The effect of pH on feeding behaviour in newt larvae (Triturus: Amphibia)

The feeding responses of three species of newt larvae were compared under circumneutral and sublethal acid conditions. Under acid conditions (pH 4.5) feeding behaviour was suppressed in palmate newts, Triturus helveticus, and smooth newts. T. vulgaris , but not in crested newts, T. cristatus. At low pH, approach and orientation towards food occurred in T. helveticus and T. vulgaris , but snapping was inhibited; T. cristatus snapped and consumed food immediately it was offered under the same conditions. These differences are not consistent with the apparent greater tolerance of T helveticus for acidified ponds. The observations suggest that the chemosensory system of T. helveticus and T. vulgaris may be impaired at low pH.     

Predation and competition within an assemblage of larval newts (Triturus)

The impact of crested newts (Triturus cristatus) on the smaller-bodied palmate and smooth newts (T helveticus and T vulgaris) was studied during the larval stages using a combination of field and laboratory experiments In pond enclosures T cristatus larvae had no effect on the two smaller species over the first four weeks of development By eight weeks, however. T cristatus had achieved a size advantage which enabled it to eliminate T helveticus and severely reduce T vulgaris by predation In laboratory trials under food-limited conditions, T helveticus and T vulgaris were slightly smaller when raised with T cristatus, suggesting that this predatory effect was complemented by interspecific competition during early development Predation of the smaller species started when T cristatus reached a threshold size of c 27 mm No reciprocal effects on T cristatus growth or survival were observed Although T cristatus may be a significant predator of congeneric species in natural ponds, other factors, such as differences in microhabitat selection, higher-order predator-prey interactions and the occasional desiccation of pond habitats may facilitate coexistence between the species     

Differential effects of pH and temperature on embryonic development in the British newts (Triturus)

Changes in the water quality and temperature relationships of ponds may affect the structure of amphibian assemblages. The survival, time to hatching, hatching size and hatching stage of newt embryos were studied in the three British species ( Triturus cristatus, T. helveticus and T. vulgaris ), at two temperatures and two pHs. All T. cristatus embryos failed to hatch at pH 4.5, whereas over 80% of T. helveticus and T. vulgaris embryos hatched successfully at the same pH. At pH 7.5, T. cristatus survival was the same as the other two species, after the 50% mortality due to the homomorphism of chromosome 1 was taken into account. Temperature had no effect on survival of embryos. Time to hatching was two to four times longer at 12°C than at 17°C. Low pH shortened development time in T. vulgaris but not in T. helveticus . Low pH, but not temperature, affected size at hatching, with T. helveticus and T. vulgaris embryos emerging at a smaller size and earlier stage of development under acidic conditions. This reduction of size at low pH affected T. vulgaris more than T. helveticus . We predict that T. cristatus embryos will be the most vulnerable of the three species to acidification in nature. Warm ponds will result in rapid embryonic development, but T. helveticus and T. vulgaris larvae hatching in acid ponds will do so at a smaller size and earlier stage of development. The pattern of vulnerability to acidification within amphibian assemblages may change during ontogeny.     

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.     

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.     

Characteristics of newt breeding sites

Breeding site characteristics have been studied for the three species of newt that occur in Britain, the Palmate ( Triturus helveticus (Razoumowski)), Smooth ( T. vulgaris (L.)) and Warty ( T. cristatus (Laurenti)). The Warty newt was seldom found in the absence of the much commoner Smooth newt, but seemed to prefer sites that were relatively large and deep and that had a high proportion of open water surface. All three species tended to breed in ponds having abundant aquatic vegetation. Smooth newts, unlike Palmate newts, were rarely encountered in water with pH <6. The Smooth newt tended to be found in water with relatively high concentrations of metals, while the reverse was true for the Palmate newt. Over Britain, Smooth and Warty newts are relatively less common in soft water areas, while the Palmate is less common in hard water areas. Possible reasons for these associations are discussed.     

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.     

