Ecological consequences of ontogenetic changes in head shape and bite performance in the Jamaican lizard Anolis lineatopus

It has been documented extensively that body size affects the physiology and musculoskeletal function of organisms. However, less well understood is how body size affects the ecology of organisms through its effects on physiology and performance. We explored the effects of body size on morphology and performance in different ontogenetic classes and sexes of a common Anolis lizard ( A. lineatopus ). Next, we tested whether these morphological and performance differences may affect functional aspects of the diet such as prey size and prey hardness. Our data showed that males, females and juveniles differ significantly in head size, head shape and bite force. Multiple regression models indicated that head shape and bite force are significantly correlated to prey size and hardness. Yet juveniles had relatively large heads and bit disproportionately hard for their size, allowing them to eat prey as large as those of females. However, for a given prey size, males and females ate more robust prey than did juveniles. Additionally, males ate relatively harder prey than did juveniles. These data suggest that: (1) body size affects the dietary ecology of animals through its effect on head size and bite force; (2) changes in head morphology independent of changes in overall size also have important effects on performance and diet.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 443–454.     

Adaptive modifications of carapace outlines in the Cytheroidea (Ostracoda: Crustacea)

Shell outlines of 202 extant cytheroidean ostracods were analysed in dorsal, lateral, and posterior views by elliptic Fourier analysis. The results obtained confirm that the exterior morphology is related to ecological factors as well as phylogenetic constraints. Phytal species living on tall seagrass and benthic species burrowing in sediments are characterized and differentiated from the species crawling on sediments by the presence of slender shells with tapered venters. With reference to reconstruction of ancestral state of outline traits on the molecular phylogeny, the hypothetical ancestor of cytheroidean ostracods is presumed to have had an average-shaped shell. Morphological plasticity of the shell outline was observed in many families. The phytal species living on tall seagrass appear to have evolved convergently with species from other habitats, acquiring slim shell outlines during the Cenozoic period. The present analysis also reveals the phylogenetic constraints on the morphological evolution of the Trachyleberididae in their adaptation to a burrowing habit.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 810–821.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Costs of carnivory: tooth fracture in Pleistocene and Recent carnivorans

Large, carnivorous mammals often break their teeth, probably as a result of tooth to bone contact that occurs when carcasses are consumed more fully, a behaviour likely to occur under conditions of food stress. Recent studies of Pleistocene predators revealed high numbers of teeth broken in life, suggesting that carcass utilization and, consequently, food competition was more intense in the past than at present. However, the putative association between diet and tooth fracture frequency was based on a small sample of large, highly carnivorous species. In the present study, a greater diversity of extant carnivorans is sampled, including insectivorous, omnivorous, and carnivorous forms, ranging in size from weasels to tigers. Species that habitually consume hard foods (bones, shells) had the highest fracture frequencies, followed by carnivores, and then insectivorous and/or omnivorous species. Predator and prey sizes were not associated with tooth fracture frequency, but more aggressive species did break their teeth more often. Comparison of the modern sample with five Pleistocene species confirms the previous finding of higher tooth breakage in the past, although some extant species have fracture frequencies that approach those of extinct species. Thus, the Pleistocene predator guild appears to have been characterized by relatively high levels of competition that are rarely observed today.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 68–81.     

Discordance in body size, colour pattern, and advertisement call across genetically distinct populations in a Neotropical anuran (Dendropsophus ebraccatus)

Patterns of intraspecific geographic variation in morphology and behaviour, when examined in a phylogenetic context, can provide insight into the microevolutionary processes driving population divergence and ultimately speciation. In the present study, we quantified behavioural and phenotypic variation among populations from genetically divergent regions in the Central American treefrog, Dendropsophus ebraccatus . Our fine-scale population comparisons demonstrated regional divergence in body size, colour pattern frequencies, and male advertisement call. None of the characters covaried with phylogenetic history or geographic proximity among sampled populations, indicating the importance of highly localized selection pressures and genetic drift in shaping character divergence among isolated regions. The study underscores how multiple phenotypic characters can evolve independently across relatively small spatial scales.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 298–313.     

