Patterns of morphological evolution in Marmota (Rodentia, Sciuridae): geometric morphometrics of the cranium in the context of marmot phylogeny, ecology and conservation

Marmots are of great interest for both sociobiologists studying the evolution of mammal societies and conservationists trying to protect them from extinction. In contrast, their phylogeny and morphological evolution are poorly understood and studied. Recently, a phylogenetic analysis using cytochrome b provided the first reconstruction of marmot evolutionary history and suggested that a high level of sociality evolved at least twice independently in the two proposed marmot subgenera. A morphological analysis of the marmot mandible supported this subgeneric classification and showed interesting, and unexpected, patterns in the evolution of marmot skeletal characters. In the present study we investigated a more complex, and potentially informative structure, the ventral cranium. Geometric morphometric techniques were applied in the first analysis of cranial morphology including all marmot species. Three main phenetic groups were found, which reflect phylogeny (subgenus Petromarmota , and Palaearctic subgenus Marmota ) or geographical distribution (Palaearctic vs. Nearctic subgenus Marmota ). Convergence in skeletal characters due to size similarities, a common finding in the sciurid skeleton according to traditional morphological analyses, did not occur in the marmot ventral cranium. Despite a genetic distance between Marmota vancouverensis and Marmota caligata similar to that among different populations of the latter species, the Vancouver Island marmot had the most atypical ventral cranium in the subgenus Petromarmota . This finding confirmed results obtained with the mandible, and emphasized the uniqueness of M. vancouverensis and the usefulness of complementing molecular analyses with morphological studies for a thorough characterization of population divergence, and a careful planning of conservation strategies.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 385–407     

Morphological evolution in marmots (Rodentia, Sciuridae): size and shape of the dorsal and lateral surfaces of the cranium

Marmots are the largest ground squirrels and have been extensively studied by sociobiologists investigating the evolution of mammal societies. Being a member of the sciurid clade, traditionally considered inclined to convergence, they are also a group on which to test the hypothesis of sciurid propensity to homoplasy of osteological characters. In the present analysis, the dorsal and lateral surfaces of the cranium of all living marmot species are compared with geometric morphometric techniques. Phenetic groups are found which reflect the subgeneric classification of marmots and are consistent with previous morphometric analyses of the mandible and ventral cranium. Two species have distinctive morphologies and phenetic relationships not congruent with phylogeny. Marmota vancouverensis is highly divergent for osteological characters, fur colour and behaviour despite its young age and close genetic similarity to Marmota caligata . Its small population may represent a rare chance to study evolutionary processes during rapid allopatric speciation in mammals, but strong conservation efforts are required to preserve this unique component of the Vancouver Island biodiversity. Also, Marmota monax has distinctive cranial traits. These are possibly related to its long separate evolutionary history and unique ecology and behaviour. Size-related convergence is not evident in Marmota . When outgroup species are included, Spermophilus , Cynomys , Tamias , and Sciurus group together on one branch, Marmota on the other. This is best explained as a retention of the ancestral morphology in the smaller members of the Marmotini ( Spermophilus , Cynomys , and Tamias ) and the evolution of derived morphology in Marmota .     

The geometry of the marmot (rodentia: sciuridae) mandible: phylogeny and patterns of morphological evolution

Marmots have a prominent role in the study of mammalian social evolution, but only recently has their systematics received the attention it deserves if sociobiological studies are to be placed in a phylogenetic context. Sciurid morphology can be used as model to test the congruence between morphological change and phylogeny because sciurid skeletal characters are considered to be inclined to convergence. However, no morphological study involving all marmot species has ever been undertaken. Geometric morphometric techniques were applied in a comparative study of the marmot mandible. The adults of all 14 living marmot species were compared, and mean mandible shape were used to investigate morphological evolution in the genus Marmota. Three major trends were observed. First, the phylogenetic signal in the variation of landmark geometry, which describes mandible morphology, seems to account for the shape differences at intermediate taxonomic levels. The subgenera Marmota and Petromarmota, recently proposed on the basis of mitochondrial cytochrome b sequence, receive support from mandible morphology. When other sciurid genera were included in the analysis, the monophyly of the genus Marmota and that of the tribe Marmotini (i.e., marmots, prairie dogs, and ground squirrels) was strengthened by the morphological data. Second, the marmotine mandible may have evolved as a mosaic of characters and does not show convergence determined by size similarities. Third, allopatric speciation in peripheral isolates may have acted as a powerful force for modeling shape. This hypothesis is strongly supported by the peculiar mandible of M. vancouverensis and, to a lesser degree, by that of M. olympus, both thought to have originated as isolated populations in Pleistocene ice-free refugia.     

