Quantitative morphometrical analysis of a North African population of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: sexual dimorphism,and comparison with European populations

Genetic variability of quantitative traits was investigated in aMoroccan population of Drosophila melanogaster, with an isofemale line design. Results were compared with data previously obtained from French populations. Although the environmental and thermal conditions are very different in France and Morocco, only two significant differences were observed: a shorter wing and a lighter abdomen pigmentation in Morocco. It is, therefore, concluded that Moroccan D. melanogaster are quite typical temperate flies, belonging to the Palaearctic region, and very different from the ancestral Afrotropical populations. Almost all traits were genetically variable, as shown by significant intraclass correlations among lines. Genetic correlations were highly significant among three size-related traits, while much lower between size and bristle numbers. Fluctuating asymmetry was greater for abdominal bristles than for sternopleural bristles. Sex dimorphism, analysed as a female/male ratio, was identical in French and Moroccan populations. Examination of the thorax length/thorax width ratio showed that the thorax is more elongated in females. Sexual dimorphism of wing length was significantly more correlated to thorax width than to thorax length. The results illustrate the value of measuring numerous quantitative traits on the same flies for characterizing the genetic architecture of a natural population. In several cases, and especially for genetic correlations, some interesting suggestions could be made, which should be confirmed, or invalidated, by more extensive investigations.     

Transcriptomic analysis of skin pigmentation variation in the Virginia opossum (Didelphis virginiana)

Skin pigmentation and coat pigmentation are two of the best‐studied examples of traits under natural selection given their quantifiable fitness interactions with the environment (e.g., camouflage) and signalling with other organisms (e.g., warning coloration). Previous morphological studies have found that skin pigmentation variation in the Virginia opossum (Didelphis virginiana) is associated with variation in precipitation and temperatures across its distribution range following Gloger's rule (lighter pigmentation in temperate environments). To investigate the molecular mechanism associated with skin pigmentation variation, we used RNA‐Seq and quantified gene expression of wild opossums from tropical and temperate populations. Using differential expression analysis and a co‐expression network approach, we found that expression variation in genes with melanocytic and immune functions is significantly associated with the degree of skin pigmentation variation and may be underlying this phenotypic difference. We also found evidence suggesting that the Wnt/β‐catenin signalling pathway might be regulating the depigmentation observed in temperate populations. Based on our study results, we present several alternative hypotheses that may explain Gloger's rule pattern of skin pigmentation variation in opossum, including changes in pathogen diversity supporting a pathogen‐resistant hypothesis, thermal stress associated with temperate environments, and pleiotropic and epistatic interactions between melanocytic and immune genes.     

PLASTICITY VERSUS ENVIRONMENTAL CANALIZATION: POPULATION DIFFERENCES IN THERMAL RESPONSES ALONG A LATITUDINAL GRADIENT IN DROSOPHILA SERRATA

The phenotypic plasticity of traits, defined as the ability of a genotype to express different phenotypic values of the trait across a range of environments, can vary between habitats depending on levels of temporal and spatial heterogeneity. Other traits can be insensitive to environmental perturbations and show environmental canalization. We tested levels of phenotypic plasticity in diverse Drosophila serrata populations along a latitudinal cline ranging from a temperate, variable climate to a tropical, stable climate by measuring developmental rate and size-related traits at three temperatures (16°C, 22°C, and 28°C). We then compared the slopes of the thermal reaction norms among populations. The 16–22°C part of the reaction norms for developmental rate was flatter (more canalized) for the temperate populations than for the tropical populations. However, slopes for the reaction norms of the two morphological traits (wing size, wing:thorax ratio), were steeper (more plastic) in the temperate versus the tropical populations over the entire thermal range. The different latitudinal patterns in plasticity for developmental rate and the morphological traits may reflect contrasting selection pressures along the tropical–temperate thermal gradient.     

