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.     

Sperm transport after insemination in the Alpine newt (Triturus alpestris, Caudata, Salamandridae)

The goal of this study was to test if sperm transport to the spermathecae in the Alpine newt (Triturus alpestris) requires active co-operation of the female. Artificial insemination of anaesthetised female newts was conducted using spermatophores collected from courting males and with sperm duct contents collected from sacrificed males. Sperm was present in the spermathecae of 9 out of 10 females inseminated with the spermatophores but in only 1 out of 8 females inseminated with sperm duct contents. The females of both groups laid some eggs after insemination, and a portion of these eggs in group of females inseminated with spermatophores were fertilized. However, the number of eggs produced by the females was much lower than typical egg-production in newts. The presence of sperm in the spermathecae of females inseminated with spermatophores and lack of sperm in the spermathecae of females inseminated with sperm duct contents suggests that sperm transport is either induced by the substances present in spermatophores and/or that sperm from the sperm duct is not fully mobile in comparison with sperm from the spermatophores.     

Natural variation in the limb skeletal pattern of the crested newt,Triturus carnifex (amphibia: Salamandridae)

The forelimb skeletal patterns of two geographically isolated populations of Triturus carnifex, one from northern Italy (Rosate, Milano) and one from central Italy (Bagnaia, Perugia) were studied. A total of 1,018 limbs were examined. Limb skeletal variants, generally consisting either of a reduced number of carpals and/or of an alternate phalangeal formula, were commonly observed, as has been reported for other amphibian species. The population from northern Italy showed a greater variability than that from central Italy. Skeletal variants were present in 34% and 12% of the limbs, respectively (i.e., in 48% and 20% of the newts). In the Rosate population, the majority of variants consisted of fusions of radiale and prepollicis and of the phalangeal formula 1-2-3-2. Severe skeletal defects were also found in 3% of the limbs of the newts from Rosate and in 1% of the limbs of those from Bagnaia. At present, no conclusion can be drawn about the evolutionary significance of the limb skeletal variation observed in the two populations, because of the paucity of data on intra- and interpopulation variations in other species of the genus Triturus. © 1992 Wiley-Liss, Inc.     

Thermal acclimation of swimming performance in newt larvae: the influence of diel temperature fluctuations during embryogenesis

1.  Thermal acclimation is one of the basic strategies by which organisms cope with thermal heterogeneity of the environment. Under predictable variation in environmental temperatures, theory predicts that selection favours acclimation of thermal performance curves over fixed phenotypes.
2.  We examined the influence of diel fluctuations in developmental temperatures on the thermal sensitivity of the maximal swimming capacity in larvae of the alpine newt, Triturus alpestris .
3.  We incubated newt eggs under three thermal regimes with varying daily amplitudes (1, 5 and 9 °C) and similar means (17·6–17·9 °C), and accordingly we measured the swimming speed of hatched larvae at three experimental temperatures (12, 17 and 22 °C), which they would normally experience in their natural habitat.
4.  Embryonic development under low and middle temperature fluctuations produced larvae with similar swimming speeds across experimental temperatures. In contrast, the most fluctuating regime induced development of phenotypes, which at 12 °C swam faster than larvae developed under moderate diel fluctuations.
5.  Our results provide evidence that diel temperature fluctuations induce acclimation of thermal dependence of locomotor performance. In ectotherms experiencing diel cycles in environmental temperatures, this plastic response may act as an important pacemaker in the evolution of thermal sensitivity.     

Cranial shape variation and molecular phylogenetic structure of crested newts (Triturus cristatus superspecies: Caudata,Salamandridae) in the Balkans

In the present study, we investigated the degree of congruence between phylogeny, as inferred from mitochondrial (mt)DNA sequences, and cranium shape variation of crested newts (Triturus cristatus superspecies) in the Balkans. These newts belong to four phylogenetic clades defined by mtDNA analysis, and significantly differed in cranial shape. Allometry explained a high percentage of shape variation in crested newts. The clade‐specific allometric slopes significantly diverged for both the ventral cranium and dorsal cranium, indicating that differences in shape between clades could not be a simple consequence of their difference in size. The analysis of hierarchical and spatial variation showed similarity in the patterns of global and spatially localized hierarchical variation of cranial shape. We also found significant congruence between the pattern of cranial shape variation and molecular phylogeny. The differences in morphology of Triturus dobrogicus in comparison to other crested newt clades, including marked differences in cranium shape, is discussed in the context of the evolution and ecology of crested newts. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 348–360.     