Resource partitioning by three species of newts during their aquatic phase

This paper describes the multidimensional niche of syntopic smooth newts Triturus vulgaris. alpine newts Triturus alpestris. and Italian crested newts Triturus carnifex. which constitute a complex community of eight components (the three species and their larvae, plus juvenile and neotenic alpine newts) All the potentially limiting resources (seasonal time, diel time, habitat and food) are analyzed as single niche dimensions, and then the real multidimensional niche is calculated Resource states are objectively defined, using an algorithm that groups the significant resource categories The larvae were segregated by prey, season, habitat, and water depth, in order of decreasing importance, their overlap was very low when their complete multidimensional niche was considered, and therefore they have few opportunities for present–day competition The adults were largely segregated by season and water depth, and to a lesser degree by habitat and prey, their multidimensional niches overlapped to a certain degree, thus leaving conditions for ongoing competition Considenng all the species and the life stages together, the order of importance for segregation was season, prey, habitat, water depth and diel activity, with a low overlap in the complete niche Differences m prey size were probably due to morphological constraints, differences in diel activity had very little effect on segregation except m separating larvae from the adults, and were likely promoted by need to avoid mutual predation and cannibalism It remains to be tested whether seasonal, habitat, and water depth differences are due to competitive pressures or to ecophysiological constraints     

Sexual size and shape evolution in European newts (Amphibia: Caudata: Salamandridae) on the Balkan Peninsula

We used a phylogenetic perspective in an examination of the direction and extent of sexual dimorphism in body size and body shape in European newts from the Balkan Peninsula (alpine newts, Mesotriton alpestris; crested newts, Triturus cristatus superspecies; smooth newts, Lissotriton vulgaris). We found a strong, female‐biased sexual size dimorphism (SSD) in the analysed clades of alpine newt, whereas within crested newts we found a less stringent female‐biased SSD in Triturus carnifex, Triturus macedonicus and Triturus karelinii, and no significant SSD in T. cristatus or Triturus dobrogicus. Among the smooth newts, we found male‐biased SSD in Lissotriton vulgaris vularis and Lissotriton vulgaris greacus and no SSD in Lissotriton vulgaris meridionalis. Most of these newts also exhibit a significant sexual dimorphism in body shape, which varied more randomly than body size, regardless of SSD level. Female and male body size as well as the degree of SSD displayed statistically significant phylogenetic signal, while sexual dimorphism in body shape was phylogenetically independent. The relationship between independent contrast data for female size and male size indicated that SSD in European newts could be driven by a disproportionate increase in female size as increase in female size was not accompanied by a proportional increase in male size.     

Limitation of similarity and feeding habits in three syntopic species of newts (Triturus, Amphibia)

Differences in body size between both species and sex were observed to be close to Hutchinsonian ratios in an assemblage of three species of newts (Triturus italicus, T alpestris and T carnifex) in southern Italy According to feeding, differences in body size were not related with a strict food niche segregation The diet of larger newt included indeed all the prey eaten by smaller newts but showed new prey types Differences between species were more marked than between sexes of the same species, despite a strong sexual dimorphism in body size Larger newts foraged in a wider range of microhabitats and exhibited a greater diversity of foraging tactics Triturus italicus fed essentially on plankton and chironomids in shallow water at the banks of the pond Triturus alpestris foraged also in deeper water, particularly on lsopods in vegetation Prey fallen on the water surface constituted the main part of T carnifex diet According to recent data on newt evolution, speciation consisted in the reduction of body size Our data show that this process has been related with a specialization of the diet and of habitat use     

Necator americanus: temperature, pH, light, and larval development, longevity, and desiccation tolerance

The effect of incubation temperature and pH on the hatch rate of eggs of Necator americanus, and the desiccation tolerance of the resulting infective stage-3 larvae were investigated in the laboratory under controlled conditions. Hatching did not occur below 15 C and above 35 C. A 21% hatch rate was obtained at 15 C while a 10.6% hatch rate was obtained at 35 C. The highest hatch rate (93.7%) was obtained at 30 C. The optimum pH for hatching was 6.0, but the larvae did not reach the infective stage. Incubation temperature of the eggs affected the longevity and desiccation tolerance of resultant infective larvae. Larvae hatched at 30 C and maintained at 26 C under bright fluorescent light had a 50% survival time (S50) of 4 days. In the dark or shade, the S50 for larvae raised at 30 C was 5 weeks, while that of larvae hatched at 20 C was 7 weeks. Incubation temperature also affected the desiccation tolerance of larvae. Larvae developed at 20 C were more resistant to desiccation at various relative humidity values than larvae hatched at 30 C.     

