Relationships among the Neotropical Candirus (Trichomycteridae, Siluriformes) and the evolution of parasitism based on analysis of mitochondrial and nuclear gene sequences.

Phylogenetic relationships among the trichomycterid catfishes are investigated for the first time using molecular sequence data. Data derived from mitochondrial and nuclear DNA sequences for representatives of 17 genera were analyzed to test previous hypotheses of relationships among trichomycterid subfamilies, the monophyly of the subfamily Stegophilinae, and the monophyly and relationships among the genera of parasitic members of the family. We analyzed 2325 aligned base-pairs from mitochondrial 12S, 16S, ND4 (tRNA(His) tRNA(Ser)), and the nuclear histone H3 gene for representatives of 10 of 12 stegophiline and 3 of 4 vandelliine genera, plus 10 outgroup taxa selected to represent the range of subfamilial diversity. Maximum parsimony and likelihood approaches resolved a monophyletic semiparasitic Stegophilinae as the sister-group of the obligate hematophagous Vandelliinae. At the level of subfamilies, the pattern of relationships of the parasitic members among the remainder of the family is fully congruent with the most recent hypothesis of relationships for trichomycterids based exclusively on morphological data. Within stegophilines, our results differ from multiple previous morphological studies in recovery of (1) Haemomaster and Ochmacanthus as sister-taxa, (2) the morphologically plesiomorphic Pareidon microps nested within a relatively distal part of the tree topology, (3) Apomatoceros as sister to Henonemus, rather than to the morphologically similar Megalocentor. These result indicate that parasitism arose once and was unreversed within the Trichomycteridae. Survey of diet and feeding morphology among trichomycterids suggests that the semiparasitic lifestyle of the members of the Stegophilinae was retained in the enigmatic Pareiodon microps, despite reversal to the generalized trichomycterid condition of the associated morphological specializations found in all other stegophilines. These results further support the reconstruction of semiparasitism, rather than blood feeding, for the shared common ancestor of the parasitic Trichomycteridae.

Relationships among characiform fishes inferred from analysis of nuclear and mitochondrial gene sequences.

Suprafamilial relationships among characiform fishes and implications for the taxonomy and biogeographic history of the Characiformes were investigated by parsimony analysis of four nuclear and two mitochondrial genes across 124 ingroup and 11 outgroup taxa. Simultaneous analysis of 3660 aligned base pairs from the mitochondrial 16S and cytochrome b genes and the nuclear recombination activating gene (RAG2), seven in absentia (sia), forkhead (fkh), and alpha-tropomyosin (trop) gene loci confirmed the non-monophyly of the African and Neotropical assemblages and corroborated many suprafamilial groups proposed previously on the basis of morphological features. The African distichodontids plus citharinids were strongly supported as a monophyletic Citharinoidei that is the sistergroup to all other characiforms, which form a monophyletic Characoidei composed of two large clades. The first represents an assemblage of both African and Neotropical taxa, wherein a monophyletic African Alestidae is sister to a smaller clade comprised of the Neotropical families Ctenolucidae, Lebiasinidae, and the African Hepsetidae, with that assemblage sister to a strictly Neotropical clade comprised of the Crenuchidae and Erythrinidae. The second clade within the Characoidei is strictly Neotropical and includes all other Characiformes grouped into two well supported major clades. The first, corresponding to a traditional definition of the Characidae, is congruent with some groupings previously supported by morphological evidence. The second clade comprises a monophyletic Anostomoidea that is sister to a clade formed by the families Hemiodontidae, Parodontidae, and Serrasalmidae, with that assemblage, in turn, the sistergroup of the Cynodontidae. Serrasalmidae, traditionally regarded as a subfamily of Characidae, was recovered as the sistergroup of (Anostomoidea (Parodontidae+Hemiodontidae)) and the family Cynodontidae was recovered with strong support as the sistergroup to this assemblage. Our results reveal three instances of trans-continental sistergroup relationships and, in light of the fossil evidence, suggest that marine dispersal cannot be ruled out a priori and that a simple model of vicariance does not readily explain the biogeographic history of the characiform fishes.

Odontode morphology and skin surface features of Andean astroblepid catfishes (Siluriformes, Astroblepidae).

Two types of odontodes, or dermal teeth, occur in the neotropical Andean astroblepid catfishes. Both odontode types conform in structure to dermal teeth of gnathostomes in having dentine surrounding a central pulp cavity covered by a superficial layer of enameloid, but differ from one another in terms of attachment and association with other epidermis features. Type I odontodes in astroblepids, also found in all representatives of the superfamily Loricarioidea, are larger (40-50 microm base diameter), generally conical and sharply pointed, occur on the fin rays, and are associated with dermal bone. Type I odontodes attach to an elevated pediment of dermal bone of the fin lepidotrich, and to dermal bone generally in loricarioids, via a ring of connective tissue. Type II odontodes of astroblepids are smaller (15-20 microm base diameter) and blunt, occur in the skin of the head, maxillary barbels, nasal flap, and lip margins, and are not associated with dermal bone. Observations based on histology and scanning electron microscopy indicate that Type II odontodes are associated with other epithelial structures to form a putative mechanosensory organ. The odontode base lies deep in the dermis. The shaft is surrounded by a dense patch of microvillous epithelium and projects from within a pit formed by an elevated ring of laminar epithelial cells bearing several columnar, knob-like putative mechanosensory structures. Type II odontode organs have thus far been observed in only three astroblepid species, Astroblepus longifilis, A. chotae, A. rosei, where they occur in especially dense arrays on the maxillary barbels, surrounded by discrete patches of microvilli and separate mechanoreceptors. Type II odontode organs are less dense elsewhere on the body, but also occur in the skin of the snout, head, and lips. Typical taste buds are absent from the barbels of these species, but present in other astroblepids. The presence of Type II odontodes and their association with specialized epithelial pit organs are unique to astroblepids among siluriforms and may be potentially important adaptations to life in torrential mountain streams.

TESTING HISTORICAL HYPOTHESES OF MORPHOLOGICAL CHANGE: BIOMECHANICAL DECOUPLING IN LORICARIOID CATFISHES.

The "decoupling hypothesis" has been proposed as a mechanistic basis for the evolution of novel structure and function. Decoupling derives from the release of functional constraints via loss of linkages and/or repetition of individual elements as redundant design components, followed by specialization of one or more elements. Examples of apomorphic decoupling have been suggested for several groups of organisms, however there have been few empirical tests of explicit statements concerning functional and morphological consequences of decoupling. Using the loricarioid catfishes, we tested one particular consequence of decoupling, the prediction that clades possessing decoupled systems having increased biomechanical complexity will exhibit greater morphological variability of associated structures than outgroups having no such decoupled systems. Morphometric procedures based on interlandmark distances were used to quantify morphological variance at three levels of design at successive nodes in the loricarioid cladogram. Additional landmark-based procedures were used to localize major patterns of shape change between clades. We report significantly greater within-group morphometric variance at all three morphological levels in those lineages associated with decoupling events, confirming our predictions under the decoupling hypothesis. Two of 12 comparisons, however, yielded significant variance effects where none were predicted. Localization of the major patterns of shape change suggests that disassociation between morphological and functional evolution may contribute to the lack of fit between variance predictions and decoupling in these two comparisons.