Trunk elongation and ontogenetic changes in the axial skeleton of Triturus newts

Body elongation in vertebrates can be achieved by lengthening of the vertebrae or by an increase in their number. In salamanders, longer bodies are mostly associated with greater numbers of vertebrae in the trunk or tail region. However, studies on the relative contribution of the length of single vertebra to body elongation are lacking. In this study, we focus on evolutionary and ontogenetic changes in differentiation of the trunk vertebrae and the relative contribution of individual vertebrae to trunk lengthening in Triturus newts, a monophyletic group of salamanders that shows remarkable disparity in body shape. We compared juveniles and adults of the most elongated T. dobrogicus , which has 17 trunk vertebrae, with juveniles and adults of two closely related species (T. ivanbureschi and T. anatolicus belonging to the T. karelinii species complex) representing a stout and robust morphotype with thirteen trunk vertebrae. We show that trunk vertebrae are uniform in size at the juvenile stage of both analyzed morphotypes. In adults, the trunk vertebrae of the elongated T. dobrogicus are largely uniform, while in those of T. anatolicus , the first two vertebrae differ from the remaining trunk vertebrae. There was no difference in the relative contribution of individual vertebrae to body lengthening between species or stages. We conclude that body elongation in Triturus newts is achieved by increasing the number of vertebrae but not their length.     

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.     

Sexual size dimorphism is not associated with the evolution of parental care in frogs

Sex differences in parental care are thought to arise from differential selection on the sexes. Sexual dimorphism, including sexual size dimorphism (SSD), is often used as a proxy for sexual selection on males. Some studies have found an association between male‐biased SSD (i.e., males larger than females) and the loss of paternal care. While the relationship between sexual selection on males and parental care evolution has been studied extensively, the relationship between female‐biased SSD (i.e., females larger than males) and the evolution of parental care has received very little attention. Thus, we have little knowledge of whether female‐biased SSD coevolves with parental care. In species displaying female‐biased SSD, we might expect dimorphism to be associated with the evolution of paternal care or perhaps the loss of maternal care. Here, drawing on data for 99 extant frog species, we use comparative methods to evaluate how parental care and female‐biased SSD have evolved over time. Generally, we find no significant correlation between the evolution of parental care and female‐biased SSD in frogs. This suggests that differential selection on body size between the sexes is unlikely to have driven the evolution of parental care in these clades and questions whether we should expect sexual dimorphism to exhibit a general relationship with the evolution of sex differences in parental care.     

The evolution of thermal performance curves in semi-aquatic newts: Thermal specialists on land and thermal generalists in water?

The position and shape of thermal performance curves (TPCs, the functions relating temperature to physiological performance) for ecologically relevant functions will directly affect the fitness of ectotherms and therefore should be under strong selection. However, thermodynamic considerations predict that relationships between the different components of the TPC will confound its evolutionary optimization. For instance, the “jack-of-all-temperatures” hypothesis predicts a trade-off between the breadth of the TPC and the maximal performance capacity; the “warmer is better” hypothesis suggests that low thermal optima will come with low absolute performances. Semi-aquatic organisms face the additional challenge of having to adjust their TPCs to two environments that are likely to differ in mean temperature and thermal variability. In this paper, we examine how parameters of the TPCs for maximal running and swimming speed have co-evolved in the semi-aquatic newt genus Triturus. We consider evolutionary relationships between the width and the height of the TPCs, the optimal temperatures and maximal performance. Phylogenetic comparative analyses reveal that in Triturus, swimming and running differ substantially in the (co-)variation of TPC parameters. Whereas evolutionary changes in the TPC for swimming primarily concern the shape of the curve (generalist versus specialist), most interspecific variation in running speed TPCs involves shifts in overall performance across temperatures.     

No trade-off between sprinting and climbing in two populations of the Lizard Podarcis hispanica (Reptilia: Lacertidae).

