MORPHOMETRIC VARIABILITY IN CONTINENTAL AND ATLANTIC ISLAND POPULATIONS OF CHAFFINCHES (FRINGILLA COELEBS)

The magnitudes of phenotypic variances in peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Azores, Madeira, and Canaries relative to their continental source populations in Iberia and Morocco have played a pivotal role in the formulation of competing hypotheses of phenotypic evolution in these isolates. Because Van Valen's niche variation hypothesis and Grant's model of island evolution were based on museum skins prone to measurement error and temporal variation, we re-examined the patterns of phenotypic variability using more precise skeletal measurements from freshly collected specimens. Levene's tests showed that univariate character variances were homogeneous in all island and continental populations, although there was a consistent trend for the magnitude of the variances to be lower for all characters in all Canary island populations. Multivariate Levene's tests, however, revealed significantly reduced total variances in the Hierro and Madeira populations compared to some Azores and continental populations. The Azores and continental populations did not differ in variability, and lower variances in the Canaries were not related to the presence or absence of the congeneric blue chaffinch (F. teydea), contrary to the predictions of Van Valen's niche variation hypothesis. Population variability was not inversely related to differentiation or isolation within the Azores or Canaries archipelagoes, opposite to the association reported by Grant. Our results also differ from both previous studies which reported much larger differences in population variabilities, and this likely reflects the use of heterogeneous samples of museum specimens, less precise external characters, and the use of tests sensitive to sample size. Differentiation among populations has been markedly greater in the Canary islands, implicating founder events and possibly historically stronger directional selection as determinants of this enhanced divergence relative to the Azores. These variance-reducing processes are unlikely to explain current lower levels of phenotypic variability because there has been sufficient time since colonization for replenishment of variability in polygenic characters. Average heterozygosities at putatively neutral allozyme loci are 1.6 times higher in the Azores compared to the Canaries, and support the view that effective population sizes are smaller in the latter archipelago. We argue that reduced variances in the Canary island populations represent lower equilibrium levels maintained by drift and mutation in populations with smaller long-term effective sizes, consistent with Lynch and Hill's neutral model of phenotypic evolution. Although episodes of selection in the past may have been interspersed with long periods of effective neutralism and drift, adaptationist hypotheses invoking a primary role for variance-reducing selection appear to be unwarranted.     

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.     

Genetic structure in Orkney island mice: isolation promotes morphological diversification

Following human occupation, the house mouse has colonised numerous islands, exposing the species to a wide variety of environments. Such a colonisation process, involving successive founder events and bottlenecks, may either promote random evolution or facilitate adaptation, making the relative importance of adaptive and stochastic processes in insular evolution difficult to assess. Here, we jointly analyse genetic and morphometric variation in the house mice (Mus musculus domesticus) from the Orkney archipelago. Genetic analyses, based on mitochondrial DNA and microsatellites, revealed considerable genetic structure within the archipelago, suggestive of a high degree of isolation and long-lasting stability of the insular populations. Morphometric analyses, based on a quantification of the shape of the first upper molar, revealed considerable differentiation compared to Western European populations, and significant geographic structure in Orkney, largely congruent with the pattern of genetic divergence. Morphological diversification in Orkney followed a Brownian motion model of evolution, suggesting a primary role for random drift over adaptation to local environments. Substantial structuring of human populations in Orkney has recently been demonstrated, mirroring the situation found here in house mice. This synanthropic species may thus constitute a bioproxy of human structure and practices even at a very local scale.Subject terms: Evolutionary ecology, Population genetics     

Layers of contingency shroud pervasive ecological divergence in a local radiation of land snails

The predictability of evolution depends on the roles that selection and historical contingency play in determining its outcomes, but the relative importance of these evolutionary mechanisms has attracted considerable debate. One view is that historical events have such a profound impact on the genetic structure of populations that patterns of phenotypic evolution are essentially unpredictable. The opposing view is that selection is so powerful that evolutionary change is primarily deterministic, and thus highly predictable. By controlling for the effects of phylogeny, geographic location and habitat, this study examined the relative roles of contingency and determinism in a local radiation of land snails, genus Rhagada, in a continental archipelago. Informed by previous studies on a single island, which revealed a strong association between low‐spired shells and rocky habitats, 28 population pairs were sampled in directly adjoining rocky and spinifex plain habitats. When considered in their respective pairs, the effect of habitat was remarkably consistent, with lower‐spired shells observed in the rocky habitat in 24 of the comparisons. However, when analyzed outside the context of those pairs, the association was obscured by broad variation in shell shape within habitat types and among lineages. These results reveal the complex nature of a morphological radiation; while the pattern of ecological divergence is highly predictable at the scale that selection acts, deterministic evolution is largely obscured by phylogenetic and population history.     

