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     

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.     

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.     

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.     

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.     

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.     

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.     

Splenic respiratory compensation and blood volume in the newt

In the newt Triturus cristatus carnifex , reversible increase in size of the spleen is linked to the respiratory state of the animal: when the newt is exposed to the air, and thus well oxygenated, the spleen hoards erythrocytes; when immersed in still water, in an hypoxic state, the spleen releases erythrocytes into the bloodstream. In chlorobutanol-anaesthetized specimens exposed to the air, the maximum size reached by the spleen diminishes with a rise in temperature up to the disappearance of all congestion at 33°C. The blood volume of newts kept in humid air at 6°C and 18°C after anaesthesia varies from about 6–9 ml per 100 g of body weight, while the red blood cell count and haematocrit value remain stable. In anaesthetized specimens kept in still water at the same temperatures the blood volume is stable, at about 7 ml per 100 g, but the red cell count and haematocrit are notably higher. At 33°C, a critical temperature for the newts, the specimens in still water succumb while those in air present the same blood volume as at 18°C, but have a higher erythrocyte count and haematocrit value.     

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.     

The newt Triturus carnifex as a model for monitoring the ecotoxic impact of the fungicide thiophanate methyl: adverse effects on the adrenal gland

The aims of this study were to propose a bioindicator organism, the newt Triturus carnifex, for the assessment of toxicological impact of thiophanate methyl in the Campania region (Italy) and the possible adverse activity on the adrenal gland. In the acute toxicity study, experimental groups of T. carnifex were exposed to 2.40, 4.80, 9.60 and 19.20 microg/L tap water of thiophanate methyl for 2 days; the LD50 was found to be 9.60 microg/L. To evaluate the effects on the adrenal gland, newts were exposed to a dose of 25% of the LD50 2 days for 8 days. The ultrastructural features of the tissues as well as the serum levels of aldosterone, corticosterone, norepinephrine (NE) and epinephrine (E) were evaluated. The number of secretory vesicles in the chromaffin cells appeared significantly decreased, whereas NE and E serum levels appeared strongly increased. Moreover, corticosterone and aldosterone serum levels appeared significantly reduced. The results suggest that: 1) T. carnifex has the features of an ideal bioindicator, due to its high sensitivity to thiophanate methyl, 2) thiophanate methyl acts as endocrine disruptor, affecting the adrenal gland at very low doses, 3) thiophanate methyl may be toxic for nontarget organisms, such as newts.     

Non-interactive multiple predator effects on tadpole survival

Interactions among and within three species of predators were estimated in terms of their effects on prey survival using short-term predation experiments. The prey were tadpoles (Rana temporaria), and the predators were dragonfly larvae (Anax imperator), newts (Triturus alpestris), and backswimmers (Notonecta glauca). Mortality rate per predator imposed by Triturus and Notonecta did not decline with predator density, whereas the predation rate of Anax was strongly reduced when the number of predator individuals increased. Impacts of all three predators were not altered by the presence of other species in pairwise combinations. This system is therefore characterized by interference between individual dragonflies but relatively independent effects of predator species. These results were largely predictable based on the natural history of the predators and are encouraging for attempts to model communities as assemblages of interacting species.     

The evolution of the adult body form of the crested newt (Triturus cristatus superspecies,Caudata, Salamandridae)

We characterized the adult body form of the crested newt (Triturus cristatus superspecies) and explored its evolution. From seven morphometric traits, we determined that body size, interlimb distance and head width define the body form. None of the morphometric traits showed a phylogenetic signal. Three body‐shape morphotypes (Triturus dobrogicus + T. cristatus, Triturus carnifex + Triturus macedonicus and Triturus karelinii + Triturus arntzeni) and three body‐size morphotypes (T. dobrogicus, T. cristatus and all other crested newts) could be recognized. The ancestral phenotype (a large body with a short trunk and a wide head) characterized T. karelinii and T. arntzeni. Triturus carnifex and T. macedonicus had a somewhat different phenotype (large body and wide head, accompanied by mild body elongation). The most derived phenotype included body size reduction and more pronounced body elongation in T. cristatus and, especially, in T. dobrogicus. Body elongation occurred by trunk lengthening but not head and tail lengthening. Additionally, contrary to other tetrapods, evolutionary axis elongation in crested newts was followed by a decrease in body size. We advocate the hypothesis that ecology drives the evolution of body form in crested newts.     

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.     

A comparative analysis of predator-induced plasticity in larval Triturus newts

Species that occupy similar habitats are expected to show convergent phenotypes. If habitats are defined by the presence of predators, then traits that modify vulnerability to predation, including predator-induced phenotypic plasticity, should be similar within habitats. We tested this idea using larvae of six syntopic newt species belonging to the two Triturus clades. Behavioural plasticity induced by odonate predators was strongly dissimilar between the two main clades but similar within them. Morphological plasticity was variable among species, even between one pair of closely related species. A predation experiment tested whether differences between clades could be caused by differences in body size. Size-specific vulnerability differed between newts in the small-bodied and large-bodied clades, indicating that similar predators may affect the two clades differently. The results showed both similarity and dissimilarity in predator-induced phenotypic plasticity in syntopic larval newts although theory suggests that divergence is unlikely in such ecologically similar species.     

