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At least 19 recordsLinked to original sources

A Bayesian approach on molecules and behavior: reconsidering phylogenetic and evolutionary patterns of the Salamandridae with emphasis on Triturus newts.

The monophyly of European newts of the genus Triturus within the family Salamandridae has for decades rested on presumably homologous behavioral and morphological characters. Molecular data challenge this hypothesis, but the phylogenetic position of Triturus within the Salamandridae has not yet been convincingly resolved. We addressed this issue and the temporal divergence of Triturus within the Salamandridae with novel Bayesian approaches applied to DNA sequence data from three mitochondrial genes (12S, 16S and cytb). We included 38 salamandrid species comprising all 13 recognized species of Triturus and 16 out of 17 salamandrid genera. A clade comprising all the "Newts" can be separated from the "True Salamanders" and Salamandrina clades. Within the "Newts" well-supported clades are: Tylototriton-Pleurodeles, the "New World Newts" (Notophthalmus-Taricha), and the "Modern Eurasian Newts" (Cynops, Pachytriton, Paramesotriton=together the "Modern Asian Newts", Calotriton, Euproctus, Neurergus and Triturus species). We found that Triturus is a non-monophyletic species assemblage, which includes four groups that are themselves monophyletic: (i) the "Large-Bodied Triturus" (six species), (ii) the "Small-Bodied Triturus" (five species), (iii) T. alpestris and (iv) T. vittatus. We estimated that the last common ancestor of Triturus existed around 64 million years ago (mya) while the root of the Salamandridae dates back to 95 mya. This was estimated using a fossil-based molecular dating approach and an explicit framework to select calibration points that least underestimated their corresponding nodes. Using the molecular phylogeny we mapped the evolution of life history and courtship traits in Triturus and found that several Triturus-specific courtship traits evolved independently.

Animals↗

Using the volumetric indices of telencephalic structures to distinguish Salamandridae and Plethodontidae: comparison of three statistical methods.

The aim of this study was to establish whether appropriate statistical analysis of 16 volumetric indices corresponding to 16 structures making up the entire telencephalon of Urodela could distinguish between two families, Salamandridae and Plethodontidae. We compared the efficiency of three statistical methods (stepwise discriminant analysis, artificial neural networks, correspondence factor analysis) and the information they provide. All three methods found the same species difficult to classify. However, only correspondence factor analysis could suggest explanations for "misclassifications" as it superimposes the two sets of variables, (sub)species and anatomical variables, thus revealing the correlations between them. The bulbus olfactorius accessorius and the caudal mitral cell layer of the bulbus olfactorius principalis were the most discriminatory structures in separating Salamandridae and Plethodontidae. The correspondence factor analysis mapped species very much in line with accepted taxonomy and highlighted several current controversies [e.g. positioning of certain newts (T. marmoratus, T. vulgaris, T. alpestris), of Salamandrina terdigitata, and of members of the genus Euproctus]. Mapping of Plethodontidae was less clear-cut than that of Salamandridae with more overlap among genera but was quite consistent with knowledge of brain structure complexification. We conclude that relationships derived from analyses of telencephalic structures provide valuable information that might help resolve ambiguities; we have coined the term "neurotaxonomy" for this approach.

Animals↗

Pigment patterns of larval salamanders (Ambystomatidae, Salamandridae): the role of the lateral line sensory system and the evolution of pattern-forming mechanisms.

In many species of salamanders, pigment cells derived from the neural crest give rise to a horizontal stripe pattern in hatchling larvae. A defining element of these horizontal stripe patterns is a region over the middle of the myotomes that is relatively free of melanophores. This study shows that formation of a "melanophore-free region" and horizontal stripe pattern in Ambystoma tigrinum tigrinum (family Ambystomatidae) correlates with the development of the trunk lateral line sensory system. Moreover, prevention of lateral line development results in greater densities of melanophores in the middle of the flank, essentially eliminating the melanophore-free region in this taxon. A phylogenetic survey also revealed that ablation of the lateral lines has qualitatively similar effects on melanophores in seven of eight additional taxa (Ambystomatidae: A. barbouri, A. maculatum, A. talpoideum; Salamandridae: Notophthalmus viridescens, Pleurodeles waltl, Taricha granulosa, T. rivularis). In Taricha torosa, however, a superficially similar melanophore-free region forms prior to lateral line development, and ablation of the lateral lines does not perturb the horizontal stripe pattern. Finally, heterospecific grafting experiments demonstrated that T. torosa lateral lines are competent to generate a melanophore-free region, and T. torosa melanophores are competent to respond to cues associated with the lateral lines. These results indicate that lateral line-dependent pattern-forming mechanisms are common and probably ancestral within the families Ambystomatidae and Salamandridae and suggest that these ancestral mechanisms have been retained in T. torosa as redundant, lateral line-dependent mechanisms for stripe formation have evolved.