A phylogeny for the Old World newts, genus Triturus: biochemical and behavioural data

A reconstruction is presented of the phylogenetic relationships withing Triturus . Two different data sets are used. Protein electrophoretic data were scored for 19 presumptive gene loci in nine species (Rafinski & Arntzen, 1987) and a set of 16 behaviour characters from the courtship of Triturus was scored for the same nine species. When these are analysed in concernt, a single, fully resolved and robust solution is found. Consequently, two subgenera are recognized: Triturus and Palaeotriton , the former including all te medium- and large-bodied species: T. alpestris, T. vittatus, T. marmoratus, T. cristatus and its three former subspecies ( T. carnifex, T. dobrogicus and T. Karelini ); the latter comprising the small-bodied species: T. bascai, T. italicus, T. helveticus, T. vulgaris and T. montandoni . Within the subgenus Triturus, T. alpestris and T. vittatus are sister taxa to the remaining members of the subgenus. Similarly, T. boscai and T. italicus are a monophyletic group, sister to T. vulgaris, T. helveticus and T. montandoni . Alternative solutions may be found on the basis of a quantitative analysis of the behavioural characters employed in this study. An alternative phylogenetic tree is included as an example.     

Body size, competitive interactions, and the local distribution of Triturus newts

1. Pairs of European Triturus newt species of similar size tend not to co-occur syntopically, suggesting that similarity in body size is associated with competitive interactions that prevent coexistence. I tested this hypothesis with an experiment involving larvae of four species in 675-L artificial ponds. 2. There were strong interactions between most species pairs. Even the small T. helveticus had a clear impact on the larger T. alpestris. Pairs of species with different body sizes did not interact less strongly. 3. A standard increase in competitor biomass (c. 2 g mass at metamorphosis) caused 42% lower expected survival from hatching to 1 year of age, regardless of whether the species were of similar or different size. In most cases this resulted from delayed metamorphosis, reduced size at emergence, and slightly lower larval survival. 4. A standard increase in competitor density (0.74 individuals m(-2)) caused a greater reduction in expected 1-year survival when the competitor was larger (18% decline) than when both species were of similar size (6% decline), primarily because the very large T. cristatus consumed the smallest species. 5. These findings suggest that species interactions during the larval stage cannot explain distribution patterns of same- and different-sized Triturus.     

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.     

Helminth parasite populations in newts and their tadpoles

Five species of helminth parasites were found in adults and tadpoles of the newts Triturus helveticus, T. vulgaris, and T. cristatus, caught at eight different localities in Somerset. These parasites were Acanthocephalus anthuris, A. ranae, Megalobatra-chonema terdentatum, Thominx filiformis and Oswaldocruzia molgeta. Mean worm burdens did not change through the year, and egg-producing females of Acantho-cephalus anthuris and Megalobatrachonema terdentatum were found throughout. There was no difference in the level of infection of male and female newts.     

High aquatic niche overlap in the newts Triturus cristatus and T. marmoratus (Amphibia,Urodela)

We studied spatial niche metrics of large-bodied newts (Triturus cristatus and T. marmoratus) in three breeding ponds in western France. Adults and larvae were sampled with underwater funnel traps. Larvae were identified to the species with diagnostic microsatellite DNA markers. The distribution of adult T. cristatus and T. marmoratus across pond regions differed in one out of six cases, no differences were observed between larvae (two ponds studied). Niche overlap and niche breadth indices across resource states defined as pond regions or individual traps were high (Schoener's C: pond regions 0.60–0.98, traps 0.35–0.71; Levins' B: pond regions 0.71–0.98, traps 0.35–0.76). Adults of large-bodied newts significantly differed in resource use from small-bodied newts (T. helveticus). The results are discussed in view of the occurrence of interspecific breeding attempts, and the unpredictable ecological characteristics of newt breeding ponds.     

