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.     

Alpine newts (Triturus alpestris) as top predators in a high-altitude karst lake: daily food consumption and impact on the copepod Arctodiaptomus alpinus

1. Population dynamics and feeding ecology of adult and larval alpine newts (Triturus alpestris, Laurenti) were investigated in a high-altitude karts lake to estimate their feeding pressure on the copepod Arctodiaptomus alpinuf (Imhof). Estimates of population size for reproducing adults ranged from 666 to 864 individuals in the lake during July and August. Total abundance of larvae before the onset of ice cover varied considerably between 4400 and 25400 individuals in different years. 2. Arctodiaptomus alpinus was an important prey item for adult and larval alpine newts. During the second half of their aquatic period, adult newts moved to deeper water where the copepod reached its highest densities near the sediment. Adults and larvae exhibited no periodic feeding pattern. The feeding rhythm was more synchronized among the larvae than among the adults. 3. Daily food consumption, estimated using the Elliott & Persson (1978) model, reached 4–21 mg dry biomass in adults. The daily ration of larvae was about 7% of body dry weight in the temperature range 6-11°C. Compared to published estimates of daily food consumption in salmonid fishes, the feeding pressure of newts appears low.     

Do Experience and Body Size Play a Role in Responses of Larval Ringed Salamanders,Ambystoma annulatum,to Predator Kairomones? Laboratory and Field Assays

Prey may experience ontogenetic changes in vulnerability to some predators, either because of changes in morphology or experience. If prey match their level of antipredator behavior to the level of predatory threat, prey responses to predators should reflect the appropriate level of threat for their stage of development. For larval salamanders, responses to predators may change with body size because larger larvae are less vulnerable to predation by gape‐limited predators or because fleeing responses by large salamanders may be more effective than for smaller salamanders. In a field experiment, small larval ringed salamanders, Ambystoma annulatum, responded to chemical stimuli (‘kairomones’) from predatory newts, Notophthalmus viridescens, with an antipredator response (decreased activity). Laboratory‐reared larvae decreased their activity following exposure to newt kairomones, indicating that larval ringed salamanders do not require experience with newts to recognize them as predators. In both experiments, larvae distinguished between chemical stimuli from newts and stimuli from tadpoles (non‐predators) and a blank control. In a third experiment, field‐caught (experienced) larvae showed a graded response to newt kairomones based on their body size: small larvae tended to decrease their activity while larger larvae showed no change or an increase in activity. This graded response was not observed for neutral stimuli, indicating that it is predator‐specific. Therefore, ringed salamander larvae exhibit threat‐sensitive ontogenetic changes in their response to chemical stimuli from predatory newts.     

Stream microhabitat selectivity,resource partitioning,and niche shifts in grazing caddisfly larvae

Microhabitat selectivity, resource partitioning, and niche shifts in five species of grazing caddisfly larvae (Glossosoma califica, G. penitum, Dicosmoecus gilvipes, Neophylax rickeri, and N. splendens) were quantified by underwater measurement of microhabitat availability and utilization in three northern California streams. The microhabitat parameters water depth and velocity and rock size, roughness, and slope were measured. Comparisons of habitat available to habitat used revealed significant selection for at least two microhabitat parameters by each population, with depth and velocity being the most important. Comparisons of habitat used by different species showed significant partitioning of at least two microhabitat parameters at each site, with depth being partitioned at all sites. Non-parametric discriminant analysis revealed significant microhabitat partitioning on a multivariate level at two sites. Comparisons of habitat used at different sites quantified a major niche shift by D. gilvipes in its preference for riffles versus pools. Size-selective predation by dippers (Cinclus mexicanus) and steelhead (Salmo gairdneri gairdneri) is proposed as a hypothesis to explain the observed resource partitioning and niche shift.     

Priority effects of the early breeding fire salamander on the late breeding banded newt

Early breeding intraguild predators may have advantages over late breeding predators via priority effects; early breeding predators may reduce shared prey resources before late breeders appear and may also prey upon the late breeders. Here we show that predatory larvae of the late-breeding predatory banded newt, Triturus vittatus vittatus, occupy the same temporary pond toward the end of the developmental period of the early-breeding predatory fire salamander, Salamandra salamandra, resulting in a large size disparity between larvae of these two species while they co-occur. We conducted outdoor artificial pool experiments to assess priority effects of large larval Salamandra at the end of their larval development period, on recently hatched larval Triturus. We also assessed how artificial vegetation may influence larval Triturus performance in the presence or absence of Salamandra Salamandra, introduced into the experimental pools two weeks prior to the newt larvae, strongly reduced invertebrate prey abundance shared by these two predatory urodeles and with only a one week period of overlap, strongly reduced abundance of Triturus larvae. The artificial vegetation had only a small ameliorating effect on Triturus survival when Salamandra was present. Triturus size at metamorphosis (snout-tail length) was significantly larger in the Salamandra pools, presumably due to a combination of a strong “thinning effect” and greater vulnerability of smaller Triturus individuals to predation by Salamandra. Time to metamorphosis was not significantly affected by Salamandra. These results have conservation implications as T. v. vittatus is listed as highly endangered and may also explain the largely negative spatial association of the two species. Handling editor: K. Martens     