Speciation on the Azores islands: congruent patterns in shell morphology, genital anatomy, and molecular markers in endemic land snails (Gastropoda, Leptaxinae)

Morphological data, in combination with molecular data, may provide invaluable insights into speciation processes on archipelagos. Land snails offer ample opportunities to evalutate adaptive and non-adaptive speciation scenarios. However, studies investigating processes of differentiation and speciation on the Azores are scarce. The present study comprises a morphometrical analysis of shell and genital characters in a group of Azorean land snails (Pulmonata, Leptaxinae). Geographical isolation appears to be an important mechanism underlying morphological and molecular differentiation in the Azorean Leptaxini, instead of adaptive radiation through ecological differentiation. Nevertheless, we could not exclude the occurrence of ecological speciation on the oldest island (Santa Maria) where two species that markedly differ in shell-shape co-occur.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 166–176.     

Ontogeny of intersexual head shape and prey selection in the pitviper Agkistrodon piscivorus

Different animal intraspecific classes commonly differ in their prey selection. Such differences in feeding ecology are thought to reduce resource competition between classes, but other factors (i.e. behavioural, morphological, and physiological differences) also contribute to this widespread phenomenon. Although several studies have correlated the size of the feeding apparatus with prey selection in many animals, few studies have examined how the shape of the feeding apparatus is related to prey selection. Furthermore, even though the dietary regimen of many animals changes during ontogeny, few studies have examined how shape changes in the feeding apparatus may be related to these ontogenetic dietary shifts. Here we address these issues by examining how head shape, head size and prey selection change over ontogeny in adult males, adult females and juveniles of the cottonmouth snake Agkistrodon piscivorus . Our scaling data for head characteristics showed that all head measurements in adult male and female A. piscivorus scaled with significant negative allometry, whereas juvenile head measurements typically scaled isometrically, except for head volume (positive) and head length (negative). Thus, juveniles have relatively broad and high, but short, heads. Large adult male and female A. piscivorus have relatively small head dimensions overall. Thus, juveniles appear to undergo a rapid change in head volume, which subsequently slows considerably as sexual maturity is achieved. However, our multivariate analysis of size-adjusted head dimensions showed that juveniles differed only slightly in their head shape compared with adult male and female A. piscivorus . In general, prey size increased with snake size across all age and sex groups, but an ontogenetic shift in prey type was not detected in either males or females.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 151–159.     

Leaf teeth in certain Salicaceae and 'Flacourtiaceae'

The leaf teeth glands in four taxa from Salicaceae and six from 'Flacourtiaceae' were examined using both light and scanning electron microscopes. There appears to be a progression from glands of simple structure in the flacourtiaceous taxa and a tendency to a more complicated development in morphology and anatomy of the salicaceous species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 241–256.     

Species' diversity in the New Caledonian endemic genera Cephalidiosus and Nobarnus (Insecta: Heteroptera: Tingidae), an approach using phylogeny and species' distribution modelling

The patterns of local endemism in New Caledonia were analysed in two endemic genera of Tingidae (Insecta, Heteroptera), Cephalidiosus and Nobarnus , through a phylogenetic analysis and species' distribution modelling. The aim was to determine the possible causes of diversification and endemism in New Caledonia. Our results show that environmental conditions are probably important for the distribution of the genus Cephalidiosus , in conjunction with other factors such as resource (host plant) distribution, but suggest that the same environmental conditions have not influenced the speciation processes and diversification in the genus.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 177–184.     

