Aqua, International Journal of Ichthyology Aqua, International Journal of Ichthyology
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aqua International Journal of Ichthyology Vol. 12 (4) 1 August 2007 Aquapress ISSN 0945-9871 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, taxonomy, 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.: +39-0382-754707 will be considered. aqua is fully refereed and aims at pub- Fax: +39-0382-754129 lishing manuscripts within 2-4 months of acceptance. In E-mail: [email protected] view of the importance of color patterns in species identi - fication and animal 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 Friedhelm Krupp meeting high scientific and technical standards. Curator of Fishes Senckenberg Research Institute and Natural History Museum Call for papers Senckenberganlage 25 The editors welcome the submission of original manu- 60325 Frankfurt am Main, Germany scripts which should be sent in digital format to the scien- Tel: +49-69-7542.1255 tific editor. Full length research papers and short notes will Fax: +49-69-7542.1253 be considered for publication. There are no page charges E-mail: [email protected] and color illustrations will be published free of charge. Authors will receive one free copy of the issue in which their paper is published and an e-print in PDF format. Editorial Board: Subscription Notice Gerald R. Allen At least one volume (4 issues) of aqua is being published Department of Aquatic Zoology, per year, each issue comprising 48 pages (includ ing cover). Western Australian Museum, Perth, Australia The subscription rate (for one volume = 4 issues) is from volume 12 on: Personal subscription: Euro 75,00 Nina G. Bogutskaya (incl. priority mail); Institutional subscription: Euro 140,00 Zoological Institute of the Russian Academy of (incl. priority mail). Subscription enquires should be sent Sciences, St. Petersburg, Russia to the publisher at the address given below or by e-mail to: [email protected] Wilson J. E. M. Costa Laboratório de Ictiologia Geral e Aplicada, aqua binder Departamento de Zoologia, Binders for Volumes of aqua are available at cost price Universidade Federal do Rio de Janeiro, Brasil Euro 12,50 (US$ 15.00) plus postage Euro 8,00 (US$ 10.00). Notice: aqua Volumes 1(1)-5(4) = 1st. binder; Axel Meyer Volumes 6(1)-9(4) = 2nd. binder; Volumes 10(1)-13(4) Lehrstuhl für Zoologie und Evolutions biologie, = 3rd. binder. Universität Konstanz, Germany Special Publication Paolo Parenti Since 2003 Aquapress publishes a series of Special Publi- Department of Enviromental Sciences, cations, which are produced at irregular intervals. All Spe- University of Milano-Bicocca, cial Publications have about 100 or more pages and are Milan, Italy available separately from regular issues of aqua. Enquiries about subscriptions and prices should be sent to the pub- Mário de Pinna lisher at the address given here above or by e-mail to: Museu de Zoologia da USP, São Paulo, Brazil [email protected] John E. Randall Bishop Museum, Honolulu, Hawaii, U.S.A. ISSN 0945-9871 Publisher: Aquapress, Redazione aqua, Richard Winterbottom I-27010 Miradolo Terme (Pavia), Italy Centre of Biodiversity & Conservation Biology, Printer: PRINTO s.r.o. – Czech Republic Royal Ontario Museum, Copyediting and layout: Rossella Bulla Toronto, Canada © 2007 aqua, International Journal of Ichthyology aqua, International Journal of Ichthyology Revision of the Genus Symphysodon Heckel, 1840 (Teleostei: Perciformes: Cichlidae) based on molecular and morphological characters Heiko Bleher1, Kai N. Stölting2,3, Walter Salzburger2,4, and Axel Meyer2 1) Aquapress, Via G. Falcone 11, I-27010 Miradolo Terme (Pavia), Italy. E-mail: [email protected] 2) Lehrstuhl für Zoologie und Evolutionsbiologie, Department of Biology, University of Konstanz, D-78457 Konstanz, Germany. E-mail: [email protected] 3) Zoologisches Museum, University Zürich, Switzerland. 