title: A test of color-based taxonomy in nudibranchs: Molecular phylogeny and species delimitation of the Felimida clenchi (Mollusca: Chromodorididae) species complex.
abstract: Traditionally, species identification in nudibranch gastropods relies heavily on body color pattern. The Felimida clenchi species complex, a group of brightly colored Atlantic and Mediterranean species in the family Chromodorididae, has a history of exceptional controversy and discussion among taxonomists. The most widely accepted hypothesis is that the complex includes four species (Felimida clenchi, F. neona, F. binza and F. britoi), each with a characteristic body color pattern. In this study, we investigated the taxonomic value of coloration in the Felimida clenchi complex, using molecular phylogenetics, species-delimitation analyses (ABGD, GMYC, PTP), haplotype-network methods, and the anatomy of the reproductive system. None of our analyses recovered the traditional separation into four species. Our results indicated the existence of three species, a result inconsistent with previous taxonomic hypotheses. We distinguished an undescribed species of Felimida and redefined the concepts of F. clenchi and F. binza, both highly polychromatic species. For the first time, molecular data support the existence of extreme color polymorphism in chromatic nudibranch species, with direct implications for the taxonomy of the group and its diversity. The polychromatism observed in the F. clenchi complex apparently correlates with the regional occurrence of similar color patterns in congeneric species, suggesting different mimicry circles. This may represent a parallel in the marine environment to the mechanisms that play a major role in the diversification of color in terrestrial and fresh-water chromatic groups, such as heliconian butterflies.
authors.affiliation: SNSB-Bavarian State Collection of Zoology (ZSM), Münchhausenstrasse 21, 81247 Munich, Germany; Biozentrum and GeoBioCenter, Ludwig-Maximilians-Universität München, Großhadernerstraße 2, 82152 Planegg-Martinsried, Germany. Electronic address: viniciuspadula@yahoo.com.
authors.forename: Vinicius
authors.initials: V
authors.lastname: Padula
authors.affiliation: SNSB-Bavarian State Collection of Zoology (ZSM), Münchhausenstrasse 21, 81247 Munich, Germany; Biozentrum and GeoBioCenter, Ludwig-Maximilians-Universität München, Großhadernerstraße 2, 82152 Planegg-Martinsried, Germany.
authors.forename: Juliana
authors.initials: J
authors.lastname: Bahia
authors.affiliation: SNSB-Bavarian State Collection of Zoology (ZSM), Münchhausenstrasse 21, 81247 Munich, Germany.
authors.forename: Isabella
authors.initials: I
authors.lastname: Stöger
authors.affiliation: Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, 11501-2060 San Pedro, San José, Costa Rica.
authors.forename: Yolanda
authors.initials: Y
authors.lastname: Camacho-García
authors.affiliation: Phylogenetic Systematics and Evolution Research Group, Section of Taxonomy and Evolution, Department of Natural History, University Museum of Bergen, University of Bergen, PB 7800, 5020 Bergen, Norway.
authors.forename: Manuel António E
authors.initials: MAE
authors.lastname: Malaquias
authors.affiliation: Departamento de Biología, Facultad de Ciencias del Mar y Ambientales, Campus de Excelencia Internacional del Mar (CEI·MAR), Universidad de Cádiz, Polígono Río San Pedro, s/n, Ap. 40, 11510, Puerto Real, Cádiz, Spain.
authors.forename: Juan Lucas
authors.initials: JL
authors.lastname: Cervera
authors.affiliation: SNSB-Bavarian State Collection of Zoology (ZSM), Münchhausenstrasse 21, 81247 Munich, Germany; Biozentrum and GeoBioCenter, Ludwig-Maximilians-Universität München, Großhadernerstraße 2, 82152 Planegg-Martinsried, Germany.
authors.forename: Michael
authors.initials: M
authors.lastname: Schrödl
mesh_terms: D000349:Africa; D000818:Animals; D001938:Brazil; D017691:Caribbean Region; D045304:Cytochromes c / Q000145:classification / Q000235:genetics / Q000378:metabolism; D004247:DNA / Q000737:chemistry / Q000302:isolation & purification / Q000378:metabolism; D030541:Databases, Genetic; D049851:Gastropoda / Q000145:classification*; D006239:Haplotypes; D006657:Histones / Q000145:classification / Q000235:genetics / Q000378:metabolism; D010802:Phylogeny; D058974:Phylogeography; D010858:Pigmentation; D012336:RNA, Ribosomal, 16S / Q000145:classification / Q000235:genetics / Q000378:metabolism; D012339:RNA, Ribosomal, 28S / Q000145:classification / Q000235:genetics / Q000378:metabolism; D016415:Sequence Alignment; D017422:Sequence Analysis, DNA
chemical_list: D006657:Histones; D012336:RNA, Ribosomal, 16S; D012339:RNA, Ribosomal, 28S; D045304:Cytochromes c; D004247:DNA
keywords: Color polymorphism; Evolution; Integrative taxonomy; Mimicry; Nudibranchia; Species delimitation