Landscape‐scale life‐history gradients in New Zealand freshwater fish

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Cranial morphology of the plagiosaurid Gerrothorax pulcherrimus as an extreme example of evolutionary stasis

Schoch, R.R. & Witzmann, F. 2011: Cranial morphology of the plagiosaurid Gerrothorax pulcherrimus as an extreme example of evolutionary stasis. Lethaia, Vol. 45, pp. 371–385. The plagiosaurid Gerrothorax pulcherrimus from the Triassic of Greenland and Germany is represented by skulls ranging from 4 to 12 cm in length and sheds light on ontogeny, individual variation, and variation in time and space. Ontogeny was remarkably stable in G. pulcherrimus, with the smallest known specimens resembling the adults closely in most features. A true ontogenetic change is evident in the ornament of dermal bones, in that the smallest specimens have ridges whereas in the successively larger ones, pustules spread over increasingly larger areas. The skull becomes proportionally longer, and the adductor chambers relatively narrower. The positive allometry of both the orbits and the interpterygoid vacuities suggests that the eye supporting musculature – rather than the jaw adductors – increased proportionally during growth. Individual, not age‐related variation in the dermal skull roof affects partial fusion of parietals, presence and extent of the interfrontoparietal, and the morphological pattern of the posterior skull table. The ventral surface of the basal plate of the parasphenoid ranges from smooth over poorly to heavily ornamented or dentigerous. Considering the impressive longevity of more than 35 Myr, the morphological changes of G. pulcherrimus are minor. Our ecological interpretation for G. pulcherrimus is that it relied on the permanent presence of water, but was flexible with respect to the size and nature of the water body as well as to changes in salinity. The unparalleled extent of evolutionary stasis may therefore be based on the ecological flexibility of this morphologically so tightly constrained temnospondyl. □Ecological flexibility, ontogeny,Temnospondyli, Triassic, variation.     

Predicting the ecological impacts of a new freshwater invader: functional responses and prey selectivity of the ‘killer shrimp’, Dikerogammarus villosus,compared to the native Gammarus pulex

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The influence of ecological and life history factors on ectothermic temperature–size responses: Analysis of three Lycaenidae butterflies (Lepidoptera)

Body size has been shown to decrease with increasing temperature in many species, prompting the suggestion that it is a universal ecological response. However, species with complex life cycles, such as holometabolous insects, may have correspondingly complicated temperature–size responses. Recent research suggests that life history and ecological traits may be important for determining the direction and strength of temperature–size responses. Yet, these factors are rarely included in analyses. Here, we aim to determine whether the size of the bivoltine butterfly, Polyommatus bellargus, and the univoltine butterflies, Plebejus argus and Polyommatus coridon, change in response to temperature and whether these responses differ between the sexes, and for P. bellargus, between generations. Forewing length was measured using digital specimens from the Natural History Museum, London (NHM), from one locality in the UK per species. The data were initially compared to annual and seasonal temperature values, without consideration of life history factors. Sex and generation of the individuals and mean monthly temperatures, which cover the growing period for each species, were then included in analyses. When compared to annual or seasonal temperatures only, size was not related to temperature for P. bellargus and P. argus, but there was a negative relationship between size and temperature for P. coridon. When sex, generation, and monthly temperatures were included, male adult size decreased as temperature increased in the early larval stages, and increased as temperature increased during the late larval stages. Results were similar but less consistent for females, while second generation P. bellargus showed no temperature–size response. In P. coridon, size decreased as temperature increased during the pupal stage. These results highlight the importance of including life history factors, sex, and monthly temperature data when studying temperature–size responses for species with complex life cycles.     

Why selection favors protandrous sex change for the parasitic isopod, Ichthyoxenus fushanensis (Isopoda: Cymothoidae)

