very large eye. It is maps, the toponyms have changed 3 times since likely that the present specimens belong to a new 1967. For example, Xavantina was renamed Min- form, owing to the differences in colour-pattern. istro Jao Alberto (1980 map). Hyphessobrycon lowae Costa & Géry, 1994 (pho- The specimen photographed is only tentatively tos only) (Fig. 14). referred to H. lowae, because it does not exist in the Not preserved; 2 photos labelled Hyphessobrycon preserved lot; only the examination of specimens aff. elachys n. sp., Station 17 (photos in situ). Rio can tell. There are at least 2 sympatric species, das Mortes or Suia Missu basin, west flowing equally rare, possibly mimetic, as shown by the unnamed córrego in Fazenda Ipuá, just north of presence of the Hyphessobrycon described below. Água Boa on the road Barra do Garça to São Felix Hemigrammus aff. micropterus n. sp.? (Fig. 15A-B, do Araguaia, H. Bleher 19.12.2005. male & female Characid sp. 1, Station 17). The photos show two males with the longest dor- ? Hemigrammus sp. aff. brevis Seegers & Géry, sal rays dark and prolonged, probably reaching adi- 1989. pose if depressed, a probably vertical humeral spot ? Hemigrammus sp. from Serra do Roncador, Géry followed by a light zone, and a caudal spot begin- (unpublished). ning above the last fourth of the anal fin and con- 7 ex., largest 29.6 mm SL, Station 17, Rio das tinuing on the lower part of the peduncle then onto Mortes or Suia Missu basin, west flowing unnamed the middle of the fin up to the tip of the middle cau- córrego in Fazenda Ipuá, just north of Agua Boa on dal fin rays, with a light spot on the upper part of the the road Barra do Garça to São Felix do Araguaia, peduncle and on the base of the lobes, a pattern Bleher 19.12.2005 (with Characidium sp., Thaye - ressembling that of Hemigrammus hyanuary or ria cf. boehlkei and Moenkhausia phaeonota, proba- Hemi grammus haraldi, for example. Paratypes of bly also Hyphessobrycon loweae, not preserved). Hyphessobrycon loweae Costa & Géry, 1994 (type locality a tributary of the Rio Culene, upper Rio Xingú basin), are similar to the photos; they have no scales on the caudal fin and thus belong to Hyphes- sobrycon in its classical sense. They were collected in the Lago do Leo and Córrego do Gato, upper Suia Missu basin, 206 km north of the ferry on the Rio das Mortes (Alto Xavantina), some minutes north of the 13th parallel, around Divinéia (ex-Cascalheira). A

B

Fig. 15A-B. Hemigrammus aff. micropterus, male (A) and Fig. 14. Hyphessobrycon lowae. Station 17 (not indicated female (B). Station 17 (not indicated on the map, certainly on the map, certainly close to site in Station 23). Rio das close to site in Station 23). Rio das Mortes or Suia Missu Mortes or Suia Missu basin, west flowing unnamed cór- basin, west flowing unnamed córrego in Fazenda Ipuá, just rego in Fazenda Ipuá, just north of Água Boa on the road north of Água Boa on the road Barra do Garça to São Felix Barra do Garça to São Felix do Araguaia (all biotopes do Araguaia (all biotopes destroyed). Photo by H. Bleher, destroyed). Photo by H. Bleher, 19.12.2005. 19.12.2005.

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

The two largest have: depth 2.90-2.97 and head 5/(10-12)/3´, 11i, predorsal, 14-15 around pedun- 3.65-3.70 in SL; dorsal fin anterior to midbody; cle; anal fin iii,22; caudal fin covered with small one specimen has first dorsal and anal rays elon- scales on its proximal third, plus a large scale sug- gate, but without filaments, the longest dorsal ray gesting an inseminating characid sensu Weitzman; not reaching adipose when depressed; pectoral fins mouth sub-superior; premaxillary teeth 4 tricus- reaching pelvic fins, pelvic fins overlapping last pid/5 pentacuspid, maxilla rather long and narrow, anal fin rays; scales 5/(9)37/3; anal fin iii,21; base reaching up to pupil’s level, with 3 broad teeth; of caudal fin scaled, with very small scales on basal mandible with 5 pentacuspid and about 10 much third of the lobes; peduncle deep; head short; oper- smaller teeth; SO3 very developed, almost entire; cle notched; SO3 developed, almost as in the gill-rakers rather long, about 15 on the inferior Astyanax paucidens group or most Hemibryconini; part of the first arch; coloration in vivo not known, mouth a little upturned, its opening at upper part but adipose and caudal fin lobes possibly red; a of pupil’s level; maxilla long and thin, reaching to broad lateral band from opercle to end of caudal pupil’s level, with 4 small but broad teeth (4 to 5 fin, much darker on peduncle; no humeral spot; a cusps); premaxilla with an outer row of 3 broad tri- distinct black line on anal fin base; this fish vaguely to pentacuspid teeth and an inner one of 5 with resembles Moenkhausia phaeonota or Hyphesso- 5-7 cusps, broad and compressed; mandible with brycon herbertaxelrodi, differing mostly in being 5 broad teeth in front, with 5-7 cusps, followed on deeper and having the “generic” characters of the sides by a series of much smaller ones. Colour-pat- latter species. It also resembles Hyphessobrycon aff. tern (in alcohol) much as that of H. micropterus: no cachimbensis from Station 20 and Station 21, dif- humeral spot; a faint band of chromatophores fering in the larger size (but 2 months in aquar- from opercle to dorsal, then a lateral narrow dark ium), more branched anal fin rays (22 vs. 19), band from dorsal fin to end of peduncle, then a more perforated LL scales (12 vs. 5), red caudal light zone and a caudal spot limited to the middle lobes (vs. probably not red), and chiefly the scaled caudal rays up to their end; base of lobes dark; caudal fin, with small scales on the proximal third when freshly unpacked, the unpaired fins were of the lobes, plus one large scale suggesting an rosy; a photo in situ (Characid sp. 2 Station 17) inseminating characid. shows the fins hyaline, not much developed, a Hemigrammus cf. rodwayi Durbin, 1909?? (Fig. golden lateral line from upper part of opercle to 16, resembling also Moenkhausia ceros Eigenmann, peduncle separating the body colour into a yellow 1908). Photos only, labeled “Characid sp. 2 Station dorsal part and a grey to bluish ventral one; in life 20” (in situ and in aquarium) and “Characid sp. 1 the caudal spot does not reach the end of the mid- Station 21” (in aquarium). dle rays. Not preserved. Station 20 and 21, dirt road on This fish has similar morphometrics and colour BR 242, 118 km before São Felix do Araguaia, small pattern as H. micropterus, but seems deeper, with córrego not destroyed among fazendas; H. Bleher more maxillary teeth. It differs from H. lowae in 19.12.2005 (probably Chapadinha, Alto Araguaia the scaled caudal fin, the dorsal fin not very elon- basin), and Rio Xavanti, into Tirapé into Araguaia, gate, the scales less numerous, the absence of a at Nova Xavantina; H. Bleher 19.12.2005. humeral spot, the caudal spot not extended onto the inferior part of the peduncle, etc. The discriminant characters of four similar Hyphessobrycon species from the region are summa- rized in the following table: Hemigrammus n. sp.? (lateral band, resembling Hyphessobrycon aff. cachimbensis) (No N.) no photo. 1 ex., 41 mm SL, preserved after 2 months in aquarium, Station 22 (no material preserved): probably Rio Araguaia basin (near mouth of Rio nd das Mortes into Araguaia), 2 river on the road Fig. 16. Hemigrammus cf. rodwayi. Station 20, dirt road on B242 from São Felix to Alto Boa Vista, H. Bleher BR 242, 118 km before Sao Felix do Araguaia, small cór- 19.12.2005. rego not destroyed among fazendas. Photo by H. Bleher Depth 2.75 and head 3.9 in SL; trunk scales 19.12.2005 (probably Chapadinha, Alto Araguaia basin).

