Shell size and shape in Madeiran land snails: do niches remain unfilled?

The distribution of shell heights and diameters in the mainly endemic Madeiran land snail fauna shows the bimodal pattern of high- and low-spired shells found in many other faunas. Field and laboratory studies show that shell shape is associated with the angle of substrate on which the snails crawl; as elsewhere, tall spired species use vertical surfaces or burrow in soft material. Flattened species predominate on horizontal surfaces, while globular species are less specific in their preferences. Detailed comparisons with the fauna of N.W. Europe show that the proportion of high-spired species in the Madeiran fauna is low, and large high-spired species associated with vertical surfaces are very few in number despite an apparent abundance of suitable habitats. Amongst low-spired species, one family, the Helicidae, dominates the Madeiran fauna. While the overall distribution of size in these species is much as in Europe, Madeiran helicids extend into smaller size classes than do those in Europe, and they appear to fill a gap in the scatter created by the absence of other families.
Non-endemic species, other than those strictly associated with man-made environments, are generally small in size. In the upper scatter, their size distribution parallels that of endemics, but in the lower scatter they constitute the whole of the smallest size classes.
The role of interspecific competition in determining these distributions is discussed. The range of helicid sizes is compatible with a relaxation of competition or predator pressure relative to other areas, but in the upper scatter there appear to be gaps in the range of size and shape expected despite a long period in which the fauna could evolve. This could indicate the existence of adaptive troughs blocking, or delaying, radiation over the full spectrum of size and shape.     

MODERN AND HISTORICAL EVIDENCE FOR NATURAL HYBRIDIZATION BETWEEN SYMPATRIC SPECIES IN MANDARINA (PULMONATA: CAMAENIDAE)

Geographical and temporal variation in gene exchange between two endemic land snail species, Mandarina aureola and Mandarina ponderosa, was studied on Hahajima Island of the Bonin Islands. Allozyme variation in modern samples, and variation in the color and shell morphology of modern and fossil samples, suggest a complex geographical and historical pattern of hybridization. These two species occur in sympatry, and their shell morphologies and protein genotypes are markedly divergent. However, many specimens of M. aureola, collected from the middle region of the island, exhibit intermediate shell morphologies and possess marker alleles of M. ponderosa. Fossil samples of the two species strongly suggest that these intermediates were hybrids with M. ponderosa that were produced since the end of the Pleistocene. Each of these species, in addition, is subdivided into two genetically and morphologically divergent parapatric races. Interspecific hybridization appears to have produced genetical and morphological admixture among these four distinctive groups of populations. The past distribution and geographic variation of M. ponderosa can be traced in the distribution of M. ponderosa-derived genotypes in current populations of M. aureola. Temporal changes of the color pattern in the fossil populations of hybrids suggest that the traits introduced from M. ponderosa to M. aureola have been affected by natural selection and could replace traits of living species when advantageous. Moreover, these introgressed genes appeared to provide novel properties that enabled M. aureola to advance into a new environment. Relatively independent change in shell color and morphology further suggests mosaic evolution following the hybridization events. Connectively, these data reveal how hybridization events may be an important source of evolutionary novelties and make it clear that the phenomenon of reticulate evolution cannot be ignored.     

Novel colour polymorphisms in a hybrid zone of Mandarina (Gastropoda: Pulmonata)

A hybrid zone of the land snails Mandarina mandarina and Mandarina chichijimana in the oceanic Bonin Islands was studied morphologically and genetically to show the potential of hybridization as a source of morphological novelties. These species are clearly distinguishable on the basis of allozymes, colour polymorphisms, shell form and genital morphology, but exhibit a hybrid zone from the northeast to southwest of Chichijima island. There is a cline in the frequency of the colour patterns characteristic of each of the species, and specimens with intermediate colour pattern on the shells appear in the hybrid zone. Not only specimens with colour patterns that are overlapping of the patterns of the two species, but also specimens with unique colour patterns appear in the hybrid populations. These unique colour patterns are not found in the pure populations of both species or other Mandarina species in Chichijima Islands, and it is suggested that these are produced by the hybridization. Because of the appearance of many types of unique colour patterns, variability of the colour polymorphism in the hybrid populations are remarkably higher than that in the pure populations. This result suggests that the novel morphology is produced by the hybridization between species with distinctive morphology. This reveals the importance of hybridization as a source of morphological variation, diversity and evolutionary novelty.     

Morphological characterization of a brown lemur hybrid zone (Eulemur rufifrons × E. cinereiceps)

Hybridization has recently been identified as a pervasive force in the evolution of primates. In this study, we characterized a hybrid zone between two species of brown lemur (Eulemur rufifrons and E. cinereiceps) in the Andringitra region of southeastern Madagascar using morphological traits. We immobilized animals along a north-south transect (～80 km), scored them for their degree of hybridity using pelage traits and measured standard morphometric variables. Results from our study suggest that hybridization between E. rufifrons and E. cinereiceps is extensive, with the hybrid zone extending over 42.6 km and being composed mostly of later generation hybrids. We also identified significant variation between ancestral groups in our study: hybrid males exhibited longer tails than both parental species and sexual dimorphism in upper canine height favoring males was documented in E. rufifrons. These patterns could suggest that gene flow between parental and hybrid populations is relatively limited. Finally, significant differences between ancestral groups in relative body mass and skin-fold thickness were absent in our study, indicating that, as measured by these proxies, hybrids are equally as fit as parental forms. Based on these preliminary findings, the Andringitra hybrid zone could conform to the bounded superiority model of hybrid zone stability (i.e., it could be being maintained by selection favoring hybrids within transitional habitats). Accordingly, hybrids in Andringitra may be an unusual case among primates, representing a stable recombinant but distinct lineage. This conclusion has important implications for evolutionary processes within the brown lemur species complex.     