A flesh burrowing parasitic isopod, Ichthyoxenus fushanensis, was found infecting the body cavity of a freshwater fish, Varicorhinus bacbatulus, in pairs. The marked sexual size dimorphism, with much larger females than males, and the presence of penes vestige on mature females suggest a protandrous sex change in I. fushanensis. Here we investigate the question of why selection favors protandrous sex change for I. fushanensis, by analyzing the interactions among clutch size, female size, male size, and their host size. The number of manca, the first free-living juvenile stage released, per brood was closely related to the size of the female. Excluding the effects of interaction among causal variables, the negative correlation of male size alone on clutch size suggests that a small male did not limit an individual's mating and fertilization success. When the effect of host size is removed statistically, there exists a significant negative relationship between the sizes of paired males and females. This indicates that the resources available from host fish are limited, and that competition exists between paired male and female resulting in a trade-off of body size. Due to the very low success rate of hunting for a host of mancas, a female with larger body size and higher fecundity has a fitness advantage. To augment the clutch size, a productive combination is a smaller male and a larger female in a host. The constraints of the limited resources and the trade-off between the sizes of paired male and female may favor I. fushanensis to adopt the reproductive strategy of protandrous sex change resulting in a larger female and hence more mancas. The pattern of the interactions among male, female, and the number of mancas, may be considered as a selective force for I. fushanensis protandrous sex change, where the available resources are constrained by the size of the host. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inducible defences in Daphnia: responses to two closely related predator species

The predator-induced responses of two species of Australian Daphnia, with contrasting distributions and life history patterns, to the kairomones of two species of Anisops predators, were measured. Daphnia longicephala produced a large crest and attained a larger size when exposed to both predators. D. carinata sl matured earlier than D. longicephala and did not produce a crest. Surprisingly, kairomones of both predators inhibited the production of ephippia in D. carinata sl. Anisops stali, the larger of the two predator species, induced a significantly larger crest size in D. longicephala, and larger brood size in both species compared with the smaller A. gratus, indicating a quantitative but not qualitative effect of predator species on inducible defences. Received: 18 August 1999 / Accepted: 14 April 2000     

Sexual dimorphism in the skull geometry of newt species of <Emphasis Type="Italic">Ichthyosaura,Triturus</Emphasis> and <Emphasis Type="Italic">Lissotriton</Emphasis> (Salamandridae,Caudata, Amphibia)

In this study, we applied geometric morphometrics to explore variations in the level and pattern of sexual size dimorphism (SSD) and sexual shape dimorphism (SShD) of the ventral cranium in three different Modern Eurasian newt taxa (Ichthyosaura alpestris, Triturus species group and Lissotriton vulgaris). The ventral cranium is the part of the skull that is more directly related to foraging and feeding. Our results indicate that the level and pattern of sexual dimorphism in the ventral cranium differ among Modern Eurasian newt taxa. Regarding sexual dimorphism in skull size, Ichthyosaura alpestris and Triturus species show female-biased patterns (females are larger than males), whereas Lissotriton vulgaris appears to be non-dimorphic in skull size. In I. alpestris and Triturus species, SShD is mostly absent, whereas in L. vulgaris, SShD is more pronounced. A high level of variation between populations in both SSD and SShD indicates that local conditions may have a profound effect on the magnitude and direction of sexual dimorphism. The significant sexual differences in ventral cranium size and shape indicate possible subtle intersexual differences in ecological demands due to diet specialisation, in spite of similar general ecological settings.     

Characteristics of pollen from natural triploid Betula hybrids

Birch has a key role in the Holocene vegetation history of northern Europe and in sub‐arctic climates dwarf birch and tree birch co‐exist. In Iceland, triploid hybrids between diploid Betula nana (dwarf birch) and tetraploid B. pubescens (downy birch) are common and therefore likely to contribute to pollen deposition. Pollen from 22 triploid trees/shrubs from ten woodlands in Iceland was examined and its size and shape compared with pollen from the parental species. The mean diameter of pollen grains from the triploid hybrids was not statistically different from that of B. nana pollen, but was significantly smaller than the mean value of B. pubescens pollen. On the contrary, the size of the vestibulum was similar to that of B. pubescens, which was significantly greater than that of B. nana, and therefore the diameter‐pore depth ratio was lower than the values from either species. The pattern of size distribution within plants indicated that triploid hybrids might have produced two sizes of triporate pollen grains, but the small B. nana size was far more prevalent than the larger B. pubescens size. Several anomalies in pollen morphology were common among the hybrid pollen grains: four or more pores were the most frequent type of abnormality. Characteristics of the pollen of triploid Betula hybrids, especially structural anomalies, may provide a means to reveal periods of interspecific hybridisation in the analysis of sub‐fossil pollen.     

