Prehistoric anthropogenic introduction of partulid tree snails in Papua New Guinean archipelagos

Aim Members of the tropical tree snail family Partulidae are endemic to Pacific high oceanic islands and typically have single‐island ranges. Two nominal Papua New Guinean species, Partula carteriensis and Partula similaris, deviate from familial norms by having extensive multi‐island ranges that include low islands. We hypothesized that undocumented anthropogenic introductions may underlie this regional biogeographical anomaly and evaluated this hypothesis with novel field distributional and genotypic data. Location Papua New Guinean archipelagos between 1.4 and 11.4° S and 146.5 and 154.2° E. Methods Ethanol‐preserved museum lots of P. carteriensis (from New Britain, Bismarck Archipelago) and P. similaris (from Woodlark, Boiaboiawaga and Goodenough islands) were genotyped for a standard mitochondrial marker, cytochrome c oxidase subunit I (COI), and the resulting haplotypes were subjected to phylogeographical analyses. Results All four genotyped populations showed very little genetic or conchological differentiation, irrespective of nominal taxonomic status, the archipelago sampled or whether the island was low, high, oceanic or continental. Partula carteriensis and P. similaris exhibit atypical distributions on larger high islands, being restricted to coastal villages and absent from native forest. Main conclusions Our results strongly indicate that P. carteriensis and P. similaris are conspecific, although a formal taxonomic revision is beyond the scope of this present study. They collectively exhibit the most heterogeneous geographical range known among partulids and their explicitly synanthropic association with high island coastal villages strongly implicates human introduction as the regional dispersal mechanism. We currently lack insights into the timeframe (apart from regional prehistory) and cultural context of these translocations. We also lack a convincing source population, and it may be necessary to survey the partulid fauna of the neighbouring Solomon Islands to identify one. Partulids are critically endangered throughout much of their range and the discovery of populations that apparently thrive in human‐altered landscapes is noteworthy. Their study may provide clues of broad relevance to partulid conservation.     

Patterns of genetic variation in Pacific island land snails: the distribution of cytochrome b lineages among Society Island Partula

The radiation of Partula land snails has produced a large array of distinct morphological, ecological and behavioural types occupying many tropical volcanic islands in the Pacific Ocean. Within the Society Islands of French Polynesia, the mode of evolution is thought to have involved a single colonization event on each island, with later speciation occurring largely in situ. The present study examines genetic variation in the mitochondrial cytochrome b gene among taxa within the Society Island archipelago. Levels of intraspecific variation are found to be high, but variation among species is sometimes small. Mitochondrial variants do not always cluster according to island and some species are found to be polyphyletic in the cytochrome b tree, despite other morphological and molecular evidence that strongly supports their monophyly. A possible explanation for the polyphyly of species is that different variants are derived from ancestral mitochondrial polymorphisms that have been retained despite speciation events. Although it is possible that there has been some gene flow among islands, the distribution of mitochondrial lineages across islands strongly indicates that their origins predate the colonization of the islands in the study, and that they are very unlikely to have evolved entirely in situ.     

Experimental simulation of environmental warming selects against pigmented morphs of land snails

In terrestrial snails, thermal selection acts on shell coloration. However, the biological relevance of small differences in the intensity of shell pigmentation and the associated thermodynamic, physiological, and evolutionary consequences for snail diversity within the course of environmental warming are still insufficiently understood. To relate temperature‐driven internal heating, protein and membrane integrity impairment, escape behavior, place of residence selection, water loss, and mortality, we used experimentally warmed open‐top chambers and field observations with a total of >11,000 naturally or experimentally colored individuals of the highly polymorphic species Theba pisana (O.F. MÜller, 1774). We show that solar radiation in their natural Mediterranean habitat in Southern France poses intensifying thermal stress on increasingly pigmented snails that cannot be compensated for by behavioral responses. Individuals of all morphs acted neither jointly nor actively competed in climbing behavior, but acted similarly regardless of neighbor pigmentation intensity. Consequently, dark morphs progressively suffered from high internal temperatures, oxidative stress, and a breakdown of the chaperone system. Concomitant with increasing water loss, mortality increased with more intense pigmentation under simulated global warming conditions. In parallel with an increase in mean ambient temperature of 1.34°C over the past 30 years, the mortality rate of pigmented individuals in the field is, currently, about 50% higher than that of white morphs. A further increase of 1.12°C, as experimentally simulated in our study, would elevate this rate by another 26%. For 34 T. pisana populations from locations that are up to 2.7°C warmer than our experimental site, we show that both the frequency of pigmented morphs and overall pigmentation intensity decrease with an increase in average summer temperatures. We therefore predict a continuing strong decline in the frequency of pigmented morphs and a decrease in overall pigmentation intensity with ongoing global change in areas with strong solar radiation.     