Ambystomatidae↗

A molecular assessment of phylogenetic relationships and lineage accumulation rates within the family Salamandridae (Amphibia, Caudata).

We examine phylogenetic relationships among salamanders of the family Salamandridae using approximately 2700 bases of new mtDNA sequence data (the tRNALeu, ND1, tRNAIle, tRNAGln, tRNAMet, ND2, tRNATrp, tRNAAla, tRNAAsn, tRNACys, tRNATyr, and COI genes and the origin for light-strand replication) collected from 96 individuals representing 61 of the 66 recognized salamandrid species and outgroups. Phylogenetic analyses using maximum parsimony and Bayesian analysis are performed on the new data alone and combined with previously reported sequences from other parts of the mitochondrial genome. The basal phylogenetic split is a polytomy of lineages ancestral to (1) the Italian newt Salamandrina terdigitata, (2) a strongly supported clade comprising the "true" salamanders (genera Chioglossa, Mertensiella, Lyciasalamandra, and Salamandra), and (3) a strongly supported clade comprising all newts except S. terdigitata. Strongly supported clades within the true salamanders include monophyly of each genus and grouping Chioglossa and Mertensiella as the sister taxon to a clade comprising Lyciasalamandra and Salamandra. Among newts, genera Echinotriton, Pleurodeles, and Tylototriton form a strongly supported clade whose sister taxon comprises the genera Calotriton, Cynops, Euproctus, Neurergus, Notophthalmus, Pachytriton, Paramesotriton, Taricha, and Triturus. Our results strongly support monophyly of all polytypic newt genera except Paramesotriton and Triturus, which appear paraphyletic, and Calotriton, for which only one of the two species is sampled. Other well-supported clades within newts include (1) Asian genera Cynops, Pachytriton, and Paramesotriton, (2) North American genera Notophthalmus and Taricha, (3) the Triturus vulgaris species group, and (4) the Triturus cristatus species group; some additional groupings appear strong in Bayesian but not parsimony analyses. Rates of lineage accumulation through time are evaluated using this nearly comprehensive sampling of salamandrid species-level lineages. Rate of lineage accumulation appears constant throughout salamandrid evolutionary history with no obvious fluctuations associated with origins of morphological or ecological novelties.

Animals↗

Molecular phylogeny of the genus Paramesotriton (Caudata: Salamandridae).

To elucidate the phylogeny of the genus Paramesotriton (Caudata: Salamandridae), we investigated three mitochondrial DNA gene fragments (1207 bp in total) of cytochrome b, ND2, and ND4 for its six recognized species. The phylogenetic relationships within Paramesotriton were reconstructed by maximum parsimony (MP) and maximum likelihood (ML) methods. Phylogenetic trees (MP and ML trees) that were constructed from the combined data set of the three gene fragments indicated that all six species of Paramesotriton formed a monophyletic group, with P. caudopunctatus as basal to the other five species. This result suggests that P. fuzhongensis is a valid species in Paramesotriton.

Animals↗

Comparative anatomy and phylogeny of the cloacae of salamanders (Amphibia: Caudata). IV. Salamandridae.

