Abstract. The patterns and rates of nucleotide substitution in mitochondrial ribosomal RNA genes are described and applied in a phylogenetic analysis of fishes of the subfamily Serrasalminae (Teleostei, Characiformes, Characidae). Fragments of 345 bp of the 12S and 535 bp of the 16S genes were sequenced for 37 taxa representing all but three genera in the subfamily. Secondary-structure models based on comparative sequence analysis were derived to characterize the pattern of change among paired and unpaired nucleotides, forming stem and loop regions, respectively. Base compositional biases were in the direction of A-rich loops and G-rich stems. Ninety-five percent of substitutions in stem regions were compensatory mutations, suggesting that selection for maintenance of base pairing is strong and that independence among characters cannot be assumed in phylogenetic analyses of stem characters. The relative rate of nucleotide substitution was similar in both fragments sequenced but higher in loop than in stem regions. In both genes, C-T transitions were the most common type of change, and overall transitions outnumbered transversions by a factor of two in 16S and four in 12S. Phylogenetic analysis of the mitochondrial DNA sequences suggests that a clade formed by the genera Piaractus, Colossoma, and Mylossoma is the sister group to all other serrasalmins and that the genera Myleus, Serrasalmus, and Pristobrycon are paraphyletic. A previous hypothesis concerning relationships for the serrasalmins, based on morphological evidence, is not supported by the molecular data. However, phylogenetic analysis of host-specific helminth parasites and cytogenetic data support the phylogeny of the Serrasalminae obtained in this study and provide evidence for coevolution between helminth parasites and their fish hosts.
Previous studies based on DNA sequences of mitochondrial (mt) rRNA genes showed three main groups within the subfamily Serrasalminae: (1) a "pacu" clade of herbivores (Colossoma, Mylossoma, Piaractus); (2) the "Myleus" clade (Myleus, Mylesinus, Tometes, Ossubtus); and (3) the "piranha" clade (Serrasalmus, Pygocentrus, Pygopristis, Pristobrycon, Catoprion, Metynnis). The genus Acnodon was placed as the sister taxon of clade (2+3). However, poor resolution within each clade was obtained due to low levels of variation among rRNA gene sequences. Complete sequences of the hypervariable mtDNA control region for a total of 45 taxa, and additional sequences of 12S and 16S rRNA from a total of 74 taxa representing all genera in the family are now presented to address intragroup relationships. Control region sequences of several serrasalmid species exhibit tandem repeats of short motifs (12 to 33 bp) in the 3' end of this region, accounting for substantial length variation. Bayesian inference and maximum parsimony analyses of these sequences identify the same groupings as before and provide further evidence to support the following observations: (a) Serrasalmus gouldingi and species of Pristobrycon (non-striolatus) form a monophyletic group that is the sister group to other species of Serrasalmus and Pygocentrus; (b) Catoprion, Pygopristis, and Pristobrycon striolatus form a well supported clade, sister to the group described above; (c) some taxa assigned to the genus Myloplus (M. asterias, M tiete, M ternetzi, and M rubripinnis) form a well supported group whereas other Myloplus species remain with uncertain affinities (d) Mylesinus, Tometes and Myleus setiger form a monophyletic group.
A new species of Serrasalmidae, Tometes camunani, is described from the upper drainages of the rio Trombetas basin, Pará State, Brazil. The new species is distinguished from its congeners by having neurocranium with a slight concavity at the level of the frontal bone (vs. concavity absent, dorsal profile of neurocranium straight). It can be further distinguished from its congeners by having teeth with central cusp taller and acute (vs. central cusp shorter and with rounded edge in T. trilobatus), a terminal mouth (vs. upturned mouth in T. lebaili), and 12-26 prepelvic spines (vs. 0-9 in T. makue). The new species is strictly rheophilic like other species of Tometes, and occurs exclusively in the rapids of shield rivers, complex and fragile biotopes that are threatened by anthropogenic activities. An identification key to the species of the Myleus group is provided. Uma espécie nova de Serrasalmidae, Tometes camunani, é descrita para as drenagens superiores da bacia do rio Trombetas, estado do Pará, Brasil. A espécie nova distingue-se dos congêneres pela presença de uma ligeira concavidade no neurocrânio na altura do frontal (vs. concavidade ausente, perfil dorsal do neurocrânio reto). Também pode ser adicionalmente distinguido dos seus congêneres por possuir dentes com a cúspide central mais alta e cume agudo (vs. cúspide central mais baixa e com cume arredondado em T. trilobatus), a boca terminal (vs. boca orientada para cima em T. lebaili), e 12-26 espinhos pré-pélvicos (vs. 0-9 em T. makue). A espécie nova é estritamente reofílica, como as demais espécies de Tometes, e ocorre exclusivamente nas zonas encachoeiradas dos rios de escudo, biótopos complexos, frágeis e ameaçados por ações antropogênicas. Uma chave de identificação para as espécies do grupo Myleus é apresentada.