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Fig. 17. Thayeria cf. boehlkei. Station 17 (not indicated on the map, certainly close to site in Station 23), Rio das Mortes or Suia Missu basin, west flowing unnamed córrego in Fazenda Ipuá, just north of Água Boa on the road Barra do Garça to São Felix do Araguaia (all biotopes destroyed). Photo by H. Bleher, 19.12.2005. The specimens from Station 20, about 30 mm 1 ex. juvenile, about 16-18 mm SL, not pre- SL, have been photographied in situ and in aquar- served, Station 25, rio Aqua Azul km 694, near to ium. A third photo from Nova Xavantina, Station Barra do Garças (not preserved) H. Bleher ca 21, possibly shot in situ in different conditions, 19.12.2005 shows two similar fishes. Depth 3.15-3.30; dorsal Depth about 3.25 in the SL; colour-pattern char- fin at midbody or slightly posterior; eye big; anal acteristic; the lower caudal lobe seems longer than fin iii, 19 or 20 (??); lateral scales more than 30 and in T. boehlkei from the upper Amazon (the aquar- less than 40, lateral line can be seen as incomplete ium form). Three photos were shot; one, in situ, or complete, depending on the photo; predorsal shows one specimen from Station 17 with the series apparently regular; caudal fin possibly scaled. upper part of the body, base of the upper caudal Colour-pattern similar to that of H. levis: caudal lobe and external rays of lower one, yellow; this is spot restricted to the fin, circumscribed by a light, about the same colour in aquarium (Station 20, yellow zone: no humeral spot; no (or very feeble) not preserved); the juvenile has the band relatively black line along anal fin base; lips black. This pat- broader (photo in situ from Station 25, fish not tern is not unusual in the Serra do Roncador (see preserved). for example Characid sp. 1 from Station 25), and the photos may represent two spp (sibling). The Inc. sed. relatively long anal fin, the dorsal fin at midbody Inc. gen. sp. U1 (photo only) “Characid sp. 1”, and the colour-pattern suggest the most common Station 25. (Fig. 18, photo only). tetra from the Serra do Roncador (300 ex. collected No specimen preserved (possibly 2 alive in aquar- by R. McConnell-Lowe, both in the upper Xingú ium). Station 25, Córrego Azul (in fact muddy) and the upper Tocantins drainages), called provi- after Nova Xavantina, km 694, near to Barra do sionally H. cf. rodwayi (Roncador U4). Garças, H. Bleher ca 19.12.2005. Thayeria cf. boehlkei Weitzman, 1957 N. 096.1- A small elongate tetra, depth 4.25 in SL, resem- 3.2006 (Fig. 17). bling Hemigrammus levis without humeral spot (or 2 ex., largest 30 mm SL, Station 17, just north of feebly pronounced in vivo) and a large caudal spot Água Boa on the road Barra do Garça to São Felix on the centre of the fin, surrounded by conspicu- do Araguaia, Bleher 19.12.2005 (mark 24 on the ous white bands; anal fin short, its base sur- map). mounted by a black line on its anterior half. 1 ex., 34.5 mm SL, Station 20, córrego 118 km According to the study of the R. Lowe McConnell before São Felix de Araguaia, H. Bleher collection in the Serra de Roncador, this photo may 19.12.2005; Alto Araguaia basin. represent a new form (represented in her collection

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

by two specimens, from Córrego do Gato, Rio humeral spot except a shiny spot after opercle (see Xingú drainage, Hemigrammus Roncador U3). also A. pedri from upper Rio das Mortes, Station Inc. gen. sp. U2 (Fig. 19, photos only) “Characid 16) but the humeral region seemingly with numer- 1 and Characid 2 lower Station 9” (aquarium); ous chromatophores; a round distinct caudal spot, (according to photos, probably only one species). not to end of rays. No specimen preserved. Station 9, Rio Formosa, Unidentifiable. Distinct from the similar tetras Alto Paraguay basin near Bonito, H. Bleher from Rio Miranda, Astyanax (Jupiaba) acantho ga - 15.12.2005. ster and Astyanax cf. luetkeni, by the lack of a Collected with Astyanax cf. asuncionensis and kept humeral spot and the caudal spot not to end of in aquarium, where two forms had been recognized middle caudal rays. The slight differences in habi- (possibly mimetic). Body rather robust. No tus may be due to age or camera flash, but the pres- ence of two sympatric species cannot be elimi- nated. To note that “mimicry” means that two species from two different clades (two families for example) resemble each other. Two close species, in the same genus for example, resemble more or less each other because they have the same ancestror. When the resemblance is almost complete, due to recent sympatric speciation, they are termed sib- ling; due to ancient allopatric speciation, they are termed vicariant (= replacing each other). The Fig. 18. Inc. gen. sp. U1. Station 25, Córrego Azul (in fact cladists have a different definition. muddy) after Nova Xavantina, km 694, near to Barra do Inc. gen. sp. U3 (Fig. 20, photo only) “Characid Garças (not preserved). Photo by H. Bleher 19.12.2005. 1 Station 24” (in situ, just collected). No specimen preserved. Station 24, road back to Barra do Garça on BR 158, km 27 south of Agua Boa, first córrego, a tiny waterflow into larger muddy córrego, Rio das Mortes basin, H. Bleher 21.12.2005. Probably less than 40 mm SL; habitus generalized, looking like an Astyanax; depth about 2.8; dorsal fin at midbody; anal fin short; a humeral and a caudal spot, a silvery band and a broad cheek and probably large SO3; maxilla not very long; eye large.