Cloacae were examined from male and female salamanders representing 12 genera and 22 species in the Salamandridae. All female salamandrids possess numerous sperm storage glands, spermathecae, in the roof of the cloaca, but intergeneric variation exists in the occurrence of additional cloacal glands. Pleurodeles and Tylototriton possess both vent and anterior ventral glands, and secondary loss has occurred of vent glands in all other genera and anterior ventral glands in Chioglossa, Cynops, Paramesotriton, and Triturus The most highly derived cloaca occurs in Euproctus asper, in which the cloacal tube extends through a conical projection, and ventral glands secrete onto the dorsolateral surface of the projection rather than into the cloaca. Marked intergeneric variation occurs in males in conformation of the cloacal cavities and in extent of the dorsal gland. In Cynops, Euproctus, Pachytriton, Paramesotriton, Taricha, and Triturus, the pseudopenis (a broad, posteriorly projecting evagination of the dorsal roof) fills much of the cavity of the anterior cloacal chamber. In most salamandrids, distal ends of the dorsal glands occur lateral to pelvic glands in the anterior end of the cloaca, and dorsal gland tubules descend to secretory sites at the posterior end of the vent. Salamandra and Mertensiella possess a unique, bifurcated dorsal gland in which distal ends of tubules lie dorsal to the other cloacal glands, and proximal ends curve ventrally in the anterior end of the cloaca to secretory sites along the cloacal orifice. Cladistic analyses indicate that the variation in presence of anterior ventral glands is due to homoplasy. The occurrence of female vent glands, bifurcated dorsal glands, and the pseudopenis supports a phylogeny based upon non-cloacal characters.

Animals↗

Retinal projections in European Salamandridae.

The retinal projections to the brain were studied in three species of European Salamandridae using anterograde transport of horseradish peroxidase and autoradiography. The results obtained were basically identical for all species and confirmed earlir findings on the fiber supply to the preoptic nucleus and the basal optic neuropil. In the anterior thalamus projections to three distinct terminal fields are clearly visible: (i) the diffusely stained corpus geniculatum thalamicum, (ii) the neuropil of Bellonci, pars lateralis, and (iii) a dorsomedial terminal field, the neuropil of Bellonci, pars medialis. Caudal to these terminal fields is an almost terminal-free region, the lateral neuropil. In the posterior thalamus a medial terminal field, the uncinate field, and a laterally located terminal field, the posterior thalamic neuropil, are distinguishable. The tectum opticum displays as many as four dense layers of retinofugal fibers and terminals in the rostral part and, in addition, a more densely stained strip of neuropil running from rostral to caudal over the tectum. The extent of ipsilateral fibers is greater than previously reported in other urodele species. They supply the medial and the lateral parts of the neuropoil of Bellonci, the uncinate field, and reach the tectum opticum via the medial optic tract. Further, they form terminals in the innermost optic fiber layer throughout the rostral half of the ipsilateral tectum. A small proportion of ipsilateral fibers contributes very sparsely to all other thalamic terminal fields, leaving only the caudal part of the tectum and several layers of the rostral tectum completely free of a direct retinofugal fiber supply.

Animals↗

Two dispersed highly repeated DNA families of Triturus vulgaris meridionalis (Amphibia, Urodela) are widely conserved among Salamandridae.

Two BamHI families of repeated sequences were characterized from the genome of the Italian smooth newt, Triturus vulgaris meridionalis (Amphibia, Urodela). The first family, which is divided into subfamilies, consists of tandemly arranged arrays whose basic repeat is around 398 bp long; these arrays are dispersed throughout the entire chromosome sets of the various species of Triturus tested. Moreover the family is widely conserved among Salamandridae, being detected by genomic DNA blotting of Notophthalmus viridescens, Taricha granulosa, Salamandrina terdigitata and Euproctus platycephalus. The second BamHI family is represented by a cloned sequence of 419 bp, which is dispersed in the chromosome set of several species of Triturus. The sequence is also conserved in S. terdigitata and in E. platycephalus but is not detectable in N. viridescens or T. granulosa. The cloned sequence is most probably only part of a longer unit interspersed within the Triturus genome.

Animals↗

Paedogenesis in european newts (Triturus: salamandridae): cranial morphology during ontogeny.

A cross-sectional analysis using different ontogenetic stages (larvae, juveniles, paedotypic, and metamorphic adults) of the smooth newt, Triturus vulgaris, and the alpine newt, T.alpestris, revealed a broad spectrum of perennibranchiation influences on cranial ontogeny in European newts, more pronounced than previously thought. These influences included marked variation in ossification levels, pronounced morphometric variability of many cranial elements, and considerable skull shape changes in the transition from larvae to the adult stage. In comparison with metamorphosed individuals, paedotypic newts had a higher level of variability in both individual cranial traits and cranial shape changes. Sexual size difference of the skull traits was mostly negligible, especially in comparison to the influence of paedogenesis. The main changes in cranial shape of the European newts occurred during metamorphosis. Cranial morphological organization in the majority of examined paedotypes corresponds to cranial organization at late larval stages prior to metamorphosis or at the onset of metamorphosis.