Realized evolvability: quantifying phenotypic evolution in a Drosophila clade

Understanding the evolutionary potential of morphological evolution is still a major problem in evolutionary biology. In this study, we tried to quantify the amount of variation of different traits among species of a Drosophila clade reared under standard conditions. Nineteen different traits have been measured on nine species of the same clade, the Neotropical saltans group of Drosophila. Measured traits can be distributed into five categories: size‐traits (wing and thorax), shape indices (ratios among the size traits), number of sternopleural bristles on the thorax, number of abdominal bristles on successive sternites, and dorsal pigmentation of abdomen. All species are of medium size with a generally dark pigmentation. A remarkable feature is the presence of numerous bristles on the 6th sternite of the males, while this segment is bare in other Drosophila species. A multivariate analysis revealed that it was possible to discriminate all the investigated species by using the complete set of measured traits. For each trait, phenotypic variability was investigated at the within‐ and between‐species levels. As a rule, the interspecific coefficient of variation (CV) was much greater than the within species CV, and it is proposed to call it realized evolvability. All possible correlations were calculated between traits across species, providing many unexpected results. Size traits were highly correlated among them, but not correlated with shape indices. Abdominal traits (bristles and pigmentation) were correlated, but often in opposite directions, with thoracic shape indices. Tergite pigmentation was negatively correlated with bristle number on sternite. For the moment, most of the correlations cannot be explained by developmental processes or parallel selective pressures. Nonetheless, mapping the evolution of the two characters on a molecular phylogeny of the studied species revealed two opposite phylogenetic trends for abdominal pigmentation and setation, respectively. Our data suggest a need for similar studies in other well‐defined Drosophila clades.     

Latitudinal variability of Drosophila melanogaster: allozyme frequencies divergence between European and Afrotropical populations

Allelic frequencies at five polymorphic loci were determined in seven European and six Afrotropical populations of Drosophila melanogaster. African populations, which may be considered as ancestral for the species, showed a greater genetic diversity as measured by the number of alleles found. Within each geographic group (Europe or tropical Africa) genetic distances between local populations were very small (D = 0.027). By contrast, the average distance between European and African populations (D = 0.389) was more than 12 times bigger. It was previously known that various morphological or physiological differences, which probably reflect genetic adaptations to different environments, exist between these temperate and tropical populations. Data presented here suggest that the divergence in allozyme frequencies also reflects some selective mechanisms.     

Genetics and the origin of human races

In the last decades, the concept of human races was considered scientifically unfounded as it was not confirmed by genetic evidence. None of the racial classifications, which strongly differ in the number of races and their composition, reflects actual genetic similarity and genealogy of human populations inferred from variability of classical markers and DNA regions. Moreover, intercontinental ("interracial") variability was shown to be far lower than that within populations: the former constitutes 7 to 10% and the latter, about 85% of the total genetic variation. It is believed that the low level of differentiation of regional population groups contradicts their race status and suggests a recent origin of humans from one ancestral population. The results of studies of various genetic systems are in agreement with last conclusion rejecting the hypothesis of regional continuity. According to this hypothesis, the populations of continents regarded as large races have developed during long evolution from local types of archaic humans, in particular, Neanderthals. Phenotypic similarity of different, sometimes unrelated, populations united into one "race" is explained by strong selection since race-diagnostic traits characterize body surface and thus are directly subjected to the influence of environmental (primarily climatic) factors. It has been recently established that variability of the most important of these traits, body and hair pigmentation, is largely controlled by one locus (MC1R), which accounts for its high evolutionary lability. Other traits used for race identification are also likely to be labile and controlled by major genes. However, the fact that the currently existing race classifications are groundless does not mean that such classifications are impossible in principle. Commonly used argumentation (races do not exist because populations are not genetically separated) does not hold water. A polytypic species is characterized by genetic continuity of allopatric populations rather than the presence of narrow genetic boundaries between them. Borderlines between races are usually conventional and arbitrary. As to intergroup variation in humans, it is indeed low but comparable with that in some other species. There are no obstacles to the development of genetic systematics of human races.     

A genomewide admixture mapping study for yield factors and morphological traits in a cultivated cocoa (Theobroma cacao L.) population

The selection of productive varieties of modern Criollo cocoa, showing fine aromatic qualities in their beans, is of major interest for some producing countries, such as Venezuela. Cultivated populations of Modern Criollo or Trinitario varieties may be suitable for admixture mapping analysis, as large blocks of alleles derived from two identified divergent ancestors, recently admixed, are still preserved, after a few generations of recombination, similar to experimental mapping progenies. Two hundred and fifty-seven individuals from a cultivated population of Modern Criollo were selected and analysed with 92 microsatellite markers distributed along the genome. This population exhibited a wide range of variability for yield factors and morphological features. Population structure analysis identified two main subgroups corresponding to the admixture from the two ancestors Criollo and Forastero. Several significant associations between markers and phenotypic data (yield factors and morphological traits) were identified by a least squares general linear model (GLM) taking into account the population structure and the percentage of admixture of each individual. Results were compared with classical QTL analyses previously reported for other cacao populations. Most markers associated to quantitative traits were very close to QTLs detected formerly for the same traits. Associations were also identified between markers and several qualitative traits including the red pigmentation observed in different organs, mainly associated to common markers in linkage group 4.     

