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.     

Multiple scale patterns of shell and anatomy variability in land snails: the case of the Sicilian Marmorana (Gastropoda: Pulmonata, Helicidae)

Certain major aspects of phenotypic diversity are still largely unexplained. When phenotypic patterns do not relate to habitat variables, fine analysis of morphological patterns and their distribution sheds light on the origin of diversity. Among invertebrates, snails are an ideal model for studying the roles of the neutral processes and selection involved in creating diversity. To understand patterns and processes of variability on different scales (regional: areas; local: sites), morphological variability of two sets of characters (shell and genitalia) was quantified in a group of rock-dwelling land snails of the genus Marmorana (Pulmonata, Helicidae). To analyse shell variability, partitioning of the overall variation into size and shape components was analysed by a principal component-based approach. Shell shape and size variability is not significantly influenced by any environmental pressure. Variability at site scale is mainly attributed to shell size, which is a trait demonstrated to have a high degree of phenotypic plasticity. No sharp changes were observed for genitalia. Moreover, allometries between shell size and genitalia measurements involve a few populations. The observed multiple scale patterns are in line with the hypothesis that genital variance may be selectively controlled to maintain function.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 359–370.     

Mosaic pattern of genital divergence in three populations of Schistocerca lineata Scudder, 1899 (Orthoptera: Acrididae: Cyrtacanthacridinae)

Sexual selection theory predicts that genital structures in isolated populations are likely to diverge, but male genitalia are often species-specific, which led to the idea that male genitalia are relatively invariable within species. Previous allometric studies collectively suggested that male genitalia are intraspecifically invariable in size compared with external body parts. We investigated whether male genitalia are invariable in shape in three populations of a grasshopper Schistocerca lineata Scudder, 1899, using two independent methods of geometric morphometric analyses. Specifically, we focused on the idea that male genitalia are complex structures consisting of many functionally different components, and studied how these individual parts diverge among three populations. Individual components of male genitalia show different population-level divergence, resulting in the mosaic pattern of genital divergence. Individual components diverge independently from each other. Body size is positively correlated with genitalia size, but is significantly correlated with the shape of only one of the three genital structures we measured. Thus, different components of male genitalia may be influenced by different evolutionary processes. This study is the first to show that components of complex genitalia evolve separately within a species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 289–301.     

Covariation between predation risk, body size and fin elaboration in the green swordtail, Xiphophorus helleri

Natural and sexual selection can have either opposing or synergistic effects on the evolution of traits. In the green swordtail Xiphophorus helleri , sexual selection arising from female choice is known to favour larger males and males with longer swords. We examined variation in male and female size and fin morphology among 15 populations that varied in their predation environments. Males and females from populations in which piscivorous fishes were present had longer and deeper bodies than did males and females from populations in which piscivorous fishes were absent. Controlling for a positive effect of body size on sword length, males from populations in which piscivores were present had relatively shorter swords than did males from populations in which piscivores were absent. The associations between morphology and predation environment may be due to direct effects of predation, indirect effects of predation, other sources of selection that covary with predator presence, or other environmental effects on trait expression. These results suggest that while sexual selection favours longer swords, natural selection may have an opposing effect on sword length in populations with predators. Natural selection on body size, however, may act synergistically with sexual selection in populations with predators; both may favour the evolution of larger body size. The body size results for X. helleri contrast with related taxa that have become model systems for the study of life history evolution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 87–100.     

Evolutionary diversification of opercle shape in Cook Inlet threespine stickleback

We investigated the evolution of a large facial bone, the opercle (OP), in lake populations of the threespine stickleback that were founded by anadromous ancestors, in Cook Inlet, Alaska. Recent studies characterized OP variation among marine and lake populations and mapped a quantitative trait locus with a large influence on OP shape. Using populations from diverse environments and independent evolutionary histories, we examined divergence of OP shape from that of the anadromous ancestor. We report preliminary evidence for divergence between benthic and generalist lake ecotypes, necessitating further investigation. Furthermore, rapid divergence of OP shape has occurred in a lake population that was founded by anadromous stickleback in the 1980s, which is consistent with divergence of other phenotypic traits and with OP diversification in other lake populations. By contrast, there has been limited evolution of OP shape in a second lake population that may have experienced a genetic bottleneck early in its history and lacks genetic variation for OP divergence. Taken together, the results obtained from these two populations are consistent with studies of other stickleback phenotypic traits that implicate ancestral variation in postglacial adaptive radiation of threespine stickleback in fresh water.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 832–844.     