Biomechanical reflections suggest that lizards that have specialized in running should differ in parts of their morphological design from lizards that have specialized in climbing. Moreover, adaptation in one direction may conflict with performance in the other. We tested this assumed trade-off by measuring climbing and running performance in two subspecies of the Mediterranean lizard Poaarcis hispanica. One subspecies, P. h. atrata , inhabits a number of small islets near the Spanish coast. It is mainly ground dwelling. The other subspecies, P. h. hispanica , was sampled on the mainland of Spain. It frequendy occupies vertical elements within its habitat. Our data do not support the notion of a trade-off between both types of locomotion. Individuals of P. h. hispanica both run and climb faster than those of P. h. atrata. There is no difference between subspecies in ability to cling onto tilted substrates. Predictions on the morphology of both subspecies, inspired by biomechanical arguments, are not supported by our measurements.     

Patterns of sexual size dimorphism in Chelonia

Macroevolutionary patterns of sexual size dimorphism (SSD) indicate how sexual selection, natural selection, and genetic and developmental constraints mold sex differences in body size. One putative pattern, known as Rensch's rule, posits that, among species with female‐larger SSD, the relative degree of SSD declines with species' body size, whereas, among male‐larger SSD species, relative SSD increases with size. Using a dataset of 196 chelonian species from all fourteen families, we investigated the correlation in body size evolution between male and female Chelonia and the validity of Rensch's rule for the taxon and within its major clades. We conclude that male–female correlations in body size evolution are high, although these correlations differ among chelonian families. Overall, SSD scales isometrically with body size; Rensch's rule is valid for only one family, Testudinidae (tortoises). Because macroevolutionary patterns of SSD can vary markedly among clades, even in a taxon as morphologically conservative as Testudines, one must guard against inappropriately pooling clades in comparative studies of SSD. The results of the present study also indicate that regression models that assume the x‐variable (e.g. male body size) is measured without statistical error, although frequently reported, will result in erroneous conclusions about phylogenetic trends in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 396–413.     

Testing the evolutionary constraints of metamorphosis: The ontogeny of head shape in Triturus newts

In vertebrates with complex, biphasic, life cycles, larvae have a distinct morphology and ecological preferences compared to metamorphosed juveniles and adults. In amphibians, abrupt and rapid metamorphic changes transform aquatic larvae to terrestrial juveniles. The main aim of this study is to test whether, relative to larval stages, metamorphosis (1) resets the pattern of variation between ontogenetic stages and species, (2) constrains intraspecific morphological variability, and (3) similar to the “hour‐glass” model reduces morphological disparity. We explore postembryonic ontogenetic trajectories of head shape (from hatching to completed metamorphosis) of two well‐defined, morphologically distinct Triturus newts species and their F1 hybrids. Variation in head shape is quantified and compared on two levels: dynamic (across ontogenetic stages) and static (at a particular stage). Our results show that the ontogenetic trajectories diverge early during development and continue to diverge throughout larval stages and metamorphosis. The high within‐group variance and the largest disparity level (between‐group variance) characterize the metamorphosed stage. Hence, our results indicate that metamorphosis does not canalize head shape variation generated during larval development and that metamorphosed phenotype is not more constrained relative to larval ones. Therefore, metamorphosis cannot be regarded as a developmental constraint, at least not for salamander head shape.     

Testing for evolutionary trade-offs in a phylogenetic context: ecological diversification and evolution of locomotor performance in emydid turtles

The evolution of ecological trade-offs is an important component of ecological specialization and adaptive radiation. However, the pattern that would show that evolutionary trade-offs have occurred between traits among species has not been clearly defined. In this paper, we propose a phylogeny-based definition of an evolutionary trade-off, and apply it to an analysis of the evolution of trade-offs in locomotor performance in emydid turtles. We quantified aquatic and terrestrial speed and endurance for up to 16 species, including aquatic, semi-terrestrial and terrestrial emydids. Emydid phylogeny was reconstructed from morphological characters and nuclear and mitochondrial DNA sequences. Surprisingly, we find that there have been no trade-offs in aquatic and terrestrial speed among species. Instead, specialization to aquatic and terrestrial habitats seems to have involved trade-offs in speed and endurance. Given that trade-offs between speed and endurance may be widespread, they may underlie specialization to different habitats in many other groups.     