Flight morphology corresponds to both surrounding landscape structure and local patch conditions in a highly specialized peatland butterfly (Lycaena epixanthe)

1. Movement mediates the response of populations and communities to landscape and habitat spatial structure, yet movement capability may itself be modified by selection pressures accompanying landscape change. Insect flight morphology can be affected by both the landscape surrounding habitat patches and the distribution of resources within habitat patches. 2. This study investigated the relative influence of local habitat patch conditions and surrounding landscape structure on variation in morphological traits associated with flight in the bog copper (Lycaena epixanthe), a butterfly endemic to temperate Nearctic peatlands. 3. Eight habitat patches were sampled to assess the influence of the surrounding landscape (connectivity of potential habitat and matrix composition) and patch size (an integrated proxy of resource density and spatial distribution) on investment into flight, measured by thorax and abdomen mass, and wing area. 4. The results revealed an effect of both local habitat conditions and landscape structure on flight‐related morphological traits. Increasing forest cover in the surrounding landscape, indicative of increased habitat patch isolation, corresponded with less mobile phenotypes in both sexes. Surrounding landscapes with more water were also generally associated with less mobile phenotypes. Investment into flight was greater in smaller peatlands in which host plant density is higher and more homogeneously distributed. 5. The present study highlights that morphological traits associated with mobility may be responding to both local habitat patch characteristics and surrounding landscape structure. It also supports the hypothesis that local habitat conditions contribute to morphological variation in butterflies.     

Latitudinal variability of Drosophila melanogaster: Allozyme frequencies divergence between European and Afrotropical populations

Allelic frequencies at five polymorphic loci were determined in seven European and six Afrotropical populations of Drosophila melanogaster. African populations, which may be considered as ancestral for the species, showed a greater genetic diversity as measured by the number of alleles found. Within each geographic group (Europe or tropical Africa) genetic distances between local populations were very small (D=0.027). By contrast, the average distance between European and African populations (D=0.389) was more than 12 times bigger. It was previously known that various morphological or physiological differences, which probably reflect genetic adaptations to different environments, exist between these temperate and tropical populations. Data presented here suggest that the divergence in allozyme frequencies also reflects some selective mechanisms.     

Divergence of the seasonal races of chum salmon, <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> Walbaum, 1792, in the Amur and Poronai rivers: Ecology,genetics, and morphology

The chum salmon of the Amur River (the mainland part of the Far East) and the Poronai River (Terpeniya Bay, Sakhalin Island) are historically related to one another, as the drainage basins of these rivers are the remnants of a formerly single river system, the Paleoamur, which existed when Sakhalin Island was a part of the continent. Both river populations of chum salmon consist of the early-run and late-run ecological forms (seasonal races), which are also referred to as the summer and autumn races. They are reproductively isolated from each other due to their spawning at different times and in different types of spawning grounds. To assess the direction, pattern, and degree of divergence between these chum salmon races in the both river fragments since the Paleoamur, it is necessary to compare them using two types of traits: selectively neutral DNA markers and morphological and physiological traits, variations in which may have an adaptive value. For this, we have studied chum salmon from both rivers in terms of microsatellite DNA markers, body counts and measurements, body weight, and fecundity. Both in the Amur River and in the Poronai River, the autumn race of chum salmon prevails over the summer race in body length and weight, fecundity, number of pyloric caeca, and several other meristic traits. The intra-basin differences between the races are much more pronounced in the Amur chum salmon. The inter-race differences in microsatellites are also greater in the Amur chum salmon compared to the Poronai chum salmon. Using microsatellites, three levels of differentiation have been revealed: (1) between the basins of the Amur and Poronai rivers, (2) between the races within each of the river basins, (3) and between population samples within each race of each basin. A hypothesis is proposed that the currently existing races of chum salmon in the Amur and Poronai rivers have evolved since the breakup of the Paleoamur, and the intra-basin divergence of the races started in the Amur River earlier than in the Poronai River. An analysis of our own data and the published data suggests that the adaptation of the seasonal races of chum salmon to the conditions of their spawning grounds is determined by a complex of morphological and physiological traits, including the number of pyloric caeca, which is an adaptive and highly heritable trait associated with the incubation temperature of the water.     

