Rare hybridization and introgression in smooth and palmate newts (Salamandridae: Triturus vulgaris and T. helveticus)

This paper describes the occurrence of hybridization and introgression in two species of amphibians (the newts Triturus helveticus and Triturus vulgaris ) in mid-Wales, northern France and western France. A single aberrant adult male with intermediate phenotype was found. The multivariate analysis of 14 morphometric and two meristic characters supported its hybrid status. Electrophoretic analysis of 42 protein loci showed a genetic distance of 0.57 ± 0.14 Nei units between the species and revealed 15 diagnostic loci. The aberrant specimen was heterozygous at 11 of those and most likely to be a F₁ hybrid. Four marker loci appeared homozygous, suggesting the presence of enzymatically non-active ('null') alleles. The analysis of (maternally inherited) mitochondrial DNA showed the hybrid to be the offspring of a T. helveticus mother (and a T. vulgaris father). This observation does not conform to expectations based on the species composition in the pond from which the hybrid was collected. No F₁ hybrids were observed in a large sample (n > 5000) of larvae, recently metamorphosed newts and adults using two diagnostic protein loci. Occasionally alleles characteristic for one species were observed in the gene pool of the other species, suggesting the presence of bidirectional introgression. However, the frequency of alien genes was low (maximally 0.07%) which renders it difficult to rule out alternative explanations conclusively. The increase in total genetic variation in T. helveticus and T. vulgaris due to gene flow between them is negligible.     

The effect of pH on embryonic and larval development in smooth and palmate newts, Triturus vulgaris and T. helveticus

The distribution of smooth newts, Triturus vulgaris , and palmate newts, Triturus helveticus , in north-west Europe is related to geology and water quality. This study compared the development of the eggs and larvae of the two species under sublethal acidic and neutral conditions. Newt embryos raised under low pH hatched at an earlier stage of development, at a smaller size, and before those raised under neutral conditions. T. vulgaris hatched at a smaller size than T. helveticus , but pH did not affect the species differentially. Larvae of both species grew to a larger size under neutral than under acid conditions. Larvae raised in heterospecific pairs grew at least as well as those raised in conspecific pairs. Feeding was depressed under acid conditions, and reduced growth may therefore be associated with changes in the behaviour of newt larvae and their prey.     

The breeding migration of Smooth and Palmate newts (Triturus vulgaris and T. helveticus) at a pond in mid Wales

The breeding population sizes of newts at Llysdinam Pond in 1981 are estimated at around 800 Palmate and 300 Smooth newts. Females were more numerous than males with sex ration of 3.0: 1 for Palamte and 2.6: 1 for Smooth newts. The movement of males of both species into the pond preceded that of females, and female Palmate newts moved out of the pond significantly later than the other classes. The number of females of both species leaving the pond was much less than the number entering, apparently as a result of mortality whilst in the pond. Females leaving the pond were significantly lighter than those entering, and for female Palmate newts the weight loss in only partly attributable to loss of egg load. Temperature and humidity were important factors influencing the number of newts active during the inward migration period, but had little effect on activity during the outward migration.     

On the biological significance of certain morphological characters in males of the Smooth newt Triturus vulgaris and of the Palmate newt Triturus helveticus (Urodela: Salamandridae)

The males of Triturus vulgaris and Triturus helveticus can be differentiated by a number of morphological characters whose biological function is unknown. It is suggested that these characters enhance the effectiveness of male displays performed during sexual behaviour. This hypothesis is discussed in relation to the taxonomic and genetic relationships between the two species.     

A natural hybrid newt, Triturus helveticus×T. vulgaris, from a pond in mid-Wales

The first unequivocal example of a natural Trilurus helvelicus × T. vulgaris hybrid is described. The specimen was a male and discriminant analysis of physical characters indicated that it was morphologically intermediate between the parent species. A karyotype confirmed that the hybrid bore a haploid set of chromosomes from T. helveticus and a haploid set from T. vulgaris. Examination of the sex chromosomes showed that it was the result of mating between a male T. helveticus and a female T. vulgaris. As numerous mature sperm bundles were observed in both testes, the hybrid was therefore potentially fertile.     