Microevolution in island forms: the roles of drift and directional selection in morphological divergence of a passerine bird

Abstract.— Theory predicts that in small isolated populations random genetic drift can lead to phenotypic divergence; however this prediction has rarely been tested quantitatively in natural populations. Here we utilize natural repeated island colonization events by members of the avian species complex, Zosterops lateralis , to assess whether or not genetic drift alone is an adequate explanation for the observed patterns of microevolutionary divergence in morphology. Morphological and molecular genetic characteristics of island and mainland populations are compared to test three predictions of drift theory: (1) that the pattern of morphological change is idiosyncratic to each island; (2) that there is concordance between morphological and neutral genetic shifts across island populations; and (3) for populations whose time of colonization is known, that the rate of morphological change is sufficiently slow to be accounted for solely by genetic drift. Our results are not consistent with these predictions. First, the direction of size shifts was consistently towards larger size, suggesting the action of a nonrandom process. Second, patterns of morphological divergence among recently colonized populations showed little concordance with divergence in neutral genetic characters. Third, rate tests of morphological change showed that effective population sizes were not small enough for random processes alone to account for the magnitude of microevolutionary change. Altogether, these three lines of evidence suggest that drift alone is not an adequate explanation of morphological differentiation in recently colonized island Zosterops and therefore we suggest that the observed microevolutionary changes are largely a result of directional natural selection.     

The influence of gene flow and drift on genetic and phenotypic divergence in two species of Zosterops in Vanuatu

Colonization of an archipelago sets the stage for adaptive radiation. However, some archipelagos are home to spectacular radiations, while others have much lower levels of diversification. The amount of gene flow among allopatric populations is one factor proposed to contribute to this variation. In island colonizing birds, selection for reduced dispersal ability is predicted to produce changing patterns of regional population genetic structure as gene flow-dominated systems give way to drift-mediated divergence. If this transition is important in facilitating phenotypic divergence, levels of genetic and phenotypic divergence should be associated. We consider population genetic structure and phenotypic divergence among two co-distributed, congeneric (Genus: Zosterops) bird species inhabiting the Vanuatu archipelago. The more recent colonist, Z. lateralis, exhibits genetic patterns consistent with a strong influence of distance-mediated gene flow. However, complex patterns of asymmetrical gene flow indicate variation in dispersal ability or inclination among populations. The endemic species, Z. flavifrons, shows only a partial transition towards a drift-mediated system, despite a long evolutionary history on the archipelago. We find no strong evidence that gene flow constrains phenotypic divergence in either species, suggesting that levels of inter-island gene flow do not explain the absence of a radiation across this archipelago.     

Temporal and spatial variation of dental pattern in the voles, Microtus arvalis, of the Orkney Islands

A sample of first lower molars of Microtus arvalis of Neolithic age from the Orkney island of Westray was compared with respect to size and shape with other Neolithic and Recent samples of the species. The results show (1) a decline in size since the Neolithic on Westray (as on Orkney Mainland) and (2) divergence in shape between populations from Westray and Orkney Mainland since the Neolithic. Comparisons were also made with continental samples, and explanations of the origin and large size of the Orkney voles are reviewed.     

Divergent evolutionary processes associated with colonization of offshore islands

Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.     

