Gill arch and hyoid arch diversity and cypriniform phylogeny: Distributed integration of morphology and web-based tools

Abstract

Using the multiple tools available to support an online collaborative environment, we surveyed 62 morphological features from the hyoid arch and gill arches of 53 species of cypriniform fishes that matched those sampled in recent molecularanalyses and two sets of outgroup species (‘Saitoh outgroups’ and ‘Basal outgroups’). This is a skeletal region whose variation is considered historically significant within fishes and Cypriniformes in particular, and we review previous work in light of our own. The clarity of description of characters was enhanced by the use of a community reference ontology, the Teleost Anatomy Ontology. Terms, synonyms, and definitions for skeletal features from this region were contributed to this ontology, and links to these terms and relationships are included in our character descriptions. One thousand two hundred sixty-three images of features from this region were linked to ontology terms and deposited in a community image repository, Morphbank; these are linked to the characters described herein. Character data were analyzed using parsimony and Bayesian methods for two sets of outgroups, one of which matched that used in previous molecular analyses. The parsimony results, using either outgroup set, indicated similar higher-level relationships, including a sister group relationship between cyprinids and loaches. A basal trichotomy among Gyrinocheilus, catostomids and cyprinids + loaches was the result of Saitoh outgroup analysis in contrast to a sister group relationship between Gyrinocheilus and catostomids discovered in the Basal outgroup analysis. Interestingly, analyses including basal outgroups recovered a monophyletic Cyprinidae, consistent with all previous studies. Character evolution that supports higher-level nodes of interest in the consensus tree is described. In some respects, it might be a surprise that 62 morphological characters from a small skeletal region for only 53 cypriniform species (out of over 3,200 total species) could provide as much resolution as it does. We expect, however, further phylogenetic resolution as morphological data from across skeleton regions are combined, species sampling is increased, and molecular data are added.