Ecophenotypic variation and phylogenetic inheritance in first lower molar shape of extant Italian populations of Microtus (Terricola) savii (Rodentia)

Seventeen extant populations of Microtus (Terricola) savii have been investigated for correlations of first lower molar shape to climatic variables by means of geometric morphometrics, and controlling for phylogenetic inheritance. Comparative methods revealed that climatic variables and phylogeography provide a very similar contribution to variation in first lower molars morphology, whereas tooth size does not appear to be affected by climatic conditions. Climate‐related changes have been recognized in the anteroconid portion of the tooth. This indicates that molar tooth variation is strongly influenced by climatic conditions, although in a complex way. Calabrian populations, often ascribed to Microtus (Terricola) brachycercus, form a distinct cluster, in agreement with the most recent genetic analyses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 632–647.     

Sex‐specific evolution of bite performance in Liolaemus lizards (Iguania: Liolaemidae): the battle of the sexes

Although differential selective pressures on males and females of the same species may result in sex‐specific evolutionary trajectories, comparative studies of adaptive radiations have largely neglected within‐species variation. In this study, we explore the potential effects of natural selection, sexual selection, or a combination of both, on bite performance in males and females of 19 species of Liolaemus lizards. More specifically, we study the evolution of bite performance, and compare evolutionary relationships between the variation in head morphology, bite performance, ecological variation and sexual dimorphism between males and females. Our results suggest that in male Liolaemus, the variation in bite force is at least partly explained by the variation in the degree of sexual dimorphism in head width (i.e. our estimate of the intensity of sexual selection), and neither bite force nor the morphological variables were correlated with diet (i.e. our proxy for natural selection). On the contrary, in females, the variation in bite force and head size can, to a certain extent, be explained by variation in diet. These results suggest that whereas in males, sexual selection seems to be operating on bite performance, in the case of females, natural selection seems to be the most likely and most important selective pressure driving the variation in head size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 461–475.     

A test of Rensch's rule in varanid lizards

In a model group of giant reptiles, we explored the allometric relationships between male and female body size and compared the effects of sexual and fecundity selection, as well as some proximate causes, on macroevolutionary patterns of sexual size dimorphism (SSD). Monitor lizards are a morphologically homogeneous group that has been affected by extreme changes in body size during their evolutionary history, resulting in 14‐fold differences among the body sizes of recent species. Here, we analysed data concerning the maximum and/or mean male and female snout–vent lengths in 42 species of monitor lizard from literary sources and supplemented these data with measurements made in zoos. There was a wide scale of SSD from nearly monomorphic species belonging mostly to the subgenus Odatria and Prasinus group of the Euprepriosaurus to apparently male‐larger taxa. The variable best explaining SSD was the body size itself; the larger the species, the higher the SSD. This pattern agrees with the currently discussed Rensch's rule, claiming that the relationship between male and female body size is hyperallometric, i.e. the allometric exponent of this relationship exceeds unity and thus SSD increases with body size in the case of male‐larger taxa. All our estimates of the reduced major axis regression slopes of this relationship ranged from 1.132 to 1.155. These estimates are significantly higher than unity, and thus unequivocally corroborate the validity of Rensch's rule in this reptilian group. In spite of our expectation that the variation in SSD can be alternatively explained by variables reflecting the strength of sexual selection (presence of male combat), fecundity selection (e.g. clutch size and mass) and/or proximate ecological factors (habitat type), none of these variables had consistent effects on SSD, especially when the data were adjusted to phylogenetic dependence and/or body size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 293–306.     