Common names: eaglebeak pacu (US), Adlerschnabel-pacu (GER), pacu-capivara, pacu-tromba-de-anta (BRA). Conservation status: Endangered. Identification: D 21-23, A 23-26, vertebrae 38. The body is ovoid. The profile of the snout is blunt. Among the serrasalmins this species is characterized by a series of synapomorphies linked to its uncommon rotation of the head plan during the early stages of life. The mouth is terminal in postlarval stage, slightly downturned in specimens of around 30 mm SL and strictly ventral in specimens larger than 50 mm SL. The lower edge of the eye is located very dorsally of the commissure of the mouth. The premaxillary has five teeth in the labial series and two in the lingual series. The two median teeth of the labial series are reduced to very short canines. The lower jaw has four incisiform teeth on a single series. The pectoral fins are placed in a very low position on the sides. There are 19-21 branched rays on the dorsal fin and 22-25 on the anal fin. The number of scales in the lateral line varies from 81 to 88 to the hypural. There are 36-42 series of circumpeduncular scales. There are 11-16 postpelvic serrae of which 5-9 pairs of spines are found around the cloaca, but there are no prepelvic serrae. The number of branchiospines varies from 9-10 on the upper branch of the first branchial arch and from 11 to 16 on the lower branch. Maximum SL about 176 mm. Distribution: The presence of Ossubtus xinguense is confirmed in Xingu basin (Para´, Brazil) but restricted to the rapids area around Altamira city. Abundance: Unknown, but apparently rare. Habitat and ecology: Ossubtus xinguense is a strictly rheophilic species. Young specimens of up to 40 mm SL could be observed in schools of 20-30 individuals, sheltered under broad stones in the rapids near Altamira. Large specimens can be captured by castnet in the rapids.
Two new species of Tometes from the Brazilian Shield rapids are described. Tometes kranponhah is endemic to the Xingu River basin, whereas Tometes ancylorhynchus occurs both in the Xingu and the Tocantins-Araguaia River basins. The two species are sympatric in the Xingu drainages and have many similarities in morphology and colouration. Major diagnostic differences are the dark pigmentation on the opercle of T. kranponhah and its distinct snout shape and arrangement of premaxillary teeth. In addition, T. kranponhah is a large fish that is abundant in the Xingu River, whereas T. ancylorhynchus is a medium-sized fish for which there are few records.
A new large serrasalmid species of Tometes is described from the Tocantins-Araguaia River Basin. Tometes siderocarajensis sp. nov. is currently found in the rapids of the Itacaiúnas River Basin, and formerly inhabited the lower Tocantins River. The new species can be distinguished from all congeners, except from T. ancylorhynchus, by the presence of lateral space between 1st and 2nd premaxillary teeth, and by the absence of lateral cusps in these two teeth. However, T. siderocarajensis sp. nov. can be differentiated from syntopic congener T. ancylorhynchus by an entirely black with mottled red body in live specimens, densely pigmented pelvic fins with a high concentration of dark chromatophores, and the presence of 39 to 41 rows of circumpeduncular scales (vs. silvery body coloration with slightly reddish overtones on middle flank during breeding period in live specimens, hyaline to slightly pale coloration on distalmost region of pelvic fins, and 30 to 36 rows of circumpeduncular scales). Additionally, molecular sequence shows that T. siderocarajensis sp. nov. is reciprocally monophyletic, and diagnosable from all congeners by having two autapomorphic molecular characters in the mitochondrial gene COI. The phylogenetic reconstruction still show that T. siderocarajensis sp. nov. is closely related to T. trilobatus. This is the first molecular study using an integrative taxonomic approach based on morphological and molecular sequence data for all described species of Tometes. These findings increase the number of formally described species of Tometes to seven. A key to the Tometes species is provided.
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