Cheirodontinae Fig. 19 Inc. gen. sp. U2. Station 9, Rio Formosa, Alto Para - Serrapinnus notomelas (Eigenmann, 1915) (Fig. guay basin, near Bonito. Photo by H. Bleher, 15.12.2005. 21 in aquarium from Station 4). 1 ex. adult male, 29.33 mm SL, Station 1, Rio Tiété basin, km 458 from São Paulo, after Araçatuba on the BR 300 “Marechal Rondon”, H. Bleher 12.12.2005 N. 394.1. 18 ex., largest 28.5 mm SL, Station 4, Alto Parana basin, km 54 on the road Tres Lagoas to Campo Grande, H. Bleher 13.12.2005 (poor condition) N. 394.2-19.2006. Depth 2.55-2.90 in SL; scales (8)35, 11 trans- verse; anal fin iii,17; 5-7 interhaemals; mouth ter- Fig. 20. Inc. gen. sp. U3. Station 24, road back to Barra do minal, teeth flat multicuspid (up to 7 cusps only) Garça on BR 158, km 27 south of Agua Boa, first córrego, on both jaws; maxilla very short, with 0-1 teeth; a tiny waterflow into larger muddy córrego. Photo by H. colour-pattern rather variable; usually the anterior Bleher, 21.12.2005. part of dorsal fin with a falciform black blotch; a

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Fig. 21. Serrapinnus notomelas. Station 4, Alto Parana Fig. 22. Serrapinnus piaba. Station 7, Alto Pardo basin? basin, km 54 on the road Tres Lagoas to Campo Grande, (into Paraná), road Campo Grande to Corumba Vazante tiny creek flowing south into cattle field. Photo by H. Ble- do km 2 just west of Campo Grande, dirt road to Pirapi- her 13.12.2005. tanga, sign: Pousada Sol. Photo by H. Bleher 14.12.2005. broad caudal spot, covering almost all of the depth Characinae of the peduncle, not extending onto the middle Phenacogaster aff. jancupa Malabarba & Z. de caudal fin rays; anal fin lobe never black, only the Lucena, 1995? Two photos alive (Station 20) in first ray marked; skin over the interhaemals dark. aquarium and Fig. 23 in situ Station 18). Originally described from the Rio Tiété basin. Col- 4 ex., largest 31.0 mm SL, Station 19, possibly lected in Parana basin by Schindler (Rio Ivin- Rio Suia Missu basin, Córrego Chega de Perto heima) and H. Schultz (Tres Lagoas), and in Rio flowing north near Posta da Mata before São Felix Machado (Madeira) by Geraldo dos Santos. do Araguaia, fazenda-land; H. Bleher 19.12.2005 A photo in aquarium shows no striking colours, N. 1072.1-4.2006. except a thin lateral golden band and a shiny spot (One photo shot in situ, labelled Phenacogaster sp. posterior to the caudal spot on each lobe base. 1 Station 18, belongs perhaps also to this species. Serrapinnus piaba (Lütken, 1874) N. 290.1-3. Depth 2.95 (the small specimens more elongate) 2006 (Fig. 22, from Station 7). and head 3.50 in SL; a humeral hiatus; pre-pelvic 3 ex., 1 male and 2 females, 33.0-36.85 mm SL, squamation of the derived type, i.e. with the lateral Station 4, Alto Parana basin, km 54 on the road scales obliquely intricated and lacking a median Tres Lagoas to Campo Grande, H. Bleher series; trunk scales 6/35-37/4 (to pelvic fins as well 13.12.2005 (not too good state) N. 290.1-3.2006. as to anal fin), the two smallest with an incomplete ? Photo in situ (not preserved, labelled “Station 7, lateral line; anal fin iv, 30; premaxilla 2+3, with a Characid sp. 1, male” just west of Campo Grande, gap between/9 tricuspid; maxilla about 24; near entrance to BR 262, “Pirapitinga”, Alto Pardo mandible more than 15; a large, conspicuous, cau- basin? (into Parana). dal spot, preceeded by a light zone and a rather Depth 2.95 in SL; scales (9)34, 10-10´ transverse; anal fin iii,20; about 9 interhaemals; mouth subin- ferior, teeth flat multicuspid (up to 7-8 cusps) on both jaws; maxilla very short, with 1(?) tooth; fins hyaline; a broad caudal spot, covering the depth of the peduncle, truncate posteriorly, not extending onto the middle caudal fin rays, pointed anteriorly; the photo in situ labelled Station 7 shows a silvery lateral band, without striking colours. Originally described from the Rio São Francisco basin and possibly endemic; a similar form, or the same one, sympatric with S. notomelas in the upper Fig. 23. Phenacogaster aff. jancupa. Station 18, a few km Parana, is often found in small samples all over the before Posta da Mata, after Cascalheira, on the road Barra Amazon basin. It seems to have a slightly narrower do Garça to São Felix do Araguaia shortly before turn to caudal spot. If two (or three) forms, no differential São Felix, possibly Suia Missu basin but biotopes character found till now (but in need of revision). destroyed. Photo by H. Bleher 19.12.2005.

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

inconspicuous longitudinal line; a prominent lowing from the R. Lowe Mcconnell Roncador humeral ocellus, the black part almost as large as expedition: 2 ex., 36-37 mm SL, Xavantina (field pupil, the light zone all around it except above, N. 118 & 124); and probably 1 ex. 23.5 mm SL, tather narrow; after photographs in vivo, from Sta- Suia Missu (night camp, field N. 146)) may belong tion 20; in aquarium and in situ, the light zones to this form. white, the unpaired fins pale maroon (?), and there is a golden line from opercle to peduncle, with the Oligosarcinae (?) upper part of eye apparently of the same colour. Oligosarcus pintoi Campos, 1945 (Fig. 25 in situ). P. jancupa is geographically near (south of 6 ex. juveniles, largest 40.0 mm SL, Station 1, Rio Cuiaba), but from an entirely different basin (Rio Tiété basin, km 458 from São Paulo after Paraguay vs. Rios Xingú and Tocantins). I have seen Araçatuba on the BR 300 “Marechal Rondon”, H. four forms with the morphometrics of P. calverti (a Bleher 12.12.2005 N. 840.1-6.2006. synonym of P. franciscoensis): one from Maranão Depth 3.2 and head 3.1 (largest) in SL; dorsal fin State (coll. Ternetz and Werner) and Ceara (coll. posterior to midbody; scales 8/38/6´ (to anal fin), Roberts), one from surroundings Porto Velho (coll. 13 predorsal, 15 around peduncle; anal fin iv,26; Brittan), one from Rio Paru (coll. Geisler) and this great suborbital large, its suture with SO4 high on one. No recent revision of the genus. jaw, not fully developed, not in contact with the Phenacogaster sp.?? (Fig. 24, photo only) subopercular canal; premaxilla with 8 tricuspid “Characid 2, Station 25” (aquarium). teeth, the first one much larger; two large similar No specimen preserved (at least one alive in teeth forming an inner row, close to the outer one; aquarium). Station 25, Córrego Azul (in fact maxilla long, reaching to middle of eye level, not muddy) after Nova Xavantina, km 694, near to forming an angle with the premaxilla, with teeth Barra do Garças, Rio das Mortes basin, H. Bleher along almost its entire margin (about 13, mostly tri- ca 19.12.2005. cuspid in the largest); mandible with 5 large teeth Habitus of a Phenacogaster; depth about 3.33; in front followed on side by about 11 smaller ones; dorsal fin anterior to midbody; anal fin long, per- ectoperygoid with ca. 10 teeth, in more than one haps about 30 branched rays; perhaps some scales row (?); eye silvery; a vertical spot high on the on caudal fin base??; eye large; maxilla long, thin, humeral region, a black midlateral line starting straight, reaching pupil’s level; humeral spot dis- beneath dorsal fin, forming a large black spot on the tinct, quadrangular, with a shiny spot in front; a peduncle and prolonged onto the middle caudal fin longitudinal line, not in contact with a purely cau- rays. The photo of a living specimen shot in situ, dal spot, probably to end of middle rays, with lat- possibly for the first time, shows the body olive-yel- eral expansions at fin base; yellow spots anterior to low with some bluish relicts(?), a large caudal spot the spot and on the lobes bases; anal fin with a line with the expansion onto the middle caudal rays on the base and the border black, and the first ray scarcely visible; the black anal fin margin seems white. extremely narrow in this eastern population. Rather distinct from the preceeding in the humeral and caudal spots, anal fin, etc. The fol-