Differential Kin Discrimination by Red-spotted Newts (Notophthalmus viridescens) and Smooth Newts (Triturus vulgaris)

In organisms lacking parental care or when eggs of more than one female are layed together, parents preferentially tend to avoid eating their own offspring. In some species of newts (Caudata: Salamandridae), there is no parental care and cannibalism of eggs and larvae occurs when philopatric adults remain in the pond throughout the breeding season where eggs and larvae develop. Kin discrimination by both adult newts and larvae would be expected to occur during the breeding season as it would enable individuals to benefit from the nutrients obtained from eating eggs and larvae while they avoid eating close relatives. The effects of kinship on cannibalism of eggs and larvae were examined in two species of newts. In separate, but similar, paired behavioral trials adult female red-spotted newts, Notophthalmus viridescens, and smooth newts, Triturus vulgaris, were allowed to consume their own egg or an egg from another female. Female red-spotted newts, but not smooth newts, preferred to eat eggs of other females rather than their own, and overall smooth newts were ‘reluctant’ to eat conspecific eggs. To test for ontogenetic shifts in discrimination abilities, additional experiments were performed with adult female red-spotted newts paired with their own larvae and a larva from another female. In addition, the effects of kinship on the attraction or repulsion of larvae was also studied in red-spotted newts. Females showed no significant discrimination abilities during acts of cannibalism on larvae. Larvae spent similar amounts of time with siblings and non-siblings. These results suggest that the ability of adult newts to discriminate among eggs varies between species and that the elicitation of a kin-discrimination response may be context-dependent. For red-spotted newts, there was no evidence of discrimination abilities within and between other life-history stages.     

Micro-habitat use within a guild of newt larvae (Trituridae) in an Alpine lake

Population and ecological parameters such as numbers of larvae, microhabitat use, niche breadth and niche overlap of three species of syntopic larval newts (Alpine newt Triturus alpestris, Italian crested newt T. carnifex, and common newt T. vulgaris) were studied for two years in a small pond at 1160 m a.s.l. in NE Slovenia. Differences in microhabitat partitioning among larval newts were small. The largest niche breadth was estimated for larval T. alpestris, and the narrowest estimate was for larval T. carnifex in both years. Ecological differences seem to be very small and quite variable among sites and years. It appears that the developmental stage and size of newt larvae are more important in explaining resource partitioning than the characteristics of each species. Because of the absence of potential invertebrate predators and adult newts in the second half of the breeding season, the injuries could only be caused by intra-and interspecific predation attempts.     

Phylogeography of two European newt species--discordance between mtDNA and morphology

The newts Triturus vulgaris and Triturus montandoni are sister species that exhibit contrasting levels of intraspecific morphological variation. Triturus vulgaris has a broad Eurasiatic distribution encompassing both formerly glaciated and unglaciated areas and shows substantial morphological differentiation in the southern part of its range, while T. montandoni, confined to the Carpathians, is morphologically uniform. We analysed sequence variation of two mtDNA fragments of the total length of c. 1850 bp in 285 individuals of both species collected from 103 localities. Phylogenetic analysis of 200 unique haplotypes defined 12 major clades, their age estimated at c. 4.5-1.0 million years (Myr). Most of the older clades were found in the southern part of the range, and also in central Europe, mainly in Romania. The distribution of mtDNA clades points to the existence of several glacial refugia, located in the Caucasus region, Anatolia, the Balkan Peninsula, Italy, and more to the north in central Europe. The concordance between mtDNA based phylogeny and the distribution of T. vulgaris subspecies was weak. Triturus montandoni haplotypes did not form a monophyletic group. Instead they were found in six clades, in five of them mixed with T. vulgaris haplotypes, most likely as a result of past or ongoing hybridization and multiple introgression of mtDNA from T. vulgaris to T. montandoni. Patterns of sequence variation within clades suggested long-term demographic stability in the southern groups, moderate and relatively old demographic growth in the populations inhabiting central Europe, and high growth in some of the groups that colonized northern parts of Europe after the last glacial maximum.