Rapid evolution and the convergence of ecological and evolutionary time

Recent studies have documented rates of evolution of ecologically important phenotypes sufficiently fast that they have the potential to impact the outcome of ecological interactions while they are underway. Observations of this type go against accepted wisdom that ecological and evolutionary dynamics occur at very different time scales. While some authors have evaluated the rapidity of a measured evolutionary rate by comparing it to the overall distribution of measured evolutionary rates, we believe that ecologists are mainly interested in rapid evolution because of its potential to impinge on ecological processes. We therefore propose that rapid evolution be defined as a genetic change occurring rapidly enough to have a measurable impact on simultaneous ecological change. Using this definition we propose a framework for decomposing rates of ecological change into components driven by simultaneous evolutionary change and by change in a non‐evolutionary factor (e.g. density dependent population dynamics, abiotic environmental change). Evolution is judged to be rapid in this ecological context if its contribution to ecological change is large relative to the contribution of other factors. We provide a worked example of this approach based on a theoretical predator–prey interaction [ Abrams, P. & Matsuda, H. (1997) . Evolution, 51, 1740], and find that in this system the impact of prey evolution on predator per capita growth rate is 63% that of internal ecological dynamics. We then propose analytical methods for measuring these contributions in field situations, and apply them to two long‐term data sets for which suitable ecological and evolutionary data exist. For both data sets relatively high rates of evolutionary change have been found when measured as character change in standard deviations per generation (haldanes). For Darwin's finches evolving in response to fluctuating rainfall [ Grant, P.R. & Grant, B.R. (2002) . Science, 296, 707], we estimate that evolutionary change has been more rapid than ecological change by a factor of 2.2. For a population of freshwater copepods whose life history evolves in response to fluctuating fish predation [ Hairston, N.G. Jr & Dillon, T.A. (1990) . Evolution, 44, 1796], we find that evolutionary change has been about one quarter the rate of ecological change – less than in the finch example, but nevertheless substantial. These analyses support the view that in order to understand temporal dynamics in ecological processes it is critical to consider the extent to which the attributes of the system under investigation are simultaneously changing as a result of rapid evolution.     

How to be an invasive gammarid (Amphipoda: Gammaroidea)–comparison of life history traits

About six life history and two ecological traits of gammarid species occurring in Central European waters were compared in order to identify the characters of successful invader. The species were (1) natives: Gammarus fossarum, G. pulex, G. lacustris, G. varsoviensis, G. balcanicus, G. leopoliensis, G. roeselii, and (2) aliens: Gammarus tigrinus, Chaetogammarus ischnus (=Echinogammarus ischnus), Pontogammarus robustoides, Obesogammarus crassus, Dikerogammarus haemobaphes, D. villosus. Generally the alien species were characterised by a combination of large brood size, high partial fecundity, early maturation and by appearance of higher number of generations per year. Also, these species presented higher tolerance towards severe environmental conditions, i.e. elevated salinity and human degradation of the environment. The above features seem to facilitate the colonisation of new areas and competition with native species -- a phenomenon that has been currently observed in various parts of Europe.     

Body Size and the Maintenance of Reproductive Isolation in Stickleback,Genus Gasterosteus

The blackspotted stickleback Gasterosteus wheatlandi and the widely studied threespine stickleback G. aculeatus are sympatric throughout the former’s range and share many aspects of life history and reproductive behaviour. These two species differ significantly in size, with G. wheatlandi of both sexes measured at approximately 60% of the standard length of their G. aculeatus counterparts. This study concentrated on G. wheatlandi courtship behaviour and investigated its role in the maintenance of reproductive isolation with G. aculeatus. Specifically, the roles that (1) female body size plays in influencing male courtship preferences and (2) male body size and behaviour play in female courtship preferences were investigated through dummy and live conspecific and heterospecific stimulus presentations. Male G. wheatlandi courtship preferences are consistent with previously described patterns for G. aculeatus. Males of both species preferentially approach and court the larger of two simultaneously presented live or dummy females. Thus, the smaller G. wheatlandi males are indiscriminate with respect to assortative mate choice; not only preferring to approach and court more fecund conspecific females but, more significantly, G. aculeatus‐sized females. In contrast, females of both species demonstrate strong assortative courtship preferences. When presented with pairs of flask‐enclosed males, females of both species preferentially orient and court the conspecific male over the heterospecific. Similarly, when presented with a conspecific male and a heterospecific male presented singly, females prefer to enter the nest of the conspecific. Systematic analysis of the interactions between these pairs of fish (one male, one female) demonstrates that the breakdown of courtship in heterospecific courtship occurs late in the courtship sequence when the widely differing forms of male leading behaviour results in drastically differing female responses. I suggest that, as previously described in G. aculeatus, the supernormality effect plays a significant role in mediating adaptive mate choice behaviour in G. wheatlandi. However, the added element of a larger sympatric species introduces a possible cost in time and energy devoted to courting heterospecific, and sympatric, females that the larger G. aculeatus do not likely incur. There is substantial evidence from many sympatric G. aculeatus species pairs that there is assortative mate choice based on size and/or courtship behaviour. Courtship trials suggest a more pervasive role for females in assortative mate choice. Whether it is male body size per se, or in combination with behaviour, morphology or other cues, is unresolved in the present study.     