Species‐specific habitat preferences do not shape the structure of a crested newt hybrid zone (Triturus cristatus x T. carnifex)

Reproductive isolation barriers maintain the integrity of species by preventing interspecific gene flow. They involve temporal, habitat or behavioral isolation acting before fertilization, and postzygotic isolation manifested as hybrid mortality or sterility. One of the approaches of how to study reproductive isolation barriers is through the analysis of hybrid zones. In this paper, we describe the structure of a hybrid zone between two crested newt species (Triturus cristatus and T. carnifex) in the southern part of the Czech Republic using morphological, microsatellite, and mitochondrial (mtDNA) markers. Specifically, we tested the hypothesis that the structure of the hybrid zone is maintained by species‐specific habitat preferences. Comparing the genetic structure of populations with geographical and ecological parameters, we found that the hybrid zone was structured primarily geographically, with T. cristatus‐like populations occurring in the northeast and T. carnifex‐like populations in the southwest. Despite T. cristatus tending to occur in deeper ponds and T. carnifex on localities with more shading, the effect of both ecological parameters on the structure of the zone was minimal. Next, we corroborated that T. carnifex individuals and some hybrids possess mtDNA of T. dobrogicus, whose nuclear background was not detected in the studied hybrid zone. Hybridization between T. carnifex and T. dobrogicus (resulting in unidirectional mtDNA introgression) had to predate subsequent formation of the hybrid zone between T. cristatus and T. carnifex. Populations of crested newts in the southern part of the Czech Republic thus represent a genetic mosaic of nuclear and mitochondrial genomes of three 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.     

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.     

Genetic variation in the Italian crested newt, Triturus carnifex, and the origin of a non-native population north of the Alps

Genetic variation over 40 protein loci and 46 populations representing three taxa of crested newts revealed moderate genetic distances between Triturus carnifex carnifex, T. c. macedonicus and T. cristatus. Two populations from the Geneva Basin (presumed to be introduced) were genetically similar to T. c. carnifex and dissimilar to T. c. macedonicus and T. cristatus, showing that they belong to T. c. carnifex and not to native T. cristatus. A significant pattern of spatial genetic variation was found within T. c. carnifex along a north to south axis, from Croatia to Calabria. The Genevan populations showed highest genetic similarity with T. carnifex from Tuscany, suggesting that the propagule originated from that area. Effects of a population genetic bottleneck associated with the introduction could not be documented. The observed high allelic variation in Genevan T. c. carnifex could not be directly explained by introgression from T. cristatus. Comparisons across the range, including zones of hybridization within the T. cristatus superspecies, indicated that some alleles typical for the Genevan population may represent the so-called `hybrizymes'.     

洮河自然保护区大峪沟林区紫果云杉群落木本植物种群生态位特征

基于群落调查数据,以样地代表多种资源的综合状态,物种重要值作为生态位的计测指标,利用Levins和Shannon生态位宽度,Schoener生态位相似和Pianka生态位重叠对洮河自然保护区大峪沟林区的紫果云杉群落木本植物种群生态位进行定量研究。结果表明：紫果云杉群落乔木层共有树种8种,隶属于4科6属,紫果云杉占绝对优势,岷江冷杉为次优势种;灌木层共有植物50种,隶属于16科25属,忍冬科的蓝靛果、葱皮忍冬和唐古特忍冬等以及蔷薇科的峨眉蔷薇和银露梅等构成了灌木层的主体,紫果云杉幼苗优势不明显,但生态位宽度最大。本研究表明,生态位宽度受物种生态特性及其分布和生境共同影响,重要值大的物种生态位宽度一般较大,生态位宽度大的物种之间生态位相似性和重叠性一般较大,并与其它物种重叠的概率高、重叠程度大,生态位宽度小的物种集中出现时也会出现较大的生态位重叠,同科或同属的物种间常有较大生态位相似和重叠。生态位宽度反映着种群的扩散潜力,生态位相似反映着群落的稳定性,生态位重叠反映着物种间存在或潜在的竞争程度。紫果云杉在乔、灌木层均有最大的生态位宽度,生态适应性最强,群落结构稳定,具有进一步扩散的可能。     

An experimental study of population regulation in the salamander,Notophthalmus viridescens dorsalis (Urodela: Salamandridae)