Comparison of discriminant function and classification tree analyses for age classification of marmots

We evaluated the predictive power of two classification techniques, one parametric – discriminant function analysis (DFA) and the other non-parametric – classification and regression tree analysis (CART), in order to provide a non-subjective quantitative method of determining age class in Vancouver Island marmots ( Marmota vancouverensis ) and hoary marmots ( Marmota caligata ). For both techniques we used morphological measurements of known-age male and female marmots from two independent population studies to build and test predictive models of age class. Both techniques had high predictive power (69–86%) for both sexes and both species. Overall, the two methods performed identically with 81% correct classification. DFA was marginally better at discriminating among older more challenging age classes compared to CART. However, in our test samples, cases with missing values in any of the discriminant variables were deleted and hence unclassified by DFA, whereas CART used values from closely correlated variables to substitute for the missing values. Therefore, overall, CART performed better (CART 81% vs DFA 76%) because of its ability to classify incomplete cases. Correct classification rates were approximately 10% higher for hoary marmots than for Vancouver Island marmots, a result that could be attributed to different sets of morphological measurements. Zygomatic arch breadth measured in hoary marmots was the most important predictor of age class in both sexes using both classification techniques. We recommend that CART analysis be performed on data-sets with incomplete records and used as a variable screening tool prior to DFA on more complete data-sets.     

Ecogeographical patterns of morphological variation in pygmy shrews Sorex minutus (Soricomorpha: Soricinae) within a phylogeographical and continental‐and‐island framework

Ecogeographical patterns of morphological variation were studied in the Eurasian pygmy shrew Sorex minutus aiming to understand the species’ morphological diversity in a continental and island setting, and within the context of previous detailed phylogeographical studies. In total, 568 mandibles and 377 skulls of S. minutus from continental and island populations from Europe and Atlantic islands were examined using a geometric morphometrics approach, and the general relationships of mandible and skull size and shape with geographical and environmental variables were studied. Samples were then pooled into predefined geographical groups to evaluate the morphological differences among them using analyses of variance, aiming to contrast the morphological and genetic relationships based on morphological and genetic distances and ancestral state reconstructions, as well as assess the correlations of morphological, genetic, and geographical distances with Mantel tests. We found significant relationships of mandible size with geographical and environmental variables, fitting the converse Bergmann's rule; however, for skull size, this was less evident. Continental groups of S. minutus could not readily be differentiated from each other by shape. Most island groups of S. minutus were easily discriminated from the continental groups by being larger, indicative of an island effect. Moreover, morphological and genetic distances differed substantially and, again, island groups were distinctive morphologically. Morphological and geographical distances were significantly correlated, although this was not the case for morphological and genetic distances, indicating that morphological variation does not reflect genetic subdivision in S. minutus. Our analyses showed that environmental variables and insularity had important effects on the morphological differentiation of S. minutus.     

Marmot phylogeny revisited: molecular evidence for a diphyletic origin of sociality

We established the phylogeny of 11 species of the genus Marmota based on the entire sequence of the mitochondrial cytochrome b ( cyt-b ) gene (1.1 kb) and a partial sequence of the NADH dehydrogenase subunit 4 ( ND4 ) gene (1.2 kb). In three species ( Marmota caligata , Marmota olympus , and Marmota bobac ) full-sized nuclear pseudogenes of the mitochondrial cyt-b were identified. The mitochondrial cyt-b genes and the three pseudogenes form separate clusters in the maximum parsimony dendrogram. This finding suggests that the pseudogenes originated from a single transfer to the nucleus that may have occurred prior to the radiation of the genus Marmota . Notably, compared with their functional mitochondrial equivalents the pseudogenes show a much lower substitution rate. In the dendrograms deduced from the mitochondrial sequences two distinct clusters become apparent: one cluster consists of the North-west American species, the other contains the Eurasian species together with the North American species Marmota monax . The position of M. monax as a member of the Eurasian clade is in accordance with the evolution of chromosome numbers. The results are of special interest with respect to the evolution of social systems in the genus that vary from solitary species ( M. monax ) to highly social species living in family groups (e.g. Marmota marmota ). The molecular phylogeny suggests a diphyletic origin of high sociality in the genus Marmota .     

