Sex-specific ecomorphological variation and the evolution of sexual dimorphism in dwarf chameleons (Bradypodion spp.)

Natural selection can influence the evolution of sexual dimorphism through selection for sex-specific ecomorphological adaptations. The role of natural selection in the evolution of sexual dimorphism, however, has received much less attention than that of sexual selection. We examined the relationship between habitat structure and both male and female morphology, and sexual dimorphism in size and shape, across 21 populations of dwarf chameleon (genus Bradypodion). Morphological variation in dwarf chameleons was strongly associated with quantitative, multivariate aspects of habitat structure and, in most cases, relationships were congruent between the sexes. However, we also found consistent relationships between habitat and sexual dimorphism. These resulted from both differences in magnitude of ecomorphological relationships that were otherwise congruent between the sexes, as well as in sex-specific ecomorphological adaptations. Our study provides evidence that natural selection plays an important role in the evolution of sexual dimorphism.     

Morphological variation in the Cape Dwarf Chameleon (Bradypodion pumilum) as a consequence of spatially explicit habitat structure differences

An organism's phenotype is to some extent influenced by costs and benefits in terms of natural and sexual selection. The intensity of natural selection can in part be driven by habitat structure, which may result in varying levels of crypsis and/or selection on traits related to maximizing performance in that habitat. This may be countered by sexual selection, which can lead to sexual dimorphism in body size and/or the expression of conspicuous ornamentation relating to maximizing reproductive success. The intensity of these forces can also be different between the sexes, resulting in complex patterns of phenotypic variation. With this in mind, we examined morphological variation within the Cape Dwarf Chameleon, Bradypodion pumilum. The species inhabits two geographically disjunct habitat types and, in the present study, we demonstrate that chameleons from the two habitats show morphological differences. Large, conspicuous individuals inhabit closed vegetation, whereas small, drab individuals inhabit open vegetation. However, when morphological traits are size‐adjusted, the open vegetation morph displays many traits that are larger for its body size than the closed vegetation morph, especially for characters related to locomotion (limbs) and bite force (head width). Sexual dimorphism is also present, although the degree and number of dimorphic characters was very different between the two morphs, with size‐adjusted male‐biased dimorphism much more pronounced in the closed morph. Overall, our findings suggest that natural selection in open habitats limits both body size and conspicuous characters, although sexual selection in closed habitats favours the development of ornamentation related to display. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 878–888.     

Phylogenetics of the southern African dwarf chameleons, Bradypodion (Squamata: Chamaeleonidae)

The taxonomic relationships within the dwarf chameleons (Bradypodion) of southern Africa have long been controversial. Although informal phenotypic groups have been suggested, the evolutionary relationships among the 15 recognised species in southern Africa have not been previously investigated. To investigate the relationships among species within this genus, fragments of two mitochondrial genes (16S ribosomal RNA and ND2) were sequenced and analysed using maximum parsimony, maximum likelihood and Bayesian inference. All analyses showed congruent topologies, revealing at least 5 well-supported clades distributed across distinct geographic regions. The mtDNA gene tree indicated that in many instances, geographic location has played a role in shaping the evolution of this group, and that the previously suggested phenotypic groupings do not adequately reflect evolutionary relationships. Furthermore, it appears that some of the currently recognised species (described on morphology) are polyphyletic for mitochondrial sequences, most notably those occurring in the isolated forest patches of north-eastern South Africa, near the Drakensberg Escarpment.     

Biogeographic patterns and phylogeography of dwarf chameleons (Bradypodion) in an African biodiversity hotspot