Predation on newt eggs (Triturus alpestris and T. helveticus): identification of predators and protective role of oviposition behaviour

Predation on newt eggs ( Triturus helveticus and T. alpestris ) was studied in both natural (pond) and control (aquaria) conditions. Eggs were laid on artificial supports and set in a pond near Bourg-en-Bresse (south-east France). Survival from segmentation to pre-hatching stages was similar in these two species and estimated at 0·16 ± 0·07. Survival of eggs protected in a closed bag of plastic netting was significantly higher (0·79 ± 0·08), suggesting predation as the major mortality risk. Eggs of T. alpestris were offered to the following potential predators: newts Triturus alpestris and T. helveticus (males and females), adult aquatic insects ( Dytiscus marginalis, Acilius suleatus. Notonecta glauca, Ranatra linearis. Ilyocoris hermanni). a snail ( Lunnaea stugnalis) and a tadpole ( Rana temporaria ). Only the newts and water beetles were observed to cat the eggs. Females wrapped their eggs in a fold made with the support during oviposition. D. marginalis fed on wrapped eggs and those we unwrapped in the same proportion. but newts (especially females) and A. sulcatus clearly ate fewer wrapped than unwrapped eggs. These results are discussed in terms of mortality risk and antipredator adaptations.     

Alternative life-history pathways: paedomorphosis and adult fitness in European newts (Triturus vulgaris and T. alpestris)

Paedomorphs and metamorphs of the smooth newt ( Triturus vulgaris ) and alpine newt ( Triturus alpestris ) were compared with respect to body size, age structure, age at sexual maturity, survivorship, and female and male fecundity. Paedomorphs were significantly smaller than metamorphs, except for the alpine newt males. Non-significant differences between morphs in both species in terms of the life span, age of sexual maturity, survival rates and male fecundity were found. The relationships concerning female-fecundity parameters were not so straightforward. The total number of oocytes was significantly higher in smooth-newt paedomorphs, while in the alpine newt the difference was insignificant. When ovary mass was compared, significant differences appeared only in the alpine newt, in favour of metamorphic females. Oviductal egg size was similar in both morphs of T. vulgaris . The maintenance of both life-history strategies in the species studied is discussed in the light of these findings.     

Growth and size of Triturus vulgaris and T. cristatus (Amphibia) in different parts of Norway

The growth and size of the newts, Triturus vulgaris (L.) and T. cristatus (Laurenti) in different parts of Norway (+ Jämtland, Sweden) were studied. Age was estimated from skeletal growth marks, size-frequency histograms and, for the males, from the number of testis lobes.
On average, the T. vulgaris Larvae in central Norway and Jämtland are smaller than those in southeastern Norway, although in good localities, at the same altitude, they are of approximately the same size and complete their development already within 2½–3 months. Larvae living in eutrophic habitats are larger than those in oligotrophic-dystrophic habitats. In bog habitats at similar times of year the T. cristatus larvae from central Norway are about the same size as those from similar habitats in southwestern and southeastern Norway, but somewhat smaller than those from eutrophic ponds in the Oslofjord area.
No significant differences in the mean size of adult T. vulgaris from southeastern Norway and from central Norway were found. The maximum lengths attained were recorded from central Norway, however. In eutrophic habitats T. vulgaris adults BK usually larger than those in oligotrophic/dystrophic habitats.
In southeastern Norway, where growth is rapid, T. vulgaris may become sexually mature when 2+ yr-old, in central Norway usually a year later, in Jämtland. on average, still later, and in their northernmost locality (Vefsn) they probably do not breed until 5+ or 6+ yr-old.
On average, T. cristatus adults from southeastern Norway are slightly larger than those from central Norway, and may become mature at 2+ yr-old. compared with 4 + yr-old, at the earliest, in the latter area. The differences in the growth rates of both larvae and metamorphosed stages, of both species, most probably represent a response to climatic differences, although biotope quality is also an important factor.     

Molecular and morphological evidence of hybridization between newts Triturus vulgaris and T. montandoni (Caudata: Salamandridae) in Slovakia

Natural hybridization and introgression is recorded between the newt species Triturus vulgaris and T. montandoni in Slovakia. To confirm a hybrid status of two putative hybrids, morphological and Randomly Amplified Polymorphic DNA (RAPD) markers specific for T. vulgaris and T. montandoni were used. The individuals found in Zbojská (Veporské vrchy Mts, Slovakia; analyzed morphologically and genetically) and Zubrohlava (Oravská kotlina basin, Slovakia; analyzed only morphologically) possessed markers of both species. Segregation of RAPD markers together with a model-based Bayesian analysis revealed that the specimen from Zbojská belonged to later generation hybrids.     

Growth Studies and Ageing Methods for Adult Triturus vulgaris L. and T. cristatus Laurenti (Urodela, Salamandridae)

Growth of adult newts in southwestern Sweden has been studied from 1969 to 1972 in two ways: (1) The age of alcohol-preserved specimens has been determined by counting dark zones, which are thought to indicate hibernations, on transverse sections of bones (femur or humerus); and average body length and head width of the different age classes have been recorded and compared; (2) Free-living individuals have been identified by their ventral pattern, and body length and head width in different years have been recorded for each specimen. There was generally a small annual increase in body length and in head width, although the individual variation was large because some do not grow at all. Differences in growth were found between years, between populations and between sexes. It is concluded from these results that size should not be used for estimating the age of adult newts.     

