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The endozoic ciliates of the family Clevelandellidae Kidder, 1938 typically inhabit the hindgut of wood-feeding panesthiine cockroaches. To assess the consistency of species delimitation in clevelandellids, we tested the utility of three sources of taxonomic data: morphometric measurements, cell geometrical information, and 18S rRNA gene sequences. The morphometric and geometrical data delimited the clevelandellid morphospecies consistently and unambiguously. However, only Paraclevelandia brevis Kidder, 1937 represented a homogenous taxon in both morphological and molecular analyses; the morphospecies Clevelandella constricta (Kidder, 1937) and C. hastula (Kidder, 1937) contained two or three distinct, more or less closely related genotypes each; and the genetic homogeneity of the morphospecies C. panesthiae (Kidder, 1937) and C. parapanesthiae (Kidder, 1937) was not corroborated by the 18S rRNA gene sequences at all. Moreover, the 18S rRNA gene phylogenies suggested the C. panesthiae-like morphotype to be the ancestral phenotype from which all other clevelandellid morphotypes arose. The only exception was the C. constricta-like morphotype, which very likely branched off before the diversification of the C. panesthiae-like progenitor. The present molecular analyses also suggested that a huge proportion of the clevelandellid diversity still waits to be discovered, since examination of only four panesthiine populations revealed 10 distinct clevelandellid genotypes/molecular species.
Four earthworm species, the endogeic Octolasion tyrtaeum (Savigny, 1826), the anecic Lumbricus terrestris Linnaeus, 1758 as well as the epigeic Eisenia fetida (Savigny, 1826) and Dendrobaena veneta (Rosa, 1886), were examined for the presence of astome ciliates. Based on the integrative taxonomic approach, five ciliate species were recognized in their gastrointestinal tracts: Metaradiophrya lumbrici (Dujardin, 1841), M. varians (de Puytorac, 1954), Anoplophrya lumbrici (Schrank, 1803), A. vulgaris de Puytorac, 1954 and A. nodulata (Dujardin, 1841). Their distinctness was assessed using the multivariate morphometric approach and molecular phylogenetic analyses. Although the two species of Metaradiophrya Jankowski, 2007 on the one hand and the two former species of Anoplophrya Stein, 1860 on the other, were not distinctly separated by the multivariate morphometric analyses, they were clearly delimited by the 18S rRNA gene sequences. Species within each genus also differed by their hosts, M. lumbrici and A. lumbrici occurred only in anecic earthworms while M. varians and A. vulgaris occured exclusively in epigeic earthworms. Only a single species, A. nodulata, was detected in endogeic earthworms. It was morphologically distinct from and did not cluster with the two other species of Anoplophrya but was nested within the paraphyletic assemblage containing other astomes from endogeic earthworms. This indicates that the evolution of endosymbiotic ciliates from earthworms has very likely proceeded through a specialization to various ecological groups of their host organisms.