Morphological and plumage colour variation in the Réunion grey white‐eye (Aves: Zosterops borbonicus): assessing the role of selection

The Réunion grey white‐eye (Zosterops borbonicus), a small passerine endemic to the island of Réunion (Mascarene archipelago), constitutes an extraordinary case of phenotypic variation within a bird species, with conspicuous plumage colour differentiation at a microgeographical scale. To understand whether natural selection could explain such variability, we compared patterns of variation in morphological and plumage colour traits within and among populations. To quantify morphological variation, we used measurements obtained by Frank Gill in the 1960s from 239 individuals collected in 60 localities distributed over the entire island of Réunion. To quantify colour variation, we measured the reflectance spectra of plumage patches of 50 males from a subset of Gill's specimens belonging to the five recognized plumage colour variants and used a visual model to project these colours in an avian‐appropriate, tetrachromatic, colour space. We found that variants occupy different regions of the avian colour space and that between‐variant differences for most plumage patches could be discriminated by the birds. Differences in morphology were also detected, but these were, in general, smaller than colour differences. Overall, we found that variation in both plumage colour and morphology among variants is greater than would be expected if genetic drift alone was responsible for phenotypic divergence. As the plumage colour variants correspond to four geographical forms, our results suggest that phenotypic evolution in the Réunion grey white‐eye is at least partly explained by divergent selection in different habitats or regions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 459–473.     

Evolution of an invasive rodent on an archipelago as revealed by molar shape analysis: the house mouse in the Canary Islands

Aim The aim of this paper is to identify the patterns in the morphological differentiation in Canary Island mice, based on fossil and modern samples. In order to achieve this, the mouse species present on the archipelago were first compared with a set of continental mice. The differences between the continental and Canary Island samples, and among the Canary Island samples, provide insights into the processes of colonization and the subsequent insular evolution. Location Canary archipelago. Methods An outline analysis based on Fourier transformation was used to quantify shape differences between lower molars. Together with the fossil and modern Canary Island samples, a reference set of genotyped continental populations of the commensal Mus musculus and the wild Mus spretus was used for comparison. Results The morphometric analysis showed that all the mouse specimens from the Canary Islands and Cape Verde belonged to Mus musculus domesticus. Lower molars of extant mice from La Gomera, El Hierro, Gran Canaria, Tenerife, and to a lesser degree from Lanzarote, were similar to those of genotyped M. m. domesticus from the continent, while teeth of extant mice from Fuerteventura were more divergent. Fossil mice from Fuerteventura were very similar to the extant representatives on this island, and similar to the fossil mice on the nearby islands of Lobos and La Graciosa. Main conclusions The mouse present on the Canary archipelago has been identified as the house mouse M. m. domesticus. Based on the shape of the lower molar, the Canary Island mice are divergent from the continental ones, but the degree of divergence varies with the geography of the archipelago. Overall, populations from eastern islands are more divergent from the continental mice than populations from western ones. Fossil populations indicate that this situation was established several centuries ago. Two main factors may have contributed to this pattern: the appearance of different types of environment on the islands since the successful settlement of the mouse, and/or the number of subsequent introductions of continental individuals via shipping.     

Rapid parallel evolution of aberrant traits in the diversification of the Gulf of Guinea white-eyes (Aves, Zosteropidae)

Archipelago-endemic bird radiations are familiar to evolutionary biologists as key illustrations of evolutionary patterns. However, such radiations are in fact rare events. White-eyes (Zosteropidae) are birds with an exceptionally high colonization and speciation potential; they have colonized more islands globally than any other passerine group and include the most species-rich bird genus. The multiplication of white-eye island endemics has been consistently attributed to independent colonizations from the mainland; the white-eyes of the Gulf of Guinea archipelago had been seen as a classic case, spanning as great a breadth of phenotypic diversity as the family worldwide. Contrary to this hypothesis, our molecular phylogenetic analysis places the Gulf of Guinea white-eyes in just two radiations, one grouping all five oceanic island taxa and the other grouping continental island and land-bridge taxa. Numerous 'aberrant' phenotypes (traditionally grouped in the genus Speirops) have evolved independently over a short space of time from nonaberrant (Zosterops) phenotypes; the most phenotypically divergent species have separated as recently as 0.22 Ma. These radiations rival those of Darwin's finches and the Hawaiian honeycreepers in terms of the extent of adaptive radiation per unit time, both in terms of species numbers and in terms of phenotypic diversity. Tempo and patterns of morphological divergence are strongly supportive of an adaptive radiation in the oceanic islands driven by ecological interactions between sympatric white-eyes. Here, very rapid phenotypic evolution mainly affected taxa derived from the youngest wave of colonization, in accordance with the model of asymmetric divergence owing to resource competition in sympatry.     