Data Concatenation,Bayesian Concordance and Coalescent-Based Analyses of the Species Tree for the Rapid Radiation of Triturus Newts

The phylogenetic relationships for rapid species radiations are difficult to disentangle. Here we study one such case, namely the genus Triturus, which is composed of the marbled and crested newts. We analyze data for 38 genetic markers, positioned in 3-prime untranslated regions of protein-coding genes, obtained with 454 sequencing. Our dataset includes twenty Triturus newts and represents all nine species. Bayesian analysis of population structure allocates all individuals to their respective species. The branching patterns obtained by data concatenation, Bayesian concordance analysis and coalescent-based estimations of the species tree differ from one another. The data concatenation based species tree shows high branch support but branching order is considerably affected by allele choice in the case of heterozygotes in the concatenation process. Bayesian concordance analysis expresses the conflict between individual gene trees for part of the Triturus species tree as low concordance factors. The coalescent-based species tree is relatively similar to a previously published species tree based upon morphology and full mtDNA and any conflicting internal branches are not highly supported. Our findings reflect high gene tree discordance due to incomplete lineage sorting (possibly aggravated by hybridization) in combination with low information content of the markers employed (as can be expected for relatively recent species radiations). This case study highlights the complexity of resolving rapid radiations and we acknowledge that to convincingly resolve the Triturus species tree even more genes will have to be consulted.     

Sexual isolation between two newt species, Triturus vulgaris and T. montandoni (Amphibia, Urodela, Salamandridae)

In the present study we investigated sexual isolation between Triturus vulgaris and 77 montandoni in mating experiments run under semi-natural conditions. The two newt species offer a suitable model for studying evolution of reproductive isolation and mating preferences because they arc genetically the most similar species within the genus and readily hybridize in nature. Separate experiments were conducted in which groups of virgin females were placed together (in artificial pools) with groups of homospecific, heterospecific or both types of males. The estimates of reproductive isolation and mating propensity were based on the numbers of females producing hybrids and/or non-hybrid progeny. The levels of reproductive isolation, isolation asymmetry (IA) and propensity asymmetry (PA) were significant only for experiments in which females were given a choice between conspecific and heterospecific males. This implies that mating experiments with no interspecific choice may reduce discrimination and affect patterns of IA and PA. Asymmetry in reproductive isolation was also significant when the analysis was confined to just inseminated females. Differences in habitat preferences and condition of females possibly contributed to the relatively high values of PA.     

Early steroid metabolism in Xenopus laevis, Rana temporaria and Triturus vulgaris embryos.

Embryos of Xenopus laevis, Rana temporaria and Triturus vulgaris exposed to radioactive pregnenolone have been found to convert it to progesterone. Incubations with radioactive progesterone showed that it was actively metabolized by oocytes and embryos. In Xenopus incubations progesterone was converted to 5alpha-pregnane-3,20-dione, 17alpha-hydroxy-4-pregnen-3-one, 4-androstene-3,17-dione and 17alpha-20alpha-dihydroxy-4-pregne-3-one, indicating 5alpha-reductase, 17alpha-hydroxylase, 19-20-desmolase and 20alpha-hydroxylase activities. In oocytes of Triturus and Rana no evidence of 19-20-desmolase was found. In Rana oocytes were also not evidence of 17alpha-hydroxylase activity. All identified activities except 20alpha-hydroxylase were common to embryos of all three species. It is suggested that the steroid enzyme activities present in the embryos are not solely derived from the oocytes but synthentized during early development. Possible meaning of this kind of metabolism during differentiation remains open.     

Microhabitat selection and feeding relations of smooth and warty newts, Triturus vulgaris and T. cristatus, at an upland pond in mid-Wales

The utilization of microhabitat and food resources by Triturus vulgaris and T. cristatus was studied in an upland pond in mid-Wales. From April to June T. vulgaris was more evenly distributed across the pond than T. cristatus , which showed a preference for the middle of the pond and the bottom of the water column. From July to September however, when most T. vulgaris had left the pond, T cristatus was evenly distributed within the water column and a much higher proportion was captured around the shorelines. Both species appear to be generalist predators with diets limited by prey size. Zooplankton were the most important prey for T. vulgaris , while leeches formed the bulk of the diet of T. cristatus . However, T. cristatus did not select larger-sized zooplankton than T. vulgaris . Large body size therefore enables T. cristatus to consume a wider range of prey sizes than T. vulgaris . Microhabitat niche overlap was greater than feeding niche overlap between the species.