Endemic evolutionary radiation of Rhagada land snails (Pulmonata: Camaenidae) in a continental archipelago in northern Western Australia

In the Dampier Archipelago of Western Australia's Pilbara Region, several locally endemic, morphologically distinctive species of Rhagada land snails occur, contrasting with the morphologically conservative species with wider distributions on the adjacent mainland. To test alternative origins of this unusual local diversity in a continental archipelago, we examined sequences of the cytochrome oxidase subunit 1 and 16S mitochondrial genes in 22 described species and eight undescribed forms, including all known morphospecies from the Pilbara Region's Dampier Archipelago and adjacent mainland. Phylogenetic analyses consistently resolved four, deep clades within the Pilbara Region, with a mean sequence divergence of 15–18%. All but one of the species from the Dampier Archipelago formed one of the major clades, indicating that the morphological radiation in the archipelago evolved locally, rather than through multiple, relictual mainland lineages. Morphological divergence spanning almost that of the entire genus was within a subclade with sequence divergence < 4%, highlighting the disconnection between morphological diversification and levels of molecular genetic divergence. This in situ morphological radiation in the Dampier Archipelago, which transcends variation seen over much larger distances on the mainland, is unusual for a continental archipelago, and may relate to local heterogeneity of land forms. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 316–327.     

Variation in morphology and functional performance across distinct evolutionary lineages of the Moorish gecko (Tarentola mauritanica) from the Iberian Peninsula

Deciphering the mechanisms that underlie morphological and functional diversity is essential for understanding how organisms adapt to their environment. Interestingly, phenotypic divergence does not necessarily correspond to the geographic and genetic separation between populations. Here, we explored the morphological and functional divergence among populations of two genetically differentiated clades of the Moorish gecko, Tarentola mauritanica. We used linear and geometric morphometrics to quantify morphological variation and investigated how it translates into biting and CLIMBING PERFORMANCE, to better understand the mechanisms potentially underlying population and lineage divergence. We found marked morphological differences between clades, both in body size and head shape. However, much of this differentiation is more strongly related to local variation between populations of the same clade, suggesting that recent ecological events may be more influential than deep evolutionary history in shaping diversity patterns in this group. Despite a lack of association between morphology and functional diversification in the locomotor system of the Moorish gecko, straightforward links are observed between head morphology and biting performance, providing more hints on the possible underlying causes. Indeed, variation in bite force is mostly determined by size variation and sexual dimorphism, and differences between the two clades concern how sexual variation is expressed, reinforcing the idea that both social and ecological factors contribute in shaping differentiation. Interestingly, the individuals from the islets off the coast of Murcia exhibit particular morphological and functional traits, which suggests that the ecological conditions related to insularity may drive the phenotypic differentiation of this population.     

Absence of founder effect and evidence for adaptive divergence in a recently introduced insular population of white‐tailed deer (Odocoileus virginianus)