GENETIC DIVERGENCE IN PERIPHERALLY ISOLATED POPULATIONS OF CHAFFINCHES IN THE ATLANTIC ISLANDS

Peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Canaries, Madeira, and Azores were compared genetically with their putative ancestral stock in Iberia and Morocco, and with a population of blue chaffinches (F. teydea) from Tenerife, using protein electrophoresis of 42 loci. The continental populations are only weakly differentiated genetically (F_ST = 0.092), despite distinctive subspecific differences in plumage and morphometrics between Iberia and Morocco populations. Estimated levels of gene flow among continental populations are high enough to account for their relative genetic homogeneity, and it is unlikely that homogenizing selection is operating to mimic the effects of gene flow. In contrast, the Atlantic island populations are well differentiated genetically (F_ST = 0.321), and have diverged considerably from their continental conspecifics. The development of significant genetic differentiation within the Canaries but not the Azores likely results from smaller population sizes, very restricted gene flow, and enhanced random drift in the former populations. There is no convincing evidence in support of stronger directional selection acting on genotypes or phenotypes to reduce within-population variability in Canaries populations as proposed by Grant (1979), although other tenets of his model of island evolution are supported by our analysis. Although genetic variability is reduced in four of the Canaries populations, only the Hierro population appears to have encountered a severe bottleneck. Yet it has not differentiated markedly from the La Palma population to which it is subspecifically allied. We conclude that gradual divergence in isolated populations of small to moderate size is the most plausible explanation for the evolution of intraspecific and interspecific diversity in Atlantic island chaffinches.     

Do assemblages of Coregonus (Teleostei: Salmoniformes) in the Central Alpine region of Europe represent species flocks?

To examine models of evolution for Coregonus from the Central Alpine region of Europe, 20 populations from nine lakes were assessed for variation at six microsatellite DNA loci. Patterns of variation were tested against three evolutionary models: phenotypic plasticity, multiple invasions of lakes by divergent forms, and within-lake radiation of species flocks. All sympatric and all but one allopatric pairs of populations were significantly divergent in allele frequencies. Pairwise F -statistics indicated reduced gene flow among phenotypically divergent sympatric populations. These results reject the hypothesis that within-lake morphological and ecological diversity reflects phenotypic plasticity within a single gene pool. Genetic similarity was higher among forms within lakes than between populations of the same form in different lakes. Among-lake divergence was primarily a product of allele size differences. Mantel tests contrasting patterns of genetic divergence against patterns predicted from the multiple invasions and species flocks models indicated that the latter is the best explanation of the observed genetic variation. Thus, reproductively isolated species diverged within lakes, with similar patterns repeatedly emerging among lakes. While this study argues for a particular mode of evolution in Central Alpine Coregonus , the taxonomy of these forms remains unresolved.     

Persistent inter‐ and intraspecific gene exchange within a parallel radiation of caterpillar hunter beetles (Calosoma sp.) from the Galápagos

When environmental gradients are repeated on different islands within an archipelago, similar selection pressures may act within each island, resulting in the repeated occurrence of ecologically similar species on each island. The evolution of ecotypes within such radiations may either result from dispersal, that is each ecotype evolved once and dispersed to different islands where it colonized its habitat, or through repeated and parallel speciation within each island. However, it remains poorly understood how gene flow during the divergence process may shape such patterns. In the Galápagos islands, three phenotypically similar species of the beetle genus Calosoma occur at higher elevations of different islands, while lowlands are occupied by a fourth species. By genotyping all major populations within this radiation for two nuclear and three mitochondrial gene fragments and seven microsatellite markers, we found strong support that the oldest divergence separates the highland species of the oldest island from the remaining species. Despite their morphological distinctness, highland species of the remaining islands were genetically closely related to the lowland population on each island and within the same magnitude as lowland populations sampled at different islands. Repeated evolution of highland ecotypes out of the lowland species appears the most likely scenario and estimates of geneflow rates revealed extensive admixture among ecotypes within islands, as well as between islands. These findings indicate that gene exchange among the different populations and species may have shaped the phylogenetic relationships and the repeated evolution of these ecotypes.     