Intraspecific Genetic Admixture and the Morphological Diversification of an Estuarine Fish Population Complex

The North-east American Rainbow smelt (Osmerus mordax) is composed of two glacial races first identified through the spatial distribution of two distinct mtDNA lineages. Contemporary breeding populations of smelt in the St. Lawrence estuary comprise contrasting mixtures of both lineages, suggesting that the two races came into secondary contact in this estuary. The overall objective of this study was to assess the role of intraspecific genetic admixture in the morphological diversification of the estuarine rainbow smelt population complex. The morphology of mixed-ancestry populations varied as a function of the relative contribution of the two races to estuarine populations, supporting the hypothesis of genetic admixture. Populations comprising both ancestral mtDNA races did not exhibit intermediate morphologies relative to pure populations but rather exhibited many traits that exceeded the parental trait values, consistent with the hypothesis of transgressive segregation. Evidence for genetic admixture at the level of the nuclear gene pool, however, provided only partial support for this hypothesis. Variation at nuclear AFLP markers revealed clear evidence of the two corresponding mtDNA glacial races. The admixture of the two races at the nuclear level is only pronounced in mixed-ancestry populations dominated by one of the mtDNA lineages, the same populations showing the greatest degree of morphological diversification and population structure. In contrast, mixed-ancestry populations dominated by the alternate mtDNA lineage showed little evidence of introgression of the nuclear genome, little morphological diversification and little contemporary population genetic structure. These results only partially support the hypothesis of transgressive segregation and may be the result of the differential effects of natural selection acting on admixed genomes from different sources.     

Complex trait differentiation between host-populations of the pea aphid Acyrthosiphon pisum (Harris): implications for the evolution of ecological specialisation

Variation in traits affecting preference for, and performance on, new habitats is a key factor in the initiation of ecological specialisation and adaptive speciation. However, habitat and resource use also involves other traits whose influence on ecological and genetic divergence remains poorly understood. In the present study, we investigated the extent of variation of life-history traits among sympatric populations of the pea aphid Acyrthosiphon pisum , which shows several host races that are specialised on various plants of the family Fabaceae plants and is an established model for ecological speciation. First, we assessed the community structure of microbial partners within host populations of the pea aphid. The effect of these microbes on host fitness is uncertain, although there is growing evidence that they may modulate various important adaptive traits of their host such as plant utilisation and resistance against natural enemies. Second, we performed a multivariate analysis on several ecologically relevant features of host populations recorded in the present and previous studies (including microbial composition, colour morph, reproductive mode, and male dispersal phenotype), enabling the identification of correlations between phenotypic traits. We discuss the ecological significance of these associations of traits in relation to the habitat characteristics of pea aphid populations, and their consequences for the evolution of ecological specialisation and sympatric speciation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 718–727.     

Variability in fruit and seed morphology among and within populations of Plathymenia (Leguminosae-Mimosoideae) in areas of the Cerrado, the Atlantic forest, and transitional sites

Plathymenia reticulata is a tree species that occurs in two different Brazilian biomes, the Cerrado (a savannah environment), and the Atlantic Forest. In the present study, we evaluated morphological variation within and among five populations located in these vegetation types and in transitional sites in order to test the hypothesis that habitat selective pressures, being different in the Cerrado and the Atlantic Forest, would cause adaptive differences in morphological traits in individuals occurring under these different circumstances. Thirteen morphological traits of fruits, seeds, and of the membranous endocarp were obtained from 30 fruits and 20 seeds from each of nine to 10 individuals per population. Significant variation was found for all traits while comparing individuals within populations, and most traits varied significantly among populations as well. Some traits differed significantly between forest and Cerrado populations, while transition sites showed intermediate patterns and higher within-population variation. Contrary to our hypothesis, variation in seed size and mass among populations from different habitats was not significant. However, as predicted, the membranous endocarp was shorter for the seeds from Cerrado populations than from the Atlantic Forest. These data suggest the existence of P. reticulata ecotypes from Cerrado and Atlantic Forest; the relevance to Plathymenia evolution and to its wide ecological distribution is discussed.     