东方蝾螈幼体热耐受性和游泳表现的热驯化响应

特定物种的热驯化能力决定着其是否能成功耐受环境温度的改变,在应对未来气候变暖的趋势中扮演重要角色。为评估有尾类两栖动物的热驯化反应,在3个恒定水温(15、20℃和25℃)中驯化东方蝾螈(Cynops orientalis)幼体4周,测定驯化后幼体在不同测试温度下的运动(游泳)表现、临界低温(CT_(Min))与临界高温(CT_(Max))。结果显示:驯化与测试温度均显著影响蝾螈的游泳速度;驯化温度亦影响蝾螈的CT_(Min)和CT_(Max),但不影响可耐受温度范围(TRR)。驯化与测试温度的交互作用对蝾螈泳速的影响显著,表明驯化温度可改变其游泳表现的热敏感性。经某一温度驯化后蝾螈泳速似乎在相同测试温度下表现最好,该结果可能支持驯化有益假说。CT_(Min)和CT_(Max)随驯化温度的升高而增加,表明:低温驯化可提高动物抗低温能力,而高温驯化提高其抗高温能力。两栖类动物热耐受性与运动表现热驯化反应的种间变异可能与栖息地热环境的差异有关。     

Acclimation is beneficial at extreme test temperatures in the Danube crested newt, Triturus dobrogicus (Caudata, Salamandridae)

Despite many studies demonstrating the effect of acclimation on behavioural or physiological traits, considerable debate still exists about the evolutionary significance of this phenomenon. One of the unresolved issues is whether acclimation to warmer temperature is beneficial at treatment or at more extreme test temperatures. To answer this question, we assessed the effect of thermal acclimation on preferred body temperatures ( T _ps), maximum swimming and running speed, and critical thermal maximum ( CT _max) in the Danube crested newt ( Triturus dobrogicus ). Adult newts were kept at 15 °C (control) and 25 °C (treatment) for 8 weeks prior to measurements. We measured T _ps in an aquatic thermal gradient over 24 h, maximum speeds in a linear racetrack at six temperatures (5–33 °C), and CT _max in a continuously heated water bath. T _ps were higher in newts kept at 15 °C than in those kept at 25 °C. The maximum swimming speed did not acclimate. The maximum running speed at 30–33 °C was substantially higher in newts kept at 25 °C than in those kept at 15 °C. CT _max increased with the treatment temperature. Hence, we conclude that the acclimation response to warm temperature is beneficial not at treatment but at more extreme temperatures in newts.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 627–636.     

Uncertain branch lengths, taxonomic sampling error, and the egg to body size allometry in temperate butterflies (Lepidoptera)

The allometry of egg to body size in temperate butterflies, and the relationships between egg size and larval host structure, taxonomy, voltinism, and duration of the egg stage, are investigated using cross-species analysis and the comparative analysis of independent contrasts. In addition, the effect of two sources of uncertainty that may affect continuous data when treated under a statistical, comparative, framework, is assessed: (1) unknown evolutionary distances, and (2) taxonomic representativity (proportion of species of a given taxon, from which data are available). It is suggested that the effects of taxonomic under-representation could be important in comparative, quantitative studies, but this problem may be tempered by means of weighted regression. Under the assumption that taxonomy represents butterfly phylogeny, egg and adult body size are related by negative allometry (i.e. the slope of the line fitted to the logarithmically transformed data is lower than 1.0). However, the precise slope (0.2-0.9) depends on the method used, branch lengths, and taxonomic sampling. There is evidence for a relationship between a species' voltinism and the number of days it spends in the egg stage, as well as between adult butterfly size and the gross structure of the plants used as larval hosts (woody plants or herbs). Egg size proves to be related to foodplant taxonomy, voltinism, and duration of the egg stage when the data are analysed using species means, but these relationships become non-significant when the comparative method is employed.     