Islands are generally colonized by few individuals which could lead to a founder effect causing loss of genetic diversity and rapid divergence by strong genetic drift. Insular conditions can also induce new selective pressures on populations. Here, we investigated the extent of genetic differentiation within a white‐tailed deer (Odocoileus virginianus) population introduced on an island and its differentiation with its source mainland population. In response to their novel environmental conditions, introduced deer changed phenotypically from mainland individuals, therefore we investigated the genetic bases of the morphological differentiation. The study was conducted on Anticosti Island (Québec, Canada) where 220 individuals were introduced 120 years ago, resulting in a population size over 160,000 individuals. We used genotyping‐by‐sequencing (GBS) to generate 8,518 filtered high‐quality SNPs and compared patterns of genetic diversity and differentiation between the continental and Anticosti Island populations. Clustering analyses indicated a single panmictic island population and no sign of isolation by distance. Our results revealed a weak, albeit highly significant, genetic differentiation between the Anticosti Island population and its source population (mean F_ST = 0.005), which allowed a population assignment success of 93%. Also, the high genetic diversity maintained in the introduced population supports the absence of a strong founder effect due to the large number of founders followed by rapid population growth. We further used a polygenic approach to assess the genetic bases of the divergent phenotypical traits between insular and continental populations. We found loci related to muscular function and lipid metabolism, which suggested that these could be involved in local adaptation on Anticosti Island. We discuss these results in a harvest management context.     

Reticulate evolution and phenotypic diversity in North American ciscoes, Coregonus ssp. (Teleostei: Salmonidae): implications for the conservation of an evolutionary legacy

Coregonine fishes are notorious taxonomicproblems due to their extreme morphological andecological variation. In North America, diversity is particularly baffling among ciscoes, and both morphological and phylogenetic analyses have resulted in major polytomy among the 8 taxa of the ``Coregonus artedi' species complex. Ciscoes arealso a devastated group, accounting for 10% ofthe fish species listed by the Committee on theStatus of Endangered Wildlife in Canada. Here,we complete the genetic characterization ofNorth American ciscoes with mitochondrial andmicrosatellites markers previously used toanalyse populations of C. artedi in orderto elucidate the evolutionary history andidentify appropriate conservation units. Ourresults revealed a complex evolutionary historymarked by postglacial reticulation eventscoupled with recent and independent evolutionof similar phenotypes (taxa). Genetic variationreflects geography rather than taxonomy, andconsequently, we recommend that a single taxon,C. artedi (sensu lato) be recognized.Local genetic differentiation is often coupledwith ecophenotypic diversification, and gillraker polymorphisms, depth-related habitatpreference and reproductive behaviour areconsidered as phenotypic traits with probableadaptive value contributing to the nicheexpansion of ciscoes. Ecomorphotypes of eachparticular locale thus represent a uniqueexpression of a diverse genetic pool stillundergoing divergence and sorting.Consequently, ciscoes from lakes with distinctecomorphotypes are recognized as ESUs, as wellas each of sympatric forms when they aregenetically differentiated. We recommend thatan ESU strategy focusing at a very local levelbe adopted for continental ciscoes as a validalternative to protect significant evolutionaryprocesses of divergence encountered inpolytypic species of newly colonized habitats.     

Population genetic structure,differentiation, and diversity in Tetrix subulata pygmy grasshoppers: roles of population size and immigration

Genetic diversity within and among populations and species is influenced by complex demographic and evolutionary processes. Despite extensive research, there is no consensus regarding how landscape structure, spatial distribution, gene flow, and population dynamics impact genetic composition of natural populations. Here, we used amplified fragment length polymorphisms (AFLPs) to investigate effects of population size, geographic isolation, immigration, and gene flow on genetic structure, divergence, and diversity in populations of Tetrix subulata pygmy grasshoppers (Orthoptera: Tetrigidae) from 20 sampling locations in southern Sweden. Analyses of 1564 AFLP markers revealed low to moderate levels of genetic diversity (PPL = 59.5–90.1; Hj = 0.23–0.32) within and significant divergence among sampling localities. This suggests that evolution of functional traits in response to divergent selection is possible and that gene flow is restricted. Genetic diversity increased with population size and with increasing proportion of long‐winged phenotypes (a proxy of recent immigration) across populations on the island of Öland, but not on the mainland. Our data further suggested that the open water separating Öland from the mainland acts as a dispersal barrier that restricts migration and leads to genetic divergence among regions. Isolation by distance was evident for short interpopulation distances on the mainland, but gradually disappeared as populations separated by longer distances were included. Results illustrate that integrating ecological and molecular data is key to identifying drivers of population genetic structure in natural populations. Our findings also underscore the importance of landscape structure and spatial sampling scheme for conclusions regarding the role of gene flow and isolation by distance.     