Extinction or survival: partulid tree snails in American Samoa

Four partulid tree snail species are known from American Samoa. In 1998, we surveyed the recently established National Park (units on three islands: Tutuila, Tau, Ofu) and neighboring areas for partulids. On Tutuila, Samoana abbreviata, previously considered probably extinct, was extremely rare (15 snails seen); Samoana conica was more common (288 snails) but still rare; Eua zebrina was the most common (1102 snails), at one locality perhaps near its natural abundance. The species have similar distributions within the Park. All three have declined dramatically since the 1920s. Before 1980, when the predatory snail Euglandina rosea was introduced, habitat destruction, and perhaps rat predation and shell collecting, probably caused the decline. The Park provides protection to the Tutuila partulids by protecting habitat, although development is still a potential problem. But predation by E. rosea may yet cause their extinction. On Ofu (only outside the Park), 12 live Samoana thurstoni were found; 31 have now been recorded by western science. Also, an apparently robust population of E. zebrina, previously considered a Tutuila endemic, was found. Euglandina rosea is not on Ofu, so these populations are important remnants of the fauna. No partulids were found on Tau; none has ever been recorded there.     

Parallel evolution of sexual isolation in sticklebacks

Mechanisms of speciation are not well understood, despite decades of study. Recent work has focused on how natural and sexual selection cause sexual isolation. Here, we investigate the roles of divergent natural and sexual selection in the evolution of sexual isolation between sympatric species of threespine sticklebacks. We test the importance of morphological and behavioral traits in conferring sexual isolation and examine to what extent these traits have diverged in parallel between multiple, independently evolved species pairs. We use the patterns of evolution in ecological and mating traits to infer the likely nature of selection on sexual isolation. Strong parallel evolution implicates ecologically based divergent natural and/or sexual selection, whereas arbitrary directionality implicates nonecological sexual selection or drift. In multiple pairs we find that sexual isolation arises in the same way: assortative mating on body size and asymmetric isolation due to male nuptial color. Body size and color have diverged in a strongly parallel manner, similar to ecological traits. The data implicate ecologically based divergent natural and sexual selection as engines of speciation in this group.     

Spire index and preferred surface orientation in some land snails

Fourteen species of land snails have been tested for their preference for surfaces at 0, 90 or 180 degrees under laboratory conditions. They range from high-spired (height/breadth = 4.1) to discoidal forms (height/breadth = 0.4). There is a positive association of spire height with tendency to adopt the 90 degree surface. Species of intermediate (globular) shape show less specificity for a particular surface than high- or low-spired species. The exception is Helix aspersa , which behaves more like one of the high-spired species than like one of its similarly shaped relatives. The differences in preference will help to reduce interaction between co-existing species in the field.     