Variation in stickleback head morphology associated with parasite infection

Parasites can affect host phenotypes, influencing their ecology and evolution. Host morphological changes occurring post-infection might result from pathological by-products of infection, or represent adaptations of hosts or parasites. We investigated the morphology of three-spined sticklebacks, Gasterosteus aculeatus , from a population naturally infected with Schistocephalus solidus , which grows to large sizes in their body cavity. We examined local effects of infection on trunk shape, which are imposed directly by the bulk of the growing parasite, and distant effects on head morphology. We show that trunk shape differed between infection classes, and was affected more severely in fish with heavier total parasite mass. We further show unexpected differences in head morphology. The heads of infected fish were reduced in size and differently shaped to those of non-infected fish, with infected fish having deeper heads. Importantly, both head size and shape were also affected more severely in fish with heavier total parasite mass. This latter result suggests that differences in morphology are caused by post-infection changes. Such changes may be incidental, evolutionarily neutral 'side effects' of infection. However, because head morphology affects foraging ecology, such changes are likely to have fitness consequences for hosts, and may constitute adaptations, either of hosts or of parasites. We discuss our finding in the context of the evolution of phenotypic plasticity, and suggest testable hypotheses examining the proximate mechanisms underlying these morphological effects and their potential evolutionary basis.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 759–768.     

Behaviour and stress response during capture and handling of the red-billed chough Pyrrhocorax pyrrhocorax (Aves: Corvidae)

We studied the effect of capture and handling on free-living red-billed choughs Pyrrhocorax pyrrhocorax . We analysed the association between bird behavioural response and individual health, breeding status, sex, and age. Active responses (aggression towards the ringer, screaming) were more common in the breeding period, and in individuals with a high heterophils to lymphocytes ratio, indicating poor physiological condition and/or high chronic stress. Adults were more aggressive than juveniles and yearlings, and females were more aggressive than males. Sex, age, and condition differences were also recorded in the spectrotemporal output of distress calls. Birds with a screaming/active response appeared to be more stress-susceptible than passive and silent individuals, and this response was stronger during the energy demanding period of reproduction. The results obtained suggest that the response of the red-billed chough during capture might primarily reflect stress-susceptibility, although a number of potential alternative explanations are discussed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 846–855.     

Galápagos' Opuntia (prickly pear) cacti: extensive morphological diversity, low genetic variability

Due to the pronounced morphological variation and geographical distribution of Galápagos' Opuntia cacti, numerous hypotheses have been advanced regarding their radiation, diversification, and classification. The currently accepted classification is based on morphology and recognizes six species and fourteen varieties, but the plasticity of many of the characteristics renders any morphological taxonomy problematic. Our analysis of previously published morphological data agrees only partially with the current classification. We present the first molecular phylogeny of these plants. Multiple DNA sequences indicate little genetic distinction among the currently identified species, despite restricted gene flow and limited long distance dispersal within the archipelago. No clear relationship exists between morphological and genetic differences. These results suggest that both molecular and morphological data should be used in conservation planning.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 451–461.     

Morphological correlates of ant eating in horned lizards (Phrynosoma)

The North American horned lizards ( Phrynosoma ) represent a morphologically specialized group of ant-eating lizards. Although variation in dietary fidelity is observed among the species, all appear to possess morphological specializations thought to be related to their ant-eating diets. Previous studies have examined morphological specialization in Phrynosoma , but they have not taken into account the phylogenetic relationships of its member species. In the present study, the morphological characteristics of the head, jaws and teeth that are thought to be important in prey capture and prey processing were examined to test whether variation in cranial morphology is associated with diet in lizards of the genus Phrynosoma . It is suggested that lizards of the genus Phrynosoma are indeed morphologically specialized and that ant-eating is associated with reduced dentition and an overall reduction in the robustness of morphological structures important in prey processing. Although this trend holds for the highly myrmecophagous species of Phrynosoma , a robust cranial morphology is apparent in the short-horned lizard clade ( Phrynosoma ditmarsi , Phrynosoma douglasii , Phrynosoma hernandesi , Phrynosoma orbiculare ), implying the ability to process a variety of dietary items. The present study suggests that additional feeding specializations exist within an already specialized clade (i.e. the short-horned lizard clade) and highlights the need for more detailed dietary and behavioural studies of feeding behaviour in this uniquely specialized group of lizards.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 13–24.     