Shell thickness in the land snail Theba pisana (Pulmonata: Helicidae)

Shell thickness in Theba pisana was measured in samples from Britain, northern France and the Mediterranean region. Mean adult thickness varied between about 0.08 mm and 0.20 mm. These values are much lower than previously published values. The considerable variation within samples was related to shell size, periods of growth, and resorption of shell material during breeding. Variation between samples was not great. This contrasts with previous work which linked shell thickness variation to large-scale climatic effects. Perhaps availability of calcium may be more important than climate in determining shell thickness in Theba Pisana .     

Robustness, evolvability, and neutrality

Biological systems, from macromolecules to whole organisms, are robust if they continue to function, survive, or reproduce when faced with mutations, environmental change, and internal noise. I focus here on biological systems that are robust to mutations and ask whether such systems are more or less evolvable, in the sense that they can acquire novel properties. The more robust a system is, the more mutations in it are neutral, that is, without phenotypic effect. I argue here that such neutral change--and thus robustness--can be a key to future evolutionary innovation, if one accepts that neutrality is not an essential feature of a mutation. That is, a once neutral mutation may cause phenotypic effects in a changed environment or genetic background. I argue that most, if not all, neutral mutations are of this sort, and that the essentialist notion of neutrality should be abandoned. This perspective reconciles two opposing views on the forces dominating organismal evolution, natural selection and random drift: neutral mutations occur and are especially abundant in robust systems, but they do not remain neutral indefinitely, and eventually become visible to natural selection, where some of them lead to evolutionary innovations.     

Rapid adaptive evolution of northeastern coyotes via hybridization with wolves

The dramatic expansion of the geographical range of coyotes over the last 90 years is partly explained by changes to the landscape and local extinctions of wolves, but hybridization may also have facilitated their movement. We present mtDNA sequence data from 686 eastern coyotes and measurements of 196 skulls related to their two-front colonization pattern. We find evidence for hybridization with Great Lakes wolves only along the northern front, which is correlated with larger skull size, increased sexual dimorphism and a five times faster colonization rate than the southern front. Northeastern haplotype diversity is low, suggesting that this population was founded by very few females moving across the Saint Lawrence River. This northern front then spread south and west, eventually coming in contact with an expanding front of non-hybrid coyotes in western New York and Pennsylvania. We suggest that hybridization with wolves in Canada introduced adaptive variation that contributed to larger size, which in turn allowed eastern coyotes to better hunt deer, allowing a more rapid colonization of new areas than coyotes without introgressed wolf genes. Thus, hybridization is a conduit by which genetic variation from an extirpated species has been reintroduced into northeastern USA, enabling northeastern coyotes to occupy a portion of the niche left vacant by wolves.     

Differences in shape and size of skull and mandible in Talpa species (Mammalia: Eulipotyphla) from Turkey

We analysed with landmark-based images morphological differences between four species of Talpa which resemble each other morphologically and are all highly adapted to underground life. Subtle shape differences of the skull and mandibular bones were found between all species. However, there is also broad overlap between all species. Talpa caucasica had the largest skull and mandibles, and Talpa levantis the smallest.     

Biochemical variation and shell shape in populations of the fresh-water snail Lymnaea peregra (Mollusca, Gastropoda, Pulmonata) from south-west Ireland

Lymnaea peregra was collected at sites, predominantly from the Caha plateau, in south-west Ireland. These included sites investigated by previous workers and found to contain populations with a distinctive pattern of variation in shell-shape. Normally spired populations and non-spired ( involuta, sub-involuta ) could occur in adjacent loughs in the same watershed. The biochemical variation in some of these populations was investigated using horizontal starch-gel zone electrophoresis. Fourteen biochemical loci were investigated. Although circumscribed by difficulties in collecting and low sample numbers, the investigation showed that the pattern of variation in shell shape was still present and was not correlated or associated with the biochemical variation. While this did not support hypotheses of geographic race formation or incipient speciation, it was in agreement with the current view of variation and speciation in fresh-water pulmonates.     

Phenotypic plasticity in leaf morphology of Crataegus monogyna (Rosaceae): an experimental study with taxonomic implications

Many leaf characters are considered in the taxonomy of Crataegus in Europe, and several have been used in studies of the extent of hybridization in populations of northwest Europe. In such analyses it is assumed that the environmental component of phenotypic variation in such characters is insignificant. We tested this assumption by analysing the variation in the size and shape of leaves borne on clone cuttings of Crataegus monogyna maintained under identical conditions apart from the availability of soil nutrients. The resulting variation among leaves from this single genotype was as great as that observed previously across populations. Furthermore, although most of this variation could not be explained, a part could be attributed to differences in nutrient availability; of nine leaf characters investigated, eight showed significant variation due to this source, and in seven, the variation covaried significantly with nutrient level. The systematic implications of this are briefly explored.