Jaw biomechanics and the evolution of biting performance in theropod dinosaurs

Despite the great diversity in theropod craniomandibular morphology, the presence and distribution of biting function types across Theropoda has rarely been assessed. A novel method of biomechanical profiling using mechanical advantage computed for each biting position along the entirety of the tooth row was applied to 41 extinct theropod taxa. Multivariate ordination on the polynomial coefficients of the profiles reveals the distribution of theropod biting performance in function space. In particular, coelophysoids are found to occupy a unique region of function space, while tetanurans have a wide but continuous function space distribution. Further, the underlying phylogenetic structure and evolution of biting performance were investigated using phylogenetic comparative methods. There is a strong phylogenetic signal in theropod biomechanical profiles, indicating that evolution of biting performance does not depart from Brownian motion evolution. Reconstructions of ancestral function space occupation conform to this pattern, but phylogenetically unexpected major shifts in function space occupation can be observed at the origins of some clades. However, uncertainties surround ancestor estimates in some of these internal nodes, so inferences on the nature of these evolutionary changes must be viewed with caution.     

Ecomorphological convergence in planktivorous surgeonfishes

Morphological convergence plays a central role in the study of evolution. Often induced by shared ecological specialization, homoplasy hints at underlying selective pressures and adaptive constraints that deterministically shape the diversification of life. Although midwater zooplanktivory has arisen in adult surgeonfishes (family Acanthuridae) at least four independent times, it represents a clearly specialized state, requiring the capacity to swiftly swim in midwater locating and sucking small prey items. Whereas this diet has commonly been associated with specific functional adaptations in fishes, acanthurids present an interesting case study as all nonplanktivorous species feed by grazing on benthic algae and detritus, requiring a vastly different functional morphology that emphasizes biting behaviours. We examined the feeding morphology in 30 acanthurid species and, combined with a pre‐existing phylogenetic tree, compared the fit of evolutionary models across two diet regimes: zooplanktivores and nonzooplanktivorous grazers. Accounting for phylogenetic relationships, the best‐fitting model indicates that zooplanktivorous species are converging on a separate adaptive peak from their grazing relatives. Driving this bimodal landscape, zooplanktivorous acanthurids tend to develop a slender body, reduced facial features, smaller teeth and weakened jaw adductor muscles. However, despite these phenotypic changes, model fitting suggests that lineages have not yet reached the adaptive peak associated with plankton feeding even though some transitions appear to be over 10 million years old. These findings demonstrate that the selective demands of pelagic feeding promote repeated – albeit very gradual – ecomorphological convergence within surgeonfishes, while allowing local divergences between closely related species, contributing to the overall diversity of the clade.     

The evolution of jumping performance in anurans: morphological correlates and ecological implications

We investigated the evolution of anuran locomotor performance and its morphological correlates as a function of habitat use and lifestyles. We reanalysed a subset of the data reported by Zug (Smithson. Contrib. Zool. 1978; 276: 1–31) employing phylogenetically explicit statistical methods (n = 56 species), and assembled morphological data on the ratio between hind-limb length and snout-vent length (SVL) from the literature and museum specimens for a large subgroup of the species from the original paper (n = 43 species). Analyses using independent contrasts revealed that classifying anurans into terrestrial, semi-aquatic, and arboreal categories cannot distinguish between the effects of phylogeny and ecological diversification in anuran locomotor performance. However, a more refined classification subdividing terrestrial species into 'fossorials' and 'non-fossorials', and arboreal species into 'open canopy', 'low canopy' and 'high canopy', suggests that part of the variation in locomotor performance and in hind-limb morphology can be attributed to ecological diversification. In particular, fossorial species had significantly lower jumping performances and shorter hind limbs than other species after controlling for SVL, illustrating how the trade-off between burrowing efficiency and jumping performance has resulted in morphological specialization in this group.     