Ecology of a shell convergence between subfamilies of polygyrid land snails

The shell convergence between Neohelix dentifera and Inflectarius ferrissi is an important key to understanding the unusual evolutionary pattern of polygyrid land snails in eastern North America. This group is characterized by conchologically distinct shell-static clades that have radiated ecologically to the extent that shell shape and habitat are not detectably correlated at the species level; furthermore, entire shell-static clades converge conchologically between anatomically distinct lineages (triodopsines and polygyrines). The triodopsine N. dentifera and the polygyrine I. ferrissi both represent unique, drastic shifts from their shell-static clades, and thus are possible founders of new shell-static clades. This study shows that, within a context of high intraspecific variance, N. dentifera and I. ferrissi exhibit ecological parallelism or convergence in addition to conchological convergence. In coarse-grained ecology, both are more associated with acidic, anionic, deep, low- density soils in talus in high-altitude, exposed non-oak-hickory forests than are their close relatives, even though the two are not very close to each other. In fine-grained ecology, N. dentifera converges on I. ferrissi by increased rock-association relative to outgroups, and I. ferrissi likewise converges on N. dentifera by decreased log association. Thus the shell shape unique to these two species appears to be adaptive, whereas their intraspecific ecological variances appear high enough to permit ecological radiation.     

Population structure, history and gene flow in a group of closely related land snails: genetic variation in Partula from the Society Islands of the Pacific

Previous studies of Partula land snails from the Society Islands, French Polynesia, have shown that populations within species are highly differentiated in terms of their morphology, behaviour, ecology and molecular genetic variation. Despite this level of variability, differences between species are sometimes small, possibly reflecting the fact that reproductive isolation is not always complete and there exists the opportunity for genetic exchange between taxa through hybridization. The present study uses sequence data from a mitochondrial gene to further investigate genetic variation in Society Island Partula. Most populations are found in this study to be highly differentiated, but within individual species there seems to be no simple relationship either between genetic distance and geographical proximity, or between variation in mitochondria and that in allozymes or morphological characteristics. Among species there appears to be no simple correlation between degrees of reproductive isolation and genetic relatedness according to mitochondrial DNA. The results suggest that past events as well as ongoing drift and selection may have been important in affecting patterns of variation. Similarities among species at specific localities suggest that there must have been some genetic exchange in the past, although this may not necessarily reflect ongoing rates of hybridization. The discrepancy between results for different markers probably reflects the differential effects of drift and selection on mitochondrial and nuclear genes.     

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.     

Rapid evolution in unstable habitats: a success story of the polymorphic land snail Cepaea nemoralis (Gastropoda: Pulmonata)

The present study reports on a natural experiment with twelve replicates in which rapid, predictable and consistent divergence of Cepaea nemoralis populations occurred in response to repeated selection gradient of adjacent open and shaded habitats. Because the frequencies of various genetically‐based phenotypes varied widely among surveyed populations, and there was a large overlap between habitat types, no overall association with habitat was apparent. In paired comparisons, however, significant changes were consistently towards higher frequencies of light morphs in the open than corresponding shaded habitats, and this result is attributable to natural selection. This shows that the knowledge of the genetic composition of reference populations is often essential for discerning selection from random processes. At each site, a different morph combination contributed to the divergence of populations, indicating that there are many genetic solutions to similar ecological problems; this likely enhances the maintenance of high levels of polymorphism. Adaptation of populations occurred in contemporary time and was fast. In one case where it was possible to follow changes, significant shifts in morph frequencies occurred within just two snail generations (selection coefficients of 0.404 and 0.518). High evolvability may be one of the factors contributing to the ecological success of Cepaea nemoralis. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 251–262.     

Parallel genotypic adaptation: when evolution repeats itself

Until recently, parallel genotypic adaptation was considered unlikely because phenotypic differences were thought to be controlled by many genes. There is increasing evidence, however, that phenotypic variation sometimes has a simple genetic basis and that parallel adaptation at the genotypic level may be more frequent than previously believed. Here, we review evidence for parallel genotypic adaptation derived from a survey of the experimental evolution, phylogenetic, and quantitative genetic literature. The most convincing evidence of parallel genotypic adaptation comes from artificial selection experiments involving microbial populations. In some experiments, up to half of the nucleotide substitutions found in independent lineages under uniform selection are the same. Phylogenetic studies provide a means for studying parallel genotypic adaptation in non-experimental systems, but conclusive evidence may be difficult to obtain because homoplasy can arise for other reasons. Nonetheless, phylogenetic approaches have provided evidence of parallel genotypic adaptation across all taxonomic levels, not just microbes. Quantitative genetic approaches also suggest parallel genotypic evolution across both closely and distantly related taxa, but it is important to note that this approach cannot distinguish between parallel changes at homologous loci versus convergent changes at closely linked non-homologous loci. The finding that parallel genotypic adaptation appears to be frequent and occurs at all taxonomic levels has important implications for phylogenetic and evolutionary studies. With respect to phylogenetic analyses, parallel genotypic changes, if common, may result in faulty estimates of phylogenetic relationships. From an evolutionary perspective, the occurrence of parallel genotypic adaptation provides increasing support for determinism in evolution and may provide a partial explanation for how species with low levels of gene flow are held together.     