Fig. 25. Oligosarcus pintoi. Station 1, Rio Tiété basin, km Fig. 24. Phenacogaster sp?? Station 25, Córrego Azul (in fact 458 from São Paulo, after Araçatuba on the BR 300 muddy) after Nova Xavantina, km 694, near to Barra do "Marechal Rondon"; small creek flowing north, in a Garças (not preserved). Photo by H. Bleher 19.12.2005. mostly destroyed farm. Photo by H. Bleher 12.12.2005.

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LEBIASINIDAE 24 lateral scales, mean 21.64. The species was col- lected by Harald Schultz near Tres Lagoas, thus Lebiasininae, Pyrrhulininae near the present locality 4. Pyrrhulina australis Eigenmann & Kennedyn Pyrrhulina sp. (aff. P. stoli?) (Fig. 27). 1903 (Fig. 26 in situ Station 7, not Station 4). 1 ex., 28.5 mm SL, Station 18, a few km before 4 ex., largest 34.55 mm SL, Station 4, Alto Parana Posta da Mata, after Cascalheira, shortly before basin, km 54 on the road Tres Lagoas to Campo turn to São Felix, possibly Suia Missu basin but Grande, H. Bleher 13.12.2005 (not too good habitus biotopes destroyed, H. Bleher 19.12.2005. state) N. 395.1-4.2006. The specimen, soft, has the depth about 4.3 in Depth 4.23 in SL; scales 5´/22 or 23 total; 12 SL. scales 5/22 or 23 total; 12 predorsal, 10 around predorsal, 10 around caudal peduncle. caudal peduncle. A photo shot at Station 7 (just west of Campo 17 ex. collected by R. Lowe-McConnell in the Grande, near entrance to BR 262, “Pirapitinga”, same region (Serra do Roncador) have a few more Alto Pardo basin? (into Parana), shows the body scales. relatively short, olivaceous with the base of dorsal, Depth 3.85 to 4.70 in SL (negative allometry?); anal and caudal fins yellowish; a black band up to predorsal length 0.72 in postdorsal length in the opercle, whereas on the preserved fish the opercu- largest; scales 5/23-25+1-2 on caudal, 14 predorsal, lar part of the band is composed of less dense chro- 10 around caudal peduncle; anal fin iii,9-10; dor- matophores, grey instead of black. sal fin spot on the anterior half except unbranched According to Zarske & Géry (2004), 76 speci- rays, occupying the medial two-thirds of its depth; mens of the species (syn. macrolepis Ahl) have 19- a predorsal line, enlarged anteriorly and in the middle; lateral band from mandible to 3rd scale, then an indistinct blotch on the 5-7th scales. In an identification key to the Pyrrhulina spp., this Roncador form seems to be phenotypically near to P. stoli, from the Guianas.

ERYTHRINIDAE

Hoplerythrinus sp. (Fig. 28). No specimens; Erythrinus sp. 1 Station 22 (not preserved), probably Rio Araguaia basin (near mouth of Rio das Mortes into Araguaia), 2nd cór- Fig. 26. Pyrrhulina australis. Station 7, Alto Pardo basin? rego on the road B 242 from São Felix to Alto Boa (into Paraná), road Campo Grande to Corumba Vazante Vista, H. Bleher 21.12.2005. do km 2 just west of Campo Grande, dirt road to Pirapi- The genus is monotypic. tanga, sign: Pousada Sol. Photo by H. Bleher 14.12.2005.

Fig. 27. Pyrrhulina sp. (aff. P. stoli?). Station 18, a few km before Posta da Mata, after Cascalheira, on the road Barra Fig. 28. Hoplerythrinus sp. Station 22, probably Rio do Garça to São Felix do Araguaia shortly before turn to Araguaia basin (near mouth of Rio das Mortes into São Felix, possibly Suia Missu basin but biotopes Araguaia), second córrego on the road B242 from São destroyed. Photo by H. Bleher 19.12.2005. Felix to Alto Boa Vista. Photo by H. Bleher 20.12.2005.