Genetic structure of Gymnures (genus Hylomys; Erinaceidae) on continental islands of Southeast Asia: historical effects of fragmentation

Eustatic sea level changes during Pleistocene climatic fluctuations produced several cycles of connection-isolation among continental islands of the Sunda shelf. To explore the potential effects of these fluctuations, we reconstructed a model of the vicariant events that separated these islands, based on bathymetric information. Among many possible scenarios, two opposite phylogenetic patterns of evolution were predicted for terrestrial organisms living in this region: one is based on the classical allopatric speciation mode of evolution, while the other is the outcome of a sequential dispersal colonization of the archipelago. We tested the applicability of these predictions with an analysis of sequence variation of the cytochrome b gene from several taxa of Hylomys. They were sampled throughout SE-Asia and the Sunda islands. High levels of haplotype differentiation characterize the different island taxa. Such levels of differentiation support the existence of several allopatric species, as was suggested by previous allozyme and morphological data. Also in accordance with previous results, the occurrence of two sympatric species from Sumatra is suggested by their strongly divergent haplotypes. One species, Hylomys suillus maxi, is found both on Sumatra and in Peninsular Malaysia, while the other, H. parvus, is endemic to Sumatra. Its closest relative is H. suillus dorsalis from Borneo. Phylogenetic reconstructions also demonstrate the existence of a Sundaic clade composed of all island taxa, as opposed to those from the continent. Although there is no statistical support for either proposed biogeographic model of evolution, we argue that the sequential dispersal scenario is more appropriate to describe the genetic variation found among the Hylomys taxa. However, despite strong differentiation among island haplotypes, the cladistic relationships between some island taxa could not be resolved. We argue that this is evidence of a rapid radiation, suggesting that the separation of the islands may have been perceived as a simultaneous event rather than as a succession of vicariant events. Furthermore, the estimates of divergence times between the haplotypes of these taxa suggest that this radiation may actually have predated the climatic fluctuations of the Pleistocene. Further refinement of the initial palaeogeographic models of evolution are therefore needed to account for these results.     

Diversity despite dispersal: colonization history and phylogeography of Hawaiian crab spiders inferred from multilocus genetic data

The Hawaiian archipelago is often cited as the premier setting to study biological diversification, yet the evolution and phylogeography of much of its biota remain poorly understood. We investigated crab spiders (Thomisidae, Mecaphesa ) that demonstrate contradictory tendencies: (i) dramatic ecological diversity within the Hawaiian Islands, and (ii) accompanying widespread distribution of many species across the archipelago. We used mitochondrial and nuclear genetic data sampled across six islands to generate phylogenetic hypotheses for Mecaphesa species and populations, and included penalized likelihood molecular clock analyses to estimate arrival times on the different islands. We found that 17 of 18 Hawaiian Mecaphesa species were monophyletic and most closely related to thomisids from the Marquesas and Society Islands. Our results indicate that the Hawaiian species evolved from either one or two colonization events to the archipelago. Estimated divergence dates suggested that thomisids may have colonized the Hawaiian Islands as early as ~10 million years ago, but biogeographic analyses implied that the initial diversification of this group was restricted to the younger island of Oahu, followed by back-colonizations to older islands. Within the Hawaiian radiation, our data revealed several well-supported genetically distinct terminal clades corresponding to species previously delimited by morphological taxonomy. Many of these species are codistributed across multiple Hawaiian Islands and some exhibit genetic structure consistent with stepwise colonization of islands following their formation. These results indicate that dispersal has been sufficiently limited to allow extensive ecological diversification, yet frequent enough that interisland migration is more common than speciation.     

Ecomorphological patterns in the Blackcap Sylvia atricapilla: insular versus mainland populations

Capsule Blackcap Sylvia atricapilla populations from the Azores archipelago show morphological differences to continental birds which are consistent with the ‘Island Rule’.