Animals↗

Topologic and connectional analysis of the dorsal thalamus of Triturus alpestris (amphibia, urodela, salamandridae).

A morphological and connectional analysis was performed on the dorsal thalamus of the alpine newt, Triturus alpestris. We have used a graphic reconstruction technique for the evaluation of the connectional (HRP) data. On the basis of these reconstructions, we propose a subdivision of the salamandrid dorsal thalamus into subhabenular, anteroventral, and posterodorsal zones. Each of these zones is defined by its telencephalic projections ("ascending thalamofugal systems"). The posterodorsal zone projects to the striatum, the anteroventral zone to the pallium. The subhabenular zone projects to the subpallial telencephalon and to the tegmentum. This zonal subdivision allows a more detailed comparison of the salamandrid dorsal thalamic features with ranid dorsal thalamic structures. We compare our dorsal thalamic zones to the ones proposed by Herrick (J. Comp. Neurol. 62:239-261, '35, The Brain of the Tiger Salamander. Chicago: The University of Chicago Press, '48). Furthermore, using the same reconstructive technique, we undertook an analysis of the spatial relations of various inputs to the salamandrid dorsal thalamus ("thalamopetal systems"). Besides the well-known retinal inputs, we identified the tectum and the tegmentum as sources of inputs to the thalamus. We provide evidence that there is no extensive multi- or unimodal overlap of these thalamopetal systems.

Afferent Pathways↗

Homology and evolutionary novelty in the deployment of extracellular matrix molecules during pigment pattern formation in the salamanders Taricha torosa and T. rivularis (Salamandridae).

Salamander larvae exhibit a diverse array of pigment patterns shortly after hatching. Previous studies have identified roles for the extracellular matrix and lateral line sensory system in promoting the development of a phylogenetically common pattern of horizontal melanophore stripes. In contrast, salamanders in the genus Taricha exhibit evolutionarily derived pigment patterns and pattern-forming mechanisms. Taricha torosa larvae exhibit compact melanophore stripes that develop via redundant, lateral line-independent mechanisms, whereas T. rivularis larvae lack stripes and instead have melanophores uniformly distributed over the flank. In this study, I test roles for candidate patterning molecules of the extracellular matrix in promoting the development of species-specific pigment patterns in Taricha. I show that tenascin deposition is negatively correlated with melanophore distributions both intraspecifically and interspecifically: this matrix molecule is present where melanophores do not localize in T. torosa and is absent from these same regions where melanophores are abundant in T. rivularis. Embryological manipulations further indicate that transient expression of tenascin in a prospective interstripe region of T. torosa reflects a phylogenetically conserved effect of lateral line development. Finally, anti-laminin immunoreactivity is negatively correlated with melanophore distributions in T. torosa, and this species exhibits a general retardation of extracellular matrix development that may allow persistent, evolutionarily novel melanophore motility in this species. Together these findings identify tenascin and laminin, or molecules co-regulated with these matrix components, as candidates for promoting early larval pigment pattern development in Taricha.

Animals↗

Ontogeny of cranial ossification in the eastern newt, Notophthalmus viridescens (Caudata: Salamandridae), and its relationship to metamorphosis and neoteny.

The ontogenetic sequence of cranial osteogenesis through adulthood is described in samples of newts from completely metamorphosing and partially neotenic populations. Cranial ossification proceeds in the same sequence in both samples. Seven stages of cranial development are described on the basis of conspicuous events that occur during ontogeny. These include four larval stages, metamorphs, efts, and adults. Neotenic adults have skulls that are metamorphosed completely and indistinguishable from the skulls of non-neotenic adults. Neoteny in these newts does not involve the skull and is limited to the postmetamorphic retention of some gill structures and, thus, is termed "limited neoteny." The evolution of limited neoteny in newts as a correlated response to the inhibition of land-drive behavior is discussed.

Aging↗

MtDNA phylogeography of the California newt, Taricha torosa (Caudata, Salamandridae).