Diet, fecundity and egg size in some polyphagous predatory carabid beetles

Egg size was measured at different rates of egg laying in three polyphagous carabid species, known to be useful predators of cereal aphids; the small Bembidion lampros Herbst and the medium-sized Pterostichus cupreus L. and P. melanarius Illiger. Variations in fecundity, as well as the ability of the medium-sized species to also build up fat reserves, were obtained when beetles were subjected to different dietary regimes consisting of aphids, or foods with a lower or higher protein content. Egg size was found to be dependent on the rate of egg laying within a species. A diet of cereal aphids appeared to be adequate for egg production in these polyphagous carabids, but female P. cupreus were unable to build up fat reserves when they ingested aphids contaminated with the aphicide pirimicarb. Beetles were able to devote resources to more and larger eggs (B. lampros), or to larger eggs and/or fat reserves (P. melanarius/P. cupreus) when given access to a carbohydrate-rich food with low protein content. The highest rate of egg laying was obtained when female P. cupreus and P. melanarius were given a more varied diet at frequent intervals; including regular shifts between unsprayed aphids, carbohydrate-rich food and protein-rich maggots. Within the varied diet treatment a negative relationship was obtained between egg size and egg number among similar-sized individuals of P. cupreus and P. melanarius; females producing the largest number also laid the smallest eggs. Egg size affected larval survival, since first instars hatching from large eggs were found to survive longer than those hatching from small eggs. The influence of differences in food intake on reproduction, maintenance metabolism, and survival of fieldinhabiting carabids is discussed.     

Influence of selected factors on the dietary compositions of three targeted and co‐occurring temperate species of reef fishes: implications for food partitioning

The dietary compositions of three medium to large targeted fish species, which co‐occur over reefs in temperate waters of south‐western Australia, were determined. These data were then used to ascertain statistically the extent to which body size, season and habitat influence the diets of these species and the degree to which food resources were partitioned among and within those species, and thus reduced the potential of interspecific and intraspecific competition. On the west coast, Bodianus frenchii (Labridae) and Epinephelides armatus (Serranidae) spent their whole life over prominent limestone reefs, as did Glaucosoma hebraicum (Glaucosomatidae) in all but juvenile life, when it lived over low‐relief, limestone substrata. The dietary composition of each species changed with increasing body size, which, in G. hebraicum, was particularly pronounced at c. 300 mm total length (L_T) and therefore at the size when this species shifts habitat. When the three species co‐occurred over the same reefs, their dietary compositions were significantly different, with that of B. frenchii being by far the most discrete, reflecting a far greater contribution by sedentary taxa. Thus, the diet of B. frenchii was distinguished from those of the other two species in containing substantial volumes of bivalve and gastropod molluscs and echinoid echinoderms and essentially no teleosts. Although the diets of G. hebraicum and particularly E. armatus were dominated by teleosts, and especially for larger individuals, the former species ingested greater volumes of cephalopods and small crustaceans. The pointed jaws of B. frenchii, with their forwardly directed and interlocking anterior incisors, are ideally adapted for biting and retaining their invertebrate prey, which are attached to or reside within reef crevices. In contrast, the mouths of G. hebraicum and E. armatus are broader and rounder and contain numerous small, slender and inward‐pointing teeth. These teeth, in conjunction with prominent backward‐curved canines in E. armatus, facilitate the capture and retention of fish prey. Observations in situ indicate that G. hebraicum is a suction feeder, while E. armatus is predominantly a ram feeder. Although reef environments on the west and south coasts differ, the diet of B. frenchii on these coasts differed only slightly. Interspecific differences in diet, combined with size‐related changes in dietary compositions and the occupation of different habitats by juvenile and adult G. hebraicum, reduce the potential for competition for food resources among and within B. frenchii, G. hebraicum and E. armatus and thus helps facilitate the coexistence of these species which historically have been abundant over reefs in south‐western Australia.
     