Do the Oaxacan Highlands represent a natural biotic unit? A cladistic biogeographical test based on vertebrate taxa

Aim  We analysed the distributional patterns of six terrestrial vertebrate taxa from the Oaxacan Highlands (Sierra Mazateca, Nudo de Zempoaltépetl and Sierra de Juárez) through a cladistic biogeographical approach, in order to test their naturalness as a biotic unit.
Location  The Oaxacan Highlands, Mexico.
Methods  The cladistic biogeographical analysis was based on the area cladograms of the Pseudoeurycea bellii species group (Amphibia: Plethodontidae), the genus Chlorospingus (Aves: Thraupidae), the genera Microtus , Reithrodontomys and Habromys , and the Peromyscus aztecus species group (Mammalia: Rodentia). We obtained paralogy-free subtrees, from which the components were coded in a data matrix for parsimony analysis. The data matrix was analysed with N ona through W in C lada .
Results  The parsimony analysis resulted in a single general area cladogram in which areas were fragmented following the sequence Sierra Madre Occidental, Trans-Mexican Volcanic Belt, Chiapas, Sierra Madre Oriental + Sierra Mazateca, Sierra Madre del Sur, Nudo de Zempoaltépetl and Sierra de Juárez.
Main conclusions  The general area cladogram shows that the Oaxacan Highlands do not constitute a natural unit. The Sierra Mazateca is the sister area to the Sierra Madre Oriental, whereas the Nudo de Zempoaltépetl and the Sierra de Juárez are closely related to the Sierra Madre del Sur. The events that might have caused these patterns include cycles of expansion and contraction of mountain pinyon, juniper and oak woodlands during the Pleistocene.     

Mode and tempo in the evolution of socio-political organization: reconciling 'Darwinian' and 'Spencerian' evolutionary approaches in anthropology

Traditional investigations of the evolution of human social and political institutions trace their ancestry back to nineteenth century social scientists such as Herbert Spencer, and have concentrated on the increase in socio-political complexity over time. More recent studies of cultural evolution have been explicitly informed by Darwinian evolutionary theory and focus on the transmission of cultural traits between individuals. These two approaches to investigating cultural change are often seen as incompatible. However, we argue that many of the defining features and assumptions of 'Spencerian' cultural evolutionary theory represent testable hypotheses that can and should be tackled within a broader 'Darwinian' framework. In this paper we apply phylogenetic comparative techniques to data from Austronesian-speaking societies of Island South-East Asia and the Pacific to test hypotheses about the mode and tempo of human socio-political evolution. We find support for three ideas often associated with Spencerian cultural evolutionary theory: (i) political organization has evolved through a regular sequence of forms, (ii) increases in hierarchical political complexity have been more common than decreases, and (iii) political organization has co-evolved with the wider presence of hereditary social stratification.     