The evolution of locomotor morphology, performance, and anti-predator behaviour among populations of Leiocephalus lizards from the Dominican Republic

Most ecomorphological studies use a comparative approach to examine adaptation by studying variation among species. A question of considerable interest is whether ecomorphological patterns observed among species also exist at the population level. We studied variation in morphology, performance, and behaviour in four populations of Leiocephalus personatus and two populations of Leiocephalus barahonensis in the Dominican Republic. We combined these data with measurements of predation intensity and habitat structure to test for convergence at the population level. We predicted that predation intensity would be higher in open habitats and that lizards in these habitats would have traits conferring higher predator evasion capacity (increased wariness, faster sprint speeds, and longer limbs). Principal components analysis suggests that sites tend to differ with respect to the abundance and spacing of low-lying vegetation (i.e. percentage of shrub cover and distance to nearest vegetation), but we did not detect any striking differences among sites in tail-break frequencies or attacks on clay lizard models. Consistent with predictions we find that in open habitats, lizards tend to have longer limbs, faster sprint speeds (relative to body size), and longer approach distances. These patterns corroborate findings in other ground-dwelling lizard species and indicate that they have evolved at least twice among populations of Leiocephalus lizards. The results of this study also suggest that these traits have evolved rapidly despite recent or ongoing gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 445–456.     

Variation in the digging apparatus of the subterranean silvery mole-rat, Heliophobius argenteocinereus (Rodentia, Bathyergidae): the role of ecology and geography

The skull of most subterranean tooth-digging rodents is markedly affected by their digging mode. In the present study, we investigated the cranial variation in a strictly subterranean, highly specialized Afrotropical tooth-digger, Heliophobius argenteocinereus (Bathyergidae, Rodentia), using a geometric morphometric approach and evaluated the effect of different factors on size and shape differences among four populations. No evidence for sexual dimorphism was found in skull size or shape. The cranial shape variation was large and influenced mainly by the type of habitat (miombo woodland versus farmland and grassland) and the latitudinal gradient. The dorsal side of the skull appears to be more plastic and adaptable to local environments, as well as more independent of size, than the ventral side. Only the shortening of the rostrum is presumably an adaptive process independent of size that leads to an increase of efficacy of the tooth-digging apparatus in Heliophobius , whereas the increase in the in-force and the more procumbent incisors both comprise size-related changes caused by ontogenetic allometric growth.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 822–831.     

Evolution of life history traits in Leporidae: a test of nest predation and seasonality hypotheses

Recently, it has been proposed that adult mortality schedules may be responsible for latitudinal patterns of life history variation in passerine birds, whereas nest predation only could explain within latitude patterns. Unfortunately, no independent test has been performed regarding the importance of nest predation with different taxa. In the present study, seasonality and nest predation hypotheses explaining variations in gestation time and litter size in 17 lagomorph species were tested. Among latitude patterns were analysed using the phylogenetic independent contrast method of Felsenstein and within latitude patterns were analysed by the pairwise comparative method. The results obtained indicate that latitudinal patterns observed in both variables are explained by different factors: seasonality for litter size and nest predation for gestation time. Litter size variations within latitudes are also explained by differences in nest predation, supporting previous hypotheses. In conclusion, the present study suggests that, when compared among latitudes, different life history traits (e.g. litter size and gestation time) may be shaped by different selective forces and that the effects of nest predation may be high both within and between latitudes.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 603–610.     

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.     

Hierarchical comparative analysis of genetic and genitalic geographical structure: testing patterns of male and female genital evolution in the scarab beetle Phyllophaga hirticula (Coleoptera: Scarabaeidae)

It is generally accepted that genitalia are among the fastest evolving characters in insects and that selection on these structures may increase speciation rates in groups with polygamous mating systems. If selection is causing genitalic divergence between or among populations of a species, one prediction is that geographical structure of genitalic morphology would be in place before genetic structure of a rapidly evolving neutral marker. The current study tests this hypothesis in the geographically widespread scarab beetle Phyllophaga hirticula by evaluating whether standing variation in male and female genitalia is more or less geographically structured than a mitochondrial genetic marker. Geographical structure of mitochondrial (mt)DNA and male and female genitalic shape were analysed using analysis of variance, multivariate analysis of variance, Mantel tests, and tests of spatial autocorrelation. The results show that, although female genitalia are more geographically structured than mtDNA, male genitalia are not. This pattern suggests that selection on female genitalic variation may be causing divergence of these structures among populations.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 135–149.     