Gene flow and melanism in Lake Erie garter snake populations

Melanistic garter snakes ( Thamnophis sirtalis ) are unusually common near Lake Erie, apparently because selection for thermoregulatory ability in cool lake-shore habitats (which favours melanistic morphs) outweighs selection for crypsis (which favours striped morphs). However, morph frequencies are highly variable among sites, suggesting that random genetic drift also influences colour pattern. In an effort to better understand the evolutionary processes influencing garter snake colour patterns, we estimated Fx and Nm (the number of migrants per generation) among island and mainland populations from patterns of allozymic variation detected using electrophoresis. Estimates of Nm were high, ranging from 2.7 to 37.6 between pairs of study sites and making it unlikely that differences in morph frequencies among sites were solely the result of random genetic drift. Furthermore, differences in F _st estimates between colour pattern (a one-locus two-allele trait) and allozyme loci suggest that colour pattern alleles are not in Hardy-Weinberg equilibrium, most likely as a result of natural selection. Comparison of allozymic data from Lake Erie with those from more distant sites suggests that gene flow occurs over long distances in T. sirtalis.     

Morphology and aestivation behaviour in some Madagascan acavid land snails

Nine species of Madagascan acavid land snails were compared in a phylogenetic context. The two most plesiomorphic, Clauator johnsoni and C. moreleti, differ from the others by their high-spired shells, short tentacles, short tails, long necks, and crawling mode of hitching the shell along the ground. In the seven more apomorphic species, the crawling mode is smooth, with the shell resting on the tail, and the relative lengths of tail and shell correlate significantly. Among these seven species, three pairs of closest relatives (Helicophanta petiti and H. uesicalis, H. farafanga and H. souuerbiana, Ampeltta decaryi and A. julii) show evidence of phylogenetic constraints on ranked shell size. Aestivation site (as tentatively inferred from rare data) does not correlate with shell shape or size: burrowers have H/D = 2.7 to 0.6 and D = 70 to 25 mm; arboreals have H/D = 0.8 to 0.5 and D = 70 to 30 mm; the species with both the highest spire and the smallest diameter (C. moreleti) is neither a burrower or an arboreal, but stays on the ground surface. Inferred aestivation sites are randomly distributed phylogenetically. Climate shows no correlation, except that the arborcals are only from humid to wet regimes. Uniform shell colouration occurs only in burrowers (C. johnsoni, H. petiti, H. uestcalis), but disruptive shell colouration occurs in all others, including burrowers (H. farafanga, A. decaryt), ground-surface aestivators (C. moreleti), arboreals (H. souuerbiana, A. julii), and semi-arboreals (Ampefita subfunebris). Among all nine species, burrowers have significantly thicker shells (than their close relatives of similar size), wider bodies, and longer snouts than non-burrowers (H. souuerbiana is exceptional in being arboreal despite its huge size and in having the broad foot and snout of a burrower). Thus, although there is some evidence for phylogenetic constraints, natural selection for aestivation and crawling behaviours seems to have dominated the evolution of external body morphology and of shell thickness (but not shell size and shape) in these snails.     