The cycle of follicular and interstitial cells (Leydig cells) in the testis of the marbled newt, Triturus marmoratus (Caudata, Salamandridae)

Ultrastructural examination of the marbled newt (Triturus marmoratus) testis throughout the annual cycle revealed that during the period of testicular quiescence (November-February), primordial germ cells proliferate within cords of filament-rich epithelial cells that will become follicular cells (FCs). Fibroblast-like cells surround the FCs and form the lobule-boundary interstitial cells (ICs). During the period of germ cell development from primordial germ cells to round spermatids (March-June), the FCs surrounding the developing germ cells contain scanty cytoplasm with abundant rough endoplasmic reticulum and scarce filaments. With spermatid elongation (July-August), the FC size grows, its nucleus becomes irregularly outlined, and its cytoplasm displays abundant smooth endoplasmic reticulum, residual bodies, lipid droplets, and large vacuoles. After spermatozoon release by the FCs (August-September), the adjacent ICs increase their size and transform into Leydig cells with abundant smooth endoplasmic reticulum, mitochondria with tubular cristae, and lipid droplets. During the period of testicular quiescence (November-February), the Leydig cells undergo involution, eventually developing the morphological attributes of mesenchymal cells. Intermingled among these cells, cords of filament-rich cells are observed. During this period of the cycle, spermatozoon cysts supported by FCs are present. At the beginning of the germ cell proliferation period (March), these spermatozoa are released, and the adjacent ICs undergo a transformation into Leydig cells similar to those observed in August-September. Maturation and involution of ICs occur when testosterone levels are known to be rising and falling, respectively.     

Eggs in autumn: responses to declining incubation temperatures by the eggs of montane lizards

Facultative hatching in response to environmental cues may increase the viability of offspring, if the cue that stimulates hatching also predicts the negative consequences of delayed emergence. Declining incubation temperatures might provide such a cue for montane lizards, because eggs that fail to hatch before winter will perish in the nest. I tested this idea by incubating eggs of an alpine scincid lizard ( Bassiana duperreyi ) in the laboratory. For the first half of the incubation period the eggs were kept at nest temperatures typical of those experienced in summer in the field (daily cycle of 18 ± 7.5°C). I then transferred eggs at weekly intervals into cooler regimes (either 15 ± 7.5°C; or with daytime temperatures unchanged but dropping to 0°C overnight). Contrary to prediction, the eggs did not hatch early. However, transfer to lower temperatures caused only a relatively short delay in hatching, because of a virtual temperature-independence of developmental rates late (but not early) in incubation. Decreasing incubation temperatures also modified hatchling running speeds and post-hatching growth rates, even if the thermal decrease occurred only shortly before the usual time of hatching. These processes plausibly affect hatchling fitness in cold-climate reptiles, and might be adaptations to montane habitats. Alternatively, they may prove to be widespread in other (warmer-climate) reptile taxa, in which case no adaptive hypothesis need be proposed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 71–77.     

Environmental maternal effects on locomotor performance in the common lizard (Lacerta vivipara)