Size and shape interspecific divergence patterns partly reflect phylogeny in an Onthophagus species‐complex (Coleoptera: Scarabaeidae)

Polyphenism has been suggested as an accelerator for morphological evolution and speciation. In the dung beetles of the genus Onthophagus, horn expression is polyphenic: large males develop horns whereas smaller males express greatly reduced or no horns. Horn static allometries seem to diverge rapidly amongst extant taxa, a process which might trigger changes in the male genital morphology, thus possibly promoting speciation as a by‐product. It can therefore be hypothesized that interspecific distances in allometries and, possibly, in other morphological traits mirror phylogenetic distances. In this study we first assessed the phylogenetic relationships amongst three closely related taxa belonging to the so‐called ‘Onthophagus fracticornis‐similis‐opacicollis’ species‐complex by sequencing the mitochondrial gene cytochrome oxidase subunit 1 (cox1). Biomolecular results indicated three independent lineages, the closest relationships being found between Onthophagus similis and Onthophagus opacicollis. Then we assessed the extent to which divergence pattern of horn static allometries and size and shape divergence patterns of one genital (paramere) and two nongenital (head and epipharynx) structures mirrored the phylogenetic relationships. Interspecific divergence patterns of horn static allometries, paramere, and head shape were found to be congruent with the evolutionary relationships inferred from biomolecular data. Nevertheless, paramere size and epipharynx shape showed patterns not consistent with the phylogeny. Furthermore, the relative size of nongenital structures showed little interspecific divergence compared to their shapes. Our results suggest that size and shape interspecific divergence mirror phylogeny only in part; they also indicate that distinct morphological traits may differ in their tendency to evolve in concert, and that size and shape of the same trait can evolve independently across species. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 482–498.     

Morphotypes,varieties, or subspecies?: genetic diversity and differentiation of four Saintpaulia (Gesneriaceae) morphotypes from the East Usambara Mountains,Tanzania

Saintpaulia (H. Wendl.) is a forest-dependent, endangered plant genus endemic to Tanzania and Kenya. The taxonomy of Saintpaulia from the Usambara Mountains and adjacent lowlands in north-eastern Tanzania is problematic because of the morphological similarity of the species and the presence of considerable intraspecific variation. Conventional molecular phylogenetic methods have failed to reveal the genetic structure of this Saintpaulia complex. In this study, we assessed the genetic composition of 12 Saintpaulia populations, representing four different morphotypes, from the East Usambara Mountains using inter-simple sequence repeat (ISSR) markers. Relatively high genetic diversities were observed within populations (mean h  = 0.320), indicating their adaptive potential. Little genetic differentiation amongst populations (mean F _st  = 0.063) and the genetic divergence of the rosulate and trailing morphotypes support the hypothesis of ongoing divergent evolution within the East Usambara metapopulation(s) of Saintpaulia .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 347–355.     

Genetic and environmental components of phenotypic variation in body shape among populations of Atlantic cod (Gadus morhua L.)

A common-garden experiment was conducted on larvae to test for genetic differences in body shape among populations of Atlantic cod ( Gadus morhua ). Offspring from four north-west Atlantic regions were reared from hatching to postmetamorphosis at two temperatures (7 ± 1 °C and 11 ± 1 °C) and two food levels (1500 and 4500 prey L⁻¹). Body shape differed between populations and treatments. Population differences were greatest between south-west Scotian Shelf cod and those further north; the former were characterized by a deeper body, larger head, and longer caudal peduncle than cod from the other populations. Significant differences were also observed between two putative populations on the south-west Scotian Shelf, suggesting genetic divergence between spawning aggregations at small spatial scales (< 100 km). Temperature and food supply also influenced body shape, with the effect of the former being more pronounced. Individuals reared at the higher temperature or food level had a deeper body and a larger head than those reared at the lower temperature or food supply. Phenotypic responses to changes in the rearing environment also differed among populations, indicating genetic differences in phenotypic plasticity. Differences between populations in morphology and in phenotypic plasticity suggest genetic divergence at both large (> 1000 km) and small (< 100 km) spatial scales. The genetic differences at large spatial scales counteracted the expected effects of temperature differences in the wild, suggesting countergradient variation in morphology among these populations.  © 2006 Her Majesty the Queen in Right of Canada. Journal compilation © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 351–365.     