Body mass and correlated ecological variables in the North American muskrat lineage: evolutionary rates and the tradeoff of large size and speciation potential

This study tracks evolutionary change in body mass (W) and correlated ecological variables over the 3.75 million year history of the North American muskrat (Ondatra zibethicus). A new model is presented suggesting muskrat body mass has been in equilibrium for most of its history. Four pulses of pronounced size increase are correlated with glacial dynamics and volcanic events. Ranges of evolutionary rates in darwins and a new metric based on percent change in W document episodic size change. Proportional size change is independent of interval length, with a background range attributed to natural selection ≤25–30%. In increasing body mass by a factor of ten to about 1 kg mass-specific metabolism was halved, home range quadrupled, population density decreased fourfold, and average biomass more than doubled. Estimates of species diversity in ancient cotton rats (Sigmodon) and muskrats are calculated from a function derived from the correlation of numbers of North American rodent species and mean W. The phyletic mode of muskrat body size increase is explained as a combination of large body size reducing speciation coupled with an aquatic lifestyle. To the ecological consequences of large size in evolving clades (Cope’s rule) we can now add reduced speciation potential.     

Ecological and Phylogenetic Correlates to Body Size in the Indriidae

I investigated ecological and phylogenetic correlates to body size variations in 10 taxa of extant Indriidae (Indri, Avahi, and Propithecus). I also tested for phylogenetic niche conservatism as a model for the evolution of indriid body size. Phylogenetic niche conservatism refers to the shared attributes that related taxa have acquired because they tend to have occupied similar niches during their evolutionary history. I collected species-specific data on body mass, climate, density, and chemical properties of food items from the literature. I used 2 phylogenies in independent contrasts methods to control for phylogenetic relationships (Indri and Propithecus as sister taxa vs. Indri basal taxa to all indriids). Multivariate models indicated that lemur density and resource quality are the strongest ecological correlates to indriid body size variations. Partitioning methods revealed that 52.4–67% of indriid body size variation is explained by phylogenetic niche conservation. Thus, indriid body size variations may be the result of stabilizing selection. Though it is possible to identify constraints on lower than average body size, there are few data on selection against larger than average body size in indriids. Large body size in subfossil lemurs further complicates identification of constraints on larger than average body size in extant indriids. Researchers using independent contrast methods to control for phylogeny should be aware that some ecology-phenotype relationships are best explained as the result of the synergistic effects of ecology and phylogeny.     

Comparisons of the food niches of three native and two introduced fish species in an Australian river

Synopsis The breadth, correspondence and overlap of the diets of the small and large size classes of the three native species (Galaxias occidentalis, Bostockia porosa and Edelia vittata) and two introduced species (Gambusia holbrooki and Perca fluviatilis) of fishes found in the shallows of the main channel and in the tributary creeks of a south-western Australian river have been compared in each season. Classification and ordination were used to examine the overall interrelationships of the diets across species, size groups, seasons and the location where the fish were caught (channel or creek). The smaller fish had a narrower dietary breadth than larger fish in the spring and summer, presumably reflecting the size limit imposed on prey size by their possession of a relatively small mouth in these seasons. Intraspecific dietary overlap between large and small size classes was usually high in G. occidentalis, but generally low in G. holbrooki and P. fluviatilis, and also in B. porosa and E. vittata when the difference between the lengths of the two size groups was greatest. Dietary overlap was least in autumn when the main prey taxa were most abundant. During winter, the diets of the three native species in the tributaries converged, probably reflecting a relatively low faunal diversity in these highly seasonal water bodies. The only relatively consistent interspecific overlap in diet was between B. porosa and E. vittata. Classification and ordination of the dietary samples separated G. occidentalis (which fed extensively on terrestrial organisms from the water surface) from the smaller P. fluviatilis (that concentrated on copepods in the plankton) and from B. porosa and E. vittata (which ingested primarily benthic organisms). Furthermore, B. porosa tended to ingest larger prey taxa than E. vittata. The diet of Gambusia holbrooki is sometimes dominated by terrestrial insects and at other times by benthic organisms, demonstrating that this species is an opportunistic carnivore. It is concluded that food partitioning by the three native and two introduced fish species in the Collie River is likely to be one of the principal factors facilitating the coexistence of substantial populations of these species in this system.     