Molecular and morphological data suggest that Spinibarbus caldwelli (Nichols) (Teleostei: Cyprinidae) is a valid species

The taxonomic problem of the cyprinid species of genus Spinibarbus, occurring in southern China and northern Vietnam, was resolved on the basis of molecular and morphological analyses. Spinibarbus caldwelli and Spinibarbus hollandi have a smooth posterior edge of the last unbranched dorsal fin ray among species in the genus. Spinibarbus caldwelli is currently regarded as a junior synonym of S. hollandi because of ambiguities in diagnostic characters. In this article, 11 mtDNA cytochrome b sequences of Spinibarbus specimens were analyzed together with Barbodes gonionotus and Puntius conchonius as outgroups. Our results showed that specimens identified as S. hollandi from Taiwan were different from those from the Asian mainland at a high level of genetic divergence (0.097–0.112), which is higher than that between the two valid species, S. sinensis and S. yunnanensis (0.089), and suggested that Taiwan specimens should be considered as a different species from the Asian mainland one. In a molecular phylogenetic analysis, the sister-group relationship between Taiwan specimens and the Asian mainland specimens was supported strongly by a high confidence level (100% in bootstrap value). Further analysis of morphological characters showed that overlap of diagnostic characters is much weaker than previously suggested. Taiwan specimens had 8 branched rays in the dorsal fin, whereas those from the mainland had almost 9–10. The molecular and morphological differences suggest S. caldwelli to be valid. The molecular divergence shows the genetic speciation of S. hollandi and S. caldwelli might have occurred 5.6–4.9 million years ago; the former could be a relict species in Taiwan, and the latter dispersed in the Asian mainland.     

Phenotypic diversification and island evolution of pipistrelle bats (Pipistrellus pipistrellus group) in the Mediterranean region inferred from geometric morphometrics and molecular phylogenetics

Aim The Mediterranean Basin is a centre of radiation for numerous species groups. To increase our understanding of the mechanisms underlying speciation and radiation events in this region, we assessed the phenotypic variability within the Pipistrellus pipistrellus–pygmaeus–hanaki species complex. Although bats form the second largest mammalian order, studies of insular evolution in this group are scarce. We approached this problem from a microevolutionary perspective and tested for the recurrence of the insular syndrome. Location The Mediterranean Basin, with a special focus on isolated populations from Corsica, the Maghreb, Cyprus, Cyrenaica and Crete. Methods Phenotypic variability was assessed by cranial morphometrics using the coordinates of 41 3D landmarks and associated geometric‐morphometric methods. We analysed 125 specimens representing all of the lineages in the species complex. Differences between taxa and between insular and continental populations in cranial size, shape, form and allometries were tested using analyses of variance and visualized using boxplots and canonical variate analysis. Relationships between molecular data from a previous study (cytochrome b sequences) and morphometric data were tested with co‐inertia analyses (RV test) and multivariate regressions. Results The three species were relatively well differentiated in cranial size and shape, and each species showed a significant amount of inter‐population variability. Comparisons of pairs of insular versus continental populations revealed heterogeneities in cranial patterns among island phenotypes, suggesting no recurrent insular syndrome. Molecular and phenotypic traits were correlated, except for molecular and lateral cranium shape. Main conclusions The Pipistrellus pipistrellus – pygmaeus – hanaki species complex exhibits phenotypic variability as a result of the fragmentation of its distribution (especially on islands), its phylogenetic and phylogeographic history and, most probably, other evolutionary factors that were not investigated in this study. We found no recurrent pattern of evolution on islands, indicating that site‐specific factors play a prevailing role on Mediterranean islands. The correlation between molecular and phenotypic data is incomplete, suggesting that factors other than phylogenetic relationships, potentially connected with feeding ecology, have played a role in shaping cranial morphology in this species complex.     