The southern African landscape appears to have experienced frequent shifts in vegetation associated with climatic change through the mid-Miocene and Plio-Pleistocene. One group whose historical biogeography may have been affected by these fluctuations are the dwarf chameleons (Bradypodion), due to their associations with distinct vegetation types. Thus, this group provides an opportunity to investigate historical biogeography in light of climatic fluctuations. A total of 138 dwarf chameleons from the Cape Floristic Region of South Africa were sequenced for two mitochondrial genes (ND2 and 16S), and resulting phylogenetic analyses showed two well-supported clades that are distributed allopatrically. Within clades, diversity among some lineages was low, and haplotype networks showed patterns of reticulate evolution and incomplete lineage sorting, suggesting relatively recent origins for some of these lineages. A dispersal-vicariance analysis and a relaxed Bayesian clock suggest that vicariance between the two main clades occurred in the mid-Miocene, and that both dispersal and vicariance have played a role in shaping present-day distributions. These analyses also suggest that the most recent series of lineage diversification events probably occurred within the last 3-6 million years. This suggests that the origins of many present-day lineages were founded in the Plio-Pleistocene, a time period that corresponds to the reduction of forests in the region and the establishment of the fynbos biome.     

The relationship between cranial morphology,bite performance,diet and habitat in a radiation of dwarf chameleon (Bradypodion)

Many animals show unique morphological and behavioural adaptations to specific habitats. In particular, variation in cranial morphology is known to influence feeding performance, which in turn influences dietary habits and, ultimately, fitness. Dietary separation is an important means of partitioning ecological niches and avoiding inter‐ and intraspecific competition. Consequently, differences in dietary resources may help explain phenotypic divergence in closely‐related species occupying different habitats, as well as sexual dimorphism. We test this hypothesis on five phenotypic forms of a recent radiation of dwarf chameleons (Bradypodion) that vary extensively in habitat use and cranial morphology. By examining stomach contents, the dietary composition of each phenotypic form is compared to investigate potential differences in feeding strategies. Overall, chameleons in the present study exhibit considerable dietary overlap (at both inter‐ and intraspecific levels), indicating that diet is not a major driver of variation in cranial morphology within this radiation. However, the stomachs of closed‐canopy females were found to contain more prey items than male stomachs, possibly indicating that females require a greater caloric intake than their male counterparts.     

Population variation in second metacarpal sexual size dimorphism

This paper contrasts levels of sexual size dimorphism in second metacarpal osteometric and geometric morphology in two bioculturally distinctive populations: 19th century Euro-Canadian settlers, and proto/historic central Canadian Inuit. Significant within-group sexual size dimorphism is found for all variables, though few show significant interpopulation differences. However, in every case the Euro-Canadian sample is more dimorphic than the Inuit sample. Notably, differences reside in geometric measures (total area, Imax) sensitive to variation in functional strain, and thus are interpretable in light of proximate causal models, i.e., activity profiles distinct from generalized mode of subsistence. Other proximate factors, such as nutritional stress acting to diminish Inuit sexual size dimorphism, may also play a role. However, models often cited to explain dimorphism, such as marriage practice (e.g., polygyny) or division of labor situated in mode of subsistence, do not. The higher sexual size dimorphism in the 19th century settler sample belies the notion that technological progress inevitably leads to reduced dimorphism.     

Mandibular ramus flexure: A new morphologic indicator of sexual dimorphism in the human skeleton

In the skeleton, male and female characteristics lie along a continuum of morphologic configurations and metric values. Size alone is not the best indicator of sex. In contrast, morphologic differences that arise from genetically sex-linked growth and development allow better separation of the sexes. This study presents a new morphologic indicator of sexual dimorphism in the human mandible. A sample of 300 mandibles from adults of known sex primarily from the Dart collection was analyzed. Of these, 100 were found to have obvious bony pathologies and/or excessive tooth loss (“pathologic” sample). Thus, the normative sample consisted of 200 individuals (116 males, 84 females). Examination of morphologic features led to the discovery of a distinct angulation of the posterior border of the mandibular ramus at the level of the occlusal surface of the molars in adult males. Flexure appears to be a male developmental trait because it is only manifest consistently after adolescence. In most females, the posterior border of the ramus retained the straight juvenile shape. If flexure was noted, it was found to occur either at a higher point near the neck of the condyle or lower in association with gonial prominence or eversion. In the normative sample, overall prediction accuracy from ramus shape was 99%. When the “pathologic” sample was analyzed separately, 91.0% were correctly diagnosed. Because the African samples were overwhelmingly black, this trait was also tested on American samples (N = 247) of whites (N = 85), Amerinds (N = 66), and blacks (N = 96) that included a mix of healthy individuals and those with extensive tooth loss and evidence of pathology. The results were nearly identical to those of the “pathologic” African sample, with accuracies ranging from about 91% in whites and blacks to over 92% in Amerinds. Total accuracy for all African and American samples combined (N = 547) is 94.2%. In conclusion, at 99%, sexing from the shape of the ramus of a healthy mandible is on a par with accuracy attainable from a complete pelvis. Moreover, there is no record that any other single morphologic or metric indicator of sex (that has been quantified from the adult skeleton) surpasses the overall accuracy attained from the more representative mixed sample produced by combining all groups assessed in this study. The usefulness of this trait is enhanced by the survivability of the mandible and the fact that preliminary investigations show that the trait is clearly evident in fossil hominids. © 1996 Wiley-Liss, Inc.     