Environmental (i.e. non-genetic) maternal effects have the potential to associate the environmental conditions faced by mothers during gestation or before egg laying with the phenotype of their offspring. For this reason, maternal effects may play a major role in determining offspring phenotype independently of the genotype of the individuals, and can thus be considered a mechanistic basis of phenotypic plasticity. Despite the ecological and evolutionary implications of environmental maternal effects, few studies have experimentally investigated this phenomenon in reptiles. Here we report the results of an experimental laboratory study on the effects of maternal feeding rate and density on offspring locomotor performance in the common lizard (Lacerta vivipara). Lacerta vivipara is a viviparous lizard, and viviparity enhances the probability of a maternal influence on offspring phenotype. We focused on a particular phenotypic trait, maximal sprint running speed, because this trait is thought to be selectively important in squamates. Sprint speed was a repeatable trait, and it varied significantly among families. Maternal feeding rate significantly affected sprint speed, whereas density had no effect on this trait. The effect of maternal feeding rate differed according to the sex of the offspring and their body size, resulting in significant two-way and three-way interactions among these factors. In other words, the maternal feeding rate changed the shape of the allometric relationship between speed and size, but differently for males and females. The complexity of such effects makes it extremely difficult to offer an adaptive interpretation, but emphasizes the role played by the environment in shaping phenotypes among generations.     

Thermal limits to survival and activity in two life stages of false codling moth Thaumatotibia leucotreta (Lepidoptera,Tortricidae)

The present study examines life stage‐related variation in the thermal limits to activity and survival in an African pest, the false codling moth Thaumatotibia leucotreta (Lepidoptera, Tortricidae). Thermal tolerance, including the functional activity limits of critical thermal maxima and minima (CT_max and CT_min respectively), upper and lower lethal temperature, and the effect of heat and cold hardening (short‐term acute plasticity), is measured across a diverse range of low or high temperature stress conditions in both larvae and adults. We also report the sum of inducible and cognate forms of the amounts of heat shock protein 70 (HSP70) as an explanatory variable for changes in thermotolerance. The results show that the larvae have high variability in CT_max and CT_min at different ramping rates and low levels of basal (innate) thermal tolerance. By contrast, the adults show high basal tolerance and overall lower variability in CT_max and CT_min, indicating lower levels of phenotypic plasticity in thermotolerance. HSP70 responses, although variable, do not reflect these tolerance or survival patterns. Larvae survive across a broader range of temperatures, whereas adults remain active across a broader range of temperatures. Life stage‐related variation in thermal tolerance is most pronounced under the slowest (most ecologically‐relevant) ramping rate (0.06 °C min^–1) during lower critical thermal limit experiments and least pronounced during upper thermal limit experiments. Thus, the ramping rate can hinder or enhance the detection of stage‐related variation in thermal limits to activity and survival of insects.     

Biparental care is predominant and beneficial to parents in the burying beetle Nicrophorus orbicollis (Coleoptera: Silphidae)

Parenting strategies can be flexible within a species and may have varying fitness effects. Understanding this flexibility and its fitness consequences is important for understanding why parenting strategies evolve. In the present study, we investigate the fitness consequences of flexible parenting in the burying beetle Nicrophorus orbicollis, a species known for its advanced provisioning behaviour of regurgitated vertebrate carrion to offspring by both sexes. We show that, even when a parent is freely allowed to abandon the carcass at any point in time, biparental post‐hatching care is the most common pattern of care adopted in N. orbicollis. Furthermore, two parents together raised more offspring than single parents of either sex, showing that the presence of the male can directly influence parental fitness even in the absence of competitors. This contrasts with studies in other species of burying beetle, where biparental families do not differ in offspring number. This may explain why biparental care is more common in N. orbicollis than in other burying beetles. We suggest how the fitness benefits of two parents may play a role in the evolution and maintenance of flexible biparental care in N. orbicollis.     

Mitochondrial DNA sequence analysis reveals multiple Pleistocene glacial refugia for the Yellow‐spotted mountain newt,Neurergus derjugini (Caudata: Salamandridae) in the mid‐Zagros range in Iran and Iraq