Why do Anatolian ground squirrels exhibit a Bergmannian size pattern? A phylogenetic comparative analysis of geographic variation in body size

A phylogenetic comparative analysis of geographic variation in body size of an obligately hibernating marmotine species (Anatolian ground squirrels, Spermophilus xanthoprymnus) is presented in relation to environmental variables that pertain to four principal hypotheses (heat conservation, heat dissipation, primary productivity, and seasonality hypotheses). Adult Anatolian ground squirrels (78 males and 90 females) were collected from ten geographic localities in Anatolia for use in morphometric analyses. First, the study tested whether significant variation in body size occurs over the geographic range of S. xanthoprymnus. Then, to understand the possible cause(s) of the observed pattern of geographic variation in body size of Anatolian ground squirrels, four hypotheses were tested, separately and simultaneously, using a phylogenetic comparative method. Overall, food availability (primary productivity hypothesis) and, especially in males, over‐winter fasting endurance (seasonality hypothesis) are likely the primary underlying mechanisms generating the observed pattern of increasing body size towards colder, more seasonal environments, with higher summer precipitation and productivity (or a Bergmannian size pattern). © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 695–710.     

Climatic oscillations shape the phylogeographical structure of Atlantic Forest fire‐eye antbirds (Aves: Thamnophilidae)

In the present study, mitochondrial (mt)DNA sequence data were used to examine the genetic structure of fire‐eye antbirds (genus Pyriglena) along the Atlantic Forest and the predictions derived from the river hypothesis and from a Last Glacial Maximum Pleistocene refuge paleomodel were compared to explain the patterns of genetic variation observed in these populations. A total of 266 individuals from 45 populations were sampled over a latitudinal transect and a number of phylogeographical and population genetics analytical approaches were employed to address these questions. The pattern of mtDNA variation observed in fire‐eye antbirds provides little support for the view that populations were isolated by the modern course of major Atlantic Forest rivers. Instead, the data provide stronger support for the predictions of the refuge model. These results add to the mounting evidence that climatic oscillations appear to have played a substantial role in shaping the phylogeographical structure and possibly the diversification of many taxa in this region. However, the results also illustrate the potential for more complex climatic history and historical changes in the geographical distribution of Atlantic Forest than envisioned by the refuge model. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 900–824.     

Comparing macroecological patterns across continents: evolution of climatic niche breadth in varanid lizards

Macroecological analyses often test hypotheses at the global scale, or among more closely related species in a single region (e.g. continent). Here, we test several hypotheses about climatic niche widths among relatively closely related species that occur across multiple continents, and compare patterns within and across continents to see if they differ. We focus on the lizard genus Varanus (monitor lizards), which occurs in diverse environments in Africa, Asia, and Australia. We address three main questions. 1) How do climatic niche breadths of species on a given niche axis change based on the position of species along that niche axis? (E.g. are species that occur in more extreme environments more narrowly specialized for those conditions?) 2) Are there trade‐offs in niche breadths on temperature and precipitation axes among species, or are niche widths on different axes positively related? 3) Is variation in niche breadths among species explained primarily by within‐locality seasonal variation, or by differences in climatic conditions among localities across the species range? We generate a new time‐calibrated phylogeny for Varanus and test these hypotheses within and between continents using climatic data and phylogenetic methods. Our results show that patterns on each continent often parallel each other and global patterns. However, in many other cases, the strength of relationships can change dramatically among closely related species on different continents. Overall, we found that: 1) species in warmer environments have narrower temperature niche breadths, but there is no relationship between precipitation niche breadth and niche position; 2) temperature and precipitation niche breadths tend to be positively related among species, rather than showing trade‐offs; and 3) within‐locality seasonal variation explains most variation in climatic niche breadths. Some of these results are concordant with previous studies (in amphibians and North American lizards), and might represent general macroecological patterns.     