Social structure and facultative mating systems of hoary marmots (Marmota caligata)

Mate-choice theory predicts different optimal mating systems depending on resource availability and habitat stability. Regions with limited resources are thought to promote monogamy. We tested predictions of monogamy in a social rodent, the hoary marmot (Marmota caligata), at the northern climatic extreme of its distribution. Mating systems, social structure and genetic relationships were investigated within and among neighbouring colonies of marmots within a 4 km(2) valley near Kluane National Park, Yukon, Canada, using 21 microsatellite loci. While both monogamous and polygynous populations of hoary marmots have been observed in the southern reaches of this species' range; northern populations of this species are thought to be predominantly monogamous. Contrary to previous studies, we did not find northern hoary marmot social groups to be predominantly monogamous; rather, the mating system seemed to be facultative, varying between monogamy and polygyny within, as well as among, social groups. These findings reveal that the mating systems within colonies of this species are more flexible than previously thought, potentially reflecting local variation in resource availability.     

Quantitative trait and allozyme divergence in the Greenfinch (Carduelis chloris, Aves: Fringillidae)

Quantitative trait divergence and variability among 12 greenfinch populations across continental Europe was examined and compared to divergence in neutral genetic markers (allozymes). The added among locality variance component for 16 skeletal traits was large (mean 28%, range 4–48%) equalling a divergence of up to three SD units. The divergence in quantitative traits (Qst = 0.04-0.48) greatly exceeded that in alloymes (F_ST= 0.01-0.07), indicating the differentiation in quantitative traits to be larger than expected by mutation and drift alone. This conclusion was consistent also with results from the multivariate approach of Rogers & Harpending. However, genetic and morphometric distances between populations were positively correlated, even when controlling for the geographic distance separating pairs of populations. In concordance with Bergmann's rule, most traits were strongly and positively correlated with latitude, indicating latitudinally ordered genetic or/and environmental effects. However, the correlation between lower mandible width and latitude was strongly negative, demonstrating an inverse relationship between beak size and body size across the populations. These results are interpreted to reflect the re-colonization of history of northern Europe (genetic and geographic distances correlated) which has been paralleled by selection acting on quantitative traits (Q_ST>F_ST)- In particular, the counter-gradient variation in beak width, a functionally important trait, is suggestive of an adaptive basis for quantitative trait divergence.     

Surviving in the High Arctic: dental variation in a casually introduced population of Microtus rossiaemeridionalis (Arvicolinae,Rodentia) on Svalbard

A human‐mediated invasion of a temperate rodent Microtus rossiaemeridionalis on the High Arctic Archipelago of Svalbard in the first half of the 20th century provides an opportunity to explore extent and rate of morphological divergence over decades of isolation. We studied dental size and morphology in 124 voles captured on Svalbard (Spitsbergen) in 1997–2005 and compared the data to mainland conspecific populations across northern Eurasia. Both dental and cranial sizes in the Svalbard population fall within the limits of natural variation of the species. Dental morphology suggests that the population experiences strong effects of isolation as indicated by the significant increase in the frequency of rare dental morphs, possibly caused by inbreeding. No evidence for directional shift towards increased or decreased complexity of the morphotype dental patterns is revealed. Although the population on Svalbard is phenotypically different from the mainland populations (due to increased frequency of rare morphs), those differences are not enough to support the idea of initial rapid evolution related to colonization. The limited spatial extent and environmental homogeneity of suitable habitats on the island allowed the species to initially multiply but not to diversify so that the species exhibits phenotypic conservatism but suffers the consequences of small population size.     

MICROGEOGRAPHIC PATTERNS OF GENETIC AND MORPHOLOGICAL VARIATION IN SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS)

Surveys of genetic population structure are often limited to large geographic scales because geographically close populations are indistinguishable. Genetic uniformity across adjacent demes can be interpreted as evidence for cohesion (panmixia) or recent divergence. However, poor genetic resolution at microgeographic scales can also arise from the use of overly conservative (slowly evolving) markers. This study examines the ability of hypervariable, minisatellite loci to discriminate among geographically close populations of Savannah sparrows (Passerculus sandwichensis) and to track morphological differentiation at a microgeographic scale (interregional distance < 55 km). Savannah sparrows breeding at five island and two mainland sites in the Bay of Fundy, New Brunswick, Canada, show concordant patterns of variation in external morphology (seven characters) and multilocus DNA fingerprinting profiles (S_xy): island sparrows are phenotypically larger and genetically more similar to each other than they are to mainland sparrows. This pattern of variation is consistent with both adaptive (natural selection) and nonadaptive (genetic drift) mechanisms of population divergence. Based on minisatellite diversity, the effective size of both island and mainland populations is 37, an estimate substantially lower than census population sizes. These data are discordant with observations of sparrow vagility and abundance and suggest a closer examination of microgeographic patterns in avian systems.     