Thirty-six individuals of the California newt, Taricha torosa, representing 22 populations from throughout the range of the two currently recognized subspecies, torosa and sierrae, were examined for sequence variation in a segment (375 bp) of the mitochondrial (mt) cytochrome b gene. The maximum sequence divergence within T. torosa is about 9%. Phylogenetic analyses used the sister taxa T. rivularis and T. granulosa as outgroups. Eighteen haploid sequence types found in T. torosa were grouped by parsimony, maximum likelihood, and neighbor-joining analyses into five mitochondrial clusters: two in torosa (the northern and southern clusters) and three in sierrae (the northern, central, and southern clusters). The southern sierrae cluster apparently shared a most recent common ancestor with the northern torosa cluster. The approximate time of sequence divergence within the current species was calibrated using the known fossil record (0.8% divergence per million years or 0.01 maximum likelihood distance per million years). Phylogenetic implications of mtDNA sequence variation for evolution and biogeography of the T. torosa species complex are discussed.

Amino Acid Sequence↗

Chromosomal variation in the northwestern American newts of the genus Taricha (Caudata: Salamandridae).

Chromosomal variation was analysed in 22 populations of newts of the salmandrid genus Taricha of western North America, and compared with that of the eastern North American newts of the genus Notophthalmus. The karyotypes of the species Taricha and Notophthalmus were very similar. However, there was considerable variation in the distribution patterns of heterochromatins (revealed by C-banding) and in the sites of the nucleolar organizing region (NOR) (revealed by fluorochrome chromomycin A3 banding) within and between species of these two genera. Chromosomal variation patterns were interpreted in relation to a phylogenetic hypothesis derived from data on mitochondrial DNA sequences and allozyme variation. This study suggests that the pattern distributions of heterochromatins in chromosomes of Taricha are more derived than those of its sister taxon Notophthalmus. Furthermore, the chromosomal NOR types found in the southernmost and northernmost populations of T. granulosa, in the northernmost populations of T. t. sierrae, and in the southern populations of T. t. torosa are recently derived. The implications of this chromosomal variation for phylogeny and biogeography are discussed.

Animals↗

Physicochemical characterization and tissue distribution of esterases in two salamandridae species (Mertensiella luschani and Salamandra salamandra).

1. Tissue- and species-specificity of the electrophoretic patterns of the multiple molecular forms of esterases were observed in the urodele amphibians Mertensiella luschani luschani, M.l. helverseni and Salamandra salamandra. All esterases--distributed into two electrophoretic mobility areas in gonads, muscles and brain and into four areas in liver, stomach and intestine--were characterized as carboxylesterases. 2. M. l. luschani and S. salamandra liver esterases were electrofocused into nine and eleven major bands with pIs ranging from 4.60 to 5.65 and from 4.40 to 6.20, respectively. 3. Two size groups of esterases were observed in liver extracts of the above three subspecies by Sephadex G-200 gel filtration. The mean values of their apparent molecular weights were 70,000 and 230,000 respectively.

Animals↗

Mitochondrial DNA phylogeography of Lissotriton boscai (Caudata, Salamandridae): evidence for old, multiple refugia in an Iberian endemic.

In Europe, southern peninsulas served as refugia during cold periods in the Pleistocene, acting both as centres of origin of endemisms and as sources from which formerly glaciated areas were recolonized during interglacial periods. Previous studies have revealed that within the main refugial areas, intraspecific lineages often survived in allopatric refugia. We analysed two mitochondrial markers (nad4, control region, approximately 1.4 kb) in 103 individuals representing the entire distribution of Lissotriton boscai, a newt endemic to the western Iberian Peninsula. We inferred the evolutionary history of the species through phylogenetic, phylogeographic and historical demographic analyses. The results revealed unexpected, deep levels of geographically structured genetic variability. We identified two main evolutionary lineages, each containing three well-supported clades. The first historical split involved populations from central-southwestern coastal Portugal and the ancestor of all the remaining populations around 5.8 million years ago. Both lineages were subsequently fragmented into different population groups between 2.5 and 1.2 million years ago. According to nested clade analysis, at lower hierarchical levels the patterns suggest restricted gene flow with isolation by distance, whereas at higher levels the clades exhibit signatures of contiguous range expansion. Bayesian Skyline Plots show recent bottlenecks, followed by demographic expansions in all lineages. The significant genetic structure found is consistent with long-term survival of populations in allopatric refugia, supporting the 'refugia-within-refugia' scenario for southern European peninsulas. The comparison of our results with other co-distributed species highlights the generality of this hypothesis for the Iberian herpetofauna and suggests that Mediterranean refuges had more relevance for the composition and distribution of present biodiversity patterns than currently acknowledged. We briefly discuss the taxonomic and conservation implications of our results.

Animals↗