Lizard scales in an adaptive radiation: variation in scale number follows climatic and structural habitat diversity in Anolis lizards

Lizard scales vary in size, shape and texture among and within species. The overall function of scales in squamates is attributed to protection against abrasion, solar radiation and water loss. We quantified scale number of Anolis lizards across a large sample of species (142 species) and examined whether this variation was related either to structural or to climatic habitat diversity. We found that species in dry environments have fewer, larger scales than species in humid environments. This is consistent with the hypothesis that scales reduce evaporative water loss through the skin. In addition, scale number varied among groups of ecomorphs and was correlated with aspects of the structural microhabitat (i.e. perch height and perch diameter). This was unexpected because ecomorph groups are based on morphological features related to locomotion in different structural microhabitats. Body scales are not likely to play an important role in locomotion in Anolis lizards. The observed variation may relate to other features of the ecomorph niche and more work is needed to understand the putative adaptive basis of these patterns. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 570–579.     

Sexual selection as a promoter of population divergence in male phenotypic characters: a study on mainland and islet lizard populations

Sexual selection is often viewed as a promoter of population divergence, although some forms of sexual selection could rather hamper divergence. In the present study, we investigated whether sexual selection promotes divergence in sexually‐selected traits. We studied population variation in sexual selection in relation to colour morph and body size in islet and mainland populations of the Skyros wall lizard (Podarcis gaigeae). Females were most likely to mate with orange‐throated males with small body sizes, and male body size and coloration were therefore subject to correlational sexual selection. By contrast, male mating probabilities were not affected by any female phenotypic character. We also found variation in a female resistance trait (escape propensity), with females being more prone to escape when exposed to males from other habitats. Sexual selection could potentially affect the frequencies of throat colour morphs in this species by favouring orange‐throated males of small body size, although there was no evidence of sexual selection for local mates or rare phenotypes. The results obtained in the present study thus do not support a role for sexual selection as a promoter of population divergence in this species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 374–389.     

More than meets the eye: a morphological and phylogenetic comparison of long‐spurred,white‐flowered Satyrium species (Orchidaceae) in South Africa

Superficial similarities among unrelated species are often a result of convergent evolution and can cause considerable taxonomic confusion. A case in point is Satyrium eurycalcaratum , described here as a new species, which has been confused with several other Satyrium spp. with similar long‐spurred, white flowers. A phylogenetic analysis, based on molecular data, indicated that S. eurycalcaratum is not closely related to any of the species with which it has been previously confused. A comparative analysis of morphological characters in the seven South African Satyrium spp. with long‐spurred, white flowers showed that each of these, including S. eurycalcaratum , is characterized by a unique combination of traits. Despite the similarity in pollination syndrome characters, such as spur length and flower colour, variation in rostellum structure was particularly pronounced and four distinctive forms were present. There was no phylogenetic signal in patterns of interspecific rostellum variation, as some closely related species had different rostella, whereas some distantly related species shared similar rostellum structures. We therefore conclude that the use of rostellum traits in conjunction with phylogenetic evidence can resolve species delimitations among orchid species that share the same pollination syndrome. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 417–430.     

Patterns of ecomorphological convergence among mainland and island Anolis lizards

Differing selective pressures on islands versus the mainland may produce alternative evolutionary outcomes among closely related lineages. Conversely, lineages may be constrained to produce similar outcomes in different mainland and island environments, or mainland and island environments may not differ significantly. Among the best‐studied island radiations are Caribbean Anolis lizards. Distinct morphotypes, or ‘ecomorphs’, have been described, and the same ecomorphs have evolved independently on each Greater Antillean island. The mainland Anolis radiation has received much less attention. We use a large morphological data set and a novel phylogenetic hypothesis to show that mainland Anolis did not evolve the same morphotypes as island Anolis, despite some island species being more closely related to mainland species than to island species that share their morphotype. A maximum of four of the six Caribbean ecomorphs were found to exist on the mainland, and just 15 of 123 mainland species are assignable to a Caribbean ecomorph. This result was insensitive to differing taxon samples and alternative phylogenetic hypotheses. Mainland convergence to a Caribbean ecomorph occurs only among species assigned to the grass‐bush ecomorph. Thus, the ecomorphs that have evolved convergently multiple times in the Caribbean have not evolved in parallel on the mainland. These results are consistent with the hypothesis that mainland and island environments offer different selective pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 852–859.     

