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Revision of the Genus Symphysodon Heckel, 1840 (Teleostei: Perciformes: Cichlidae) Based on Molecular and Morphological Characters

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Revision of the Genus Symphysodon Heckel, 1840 (Teleostei: Perciformes: Cichlidae) Based on Molecular and Morphological Characters 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 Arten wurde durch umfangreiche qui a révélé trois clades de Symphysodon génétiquement Freilandstudien während der letzten 40 Jahre abgesichert. distincts. Un de ces groupes génétiques se compose de Dazu gehörten Untersuchungen der Verteilungsmuster spécimens qui sont morphologiquement des S. discus, mais und die Dokumentation von Anpassungen der drei Arten aussi de S. haraldi et d’hybrides naturels de S. discus x S. an unterschiedliche Gewässertypen mit distinkten chemis- haraldi. Ceci révèle soit que le clade “discus” se compose, chen Parametern. Hinzu kommt, dass zwei der drei Arten du moins en partie, d’hybrides, soit, alternativement, (S. discus und S. aequifasciatus) gut unterscheidbare qu’un phénotype “haraldi” a évolué (ou a été retenu) Grundfarben und Farbmuster aufweisen. Die Vertreter indépendamment dans ce clade. Les deux autres clades dieser beiden Arten besitzen jeweils neun senkrechte sont constitués de S. aequifasciatus et de S. haraldi. Streifen, aber bei S. discus sind der erste, besonders aber der La délimitation de ces trois espèces s’appuie sur des fünfte und neunte Streifen besonders auffällig und/oder études de terrain extensives, au long de ces 40 dernières breit. Bei S. aequi fasciatus hingegen sind die neun Streifen années, qui ont porté sur leur mode de distribution et sur typischerweise gleich breit; außerdem lässt sich diese Art l’adaptation de chaque espèce à un type d’eau particulier, an ihren rostbraunen bis dunkelroten verstreuten Flecken caractérisé par des paramètres chimiques propres. En erkennen, von denen nur wenige vorhanden sein können, outre, S. discus et S. aequifasciatus ont des couleurs et des die sich aber auch über den gesamten Körper erstrecken patrons de coloration différents, avec neuf barres verticales können oder in seltenen Fällen gar rote Linien aus Flecken sur les flancs. Pour le S. discus, la première et surtout la bilden oder auf den Analflossenbereich beschränkt sind. cinquième et la neuvième barre sont accentuées et/ou plus Die dritte Art (S. haraldi) kann die unterschiedlichsten larges, alors que toutes les barres sont typiquement de Farben und Muster zeigen. Die Zahl der senkrechten même largeur pour S. aequifasciatus. Cette dernière espèce Streifen kann stark variieren (8 bis zu 16), und die se distingue aussi par ses taches brun rouille ou rouges sur Streifenbreite kann ganz unterschiedlich sein, von breit bis le corps, pouvant varier de quelques-unes à une répartition sehr schmal. Diese Art ist von den anderen beiden gut zu dense sur tout le corps, pouvant former rarement des lignes unterscheiden (die Hybriden zwischen S. haraldi x S. dis- pointillées rouges ou présentes seulement dans la région de cus unberücksichtigt). l’anale. Symphysodon haraldi arbore une large gamme de Im zentralen und unteren Amazonasbecken wurden die couleurs, de patrons de coloration et un plus grand nom- geografische Verteilung der drei Arten eingehend unter- bre de barres verticales (8 jusqu’à 16), dont la forme peut sucht: am westlichen Amazonas nahezu in allen Nebenar- varier notablement. Hormis les hybrides de S. discus x men des Solimões und des Marañon bis Iquitos und im S. haraldi, S. haraldi ne ressemble pas aux deux autres östlichen Teil des Amazonas in den meisten Nebenflüssen espèces. Une étude des modes de di stribution géo- des Amazonas bis hin zur Mündung. graphique des trois espèces a été menée tout au long du Die gültigen Namen der drei Arten sind: S. discus – bassin central et inférieur de l’Amazone: dans l’Amazone Heckel-Discus; S. aequifasciatus – Grüner Discus; und S. occidental, dans presque chaque tributaire du Solimões et haraldi – Blauer Discus. Bei dem im Aquarienhandel du Marañon
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