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Hoplias sp. 1 (Fig. 29) CHARACIDIIDAE No specimens; Hoplias sp. 1. Station 25 (not pre- served), Córrego Azul (in fact muddy) after Nova Characidium sp, U1 (Fig. 31 in situ labeled Xavantina, km 694, road back to Barra do Garças Characidium 1, Station 7). and close to it (not preserved), H. Bleher 2 ex., largest 32.7 mm SL, Station 7, Alto Pardo 19.12.2005. basin? (into Parana), road Campo Grande to The identification of a Hoplias needs not only the Corumba Vazante do km 2 just west of Campo examination of a specimen, but also DNA analysis. Grande, dirt road to Pirapitanga, sign: Pousada Hoplias sp. 2 (Fig. 30). Sol. H. Bleher 14.12.2005 (not too good state). No specimens; Hoplias sp. 1 Station 23 (not pre- Depth 4.6 and head 4.15 in SL; width 1.84 in served): Road back to Barra do Garça on BR 158 greatest depth; postdorsal length/predorsal length at Fazenda Ipua (soybeans or cattle of course), nice 1.15; isthmus completely scaled; scales 4/33/3, 10 córrego before Agua Boa (near Photo 17, only on in predorsal, 12 around caudal peduncle; pectoral the other side of the road). Possibly Suia Missu fins iii,9, almost reaching pelvic fins; pelvic fins i,8, basin. H. Bleher 21.12.2005. reaching anal fin; anal fin ii, 7; dorsal fin ii, 9, ante- The identification of a Hoplias needs not only the rior to midbody; a small, anterior supraorbital; no examination of a specimen, but also DNA analysis. circumorbitals; eyes lateral; mouth small, inferior, Certainly different from the preceeding, despite its opening below eye level in profile; teeth tricus- the usual variability of the Hoplias-pattern. pid, at least 5/5 on each side; pterygoid teeth not observed; second dentary row not observed (speci- mens not dissected). A longitudinal broad line (or thin band, half scale high) from opercle to end of caudal peduncle, plus a distinct small spot at base of caudal (characteristic for a group of Characi - dium); 10-11 indistinct bands across dorsum, scarcely visible on flanks; a colour-photo shot in situ shows the body and fins yellowish, without striking colour. Characidium cf. zebra Eigenmann, 1909 (Fig. 32 Fig. 29. Hoplias sp. 1. Station 25, Córrego Azul (in fact “sp. 2, Station 8” in aquarium). muddy) after Nova Xavantina, km 694, near to Barra do 6 ex., largest 45.0 mm SL, female, Station 8, Garças (not preserved). Photo by H. Bleher 19.12.2005. Aquidauna on the BR 262 Rio Grande to

Fig. 30. Hoplias sp. 2. Station 23, (not preserved, near Station 17, only on the other side of the road), road back to Barra do Garça on BR 158 at Fazenda Ipuá (soybeans of course), nice córrego before Agua Boa, possibly Suia Missu basin near to Serra Dourada. Photo by H. Bleher 21.12.2005

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Jacques Géry

Corumba, rio of the same name at km 468, cgo below, forming a net; a longitudinal band, half córrego? flowing north over step waterfall, Alto scale high, from snout to end of peduncle, plus a Paraguay basin, coll. H. Bleher 14.12.2005) N. distinct small spot at base of caudal and a humeral 518.1-6.2006. spot just posterior to opercle; a quadrangular Depth 4.25 (female)-5.1 (male) and head 3.6 in blotch on peduncle similar to that of Ch. SL; body width 1.65-1.8 in greatest depth; post- etheostoma Cope, 1871; a spot just anterior to dor- dorsal length/predorsal length 1.15-1.20; isthmus sal fin; about 9 fasciae across dorsum; a colour- completely scaled; scales 4/36/3´, 10 in predorsal, photo shot in aquarium shows the body and fins 14 around caudal peduncle; pectoral fins iii,12, not yellowish, without striking colour, except the pos- reaching pelvic fins, the unbranched rays not terior part of peduncle which may be bright lemon enlarged; pelvic fins i,8 (probably i,7,i), the in certain circumstances; isthmus silvery, covered unbranched ray not enlarged, not reaching anal with very thin scales. fin; anal finii, 7; dorsal fin ii, 9, anterior to mid- Discussion: this Paraguayan form belongs to the body; a small, anterior supraorbital; no circumor- generalized Ch. zebra-group and is quite similar in bitals; eyes on top of the head, slightly oblique, the morphometrics and pattern to the upper Amazon- interorbital narrow; snout curved; mouth small, ian species Ch. etheostoma, through much less com- subinferior, its opening just below eye level in pro- pressed. file; teeth tricuspid, on each side at least 7 on pre- Characidium sp. U2 (Fig. 33 “sp. 1, Station 15”). maxilla, the 1st one enlarged, and 8 on dentary, the Not preserved, Station 15, Pantanal do Norte, 1st one not enlarged; pterygoid teeth not observed; Córrego do Bororo, near Poxoreu, on the road second dentary row well visible (specimens not dis- north of Rondonopolis (fast moving current), H. sected). A crescent-shaped spot on apex of each Bleher 18.12.2005. Unidentifiable. scale of the flanks above median line to one row Characidium sp. U3 (Fig. 34 in situ “sp. 1, Sta- tion 22”). Not preserved, Station 22, Rio Araguaia basin (near mouth of Rio das Mortes into Araguaia), 2nd córrego on the road B242 from São Felix to Alto Boa Vista, H. Bleher 20.12.2005. Belongs apparently to the nominotypical genus; a characteristic lateral line or thin band from snout to beginning of caudal, broader in its anterior part (up to about the sixth scale – anterior pattern as in some Pyrrhulina) and the absence of fasciae (at least in vivo). Fig. 31. Characidium sp. U1. Station 7, Alto Pardo basin? Characidium (sensu lato) sp. U4 (no photo). (into Paraná), road Campo Grande to Corumba Vazante 1 ex., 38.5 mm SL, Station 17, Rio das Mortes or do km 2 just west of Campo Grande, dirt road to Pirapi- Suia Missu basin, west flowing unnamed corrego tanga, sign: Pousada Sol. Photo by H. Bleher 14.12.2005. in Fazenda Ipuá, just north of Agua Boa on the

Fig. 32. Characidium cf. zebra. Station 8, Aquidauna on Fig. 33. Characidium sp. U2. Station 15, Pantanal do the BR 262 Rio Grande to Corumba, rio of the same name Norte, Córrego do Bororo, into Rio Vermelho, near Poxo- at km 468, córrego flowing north over step waterfall, Alto reu, on the road north of Rondonopolis (fast moving cur- Paraguay basin, coll. Photo by H. Bleher 14.12.2005. rent). Photo by H. Bleher 18.12.2005.

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Characiform fishes collected by Heiko Bleher on the road São Paulo-Corumba-Araguaia, December 2005