Phylogeography is often used to investigate the effects of glacial cycles on current genetic structure of various plant and animal species. This approach can also identify the number and location of glacial refugia as well as the recolonization routes from those refugia to the current locations. To identify the location of glacial refugia of the Yellow‐spotted mountain newt, Neurergus derjugini, we employed phylogeography patterns and genetic variability of this species by analyzing partial ND4 sequences (867 bp) of 67 specimens from 15 sampling localities from the whole species range in Iran and Iraq. Phylogenetic trees concordant with haplotype networks showed a clear genetic structure among populations as three groups corresponding to the populations in the north, center, and south. Evolutionary ages of clades north and south ranging from 0.15 to 0.17 Myr, while the oldest clade is the central clade, corresponding to 0.32 Myr. Bayesian skyline plots of population size change through time show a relatively slight increase until about 25 kyr (around the last glacial maximum) and a decline of population size about 2.5 kyr. The presence of geographically structured clades in north, center, and south sections of the species range signifies the disjunct populations that have emerged in three different refugium. This study illustrates the importance of the effect of previous glacial cycles in shaping the genetic structure of mountain species in the Zagros range. These areas are important in terms of long‐term species persistence and therefore valuable areas for conservation of biodiversity.     

Morphological plasticity in the kelp Nereocystis luetkeana (Phaeophyceae) is sensitive to the magnitude,direction, and location of mechanical loading

Nereocystis luetkeana is a canopy-forming kelp that exhibits morphological plasticity across hydrodynamic gradients, producing broad, undulate blades in slow flow and narrow, flattened blades in fast flow, enabling thalli to reduce drag while optimizing photosynthesis. While the functional significance of this phenomenon has been well studied, the developmental and physiological mechanisms that facilitate the plasticity remain poorly understood. In this study, we conducted three experiments to characterize how the (1) magnitude, (2) direction, and (3) location of plasticity-inducing mechanical stimuli affect the morphology of Nereocystis blades. We found that applying a gradient of tensile force caused blades to grow progressively longer, narrower, less ruffled, and heavier in a linear fashion, suggesting that Nereocystis is equally well adapted for all conditions within its hydrodynamic niche. We also found that applying tension transversely across blades caused the growth response to rotate 90°, indicating that there is no substantial separation between the sites of stimulus perception and response and suggesting that a long-distance signaling mechanism, such as a hormone, is unlikely to mediate this phenomenon. Meristoderm cells showed morphological changes that paralleled those of their respective blades in this experiment, implying that tissue-level morphology is influenced by cell growth. Finally, we found that plasticity was only induced when tension was applied directly to the growing tissue, reinforcing that long-distance signaling is probably not involved and possibly indicating that the mechanism on display generally requires an intercalary meristem to facilitate mechanoperception.     

Functional response of Telenomus busseolae (Hym.: Scelionidae) an egg parasitoid of the sugarcane stem borer,Sesamia nonagrioides (Lep.: Noctuidae) at different temperatures

Telenomus busseolae Gahan is the most important egg parasitoid of sugarcane stem borers in Iran. Laboratory experiments were conducted to evaluate the functional response of T. busseolae to egg densities of Sesamia nonagrioides (Lefebvre) under four different temperatures (20, 25, 30 and 35°C). Results of logistic regression revealed a type III functional response for all temperatures tested and type of functional response was not affected by temperature. Roger's random parasite equation was fitted to observed data at 35°C. In the other data sets (collected data under 20, 25 and 30 °C) Roger's equation could not provide meaningful parameters, therefore Holling's disk equation was used to fit the data. The attack rate for T. busseolae varying from 0.001784 to 0.10704, 0.0026 to 0.152, 0.0024 to 0.144 and 0.000866 to 0.05196 per h and estimated handling times were 0.59, 0.50, 0.53 and 0.29 h at 20, 25, 30 and 35°C, respectively. Based on asymptotic 95% confidence intervals, functional response parameters did not differ significantly from 20 to 30°C, which indicates that handling time and attack rate are least affected by the temperature changes. The results suggest a similar performance of T. busseolae against S. nonagrioides eggs at temperatures of 20–30°C. These results show that T. busseolae is well adapted to temperature changes during the sugarcane growing season.     