Ecological change predicts population dynamics and genetic diversity over 120 000 years

While ecological effects on short‐term population dynamics are well understood, their effects over millennia are difficult to demonstrate and convincing evidence is scant. Using coalescent methods, we analysed past population dynamics of three lizard species (Psammodromus hispanicus, P. edwardsianus, P. occidentalis) and linked the results with climate change data covering the same temporal horizon (120 000 years). An increase in population size over time was observed in two species, and in P. occidentalis, no change was observed. Temporal changes in temperature seasonality and the maximum temperature of the warmest month were congruent with changes in population dynamics observed for the three species and both variables affected population density, either directly or indirectly (via a life‐history trait). These results constitute the first solid link between ecological change and long‐term population dynamics. The results moreover suggest that ecological change leaves genetic signatures that can be retrospectively traced, providing evidence that ecological change is a crucial driver of genetic diversity and speciation.     

Sexual size dimorphism in caecilian amphibians: analysis, review and directions for future research

Sexual dimorphism, widespread in the animal kingdom, describes differences between the sexes in size, shape and many other traits. Sexual size dimorphism (SSD) plays a significant role in understanding life history evolution and mating systems. The snakelike morphology of limbless caecilian amphibians lacking obvious secondary sexual characters (in contrast to frogs and salamanders) impedes accurate intrasexual comparisons. In this study, sexual size dimorphism in the oviparous caecilian Ichthyophis cf. kohtaoensis, a phylogenetically basal caecilian, was analysed. Females were larger in all body and head characters tested. However, when adjusted to body size (total length), females differed only in their cloacal shape. Clutch volume was positively correlated to female body size, thus female fecundity increased with body size supporting the hypothesis of a fecundity-selected SSD in the oviparous Ichthyophis cf. kohtaoensis. A review of the present SSD data for caecilians shows that many species are monomorphic for body size but show dimorphism in head size, while other species demonstrate female-biased SSD. Male-biased SSD has not been reported for caecilians. To understand life history evolution in caecilians, further studies on the reproductive biology of other taxa are urgently needed, in particular for rhinatrematids and uraeotyphlids. New data will allow phylogenetically controlled comparative analyses to fully explore the pattern of SSD among caecilian lineages.     

Reflectance of sexually dichromatic UV‐blue patches varies during the breeding season and between two subspecies of Gallotia galloti (Squamata: Lacertidae)

Body coloration is sexually dimorphic in many vertebrate species, including lizards, in which males are often more conspicuous than females. A detailed analysis of the relative size of coloured patches and their reflectance, including the ultraviolet (UV) range, has rarely been performed. In the present work we quantified sexual dimorphism in body traits and surface area of all lateral patches from adult females and males of two subspecies of Gallotia galloti (G. g. galloti and G. g. eisentrauti). We also analysed the magnitude of sexual dichromatism in the UV‐visible reflectance of such patches and the changes in patch size and brightness during the reproductive season (April–July). Males had significantly larger patch areas (relative to their snout‐vent length) and higher brightness (mainly in the UV‐blue range) than did females in both subspecies. The comparison of relative patch areas among months did not reach statistical significance. However, patch brightness significantly changed during the breeding season: that of the UV‐blue (300–495 nm) range from lizards of the two subspecies was significantly larger in June than in April, while brightness in the 495–700 nm range in G. g. galloti was larger in May, June, and July than in April. A different pattern of dichromatism was also detected in the two populations, with G. g. eisentrauti being more sexually dichromatic than G. g. galloti. We discuss the results in terms of possible evolutionary causes for the sexual dichromatism related to different ecological characteristics of the habitats where each subspecies live. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 556–569.