high and, probably, by the peculiar colour-pattern just described. The presence of a supraorbital excludes it from Melanocharacidium. The following key to the five studied Characid- ium (supraorbital present; isthmus scaled; dorsal fin ii,9; anal fin ii,7) may be of use for future work (the collections from Paraguay and probably from Roncador are in the possession of P. Buckup): 1a. Dorsal fasciae; longitudinal band, if any, not well delimited in alcohol. Fig. 34. Characidium sp. U3. Station 22, probably Rio 1b. No dorsal fasciae; a well-delimited longitudinal Araguaia basin (near mouth of Rio das Mortes into band in life (morphometrics not known) ...... Araguaia), second córrego on the road B242 from São Characidium sp. U3, photo only, Rio Araguaia Felix to Alto Boa Vista. Photo by H. Bleher 20.12.2005. or das Mortes Station 22. 2a. Pectoral fins iii,9-12; pelvic fins i,8; teeth tri- road Barra do Garça to São Felix do Araguaia, H. cuspid; predorsal scales 10; dorsal hyaline. Bleher 19.12.2005. 2b. Pectoral fins iv,7; pelvic fins i,8i; premaxillary Depth 4.6 and head 4.0 in SL; body width xx in teeth conical, recurved; predorsal scales 12; largest depth; postdorsal length/predorsal length xx; dorsal barred ...... isthmus completely scaled; scales 4/37/3´, xx in pre- Characidium sp. U4 (possibly a new species) 1 dorsal, 12 around caudal peduncle; pectoral fins ex., Rio das Mortes, Station 17. iv,7, reaching pelvic fins, the unbranched rays 3a. Pectoral fins iii,9; scales 4/33/3, 12 around somewhat enlarged; pelvic fins i,8i, the unbranched caudal peduncle; 5 premaxillary teeth, the ray not enlarged, not reaching anal fin; anal fin ii,7; inner one not enlarged; depth 4.6; head 4.15; dorsal fin ii,9, anterior to midbody; caudal fin i9/8i; body width 1.84 in body depth (predorsal supraorbital present; circumorbitals reduced to the 1.15) ...... canal, cheek broad and naked; eyes vertical; nostrils Characidium sp. U1, 2 ex. Alto Parana, Station well-separated, on top of the snout; a small occipi- 7. tal triangular fontanel; snout not very elongate, 3b. Pectoral fins iii,12; scales 4/36/3´, 14 around roundish, plunging; mouth very small, inferior, its caudal peduncle; 7 premaxillary teeth, the opening below eye level in profile, maxilla short and inner one enlarged; depth 5.1; head 3.6; body broad, visible from below; teeth curved, conical on width 1.65-1.70 in body depth (predorsal premaxilla, possibly tricuspid on dentary, formula 1.14-1.18; about 10 transverse bands)...... about 7/7; pterygoid teeth not observed; second Characidium zebra-group, 6 ex. Alto Paraguay, dentaary row not visible (specimens not dissected). Station 8. A weakly delimited narrow lateral band from oper- 3c. About 14 transverse bands (morphometrics not cle to middle of caudal peduncle; an indistinct known; rather similar to C. zebra-group) ...... small spot at the base of caudal fin (not as apparent Characidium sp. Photo only, Alto Paraguay, as in the above-mentioned Characidium-group); Station 15. dorsum with a black blotch just in front of dorsal fin and less distinct fasciae (4 or 5) after dorsal fin; ANOSTOMIDAE dorsal fin with a rather distinct oblique bar at about mid-fin; tip of snout black, the black extending Leporinus juvenile (Fig. 35 shot in aquarium). somewhat up to eye; anterior and superior part of No specimens; Leporinus sp. 1 Station 22 (not eye black, with an extension on the top of the head, preserved), Rio Araguaia basin (near mouth of Rio not joining the other side; a large, oblique blotch on das Mortes), 2nd córrego on road B242 from São opercle. Felix to Alto Boa Vista, H. Bleher 20.12.2005. Comments: this specimen may represent a new According to the basic colouration (three round species; it differs from a Characidium in the high spots on flanks), it would belong to the L. friderici- number of unbranched rays on pectoral and the group; the transverse fasciae (10 bands, some split) few unbranched ones (ratio 4/7 vs. usually 3/10- may be juvenile features (to be compared with the 12), as well as the peculiar nostrils situated vwery few from Roncador collection).

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Jacques Géry

REFERENCES COSTA, W. J. M., 1994. Description of two new species of the genus Moenkhausia (: Characidae) from the central Brazil. Zoologischer Anzeiger, Jena, 232 (1/2): 21-29. (M. nigromarginata & pyrophthalma). COSTA, W. J. M., 1995. Description of a new species of the genus Astyanax (Characiformes: Characidae) from the rio Araguaia basin, Brazil. Revue Suisse de Zoologie, 102 (1): 257-262. (Astyanax kullanderi). COSTA, W. J. E. M. & GÉRY, J., 1994. Two new species of the genus Hyphessobrycon (Characiformes: Characidae) Fig. 35. Leoporinus sp. juvenile. Station 22, probably Rio from the rio Xingu basin, central Brazil. Revue française Araguaia basin (near mouth of Rio das Mortes into d’aquariologie, 20 (1993) (3): 71-76. Araguaia), second córrego on the road B242 from São Felix COSTA, W. J. E. M. & VICENTE, E. O. 1994. Une nouvelle to Alto Boa Vista. Photo by H. Bleher 20.12.2005. espäce du genre Melanocharacidium (Characiformes: ) du bassin du rio Araguaia, Brésil central. Revue française d’Aquariologie, 20 (1993) 3: 67-70 (M. PARODONTIDAE auroradiatum). LIMA, F. C. T. & MOREIRA, C. R. 2003. Three new species Parodon nasus Kner, 1859 (Fig. 36 “sp. 1” in of Hyphessobrycon (Characiformes: Characidae) from the aquarium). upper Rio Araguaia in Brazil. Neotropical Ichthyology, 2 ex., largest 90.0 mm SL, Station 8, Aquidauna SZUSP, 1 (1): 21-33 (langeanii, eilyos, weitzmanorum). on the BR 262 Rio Grande to Corumba, rio of the LUCENA, C. A. S. DE, 2003. New characid fish, Hyphesso- brycon scutulatus, from the rio Teles Pires drainage, upper same name at km 468, córrego flowing north over rio Tapajos system (Ostariophysi: Characiformes: step waterfall, Alto Paraguay basin, coll. H. Bleher Characidae). aff minimus? 14.12.2005. MOREIRA, C. R., LANDIM, M. I. & COSTA, W. J. E. M. Depth 4.1-4.5 in SL; scales 4/38 tot./3, 14 2002. Hyphessobrycon heliacus. A New Characid Fish around peduncle; pectoral fins i,15, anal fin ii,7; 4 (Ostyariophysi: Characiformes) from the Upper Rio premaxillary teeth on a straight line on each side, Tapajos Basin, Central Brasil. Copeia, 2002 (2): 428-432 the broadest with up to 16 cusps almost in a (close to H. loweae). straight line; 2 maxillary teeth; three rounded teeth on the side of the mandible; no dorsal marking; a longitudinal stripe formed by about 21 closely placed lozenge-shaped lateral spots.

Fig. 36. Parodon nasus. Station 8, Aquidauna on the BR 262 Rio Grande to Corumba, rio of the same name at km 468, córrego flowing north over step waterfall, Alto Paraguay basin, coll. Photo by H. Bleher 14.12.2005.