Adaptive significance of food income in European snakes: body size is related to prey energetics

Feeding strategies and diet patterns have been extensively investigated in vertebrates and, more specifically, in snakes. Although it has been hypothesized that prey species may differ in terms of energy content, almost no theoretical or practical study has been carried out to determine actual nutritional values of the common prey types of wild snakes. Our model taxa were a selection of widely distributed and well known European snake species, which have all been studied in depth: approximately 76% of their diet is composed of mammals, reptiles, and insects. We therefore selected a single model species for each of these categories and proceeded with the analyses. Nutritional values were determined using a standard procedure: lizards and mice were richer in proteins than insects (crickets); insects and mice were richer in lipids than lizards, and mice and crickets have a higher energy content than lizards; lizards were rich in ashes. We then applied our experimental results to a selected sample of European terrestrial snakes (11 populations, ten species, seven genera, two families) characterized by different body size (50–160 cm total length) and reproductive strategies (oviparous versus viviparous), aiming to correlate these parameters with patterns of energy income. A direct relationship was found between body mass/body length ratio (BCI, body condition index) and meal energetics: the higher the BCI, the higher was the metabolic requirement, whereas BCI was independent of species or of reproductive system effect. Large‐sized snakes thus need a highly diversified and more energy‐rich diet than smaller snakes, supporting previous hypotheses. The simple applicability of this method could be of valuable support in further comparative research work, reducing experimental costs and stimulating further ecological, behavioural, and, possibly, phylogenetic comparisons. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 307–317.     

Latitudinal clines in body size,but not in thermal tolerance or heat‐shock cognate 70 (HSC70), in the highly‐dispersing intertidal gastropod Littorina keenae (Gastropoda: Littorinidae)

Natural populations of widely‐distributed animals often exhibit clinal variation in phenotypic traits or in allele frequencies of a particular gene over their geographical range. A planktotrophic intertidal snail, Littorina keenae is broadly distributed along the north‐eastern Pacific coast through a large latitudinal range (24°50′N–43°18′N). We tested for latitudinal clines in two complex phenotypic traits – thermal tolerance and body size – and one single locus trait – heat shock cognate 70 (HSC70) – in L. keenae along almost its entire geographical range. We found only weak evidence for a latitudinal cline in the thermal tolerance and no evidence for a cline in allele frequencies at HSC70. However, as predicted by Bergmann's rule, we detected a strong latitudinal cline that accounted for 60% of the variance in body size (R² = 0.598; P < 0.001). In contrast, body size did not significantly affect thermal tolerance. HSC70 showed no genetic differentiation among the populations, supporting our previous mitochondrial gene‐based estimate of high gene flow during this snail's free‐swimming larval stage. Given that L. keenae experiences panmixia along its species range, the observed size cline may be partially or entirely caused by a phenotypically plastic response to local thermal environments rather than by genetic divergence in body size among populations in response to locally optimizing natural selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 494–505.     

The origin of snakes (Serpentes) as seen through eye anatomy

Snakes evolved from lizards but have dramatically different eyes. These differences are cited widely as compelling evidence that snakes had fossorial and nocturnal ancestors. Their eyes, however, also exhibit similarities to those of aquatic vertebrates. We used a comparative analysis of ophthalmic data among vertebrate taxa to evaluate alternative hypotheses concerning the ecological origin of the distinctive features of the eyes of snakes. In parsimony and phenetic analyses, eye and orbital characters retrieved groupings more consistent with ecological adaptation rather than accepted phylogenetic relationships. Fossorial lizards and mammals cluster together, whereas snakes are widely separated from these taxa and instead cluster with primitively aquatic vertebrates. This indicates that the eyes of snakes most closely resemble those of aquatic vertebrates, and suggests that the early evolution of snakes occurred in aquatic environments.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 469–482.     