Variation in guenon skulls (II): sexual dimorphism

Patterns of size and shape sexual dimorphism in adult guenons were examined using a large sample of skulls from almost all living species. Within species, sexual dimorphism in skull shape follows the direction of size-related shape variation of adults, is proportional to differences in size, and tends to be larger in large-bodied species. Interspecific divergence among shape trajectories, which explain within species sex differences, are small (i.e., trajectories of most species are nearly parallel). Thus, changes in relative proportions of skull regions that account for the distinctive shape of females and males are relatively conserved across species, and their magnitude largely depends on differences in size between sexes. A conservative pattern of size-related sexual dimorphism and a model of interspecific divergence in shape which strongly reflects size differences suggest a major role of size and size-related shape variation in the guenon radiation. It is possible that in the guenons, as in the neotropical primates (with whom they have obvious parallels), size has helped to determine morphological change along lines of least evolutionary resistance, influencing sexual dimorphism. In Miopithecus and Erythrocebus, the smallest and largest guenon genera, it is likely that the interaction of ecology and size contributes significantly to patterns of sexual dimorphism. The results of this study thus emphasise the need to consider allometry and size alongside ecology and behaviour when examining primate sexual dimorphism.     

Diversity and divergence patterns in regulatory genes suggest differential gene flow in recently derived species of the Hawaiian silversword alliance adaptive radiation (Asteraceae)

The impact of gene flow and population size fluctuations in shaping genetic variation during adaptive radiation, at both the genome-wide and gene-specific levels, is very poorly understood. To examine how historical population size and gene flow patterns within and between loci have influenced lineage divergence in the Hawaiian silversword alliance, we have investigated the nucleotide sequence diversity and divergence patterns of four floral regulatory genes (ASAP1-A, ASAP1-B, ASAP3-A, ASAP3-B) and a structural gene (ASCAB9). Levels and patterns of molecular divergence across these five nuclear loci were estimated between two recently derived species (Dubautia ciliolata and Dubautia arborea) which are presumed to be sibling species. This multilocus analysis of genetic variation, haplotype divergence and historical demography indicates that population expansion and differential gene flow occurred subsequent to the divergence of these two lineages. Moreover, contrasting patterns of allele- sharing for regulatory loci vs. a structural locus between these two sibling species indicate alternative histories of genetic variation and partitioning among loci where alleles of the floral regulatory loci are shared primarily from D. arborea to D. ciliolata and alleles of the structural locus are shared in both directions. Taken together, these results suggest that adaptively radiating species can exhibit contrasting allele migration rates among loci such that allele movement at specific loci may supersede genetic divergence caused by drift and that lineage divergence during adaptive radiation can be associated with population expansion.     

Effects of migration on the genetic covariance matrix

In 1996, Schluter showed that the direction of morphological divergence of closely related species is biased toward the line of least genetic resistance, represented by g_max , the leading eigenvector of the matrix of genetic variance–covariance (the G -matrix). G is used to predict the direction of evolutionary change in natural populations. However, this usage requires that G is sufficiently constant over time to have enough predictive significance. Here, we explore the alternative explanation that G can evolve due to gene flow to conform to the direction of divergence between incipient species. We use computer simulations in a mainland–island migration model with stabilizing selection on two quantitative traits. We show that a high level of gene flow from a mainland population is required to significantly affect the orientation of the G -matrix in an island population. The changes caused by the introgression of the mainland alleles into the island population affect all aspects of the shape of G (size, eccentricity, and orientation) and lead to the alignment of g_max with the line of divergence between the two populations' phenotypic optima. Those changes decrease with increased correlation in mutational effects and with a correlated selection. Our results suggest that high migration rates, such as those often seen at the intraspecific level, will substantially affect the shape and orientation of G , whereas low migration (e.g., at the interspecific level) is unlikely to substantially affect the evolution of G .