体温、摄食、断尾和雌体繁殖状态对原尾蜥虎运动表现的影响

设计4项实验研究4种内外因素(环境温度、摄食、断尾和怀卵状态)导致的肇庆原尾蜥虎运动表现种群内变异.环境温度通过影响体温而影响原尾蜥虎运动表现.两性成体疾跑速均具有在低体温范围内随体温升高而加快、在高体温范围内随体温升高而降低的一般模式.在任何体温下成年雌体的疾跑速均略大于雄体,但两者平均值在统计上无显著差异.体温对原尾蜥虎最大持续运动距离的影响存在两性间差异.成年雌体的最大持续运动距离在低体温范围(从17 ℃到27 ℃)内随体温升高而增加,而在相对较高的体温范围(从27 ℃到37 ℃)内无显著变化,成年雄体在实验体温范围(从17 ℃到37 ℃)内无显著变化;在任何体温下成年雌体的最大持续运动距离均显著大于雄性.性别与体温相互作用对最大持续运动距离有显著影响.两性个体的平均疾跑速和最大持续运动距离呈显著的正相关.当去除最大持续运动距离差异的影响后发现,疾跑速两性差异统计上仍不显著.摄食、尾自切和雌体怀卵在两个实验温度下(27 ℃和31 ℃)均减小壁虎疾跑速;而仅尾自切减小壁虎的最大持续运动距离,摄食和雌体怀卵对其无影响.以上3种因子分别与体温、性别的交互作用均不显著.这些结果说明生理限制和尾自切是原尾蜥虎运动能力变异的重要来源.一些内外因素能近因性诱导蜥蜴功能表现发生一定程度的种群内变异.     

Stomatal density responses of temperate woodland plants over the past seven decades of CO2 increase: a comparison of Salisbury (1927) with contemporary data

We investigated the possible effect of recent (1927-1995) increases in the concentration of atmospheric CO2 on the stomatal densities of leaves of a wide range of tree, shrub, and herb species (N = 60) by making new measurements for comparison with corresponding data reported by E. J. Salisbury in 1927--a time when ice core studies indicate CO2 concentrations ~55 mL/L lower than present. A detailed intraspecific study of the herb Mercurialis perennis showed plants of M. perennis in a Cambridgeshire woodland in 1994 had significantly lower stomatal densities, irrespective of leaf insertion point, compared with their 1927 counterparts. Comparisons made across species using evolutionary comparative methods (independent contrasts) revealed a significant (P 2 increases have influenced leaf morphology in a manner consistent with recent experiments and the palaeoecological record. Further analyses suggested that the strength of the stomatal density response was independent of life form but dependent on "exposure" and the initial leaf stomatal density. Consequently, firmer predictions for future changes in stomatal density across all species, expected as a possible result of anthropogenically related CO2 increases, may now be possible.     

Early evolutionary differentiation of morphological variation in the mandible of South American caviomorph rodents (Rodentia, Caviomorpha)

Caviomorphs are a clade of South American rodents recorded at least since the early Oligocene (> 31.5 Ma) that exhibit ample eco-morphological variation. It has been proposed that phylogenetic structure is more important than ecological factors for understanding mandibular shape variation in this clade. This was interpreted as a result of the long-standing evolutionary history of caviomorphs and the early divergence of major lineages. In this work, we test this hypothesis through the analysis of morphological variation in the mandible of living and extinct species and compare this information with that obtained through comparative phylogenetic analyses. Our results support the hypothesis of early origin of mandibular variation; moreover, they suggest the conservation of early differentiated morphologies, which could indicate the existence of constrained evolutionary diversification.     

A Bayesian extension of phylogenetic generalized least squares: Incorporating uncertainty in the comparative study of trait relationships and evolutionary rates

Phylogenetic comparative methods use tree topology, branch lengths, and models of phenotypic change to take into account nonindependence in statistical analysis. However, these methods normally assume that trees and models are known without error. Approaches relying on evolutionary regimes also assume specific distributions of character states across a tree, which often result from ancestral state reconstructions that are subject to uncertainty. Several methods have been proposed to deal with some of these sources of uncertainty, but approaches accounting for all of them are less common. Here, we show how Bayesian statistics facilitates this task while relaxing the homogeneous rate assumption of the well-known phylogenetic generalized least squares (PGLS) framework. This Bayesian formulation allows uncertainty about phylogeny, evolutionary regimes, or other statistical parameters to be taken into account for studies as simple as testing for coevolution in two traits or as complex as testing whether bursts of phenotypic change are associated with evolutionary shifts in intertrait correlations. A mixture of validation approaches indicates that the approach has good inferential properties and predictive performance. We provide suggestions for implementation and show its usefulness by exploring the coevolution of ankle posture and forefoot proportions in Carnivora.