Summary The roles of density-dependent larval survival and cannibalism of larvae as potential mechanisms of population regulation in the newt (Notophthalmus viridescens dorsalis) were evaluated in laboratory and field experiments. In laboratory containers, adults cannibalized larvae and large larvae cannibalized smaller larvae. In artificial ponds, larval survival did not depend on initial larval density. No cannibalism could be demonstrated in the complex environment, although the experiment was powerful enough to detect an ecologically relevant difference in survival. Adult growth was negatively correlated with the final biomass of larval newts, suggesting that the two life stages competed for resources. Larval growth rates were negatively correlated with final larval density, suggesting that larvae competed with each other. The proportion of larvae that became sexually mature at age 7 months (paedomorphs and adults that skipped the eft stage) varied inversely with larval density. Therefore, the potential regulatory mechanisms identified in this study are competition within and between life stages.     

No trade‐offs in interspecific interference ability and predation susceptibility in newt larvae

Coexistence of species with similar requirements is allowed, among others, through trade‐offs between competitive ability and other ecological traits. Although interspecific competition is based on two mechanisms, exploitation of resources and physical interference, trade‐off studies largely consider only species’ ability to exploit resources. Using a mesocosm experiment, we examined the trade‐off between interference competition ability and susceptibility to predation in larvae of two newt species, Ichthyosaura alpestris and Lissotriton vulgaris. In the presence of heterospecifics, L. vulgaris larvae slowed somatic growth and developmental rates, and experienced a higher frequency of injuries than in conspecific environments which suggests asymmetrical interspecific interference. During short‐term predation trials, L. vulgaris larvae suffered higher mortality than I. alpestris. Larvae of the smaller species, L. vulgaris, had both lower interference and antipredator performance than the larger I. alpestris, which suggests a lack of trade‐off between interference competition ability and predator susceptibility. We conclude that interference competition may produce a positive rather than negative relationship with predation susceptibility, which may contribute to the elimination of subordinate species from common habitats.     

Niche partitioning among three montane ground‐dwelling pheasant species along multiple ecological dimensions

Pheasants (order Galliformes) are typical ground‐dwelling birds, having a large body size and weak flight abilities. Sympatric pheasants are expected to share narrower niche space and face more extensive interspecific competition. However, little work has been undertaken simultaneously to investigate niche partitioning among sympatric pheasant species across multiple ecological dimensions. We compared microhabitat use, activity pattern and foraging strategy of sympatric Blood Pheasant Ithaginis cruentus, Buff‐throated Partridge Tetraophasis szechenyii and White Eared‐pheasant Crossoptilon crossoptilon on the Qinghai‐Tibet Plateau, China, to identify potential interspecific niche partitioning along different ecological dimensions in the breeding season. We found that the Buff‐throated Partridge differed significantly from the other two species in microhabitat use, and the three species showed different foraging strategies. It is likely that niche partitioning reduced potential interspecific competition, thus facilitating the species’ stable coexistence. Our study provides practical evidence of multidimensional niche theory within sympatric ground‐dwelling pheasant species, emphasizing that species interactions and coexistence within a guild are often not uni‐dimensional. Given global conservation concern for maintaining bird diversity, we recommend further restriction of yak grazing in these species’ habitat.     

Keystone predators (eastern newts, Notophthalmus viridescens) reduce the impacts of an aquatic invasive species

Predation, competition, and their interaction are known to be important factors that influence the structure of ecological communities. In particular, in those cases where a competitive hierarchy exists among prey species, the presence of certain keystone predators can result in enhanced diversity in the prey community. However, little is known regarding the influence of keystone predator presence on invaded prey communities. Given the widespread occurrence of invasive species and substantial concern regarding their ecological impacts, studies on this topic are needed. In this study I used naturalistic replications of an experimental tadpole assemblage to assess the influence of predatory eastern newts, Notophthalmus viridescens, on the outcome of interspecific competition among native and nonindigenous tadpoles. When newts were absent, the presence of the tadpoles of one invasive species, the Cuban treefrog, Osteopilus septentrionalis, resulted in decreased survival and growth rate of the dominant native species, Bufo terrestris, and dominance of the tadpole assemblage by O. septentrionalis. However, the presence of one adult newt generally reduced or eliminated the negative impacts of O. septentrionalis tadpoles, resulting in comparable survival and performance of native species in invaded and noninvaded treatments. Differential mortality among the tadpole species suggests that newts preyed selectively on O. septentrionalis tadpoles, supporting the hypothesis that newts acted as keystone predators in the invaded assemblage. The presence of nonindigenous larval cane toads, Bufo marinus, did not significantly affect native species, and this species was not negatively affected by the presence of newts. Collectively, these results suggest that eastern newts significantly modified the competitive hierarchy of the invaded tadpole assemblage and reduced the impacts of a competitively superior invasive species. If general, these results suggest that the presence of certain species may be an essential factor regulating the ecological impacts of biological invasions.     