Ecomorphological variation in male and female wall lizards and the macroevolution of sexual dimorphism in relation to habitat use

Understanding how phenotypic diversity evolves is a major interest of evolutionary biology. Habitat use is an important factor in the evolution of phenotypic diversity of many animal species. Interestingly, male and female phenotypes have been frequently shown to respond differently to environmental variation. At the macroevolutionary level, this difference between the sexes is frequently analysed using phylogenetic comparative tools to assess variation in sexual dimorphism (SD) across taxa in relation to habitat. A shortcoming of such analyses is that they evaluate the degree of dimorphism itself and therefore they do not provide access to the evolutionary trajectories of each sex. As such, the relative contribution of male and female phenotypes on macroevolutionary patterns of sexual dimorphism cannot be directly assessed. Here, we investigate how habitat use shapes phenotypic diversity in wall lizards using phylogenetic comparative tools to simultaneously assess the tempo and mode of evolution in males, females and the degree of sexual dimorphism. We find that both sexes have globally diversified under similar, but not identical, processes, where habitat use seems to drive macroevolutionary variation in head shape, but not in body size or relative limb length. However, we also observe small differences in the evolutionary dynamics of male and female phenotypes that have a marked impact on macroevolutionary patterns of SD, with important implications for our interpretation of what drives phenotypic diversification within and between the sexes.     

Macroevolutionary pattern of sexual size dimorphism in geckos corresponds to intraspecific temperature‐induced variation

Many animal lineages exhibit allometry in sexual size dimorphism (SSD), known as ‘Rensch’s rule’. When applied to the interspecific level, this rule states that males are more evolutionary plastic in body size than females and that male‐biased SSD increases with body size. One of the explanations for the occurrence of Rensch’s rule is the differential‐plasticity hypothesis assuming that higher evolutionary plasticity in males is a consequence of larger sensitivity of male growth to environmental cues. We have confirmed the pattern consistent with Rensch’s rule among species of the gecko genus Paroedura and followed the ontogeny of SSD at three constant temperatures in a male‐larger species (Paroedura picta). In this species, males exhibited larger temperature‐induced phenotypic plasticity in final body size than females, and body size and SSD correlated across temperatures. This result supports the differential‐plasticity hypothesis and points to the role phenotypic plasticity plays in generating of evolutionary novelties.     

Morphological variation and sexual dimorphism of the cephalic scales in Lacerta bilineata

The Western green lizard (Lacerta bilineata) is a lacertid distributed throughout Mediterranean and Central Europe. Little is known about the morphological variability and sexual shape differences in this species. In this paper, the variation of the cephalic scales in L. bilineata is analysed by means of a geometric morphometric approach. A main structural pattern is characterized by negative allometry of the frontal and interparietal areas, and positive allometry of the parietal and frontoparietal scales. Sexual differences are described both in size and shape. In males, the scales are generally larger, with relative shortening of the frontal area, frontoparietal enlargement, occipital lengthening and bulging of the parietal scales, which compresses and narrows the interparietal and occipital areas midsagittally. This pattern is based on a shared allometric trajectory, with males displaying a peramorphic morphotype. However, males show some shape differences in the occipital area that are not size‐related, and cannot be interpreted in terms of general head enlargement. This structural trajectory can be related to the development of the skull, but the role of soft tissues (temporal, nuchal and masticatory muscles) must also be considered. The development of the jaw and nuchal muscles involved in intra‐ and intersexual behaviours could have played a pivotal role in the evolution of this pattern.     