BRINGING HABITAT INFORMATION INTO STATISTICAL TESTS OF LOCAL ADAPTATION IN QUANTITATIVE TRAITS: A CASE STUDY OF NINE‐SPINED STICKLEBACKS

Detection of footprints of historical natural selection on quantitative traits in cross‐sectional data sets is challenging, especially when the number of populations to be compared is small and the populations are subject to strong random genetic drift. We extend a recent Bayesian multivariate approach to differentiate between selective and neutral causes of population differentiation by the inclusion of habitat information. The extended framework allows one to test for signals of selection in two ways: by comparing the patterns of population differentiation in quantitative traits and in neutral loci, and by comparing the similarity of habitats and phenotypes. We illustrate the framework using data on variation of eight morphological and behavioral traits among four populations of nine‐spined sticklebacks (Pungitius pungitius). In spite of the strong signal of genetic drift in the study system (average F_ST = 0.35 in neutral markers), strong footprints of adaptive population differentiation were uncovered both in morphological and behavioral traits. The results give quantitative support for earlier qualitative assessments, which have attributed the observed differentiation to adaptive divergence in response to differing ecological conditions in pond and marine habitats.     

Developmental constraints on an adaptive plasticity: reaction norms of pigmentation in adult segments of Drosophila melanogaster

SUMMARY Variation of dark pigmentation according to developmental temperature was investigated in two geographic populations (France and India) with the isofemale line technique (20 lines for each population). The response curves called the reaction norms, were established in females for seven different segments: the mesothorax and abdomen segments 2–7 (Abd 2–7). In all cases the response curves were non-linear and had to be described either by a quadratic convex polynomial for thorax and Abd 2–5, or by a cubic polynomial for Abd 6 and 7. Among abdomen segments, increasing antero-posterior gradients were observed for several traits, including average pigmentation, overall phenotypic plasticity, the temperature of minimum pigmentation, and the curvature parameter of quadratic norms. Genetic correlations between abdomen segments were high when adjacent segments were considered, but became nil when more distant segments were correlated, suggesting that different pigmentation genes are expressed in the anterior and the posterior part of the abdomen. Characteristic values of reaction norms provided information either on trait value (i.e., the extension of pigmentation) or on plasticity. Correlations between plasticity and pigmentation were generally low and non-significant, suggesting their genetic independence. The overall darker pigmentation which is observed at low temperatures is assumed to be an adaptive plasticity. However, the differences which are evidenced among segments reveal strong interactions with developmental genes. These interactions are less likely to be a consequence of natural selection and are better interpreted as developmental constraints. The reaction norms analysis reveals the complexity of these interactions and should help, in the future, in the identification of the responsible thermosensitive genes.     

Light body pigmentation in indianDrosophila melanogaster: A likely adaptation to a hot and arid climate

We analysed reaction norms of pigmentation (thorax and abdomen) according to growth temperature for 20 isofemale lines collected near Delhi (India) and compared them to results obtained for two French populations. The climatic conditions of the two locations were strongly different, with monthly average temperature ranging between 4.2°C and 20.5°C in France and between 14.3°C and 34.3°C in India. For each segment, a decrease of the pigmentation was observed with increasing temperature and the shapes of the reaction norms were more or less parallel. On average Indian flies were lighter than French ones, in agreement with the thermal budget hypothesis. We further investigated the shapes of reaction norms by polynomial adjustment and observed significant differences. In several cases, a maximum divergence was observed at high temperature, implying a change in the shape of the norm. Characteristic values related to the thermal reactivity were also significantly different between populations but no general tendency was found. Genetic variability, estimated by the coefficient of intraclass correlation, was significantly lower in India (0.27 ±0.026) than in France (0.39 ±0.028), and we discuss the significance of this difference.     