59 aqua vol. 19 no. 1 - 21 January 2013 AQUA19(1):AQUA 24/01/13 12:38 Pagina 60

Index of aqua Vol. 18 (1-4) (Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews) Author(s): Allen, Gerald R. and Drew, Joshua A.: A new species of Damselfish (Pomacentrus: Pomacentridae) from Fiji and Tonga. aqua 18 (3): 171-180, 15 July 2012 Allen, Gerald R. and Erdmann, Mark V.: A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia. aqua 18 (1): 9-14, 15 January 2012 Allen, Gerald R. and Unmack, Peter J.: A new species of Rainbowfish (Chilatherina: Melanotaeniidae), from the Sepik River System of Papua New Guinea. aqua 18 (4): 227-237, 15 October 2012 Anderson,William D. Jr. and García-Moliner, Graciela: A new species of Odontanthias Bleeker (Perciformes: Serranidae: ) from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean. aqua 18 (1): 25-30, 15 January 2012 Bakhshalizadeh, Shima, Abdolmalaki, Shahram and Bani, Ali: Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea. aqua 18 (2): 103-112, 15 April 2012 Bierbach, David, Ziege, Madlen, Zimmer, Claudia, Riesch, Rüdiger, Schlupp, Ingo, Arias-Rodriguez, Lenin and Plath, Martin: Male Grijalva mosquitofish (Heterophallus milleri Radda, 1987) increase individual mating preferences in front of an audience. aqua 18 (4): 199-208, 15 October 2012 de Oliveira Armesto, Leonardo and Freret-Meurer, Natalie Villar: Testing for camouflage of the Brazilian seahorse Hippocampus reidi (Syngnathidae) using the territorial damselfish Stegastes fuscus (Cuvier) (Pomacentridae). aqua 18 (3): 165-170, 15 July 2012 Foster, Ralph, Bridge, Thomas C. L. and Bongaerts, Pim: The first record of Hippocampus denise (Syngnathidae) from Australia. aqua 18 (1): 55-57, 15 January 2012 Gaither, Michelle R. and Randall, John E.: On the validity of the cirrhitid fish genus Itycirrhitus. aqua 18 (4): 219-226, 15 October 2012 Heemstra, Phillip C. and Akhilesh, K. V.: A review of the anthiine fish genus Pseudanthias (Perciformes: Serranidae) of the western Indian Ocean, with description of a new species and a key to the species. aqua 18 (3): 121-164, 15 July 2012 Hoyos-Padilla, E. Mauricio, Ceballos-Vázquez, B. Patricia and Galván-Magaña, Felipe: Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico. aqua 18 (1): 15-24, 15 January 2012 McCosker, John E. and Allen, Gerald: Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines. aqua 18 (1): 35-40, 15 January 2012 Poursaeid, Samaneh and Falahatkar, Bahram: Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae). aqua 18 (1): 31-34, 15 January 2012 Randall, John E. and Arnold, Rachel J.: Uranoscopus rosette, a new species of stargazer (Uranoscopidae: Trachinoidei) from the Red Sea. aqua 18 (4): 209-218, 15 October 2012 Randall, John E., Bogorodsky, Sergey V. and Rose, Jean Michel: Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider). aqua 18 (1): 41-54, 15 January 2012 Riesch, Rüdiger, Martin, Ryan A., Bierbach, David, Plath, Martin, Langerhans, R. Brian and Arias-Rodriguez, Lenin: Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae). aqua 18 (2): 95,102, 15 April 2012 Robert, Tyson R.: Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes). aqua 18 (2): 113-118, 15 April 2012 Vermeulen, F. B. M., Suijker, W. H. and Collier, G. E.: Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana. aqua 18 (4): 181-190, 15 October 2012 Valdesalici, Stefano: Nothobranchius kardashevi and Nothobranchius ivanovae (Cyprinodontiformes: Nothobranchiidae): two new annual killifishes from the Katuma River drainage, western Tanzania. aqua 18 (4): 191-198, 15 October 2012 Van Tassell, James L., Tornabene, Luke and Colin, Patrick L.: Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species. aqua 18 (2): 61-94, 15 April 2012 Walsh, Fenton and Tanaka, Hiroyuki: Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific. aqua 18 (1): 1-8, 15 January 2012 New Taxa: Antilligobius, n. gen. Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species. aqua 18 (2): 61-94, 15 April 2012 Antilligobius nikkiae, n. sp. Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species. aqua 18 (2): 61-94, 15 April 2012 Chilatherina pagwiensis, n. sp. A new species of Rainbowfish (Chilatherina: Melanotaeniidae), from the Sepik River System of Papua New Guinea. aqua 18 (4): 227-237, 15 October 2012 Cirrhilabrus nahackyi, n. sp. A new wrasse (Perciformes; Labridae) from the South Pacific. aqua 18 (1): 1-8, 15 January 2012 Laimosemion paryagi, n. sp. A new species (Cyprinodontiformes: Aplocheiloidei: Rivulidae), from the upper Mazaruni river drainage of Western Guyana. aqua 18 (4): 181-190, 15 October 2012 Myrichthys paleracio, n. sp. Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines. aqua 18 (1): 35-40, 15 January 2012 Nothobranchius ivanovae, n. sp. Two new annual killifishes (Cyprinodontiformes: Nothobranchiidae) from the Katuma River drainage, western Tanzania. aqua 18 (4): 191-198, 15 October 2012 Nothobranchius kardashevi, n. sp. Two new annual killifishes (Cyprinodontiformes: Nothobranchiidae) from the Katuma River drainage, western Tanzania. aqua 18 (4): 191-198, 15 October 2012 Odontanthias hensleyi, n. sp. A new species of Odontanthias Bleeker (Perciformes: Serranidae: Anthiinae) from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean. aqua 18 (1): 25-30, 15 January 2012 Pseudanthias pillai n. sp. A review of the anthiine fish genus Pseudanthias (Perciformes: Serranidae) of the western Indian Ocean, with description of a new species and a key to the species. aqua 18 (3): 121-164, 15 July 2012 Pomacentrus mafu, n. sp. A new species of Damselfish (Pomacentrus: Pomacentridae) from Fiji and Tonga. aqua 18 (3): 171-180, 15 July 2012 Scleropages inscriptus, n. sp. A new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes). aqua 18 (2): 113-118, 15 April 2012 Synchiropus tudorjonesi, n. sp. A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia. aqua 18 (1): 9-14, 15 January 2012 Uranoscopus rosette, n. sp. A new species of stargazer (Uranoscopidae: Trachinoidei) from the Red Sea. aqua 18 (4): 209-218, 15 October 2012 Biology/Ecology/Biography/Reviews: A review of the anthiine fish genus Pseudanthias (Perciformes: Serranidae) of the western Indian Ocean, with description of a new species and a key to the species. aqua 18 (3): 121-164, 15 July 2012 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea. aqua 18 (2): 103-112, 15 April 2012 Book review: The Banggai Cardinalfish: Natural History, Conservation and Culture of Pterapogon kauderni by Alejandro A. Vagelli. aqua 18 (2): 119-120, 15 April 2012 Book review: The Biology of Gobies by R. A. Patzner, J. L Van Tassell, M. Kova ié and B. G. Kappor (editors). aqua 18 (4): 239-240, 15 October 2012 Book review: Shore Fishes of Easter Island by John E. Randall and Alfred Cea. aqua 18 (4): 2387, 15 October 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider). aqua 18 (1): 41-54, 15 January 2012 Male Grijalva mosquitofish (Heterophallus milleri Radda, 1987) increase individual mating preferences in front of an audience. aqua 18 (4): 199-208, 15 October 2012 Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae). aqua 18 (2): 95,102, 15 April 2012 On the validity of the cirrhitid fish genus Itycirrhitus. aqua 18 (4): 219-226, 15 October 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico. aqua 18 (1): 15-24, 15 January 2012 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species. aqua 18 (2): 61-94, 15 April 2012 Testing for camouflage of the Brazilian seahorse Hippocampus reidi (Syngnathidae) using the territorial damselfish Stegastes fuscus (Cuvier) (Pomacentridae). aqua 18 (3): 165-170, 15 July 2012 The first record of Hippocampus denise (Syngnathidae) from Australia. aqua 18 (1): 55-57, 15 January 2012 Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae). aqua 18 (1): 31-34, 15 January 2012