Niche position, but not niche breadth, differs in two coexisting amphibians having contrasting trends in Europe

Aim We explored general habitat‐related explanations (niche breadth and niche position) to the contrasting status of two amphibian species that have largely overlapping ranges and habitats – the rare and declining crested newt (Triturus cristatus), and the more common smooth newt (Triturus vulgaris). These closely related and ecologically similar species provide an excellent opportunity to study those methodologically challenging hypotheses, and this is the first such study on amphibians. Location Denmark. Methods We derived multivariate habitat models from 27 characteristics of 210 ponds and their surroundings, and their occupation by newts. In addition to the model performance, niche breadths were compared using the mean beta diversity of amphibian communities in the presence of each newt species. Results For each newt species, the habitat models comprised three variables and correctly classified 74% of observations. Diverse invertebrate fauna (prey base) and shorter distances to other ponds inhabited by conspecifics were positive for both species, while the surrounding habitat (notably dry grasslands with forests) was important for the crested newt and the sediment type of the pond for the smooth newt only. Beta diversity of the amphibian communities of occupied ponds did not differ between the two newt species. Hence, in an area of frequent coexistence, habitat requirements of the species differed in key variables, not in the extent of specialization. Main conclusions Our study supported the niche position rather than the niche‐breadth hypothesis of rarity. We suggest that the rarity and/or continuing decline of the crested newt is related to the degradation of (semi)natural terrestrial habitats around suitable water bodies in Europe. Consequently, special restoration of such habitats has a high potential for the recovery of this rare species, while general pond management appeared more beneficial for the common smooth newt.     

Thermal behaviour of A4 lactate dehydrogenase purified from the heterothermic and sympatric vertebrate species Brook lamprey (Lampetra planeri), tench (Tenca tenca), smooth (Triturus vulgaris) and alpine newt (Triturus alpestris)

1. The A4 lactate dehydrogenase isozyme was purified to homogeneity from the tissues of Brook lamprey (Lampetra planeri), tench (Tenca tenca), smooth newt (Triturus vulgaris) and alpine newt (T. alpestris). 2. These four species share their geographical distribution in the same freshwater habitats, often live together in the same station and two of them are congeneric. Steady-state kinetic investigations have shown that: 3. Km (apparent) for pyruvate vs. temperature and (apparent) product Ki (Pyruvate) and Ki (Lactate) are fairly similar among species; 4. kcat/Km decreases with temperature in the case of the newts but increases in the case of both lamprey and tench; 5. Thermostability does not correlate to preferred ambient temperature and, in particular, tench LDH starts being inactivated up to 65 degrees C. 6. Thermostability does not correlate with activation energy either; 7. No clear relationships can be demonstrated either between activation energy and conformational transitions in the molecule (these latter indicated by breaks in the Arrhenius plots) nor between activation energy and molecular flexibility, investigated by melting experiments.     

A morphometric study of a hybrid newt population (Triturus cristatus) T. carnifex): Beam Brook Nurseries, Surrey, U.K.

The crested or great crested newt ( Triturus cristatus spp) is declining and now considered threatened in many of the countries where it is present. This has resulted in the four members of the superspecies being afforded protection under local, national and international law. This study looks at a possible threat to T. cristatus in southern England through hybridization, by the introduction of a related alien species ( T. carnifex ). The study used multivariate morphometries to discriminate closely related species, and their hybrids. The character set involved both continuous and meristic data, collected through body measurements and colour pattern. The identification of the species and/or hybrids at the introduction site and surrounding areas was mapped. From the results it can be inferred that hybridization has taken place at the introduction site, but there is no morphological evidence for the spread of hybrids/aliens in to the surrounding areas.     

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.     

The ontogenetic shift in thermoregulatory behaviour of newt larvae: testing the 'enemy-free temperatures' hypothesis

Growth and development affect life-history traits, and consequently organismal fitness. The inevitable increase in body size during ontogeny is associated with changes in both resource use and predation risk, which results in the ontogenetic shift in habitat preferences. In this study, we examined whether the shift in preferred body temperatures ( T _ps) of newt larvae Triturus alpestris increases the T _ps deviation of the most vulnerable stages (after hatching and during metamorphosis) from the T _ps range of their main predator, dragonfly larvae Aeshna cyanea – the 'predator-free temperatures' hypothesis. Analyses of thermoregulatory behaviour in the laboratory thermal gradient showed that freshly hatched newt larvae maintained lower water temperatures than larvae in later stages, whose T _ps largely fell into the T _ps range of dragonfly larvae. With respect to the thermal quality of natural habitat, the anti-predator effectiveness of the T _ps shift decreased during development. Water temperatures in natural habitats were located largely below the preferred body temperature range of both newt and dragonfly larvae, which limits their potential thermal niche partitioning. We conclude that factors other than predator avoidance drive the ontogenetic T _ps shift in our model system.