Morphological divergence within and between the Mexican sailfin mollies, Poecilia velifera and Poecilia petenensis

This study examined the patterns of morphological variation both between species, and between sexes and among populations within each species of the Mexican sailfin mollies, Poecilia velifera and Poecilia petenensis , using geometric morphometrics and linear measurements of morphological traits. While sexes within each species differed in characteristics that may be important in sexual selection, such as length of the dorsal fin, species differed in traits, such as body depth, that may also be influenced by natural selection due to differences in habitats. Within each species, many morphological traits were similar among populations, but important differences, including caudal peduncle depth in P. petenensis (but not in P. velifera ), suggested that habitat differences may also be important in shaping population divergence independently within each species. Indeed, the evolutionary vectors of male morphological population divergence for each species differed by an angle of 98·5°, representing nearly orthogonal vectors and suggesting independent shape divergence between these two molly species. Finally, geographic isolation did not explain the morphological differentiation seen among populations, suggesting that natural and sexual selection were strong forces promoting morphological diversification within these two species, despite the potential for a high degree of population connectivity and gene flow.     

Differentiation in morphology and habitat partitioning of genetically characterized Gammarus fossarum forms (Amphipoda) across a contact zone

Evolutionary processes do not affect all character types equally. Depending on environmental effects, morphological and ecological evolution may be uncoupled from molecular evolution. A natural contact zone between two forms of the amphipod Gammarus fossarum was used to assess the levels of differentiation in genetic, morphological and habitat characters. About 50 individuals per population were analysed for six enzyme loci in 72 populations and four size-standardized morphometric traits in a subset of 32 populations. Nine environmental parameters per population were used to characterize the habitat. All three character sets revealed significant divergences between G. fossarum forms. The eastern form A of G. fossarum differs from the western form B by longer setae at the second antennae (males) and longer exopodits at die third uropods (males and females). The variance partitioning within populations, among populations and between forms, was compared within the three character sets. The relative variance component between forms in genetic characters appeared to be 10-fold greater in comparison to morphological traits, indicating morphological stasis during speciation. The inter-form environmental differentiation was shown to be low, but consistent between pure populations. The eastern form A seems to prefer higher altitudes, a substrate type of gravel, stones and leaves, shading, and water with higher conductivity and pH man the western form B.     

Many ways to make darker flies: Intra‐ and interspecific variation in Drosophila body pigmentation components

Body pigmentation is an evolutionarily diversified and ecologically relevant trait with substantial variation within and between species, and important roles in animal survival and reproduction. Insect pigmentation, in particular, provides some of the most compelling examples of adaptive evolution, including its ecological significance and genetic bases. Pigmentation includes multiple aspects of color and color pattern that may vary more or less independently, and can be under different selective pressures. We decompose Drosophila thorax and abdominal pigmentation, a valuable eco‐evo‐devo model, into distinct measurable traits related to color and color pattern. We investigate intra‐ and interspecific variation for those traits and assess its different sources. For each body part, we measured overall darkness, as well as four other pigmentation properties distinguishing between background color and color of the darker pattern elements that decorate each body part. By focusing on two standard D. melanogaster laboratory populations, we show that pigmentation components vary and covary in distinct manners depending on sex, genetic background, and temperature during development. Studying three natural populations of D. melanogaster along a latitudinal cline and five other Drosophila species, we then show that evolution of lighter or darker bodies can be achieved by changing distinct component traits. Our results paint a much more complex picture of body pigmentation variation than previous studies could uncover, including patterns of sexual dimorphism, thermal plasticity, and interspecific diversity. These findings underscore the value of detailed quantitative phenotyping and analysis of different sources of variation for a better understanding of phenotypic variation and diversification, and the ecological pressures and genetic mechanisms underlying them.     

Phenotypic and genetic divergence among harbour porpoise populations associated with habitat regions in the North Sea and adjacent seas

Determining the mechanisms that generate population structure is essential to the understanding of speciation and the evolution of biodiversity. Here, we investigate a geographical range that transects two habitat gradients, the North Sea to North Atlantic transition, and the temperate to subpolar regions. We studied the harbour porpoise (Phocoena phocoena), a small odontocete inhabiting both subpolar and temperate waters. To assess differentiation among putative populations, we measured morphological variation at cranial traits (N = 462 individuals) and variation at eight microsatellite loci for 338 of the same individuals from Norwegian, British and Danish waters. Significant morphological differentiation reflected the size of the buccal cavity. Porpoises forage in relatively shallow waters preying mainly on benthic species in British and Danish waters, and on mesopelagic and pelagic fish off the coast of Norway. We suggest that the observed differentiation may be explained by resource specialization and either adaptation or developmental responses to different local habitats.