Is resource partitioning the key? The role of intra‐interspecific variation in coexistence among five small endemic fish species (Characidae) in subtropical rivers

This study focused on intra and interspecific variations in the diet of small species in the Pelotas River, upper Uruguay River ecoregion. We tested the hypothesis that five small characids coexist via resource partitioning. Samplings were performed quarterly between August 2013 and May 2014 at 14 sites in the Pelotas River basin, using gillnets of different mesh sizes, seines, cast nets and electrofishing. In this study, the contents of 1,253 stomachs were analysed from Astyanax dissensus, A. paris, A. saguazu, A. xiru and Bryconamericus patriciae. The main food resources consumed were aquatic insects, leaves, seeds and detritus. There were significant seasonal intra and interspecific variations in the use of food resources by the five species. Permutational analysis of multivariate dispersions results indicated significant differences in individual intraspecific variability between the sampled periods, with the highest values found in November and February because of an increase in allochthonous resources (leaves and seeds). Besides this, significant differences in the proportions of consumption of allochthonous and autochthonous were observed. The trophic niche breadth was significantly different among the species, such that in August, the species had narrower niche breadths. In the other periods, the highest niche breadth values were due to an increase in the consumption of allochthonous items. Diet overlap was low (< 0.4) between 62% of the species pairs. The small characid species showed pronounced changes in resource use and intra and interspecific variations indicating resource partitioning. These factors appear to be essential in species interactions and for coexistence.     

Microhabitat analyses support relationships between niche breadth and range size when spatial autocorrelation is strong

Multiple evidence of positive relationships between nice breadth and range size (NB–RS) suggested that this can be a general ecological pattern. However, correlations between niche breadth and range size can emerge as a by-product of strong spatial structure of environmental variables. This can be problematic because niche breadth is often assessed using broad-scale macroclimatic variables, which suffer heavy spatial autocorrelation. Microhabitat measurements provide accurate information on species tolerance, and show limited autocorrelation. The aim of this study was to combine macroclimate and microhabitat data to assess NB–RS relationships in European plethodontid salamanders (Hydromantes), and to test whether microhabitat variables with weak autocorrelation can provide less biased NB–RS estimates across species. To measure macroclimatic niche, we gathered comprehensive information on the distribution of all Hydromantes species, and combined them with broad-scale climatic layers. To measure microhabitat, we recorded salamander occurrence across > 350 caves and measured microhabitat features influencing their distribution: humidity, temperature and light. We assessed NB–RS relationships through phylogenetic regression; spatial null-models were used to test whether the observed relationships are a by-product of autocorrelation. We observed positive relationships between niche breadth and range size at both the macro- and microhabitat scale. At the macroclimatic scale, strong autocorrelation heavily inflated the possibility to observe positive NB–RS. Spatial autocorrelation was weaker for microhabitat variables. At the microhabitat level, the observed NB–RS was not a by-product of spatial structure of variables. Our study shows that heavy autocorrelation of variables artificially increases the possibility to detect positive relationships between bioclimatic niche and range size, while fine-scale data of microhabitat provide more direct measure of conditions selected by ectotherms, and enable less biased measures of niche breadth. Combining analyses performed at multiple scales and datasets with different spatial structure provides more complete niche information and effectively tests the generality of niche breadth–range size relationships.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Sexual dimorphism in the skull geometry of newt species of <Emphasis Type="Italic">Ichthyosaura,Triturus</Emphasis> and <Emphasis Type="Italic">Lissotriton</Emphasis> (Salamandridae,Caudata, Amphibia)

In this study, we applied geometric morphometrics to explore variations in the level and pattern of sexual size dimorphism (SSD) and sexual shape dimorphism (SShD) of the ventral cranium in three different Modern Eurasian newt taxa (Ichthyosaura alpestris, Triturus species group and Lissotriton vulgaris). The ventral cranium is the part of the skull that is more directly related to foraging and feeding. Our results indicate that the level and pattern of sexual dimorphism in the ventral cranium differ among Modern Eurasian newt taxa. Regarding sexual dimorphism in skull size, Ichthyosaura alpestris and Triturus species show female-biased patterns (females are larger than males), whereas Lissotriton vulgaris appears to be non-dimorphic in skull size. In I. alpestris and Triturus species, SShD is mostly absent, whereas in L. vulgaris, SShD is more pronounced. A high level of variation between populations in both SSD and SShD indicates that local conditions may have a profound effect on the magnitude and direction of sexual dimorphism. The significant sexual differences in ventral cranium size and shape indicate possible subtle intersexual differences in ecological demands due to diet specialisation, in spite of similar general ecological settings.