Ultrastructure of the peduncular aesthetascs of Speleonectes tanumekes (Crustacea, Remipedia) in comparison with crustaceans from stressful environments

Abstract. Osmotic stress associated with the freshwater environment and desiccation stress associated with the terrestrial environment may have a shortening effect on the length of the innervation of crustacean aesthetascs. Physical stress of the littoral environment may have a similar effect on the length of the cuticular portion of aesthetascs. The aesthetascs of crustaceans that inhabit these environments share a similar ultrastructural feature, which may help animals cope with these environmental stresses. This ultrastructural feature, the position of the basal bodies proximal to the lumen of the aesthetasc, is absent from the aesthetascs of crustaceans that occur in the typical marine environment. Interestingly, the ultrastructural feature associated with these stressful habitats is present in the peduncular aesthetascs of the remipede Speleonectes tanumekes , even though the environmental stresses that may invoke the reduction of aesthetascs are absent in the marine-cave environment where this animal occurs. The importance of the sensitivity of aesthetascs for survival in this lightless environment may result in a selective pressure that favors basal bodies to be positioned proximal to the lumen of the aesthetasc.     

A role for ecology in the evolution of colour variation and sexual dimorphism in Hawaiian damselflies

Variation in traits that are sexually dimorphic is usually attributed to sexual selection, in part because the influence of ecological differences between sexes can be difficult to identify. Sex‐limited dimorphisms, however, provide an opportunity to test ecological selection disentangled from reproductive differences between the sexes. Here, we test the hypothesis that ecological differences play a role in the evolution of body colour variation within and between sexes in a radiation of endemic Hawaiian damselflies. We analysed 17 Megalagrion damselflies species in a phylogenetic linear regression, including three newly discovered cases of species with female‐limited dimorphism. We find that rapid colour evolution during the radiation has resulted in no phylogenetic signal for most colour and habitat traits. However, a single ecological variable, exposure to solar radiation (as measured by canopy cover) significantly predicts body colour variation within sexes (female‐limited dimorphism), between sexes (sexual dimorphism), and among populations and species. Surprisingly, the degree of sexual dimorphism in body colour is also positively correlated with the degree of habitat differences between sexes. Specifically, redder colouration is associated with more exposure to solar radiation, both within and between species. We discuss potential functions of the pigmentation, including antioxidant properties that would explain the association with light (specifically UV) exposure, and consider alternative mechanisms that may drive these patterns of sexual dimorphism and colour variation.     

Ecomorphological correlates in tropical stream fishes of southern India

Synopsis I studied ecomorphological correlations in 18 stream fish species (belonging to five families) commonly occurring in diversity rich habitats of rivers in the central Western Ghats (Southern India). Univariate and multivariate analyses indicated a clear segregation of species in morphological and ecological (habitat) space. Cluster analyses using morphological features segregated species based mainly on their body shape and feeding parameters while habitat parameters segregated species according to their position along the water column and preference to runs or pool habitats. Principal component analyses showed that body size, head dimensions, mouth position and eye diameter were important characters for species segregation in morphological space while run or pool habitat type and the position of the fish in the water column best explained segregation of species in habitat space. Consistencies in some of the species clusters obtained from analyses on morphological characteristics and habitat preferences warrant closer examination of the relationships between them. These were studied employing multivariate methods like canonical correspondence analysis and the Mantel test. The tests revealed weak correlations indicating that in addition to morphology, other factors like behavioural, physiological and evolutionary constraints together are likely to play an important role in ecosystem structuring of these complex tropical assemblages.     