Epidermal glands in cordylid lizards,with special reference to generation glands

We investigated inter‐ and intraspecific variation in epidermal gland characteristics in cordylid lizards. Our particular interest was whether a causal relationship exists between the presence of generation glands and lifestyle, i.e. in rock‐dwelling or ground‐dwelling lizards. We established that both femoral and generation glands are always present in adult males, but that species can be divided into three categories on the basis of the presence of these glands in adult females: both femoral and generation glands absent; femoral glands present, but generation glands absent; and both femoral and generation glands present. The absence of epidermal glands in females appears to be the basal condition in the Cordylidae. All cordylid species are sexually dimorphic as far as generation gland number is concerned, with females having consistently less glands than males. Gland number in females is strongly affected by climate, being lower in cooler compared with warmer environments. In males, there is no clear geographical pattern, although in some clades males of high‐altitude species have more generation glands than males of species occurring at lower altitudes. At least four of the five ground‐dwelling species in the family display unique generation gland characteristics, compared with rock‐dwelling species, reflecting the continued importance of generation glands in terrestrial environments. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 312–324.     

Lack of taxonomic information from parietal spine size invalidates subspecies in the Atlantic hookear sculpin Artediellus atlanticus

An analysis of 107 individuals of Atlantic hookear sculpin Artediellus atlanticus from the Barents Sea–Svalbard region and from north‐east Greenland shows that the state of the parietal spines, although not randomly distributed geographically, occurs sympatrically. They do not diagnostically describe populations. They are therefore inept for subspecies delimitation into Artediellus atlanticus atlanticus, Artediellus atlanticus corniger and Artediellus atlanticus europaeus. Sympatric occurrence of parietal spine states called for a molecular approach to look for differences at a species level. Sequencing the mitochondrial genes, cytochrome oxidase c subunit I (coI) and cytochrome b (cytb), shows no difference between individuals with different parietal spine size. Thus, there is no evidence of the spine states representing species‐level variation. The study shows that parietal spine size has no taxonomic information for this species. The patterns of variation in parietal spine size differ to some degree between the sexes.     

Morphology of the Podarcis wall lizards (Squamata: Lacertidae) from the Iberian Peninsula and North Africa: patterns of variation in a putative cryptic species complex

Cryptic species complexes represent groups that have been classified as a single species, because of the difficulty in distinguishing its members morphologically. Morphological investigation following the discovery of cryptic diversity is crucial for describing and conserving biodiversity. Here we present a detailed account of morphological variation in a group of Iberian and North African Podarcis wall lizards of the family Lacertidae, trying to elucidate the morphological patterns observed between known mitochondrial lineages. Our results reveal very high morphological variation within lineages, considering both biometric and pholidotic traits, but also indicate that lineages are significantly different from each other. The main sources of variation, both globally and between lineages, arise from body size, head dimensions, and limb length, possibly pointing to underlying ecological mechanisms. A combination of body size, body shape, and continuous pholidotic traits allows a relatively good discrimination between groups, especially when comparing one group with the rest or pairs of groups. However, ranges of variation greatly overlap between groups, thereby not allowing the establishment of diagnostic traits. The high morphological variation observed indicates that external morphology is not particularly useful for species delimitation in this group of lizards, as local adaptation seems to play a major role in within‐ and between‐group differentiation. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 173–193.     

Sexual dimorphism and interpopulation differences in lizard hind limb length: locomotor performance or chemical signalling?