aqua vol. 19 no. 1 - 21 January 2013 60 AQUA19(1):AQUA 24/01/13 12:38 Pagina 61

Guidelines for Authors 1. Manuscript preparation: manuscripts must be submit- Examples of correct reference formats: ted in English. In exceptional cases aqua may provide BLABER, S. J. M. 1980. Fish of the Trinity inlet system of translations of manuscripts written in French, German, North Queensland, with notes on the ecology of fish Italian, or Spanish. faunas of tropical Indo-Pacific estuaries. Australian Manuscripts must be word-processed in Microsoft Journal of Marine and Freshwater Research 31:137-46. WORD and submitted in an electronic form. Generic, specific, and sub-specific names must be italicised. All DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981. papers must conform to the International Code of Zoo- Estuarine fishes. In: Estuarine Ecology with Particular logical Nomenclature. Authors are strongly advised to fol- Reference to Southern Africa. (Ed. J.H. Day.): low the format set out in previous publications of aqua. 197-221. A. A. Balkema, Rotterdam. 2. Title: the title must be short and should precisely iden- DIMMICH, W. W. 1988. Ultrastructure of North Ameri- tify the main topic of the article. Names of genera or can cyprinid maxillary barbels. Copeia 1988 (1): species are followed by the systematic group to which they 72-79. belong. Author name(s) are given in full beneath the title, TREWAVAS, E. 1983. Tilapiine Fishes of the Genera followed by the complete mailing and e-mail address(es). Sarotherodon, Oreochromis and Danakilia. British 3. Abstract: the abstract should not exceed 250 words and Museum (Natural History), London, 583 pp. draw attention to the principal conclusions. It should not contain any uncommon abbreviations or literature cita- 6. Submission of manuscript and illustrations: The man- tions. The inclusion of abstracts in other languages is uscript and illustrations must be submitted digitally to the optional. Scientific Editor: 4. Subject matter: the text of the manuscript is usually Dr Frank L. Pezold arranged in four main sections: Introduction, Materials College of Science & Engineering and methods (including a key to abbreviations), Results, Texas A&M University – Corpus Christi and Discussion. Other subdivisions may be chosen 6300 Ocean Drive depending on the material presented. Acknowledgements Corpus Christi, TX 78412-5806 should be placed between the text and references. Email: [email protected] Scientific names of genera, species, and subspecies should be followed by the name(s) of author(s) and the year of to whom all subsequent correspondence shall be addressed. publication on first mention. A description of a new taxon Texts, tables, and graphs should be in Microsoft-compati- must contain the following sections: Material, Diagnosis, ble electronic form. Description, and Affinities. Synonyms must be arranged After the paper has been accepted for publication, illustra- in chronological order. Identification keys must be tions as high-resolution TIF files or original photographs dichotomous. (ideally transparencies; otherwise glossy prints, preferably Holotype and paratypes must be clearly identified, the in the size in which they will appear - the type area of aqua institution in which they have been deposited named, and is 158 x 224 mm, one column is 76 mm wide) must be the catalogue numbers given. Private collections are not sent to: acceptable as repositories for holotypes. Aquapress, The Managing Editor DNA sequences must be archived in GenBank, DNA Via G. Falcone 11, Databank of Japan or European Molecular Biology Labora- 27010 Miradolo Terme (Pavia), Italy tory. Voucher specimens associated with DNA sequences E-mail: [email protected] must be deposited in a recognized research collection acces- Authors should retain copies of all materials for reference. sible to the professional community. Accession numbers for sequences and catalog number for vouchers must be Proofs of accepted papers will be sent as PDF files by e- included as a table in an appendix to the final manuscript. mail attachment to the corresponding author. The metric system and SI units must be used. Tempera- 7. Evaluation of manuscripts: manuscripts will be evalu- tures are given in °C. Fractions should not be used. ated by the editors and referees. Papers are accepted on the understanding that they have not and will not be pub- 5. References to literature: the name-year system must be lished elsewhere. used. The list of references should be placed at the end of the paper, alphabetically arranged according to author 8. Reprints: Authors will receive one free copy of the issue name. Only those publications cited in the paper may be in which their paper appears and an e-print in PDF for- included. Titles of journals must be given in full. mat. Additional copies may be ordered from Aquapress. AQUA19(1):AQUA 24/01/13 12:38 Pagina 62

aqua International Journal of Ichthyology Vol. 19 (1), 21 January 2013

Contents:

John E. Randall and Joseph D. DiBattista: A new species of damselfish (Pomacentridae) from the Indian Ocean ...... 1-16

Matthew L. Wittenrich and Adeljean L.F.C. Ho: Social structure and reproductive behavior of the callionymid fish Callionymus bairdi (Gobiesociformes: Callionymidae): with notes on male alternative reproductive tactics ...... 17-28

Jose Tavera and Arturo Acero P.: Description of a new species of Hypoplectrus (Perciformes: Serranidae) from the Southern Gulf of Mexico ...... 29-38

Jacques Géry: Characiform fishes collected by H. Bleher on the road São Paulo-Corumba-Araguaia, December 2005 ...... 39-59

Index of aqua Vol. 18 (1-4) ...... 60

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: Territory-holding male Callionymus bairdi displaying to a female within his territory. Photo by M. L. Wittenrich.

Rio das Mortes or Suia Missu basin, west flowing unnamed córrego in Fazenda Ipuá, just north of Água Boa on the road Barra do Garça to São Felix do Araguaia (all biotopes destroyed). Station 17 (not indicated on the map, certainly close to site in Station 23). See pages 39-59. Photo by H. Bleher 19.12.2005.