Sympatric occurrence of diverged mtDNA lineages of Pityogenes chalcographus (Coleoptera, Scolytinae) in Europe

Differentiation among the European populations of the six-toothed spruce bark beetle Pityogenes chalcographus was observed for the first time in the 1970s as mating males from Northern Europe with females from Central Europe led to a significant decline in fecundity. Morphological examination revealed that P. chalcographus can be separated into two European races. Here, we investigated the genetic background of this separation by analysing 39 populations ( n  = 695), sequencing almost the complete cytochrome oxidase I gene (1543 bp) and applying single-strand conformation polymorphism. Phylogenetic analysis of 58 haplotypes yielded three major clades with a maximum sequence divergence of 2.33%, indicating that the demographic events took place in the late Pleistocene. These results support the hypothesis of allopatric divergence of the mtDNA lineages, which postglacially came into sympatric existence in Europe. However, as a result of partial crossing incompatibility the diverged lineages retained their genetic identity during postglacial times.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 331–340.     

The interplay between multiple predators and prey colour divergence

Evolutionary divergence in the coloration of toxic prey is expected when geographic variation in predator composition and behavior favours shifts in prey conspicuousness. A fundamental prediction of predator‐driven colour divergence is that the local coloration should experience lower predation risk than novel prey phenotypes. The dorsal coloration of the granular poison frog varies gradually from populations of conspicuous bright red frogs to populations of dull green and relatively cryptic frogs. We conducted experiments with clay models in four populations to examine the geographic patterns of taxon‐specific predation. Birds avoided the local phenotype while lizards consistently selected for decreased conspicuousness and crab predation did not depend on frog coloration. Importantly, birds and lizards favoured low conspicuousness in populations where relatively cryptic green morphs have evolved. This study provides evidence for the interplay among distinct selective pressures, from multiple‐predator taxa, acting on the divergence in protective coloration of prey species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 580–589.     

Risk of predation as a promoting factor of species divergence in threespine sticklebacks (Gasterosteus aculeatus L.)

Icelandic freshwater systems are geologically young and contain only six species of freshwater fish. As these species colonized Icelandic fresh waters they were presented with a diversity of unique, uncontested habitats and food resources, promoting the evolution of new behaviour strategies crucial to the formation of new morphs and speciation. To determine the likelihood that predation threat could affect the antipredator behaviour and possibly the sympatric divergence of prey populations, we analysed antipredator behaviour of seven groups of Icelandic threespine sticklebacks ( Gasterosteus aculeatus ): two marine groups, one group from a lake without piscine predators, and two polymorphic lake populations, each with two groups occupying unique habitats. Shoaling cohesion, school formation and duration, and vigilance in predator inspection/avoidance behaviour varied greatly among groups. The differences appeared to be related to the risk of predation as well as to opportunities and constraints set by the different habitats. Antipredator behaviour was especially pronounced and differed extensively in two polymorphic forms from the lake Thingvallavatn, where predation risk is very high. By keeping the two morphs separate in their respective habitats, high predation risk may be a contributing factor in promoting the habitat-specific divergence of G. aculeatus seen in the lake. This suggests that in situations where refuge habitats are spatially separated, the risk of predation may contribute to the evolution of separate sympatric forms of small fish such as G. aculeatus .  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 189–203.     

Combining geometric morphometrics and pattern recognition to identify interspecific patterns of skull variation: case study in sympatric Argentinian species of the genus Calomys (Rodentia: Cricetidae: Sigmodontinae)

Sympatric species of vesper mice Calomys laucha and Calomys musculinus are difficult to discriminate, especially in natural history collections where they are identified by external body measurements and cranial characteristics. Accurate identification of these two species can be important because only one of them, C. musculinus , is a Junin virus reservoir, the aetiological agent of the Argentine Hemorragic Fever. Research has focused into the development of molecular techniques to unambiguously identify these species. We apply statistical procedures from the field of pattern recognition to three-dimensional geometric morphometric data based on skull landmarks to identify sympatric species C. laucha , C. musculinus and Calomys venustus . Pattern recognition techniques correctly identified the three species without any prior information on specimen identity. By contrast to expectations, C. venustus differed from the other two species mainly on the basis of shape and not by its centroid size. The main sources of difference between C. laucha and C. musculinus were of shape, specifically localized at the landmarks defined by: (1) the sutures between the premaxillaries, the nasals and the frontals; (2) the sutures between the parietals, the frontals and the squamosals; and (3) the suture between the parietals and the interparietal. Nevertheless, allometries dominate the patterns of interspecific variation between these latter species and may partly explain past identification difficulties. Morphological evolution is discussed. The need for objective methods to define phenotypic clusters is highlighted with respect to the need for fast and precise biodiversity assessments.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 365–378.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.