Aims The morphology of insular vertebrates is usually the result of the evolution in their particular environment and leads to predictable morphological patterns, according to the Island Rule. We test the predictions of the Island Rule, using the Blackcap of the Azores archipelago as our model.

Methods We compared morphological variation (body size and wing shape) of populations from the nine islands of the Azores to continental birds, using multivariate indexes. Also, we looked at the relationship between these patterns and possible insular ecological drivers of morphological divergence.

Results Our findings are concordant with Island Rule predictions, as in general birds from the Azores are larger than continental populations, especially birds from the most distant islands. Wing shape also differs significantly, as Azorean Blackcaps tend to have rounder wings than continental birds with a migratory-like phenotype.

Conclusion Overall, we conclude that the observed morphological patterns in Blackcap in the Azores conform in general to the Island Rule predictions.     

HOMOGENEITY OF THE GENETIC VARIANCE-COVARIANCE MATRIX FOR ANTIPREDATOR TRAITS IN TWO NATURAL POPULATIONS OF THE GARTER SNAKE THAMNOPHIS ORDINOIDES

Quantitative genetic models of evolution rely on the genetic variance-covariance matrix to predict the phenotypic response to selection. Both prospective and retrospective studies of phenotypic evolution across generations rely on assumptions about the constancy of patterns of genetic covariance through time. In the absence of robust theoretical predictions about the stability of genetic covariances, this assumption must be tested with empirical comparisons of genetic parameters among populations and species. Genetic variance-covariance matrices were estimated for a suite of antipredator traits in two populations of the northwestern garter snake, Thamnophis ordinoides. The characters studied include color pattern and antipredator behaviors that interact to facilitate escape from predators. Significant heritabilities for all traits were detected in both populations. Genetic correlations and covariances were found among behaviors in both populations and between color pattern and behavior in one of the populations. Phenotypic means differed among populations, but pairwise comparisons revealed no heterogeneity of genetic parameters between the populations. The structure of the genetic variance-covariance matrix has apparently not changed significantly during the divergence of these two populations.     

THE CONTINENTAL ASIAN ELEMENT IN THE FAUNA OF THE PHILIPPINES AS EXEMPLIFIED BY COLADENIA MOORE, 1881 (LEPIDOPTERA: HESPERIIDAE)1

Abstract — The Philippines are an archipelago composed of oceanic islands originating from different island arcs, and a few pieces of continental origin. The continental fragments can be found in N Palawan, the Calamian Islands, W Mindoro and W Panay. This study aims at answering the question whether traces can be found of a terrestrial fauna travelling on these fragments to the east to become part of the Philippine fauna. A phylogenetic analysis of the genusColadenia(Lepidoptera: Hesperiidae) based on morphological characters has been carried out. Three methods for detecting distribution patterns have been applied and compared, viz. Page's Tree Reconciliation, Brooks Parsimony Analysis, and a method similar to character optimization and here called Area Optimization. It is concluded that there is evidence of a continental element in the Philippine fauna. The (expected) generality of this pattern can only be demonstrated when more phylogenetic analyses of relevant groups become available.     

Testing multivariate patterns of within-island differentiation in Podarcis dugesii from Madeira

Podarcis dugesii is a polymorphic species found on the ecologically-heterogeneous Atlantic island of Madeira. Patterns of geographic variation in size-free body dimensions (body shape), scalation, and size were portrayed on maps with isophene contours. There was no dominant pattern of among-site variation as seen in several other island lizards, although some linear combinations of morphological variables showed evidence of clines towards the east coast. Multivariate statistical techniques were developed to evaluate whether morphological variation contained an ecology-related spatial pattern, after removal of the effects of statistical non-independence of studied populations. Specifically, a partial correlation matrix comparison method provided evidence to support a proposed elevation model of among-site divergence in scalation and body size, with both an alternative rainfall model and spatial non-independence held constant. Similar divergence in scalation has been observed in other lizard species on other island archipelagos as well on some continental areas. The rainfall model was rejected for both body shape and scalation. The degree of within-population polymorphism in body shape and scalation showed no clear geographic trends in general, although the elevation and rainfall models could not be rejected for female scalation.