Biological and environmental degradation of gorilla hair and microsatellite amplification success

Naturally shed hairs are an important source of genetic material for both conservation and forensics but are notoriously poor sources of DNA. DNA degradation in hair roots is caused by apoptosis as part of the cycle of hair growth and by autolysis in decomposing animals. Shed hairs are additionally exposed to degenerative environmental processes. However, genetic studies rarely examine hair root morphologies or refer to root growth phases prior to analysis, and detailed knowledge of the rapidity of DNA degradation amongst shed hairs is lacking. We examined the effects of biological and environmental processes on western lowland gorilla ( Gorilla gorilla gorilla Savage and Wyman) hair roots with respect to morphological characteristics and polymerase chain reaction (PCR) success at eight nuclear loci. Root type frequencies indicate that gorilla body hairs may exhibit a longer telogen phase than human head hairs. All plucked hair root types amplified more efficiently than shed hairs, and only 41% of shed hairs had root types considered suitable for genotyping. Telogen hairs from fresh nests were four-fold more useful for genotyping if the roots were associated with translucent epithelial tissue, and preselection of these root types doubled the overall data-yield to 58%. Nest age correlated with root morphology and PCR success, and PCR success was almost halved after 3 days of exposure. Finally, an association between postmortem interval, root morphology, and PCR success was observed that was consistent with postmortem changes reported in human head hairs.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 281–294.     

Changes in ventral head width, a discriminating shape factor among African cichlids, can be induced by chronic hypoxia

A massive enlargement of the gill surface proved to be an important factor in the hypoxia survival of young cichlids. Because the heads of cichlids are densely packed with structures related to both feeding and breathing, we hypothesized that the extra space needed for gill enlargement requires such large structural reorganizations that outer head shape is affected. We used a three-dimensional model to describe changes in the outer head shape of cichlids. Broods of cichlids of different phylogenetic lineages, habitats, and trophic specialization were split and raised at either 10% or 80–90% air saturation. Despite the above-mentioned differences between the species that were used, all hypoxia raised groups showed similar volume enlargements. Volume increases were most prominent in the ventral suspensorial and ventral opercular subcompartments. A relation with the enlarged gills of hypoxia raised fish is likely because the gills are mainly located in these compartments. The differences in ventral width correspond to those found in other studies comprising a wide variety of genotypic and phenotypic variations. The present study shows that such variation in the ventral width is conceivable by phenotypic plasticity alone.   © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 608–619.     

New evolutionary evidence of Grylloblattida from the Middle Jurassic of Inner Mongolia, north-east China (Insecta, Polyneoptera)

The small modern insect order Grylloblattida has an abundant fossil record during the Late Palaeozoic and the Mesozoicirca. The relationships between these fossil taxa and the modern grylloblattids remain unclear because most of them are based on isolated wings or have poorly preserved body features. Modern grylloblattids are wingless insects. The new grylloblattid family Plesioblattogryllidae fam. nov. is erected for the new genus and species Plesioblattogryllus magnificus gen. nov., sp. nov. , from the Middle Jurassic of north-eastern China. The well-preserved specimen provides further evidence that could support its close relationships with the modern grylloblattids: (1) several very similar head structures, e.g. developed laciniae with inner row of setae, maxillary palps segmented into five, labial palps segmented into three, large labrum, and morphology of antenna; (2) paired eoplantulae on tarsomeres 1–4; (3) long ovipositor and large eggs comparable with those of modern taxa. The new genus has strongly developed mandibles with sharp pointed apical teeth and strong marginal teeth, and strong hook-like fore claws with basal teeth, suggesting it was carnivorous. The major differences between the extinct and extant Grylloblattida, such as the lack of wings, the eyes and ocelli either degenerated or absent, and the thorax degenerated in the modern forms, are probably related to their adaptation to their life under rocks and rock-crawler habits.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 17–24.     