Intraspecific variation in morphology has often been related to fitness differences through its effects on performance. In lizards, variation in hind limb length can be shaped by natural selection for increased locomotor performance, sexual selection on the number or size of femoral pores involved in chemical signalling, or both. Here, we analyse the selective forces involved in sexual dimorphism and differences in hind limb length between two populations of Psammodromus algirus living at different elevation. Males were more robust and had longer hind limbs and limb segments than females, and low‐elevation lizards had longer limbs than high‐elevation lizards. However, differences in locomotor performance were small and non‐significant, making natural selection for faster runs an unlikely explanation for the observed pattern. On the other hand, males had more femoral pores than females, and lizards had more pores at lower elevation, although the difference was significant only for males (which invest more in chemical signalling). In males, the number of pores, which remains constant along a lizard's life, was not correlated with hind limb length. However, femur length was positively correlated with mean pore size, allowing low‐elevation males to have larger than expected pores, which could increase the effectiveness with which they spread their signals in a dry and warm habitat where chemicals become volatile rapidly. Also, saturation of the sexual coloration of the head was higher for low‐elevation males, suggesting that sexual selection pressures may be more intense. Overall, our results indicate that sexual selection plays a significant role in shaping intraspecific variation in hind limb length. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 318–329.     

Disentangling the effects of intraspecies variability,phylogeny, space,and climate on the evolution of shell morphology in endemic Greek land snails of the genus Codringtonia

Extensive variation in land snail shell morphology has been widely documented, although few studies have attempted to investigate the ecological and evolutionary drivers of this variation. Within a comparative phylogenetic framework, we investigated the temporal and spatial evolution of the shell morphology of the Greek endemic land snail genus Codringtonia. The contribution of both inter‐ and intraspecies shell differentiation in the overall shell variability is assessed. The effect of climate, space, and evolutionary history on the shell variability was inferred using a variance partitioning framework. For Codringtonia species, intraspecies divergence of shell traits contributes substantially to the overall shell variability. By decomposing this variability, it was shown that the overall shell size of Codringtonia clades is phylogenetically constrained, related to early speciation events, and strongly affected by large‐scale spatial variability (latitudinal gradient). The effect of climate on shell size cannot be disentangled from phylogeny and space. Shell and, to a larger extent, aperture shape are not phylogenetically constrained, and appear to be mostly related to conspecific populations divergence events. Shell shape is substantially explained by both climate and space that greatly overlap. Aperture shape is mainly interpreted by medium to small‐scale spatial variables. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 796–813.     

Large‐sized insects show stronger seasonality than small‐sized ones: a case study of fruit‐feeding butterflies

Animal species have a restricted period during the year when conditions for development are optimal, and this is known as the temporal window. Duration of the temporal window can vary among species, although the causes of variation are still poorly understood. In the present study, examining butterflies, we assume that the temporal window duration is correlated with the seasonal period of flight (termed seasonality). To understand how species characteristics are correlated with this, we examine whether there is a relationship between body size and length of flight period of fruit‐feeding butterflies in forest fragments, and whether these two parameters have a phylogenetic signal. Using wing size as a measure of body size and the period of adult flight as a measure of seasonality, we found significant positive correlations between body size and seasonality among subfamilies but not within subfamilies. We also found a clear phylogenetic signal in size but not in seasonality. The results obtained suggest the existence of a trade‐off between insect size and seasonality, with size limiting flight period length. The relationship between body size and seasonality and the synchrony with their resources may be one factor explaining the vulnerability of large insects to forest fragmentation. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 820–827.     

Climate niche conservatism does not explain restricted distribution patterns in Tynanthus (Bignonieae,Bignoniaceae)

Studies on niche evolution allow us to establish how species niches have changed over time and to identify how long‐term evolutionary processes have led to present‐day species distributions. Here, we investigate the patterns of climatic niche evolution in Tynanthus (Bignonieae, Bignoniaceae), a genus of narrowly distributed species. We test the hypothesis that niche conservatism has played an important role in the history of this group of Neotropical lianas. We perform univariate and multivariate comparisons between climatic niches of species and associated environmental data with information on phylogenetic relationships. We encountered considerable divergence in niches among species, indicating that niche conservatism in climatic variables does not seem to have played a key role in the history of the genus. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 95–109.