4) Department of Ecology and Evolution (DEE), University of Lausanne, UNIL Sorge, Le Biophore, CH-1015 Lausanne, Switzerland. Received: 27 March 2007 – Accepted: 03 July 2007 Abstract which may differ substantially in shape. Except for hybrids Systematics of the cichlid genus Symphysodon has been of S. discus x S. haraldi, S. haraldi does not resemble the investigated and three species are recognised: S. discus other two species. A study of geographic distribution pat- Heckel, 1840 (synonym: S. discus willi schwartzi Burgess, terns of the three species was carried out throughout the 1981); S. aequifasciatus Pellegrin, 1904 (synonyms: S. dis- central and lower Amazon basin: in the western Amazon in cus var. aequifasciata Pellegrin, 1904; S. aequifasciata almost every tributary of the Solimões and the Marañon aequifasciata sensu Schultz, 1960; S. Discus Tarzoo – sic – Rivers to Iquitos, and in the eastern part in most tribu- Lyons, 1960); and S. haraldi, Schultz, 1960 (synonyms: S. taries of the Amazon River down to its mouth. The valid aequifasciata haraldi Schultz, 1960; S. aequifasciata axel- names of the three species are: S. discus – the Heckel dis- rodi Schultz, 1960). The present revision is based on DNA cus; S. aequifasciatus – the green discus; and S. haraldi – sequences of partial mitochondrial control regions of 48 the blue discus. The “brown” or “common” discus of the specimens of Symphysodon from 20 different locations in aquarium trade is the same as the “blue” discus. the central and lower Amazon basin, which revealed three In S. aequifasciatus a congruence of genetic and morpho- genetically distinct clades of Symphysodon. One of these logical (colour) characters has been found, whereas some genetic clusters is composed of specimens that morpholog- specimens that would phenotypically be assigned to S. ically are S. discus, but also of S. haraldi and natural haraldi, genetically group also with the S. discus clade. hybrids of S. discus x S. haraldi. This indicates that either Only future studies using nuclear DNA markers will allow the “discus” clade is composed, at least partially, of hybrids untangling the evolutionary history of the phenotypcially or, alternatively, that a “haraldi” phenotype evolved (or was heterogeneous S. “discus” clade. retained) independently in this clade. The other two clades consist of S. aequifasciatus and S. haraldi. Zusammenfassung The definition of the three species is supported by exten- Eine Untersuchung der Discus-Gattung Symphysodon sive field studies over the last 40 years, investigating distri- zeigte, dass drei Arten gültig sind: S. discus Heckel, 1840 butional patterns and documenting adaptation of each (Synonym: S. discus willi schwartzi Burgess, 1981); S. species to a distinct type of water, characterised by unique aequifasciatus Pellegrin, 1904 (Synonyme: S. discus var. chemical parameters. In addition, S. discus and S. aequifas- aequifasciata Pellegrin, 1904; S. aequifasciata aequifasciata ciatus display distinct colorations and colour patterns, with sensu Schultz, 1960, S. Discus Tarzoo – sic – Lyons, nine vertical bars on each flank. In S. discus the first, and 1960); sowie S. haraldi Schultz, 1960 (Synonyme: S. particularly the fifth and ninth bars are prominent and/or aequifasciata haraldi Schultz, 1960; S. aequifasciata axel- wider, while all bars are typically of equal width in S. rodi Schultz, 1960). Die vorliegende Revision basiert auf aequi fasciatus. The latter species is also recognised by its DNA-Sequenzierungen partieller mitochondrialer Kon- rust-brown or red dots on the body, ranging from a few trollabschnitte bei 48 Exemplaren von Symphosodon von spots to a dense cover all over, rarely forming red spotted 20 verschiedenen Fundorten im mittleren und unteren lines or being present in the anal fin region only. Sym- Amazonasbecken, bei denen sich drei genetisch getrennte physodon haraldi displays a wide range of colours, colour Kladen der Gattung Symphysodon zeigten. Eines dieser patterns and a larger number of vertical bars (8 up to 16), genetischen Cluster umfasst Exemplare, die morpholo- 133 aqua vol. 12 no. 4 - 1 August 2007 Revision of the Genus Symphysodon Heckel, 1840 (Teleostei: Perciformes: Cichlidae) based on molecular and morphological characters gisch der zuerst beschriebenen Discus-Art entsprechen: S. S. aequifasciata Pellegrin, 1904 (synonymes: S. discus var. discus, aber auch Exemplare, die S. haraldi und Naturhy- aequifasciata Pellegrin, 1904; S. aequifasciata aequifasciata briden aus S. discus x S. haraldi ähnlich sind. Dies deutet sensu Schultz, 1960; S. Discus Tarzoo – sic – Lyons, darauf hin, dass entweder die Klade „discus” zumindest 1960); et S. haraldi Schultz, 1960 (synonymes: S. aequifas- teilweise aus Hybriden besteht oder dass sich in dieser ciata haraldi Schultz, 1960; S. aequifasciata axelrodi Klade unabhängig ebenfalls ein „haraldi”-Phänotyp Schultz, 1960). Cette révision se base sur des séquençages entwickelt hat (bzw. erhalten blieb). Die anderen beiden ADN mitochondriaux partiels de régions de contrôle de Kladen werden von den bekannten Formen von S. 48 spécimens de Symphysodon de 20 localités différentes aequifasciatus bzw. S. haraldi gebildet. situées dans le bassin central et inférieur de l’Amazone, ce Die Diagnosen der drei