The adult body size variation of Dryas iulia (Lepidoptera,Nymphalidae, Heliconiinae) in different populations is more influenced by temperature variation than by host‐plant availability during the seasons

The body size of insects is affected by environmental conditions during development and can present considerable intraspecific variations, which can be seen as an ultimate consequence/adaptation to environmental conditions. This paper evaluated whether the body size of the butterfly Dryas iulia from subtropical populations was influenced by changing climate conditions and food source availability during the seasons. The likely reasons behind body size variation were also investigated. First, field data on body size variation, host‐plant availability and climate fluctuation throughout the seasons were recorded. Then, the effects of host‐plant species and temperature on body size were analyzed by controlled experiments. Field data revealed that body size and host‐plant availability varied significantly through the seasons. Populations had the smallest body size during the spring and the biggest size during summer, whereas host‐plant availability was lower during winter and higher during spring. The controlled experiments revealed that both temperature and host‐plant had significant effect on the plasticity of body size. Larvae subjected to winter temperature treatment led to smaller butterflies when compared to immatures reared under summer temperature treatment, and larvae fed with Passiflora misera produced bigger adults when compared to larvae reared on Passiflora suberosa. The combination of data gathered in the field and in the laboratory suggests that seasonal body size variation in D. iulia is related mainly to differences in the temperatures to which larvae are subjected during development, while host‐plant shifts caused by differential availability of food through the seasons had slightly effects on the variation observed.     

Diet of Anguilla mossambica (Teleostei,Anguillidae) elvers in the Sundays River,Eastern Cape,South Africa

The southern African crab Hymenosoma orbiculare was recently split into five distinct species, of which three are estuarine/coastal and have peripatric distributions that are linked to temperature-defined marine bioregions. This suggests that the species’ ranges may be limited by physiological adaptations to their thermal environment. We explored this hypothesis by rearing the larvae of the warm-temperate lineage of H. orbiculare and the warm-temperate/subtropical H. longicrure at a range of water temperatures, and found clear temperature-dependent differences in the duration of larval development. Our study contributes to the growing body of evidence that stresses the importance of adaptation to regional environmental conditions, rather than physical dispersal barriers on their own, in limiting the mixing of marine species between temperature-defined biogeographic regions.     

Moose (Alces alces) reacts to high summer temperatures by utilizing thermal shelters in boreal forests – an analysis based on airborne laser scanning of the canopy structure at moose locations

The adaptation of different species to warming temperatures has been increasingly studied. Moose (Alces alces) is the largest of the ungulate species occupying the northern latitudes across the globe, and in Finland it is the most important game species. It is very well adapted to severe cold temperatures, but has a relatively low tolerance to warm temperatures. Previous studies have documented changes in habitat use by moose due to high temperatures. In many of these studies, the used areas have been classified according to how much thermal cover they were assumed to offer based on satellite/aerial imagery data. Here, we identified the vegetation structure in the areas used by moose under different thermal conditions. For this purpose, we used airborne laser scanning (ALS) data extracted from the locations of GPS‐collared moose. This provided us with detailed information about the relationships between moose and the structure of forests it uses in different thermal conditions and we were therefore able to determine and differentiate between the canopy structures at locations occupied by moose during different thermal conditions. We also discovered a threshold beyond which moose behaviour began to change significantly: as day temperatures began to reach 20 °C and higher, the search for areas with higher and denser canopies during daytime became evident. The difference was clear when compared to habitat use at lower temperatures, and was so strong that it provides supporting evidence to previous studies, suggesting that moose are able to modify their behaviour to cope with high temperatures, but also that the species is likely to be affected by warming climate.     

A laboratory-based comparison of osmoregulatory ability at different water temperatures and salinities in the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea

The aim of the investigation was a comparison of osmoregulatory ability at different water temperatures and salinities by the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea. The experiments were performed under laboratory conditions at different water temperatures (5, 10 and 15?°C) and salinities (3, 6.6, 15 and 25 PSU for S. entomon and 3, 6.3, 15 and 25 PSU for C. volutator). The osmotic concentrations of the crustaceans’ haemolymph were determined using the melting point method. Temperature had a statistically significant effect on the osmoregulation in S. entomon, but had generally no statistical influence on the osmoregulation in C. volutator. This physiological information regarding adaptation provides a basis for predicting the distribution of these species in changing environments. The implications of the results for the comparative adaptability of the two species are considered.