Using image analysis on the ventral colour pattern in Salamandrina perspicillata (Amphibia: Salamandridae) to discriminate among populations

The ventral colour pattern in Salamandrina perspicillata is characteristic of each individual and remains invariant throughout the salamanders' life. The present study aimed to determine, using statistical methods applied to quantitative image analysis, whether the coloration pattern is population-specific. The images of ventral colour pattern of 180 salamanders belonging to five populations were analysed. The images were previously warped by geometric morphometry. To our knowledge, this is the first study to use an approach based on geometric morphometrics to standardize ventral shapes. This technique is useful for eliminating form distortions. The results of an analysis of similarities recognized more coloration similarity among individuals of the same population than among individuals from different populations. Analysis by partial least squares correctly classified the 88.88% of individuals into the correct population. The similarities or differences among individuals of different populations are not related to the geographical distances. The results obtained showed that the coloration of the ventral side of the head of the salamanders can be used for discriminating among populations.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 35–43.     

Convergent body flattening in a clade of tropical rock-using lizards (Scincidae: Lygosominae)

The repeated occurrence of similar morphologies in organisms from similar habitats provides good evidence of convergent selection, and convergent patterns of evolutionary change. In lizards, a flattened morphology has often been noted; however, whether this trait is convergent in specific habitats has never been tested using phylogenetic methods. The present study examined patterns of morphological convergence in 18 species of tropical Lygosomine skinks from three broad habitat categories (generalist, leaf litter-dwelling, and rock-using species). In general, although there where relatively few morphological differences of species from different habitats, phylogenetic analyses revealed that rock-using species have consistently and repeatedly evolved a dorsoventrally flattened head and body. The adaptive basis of this flattened morphology is consistent with both biomechanical predictions of performance (e.g. climbing locomotion) and ecology (e.g. use of rock crevices, camouflage) of species that occupy rocky habitats.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 399–411.     

The relationship between limb morphology, kinematics, and force during running: the evolution of locomotor dynamics in lizards

Terrestrial locomotion occurs via the hierarchical links between morphology, kinematics, force, and center-of-mass mechanics. In a phylogenetically broad sample of seven lizard species, we show that morphological variation drives kinematic variation, which, in turn, drives force variation. Species with short limbs use a short stride–high frequency strategy when running at steady-speed and to change speeds. This link between morphology and kinematics results in relatively small vertical forces during the support phase of the stride cycle. Conversely, species with long limbs use a long stride–low frequency strategy, resulting in large vertical forces during the support phase. In view of these findings, we suggest that limb length may predict locomotor energetics in lizards because energetics are largely determined by vertical forces and stride frequency. Additionally, we propose an energetic trade-off with both long- and short-limbed species paying the most energy to move, whereas intermediate-limbed species move using less energy. Finally, when these traits are mapped onto a lizard phylogeny, we show that locomotor functional morphology exhibits both deep phylogenetic effects and contemporary patterns of evolutionary convergence. Overall, the present study provides a foundation for testing hypotheses regarding the integration and evolution of functional traits in lizards and animals in general.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 634–651.     

The evolution of venom-delivery systems in snakes

The Colubroidea represents approximately 2300 of the 2700 species of living snakes and includes all venomous taxa. Although many morphological studies of colubroid snakes have been carried over the last hundred years, the phylogenetic relationships within this group are poorly known. In this study, components of the venom-delivery system (VDS) were examined within the context of two conflicting phylogenetic hypotheses proposed in 1988 by Cadle and in 1998 by Kraus & Braun. The results suggest that several major morphological changes occurred early in colubroid evolution: a Duvernoy's gland evolved, the posterior maxillary teeth became specialized relative to the anterior maxillary teeth, and the attachment of the pterygoideus muscle moved forward to a position associated with the posterior maxillary teeth. These innovations may have allowed the great radiation of colubroid snakes that led to the Colubroidea representing such a large percentage of living snakes. More recently, three separate lineages of colubroids have independently evolved highly specialized front-fanged VDSs with large and complex venom glands, venom gland compressor muscles, and tubular fangs.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 337−354.