Four New Species of Litter Inhabiting Scytodes Spiders (Araneae,Scytodidae) from Amazonia

Four new species of spitting spiders of the genus Scytodes are described: S. panguana from Peruvian Amazonia and S. paarmanni, S. martiusi and S. janauari from central Amazonia in Brazil. All are small, short legged scytodids. The specimens were collected in the course of ecological studies in the leaf litter layer on the ground of Amazonian terra firme forests. Quatro novas espécies de Scytode s são descritas: S. panguana da Amazônia peruana e S. paarmanni, S. martiusi e S. janauari da Amazônia central do Brasil. Todas são espécies pequenas, de pernas curtas, que foram coletadas durante estudos ecológicos em liteira no chão de florestas de terra firme da Amazônia.     

Ecomorphometric variation and sexual dimorphism in the common shrew (Sorex araneus)

We investigated the evolution of the biomechanics of the mandible in island and mainland populations of the common shrew on the west coast of Scotland. We predicted that climatic differences between populations should cause differences in prey composition leading to changes in the mechanical potential (MP) of the mandible. In females, MP was correlated with climate, with greater MP in warmer and drier habitats. In males, MP was significantly greater than in females but there was no relationship between male MP and climate. This led to increased sexual dimorphism in colder and wetter climates. The same pattern was found after a phylogenetic least squares analysis was conducted to account for shared phylogenetic history. We discuss possible reasons for this pattern, including male–male combat and the greater necessity of females to feed as efficiently as possible to meet their extremely high energy requirements during lactation.     

A new phylogenetic classification for the gymnophthalmid genera Cercosaura , Pantodactylus and Prionodactylus (Reptilia: Squamata)

Because of the poor state of knowledge of many of the gymnophthalmid genera, systematic revision is necessary to render the classification consistent with evolutionary history. To that end, I conducted a review of the species of three genera of the Cercosaurinae which appear to form a monophyletic group: Cercosaura , Pantodactylus , and Prionodactylus . Phylogenetic analysis of 61 morphological characters was conducted after specimens of all species were examined to evaluate the composition of each taxon. The phylogenetic reconstruction suggested that the genus Prionodactylus was paraphyletic. A new phylogenetic classification is proposed that synonymizes Pantodactylus and Prionodactylus with Cercosaura. Cercosaura is redefined to include 11 species and seven subspecies. A key is provided to distinguish among species.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 101−115.     

Sexual dimorphism in lizard body shape: the roles of sexual selection and fecundity selection

Sexual dimorphism is widespread in lizards, with the most consistently dimorphic traits being head size (males have larger heads) and trunk length (the distance between the front and hind legs is greater in females). These dimorphisms have generally been interpreted as follows: (1) large heads in males evolve through male-male rivalry (sexual selection); and (2) larger interlimb lengths in females provide space for more eggs (fecundity selection). In an Australian lizard (the snow skink, Niveoscincus microlepidotus), we found no evidence for ongoing selection on head size. Trunk length, however, was under positive fecundity selection in females and under negative sexual selection in males. Thus, fecundity selection and sexual selection work in concert to drive the evolution of sexual dimorphism in trunk length in snow skinks.     

Development and sexual dimorphism of branchiae in Streblospio benedicti

The Spionidae is one of the largest and most studied annelid families, but to date, the development and differentiation of post-metamorphic anatomy have not been documented. This study used scanning electron microscopy to examine the development of the branchiae, presumed respiratory organs, in Streblospio benedicti. Branchiae in this species are prominent, paired head structures and first appear around the time of metamorphosis, but do not complete their development until the worm reaches the older juvenile or adult stages. We observed that as the branchiae grew, their overall morphology changed through four different shapes: small bud, tubular, tapered, and, finally, bilimbate. In addition, the abfrontal and frontal surfaces each possessed a unique set of cilia patterns, which we named, and these arose in a particular sequence between the 8- and 35-chaetiger stages. This detailed examination of every stage of branchial development led us to discover that branchia in Streblospio benedicti was a sexually dimorphic organ. Streblospio benedicti is one of approximately eight Spionidae in which there is any type of structural sexual dimorphism, and it is the only species in which sexually dimorphic branchiae are found. The male's frontal surface had four unique cilia patterns, and we hypothesize that those located around the medial protrusion capture and control the release of the spermatophores. This first documentation of a spionid's branchial developmental sequence revealed that not only is this respiratory organ involved in reproduction, but it significantly differentiates after metamorphosis through adulthood.