Allozyme variation in sympatric populations of British grasshoppers,—evidence of natural selection

Seven species of British grasshopper were analysed for variation at 15 enzyme loci. Where possible, sympatric populations were collected so that direct comparisons could be made. Allozyme frequencies in Chorthippus brunneus and C. albomarginatus were compared using correlation analysis. A positive correlation was noted for a phosphoglucomutase allozyme and a negative correlation for an isocitrate dehydrogenase allozyme. It was suggested that natural selection may affect allozyme frequencies in the wild.
Using cluster analysis and principal component analysis, evolutionary relationships of grasshoppers have been examined and compared with interpretations based on morphology.     

Character displacement, frequency-dependent selection, and divergence of shell colour in land snails Mandarina (Pulmonata)

Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an exceptional example of habitat and character divergence among closely related species. In this study, microhabitat differences between sympatric ground-dwelling species were studied by distinguishing habitats on the basis of vegetation and types of litter. In all sites where two ground species coexisted, segregation occurred with each species showing preference for the microhabitat in which they were found. When they were in sympatry, one species was predominant in relatively wet and sheltered sites and the other in relatively dry and exposed sites. Although most species can live in both types of habitat, occupation by one species is inhibited by occupation by another. This suggests that competitive interaction between sympatric species caused segregation. Except for populations that have undergone interspecific hybridization, no examples were found of sympatric populations of two ground species sharing a similar shell colour. Species that were predominant in relatively wet and sheltered sites possessed shells with dark coloration and their colour patterns were mostly of one type. Species that were predominant in relatively dry and exposed sites possessed shells with bright coloration and their color patterns were polymorphic. Most populations from areas in which single species were distributed had shells with medium coloration. Microhabitat differentiation between sympatric species possibly caused diversification of shell colour, because bright shells are advantageous in sites where snails are largely exposed, and dark shells are advantageous in sites in where they are mostly sheltered from sunlight. In addition, frequency-dependent selection by predators hunting by sight may have operated to maintain colour polymorphism in the populations which are restricted to exposed habitats by competition with other sympatric species. This reveals the importance of interaction among closely related species as a cause of diversification in ecological and morphological traits.     

Allozyme variation in Atlantic salmon from the British Isles: associations with geography and the environment

Data on geographical variation in allele frequencies at enzyme coding loci in Atlantic salmon from the British Isles were collated from published and unpublished sources. Statistically significant differences in allele frequencies were found among samples both within and among river systems, suggesting that the Atlantic salmon in the British Isles is not a panmictic population and that even within major river systems it cannot be treated as a single genetic stock for fisheries management purposes. Although there was some evidence of regional differences in the frequency of some rare alleles, most single‐locus variation did not show strong geographic patterns, with the exception of the AAT‐4 * locus at which allele frequencies had a significant latitudinal cline. There was some evidence for the existence of genetically‐distinct celtic and boreal races of Atlantic salmon in the British Isles as previously has been suggested. Multiple regression analyses revealed associations between genetic variation and local environmental conditions ( i.e . between variation at MEP‐2 * and both temperature and local river gradient), providing additional evidence for adaptive population divergence in the species.     

Reviews of publications

Book reviewed in this article:
Island ecology (Outline studies in ecology), by M. Gorman
Comparative Morphology of Recent Crustacea, by Patsy A. McLaughlin. W. H. Freeman
Genetic Models of Sexual Selection, by Peter O'Donald
Pulmonates, edited by Vera Fretter and J. Peake
Weeds (2nd Edition), by W. C. Muenscher
Orang-Utan Behavior, by T. L. Maple
Aquatic Oligochaete Biology, edited by R. O. Brinkhurst & D. G. Cook
Studies in Mathematical Biology, Part II: Populations and Communities, edited by S. A. Levin
Atlas of the Netherlands Flora. I, Extinct and very rare species, edited by J. Mennema, A. J. Quene-Botorenbrood & C. L. Plate. Dr W. Junk     

Simultaneous character convergence and divergence in Western Australian land snails

Three distantly related genera of land snails, Amplirhagada Iredale, 1933, Quistrachia Iredale, 1939, and Westraltrachia Iredale, 1933, overlap in distributions only in the Napier and Oscar Ranges, east of Derby, Western Australia. The first two remain allopatric, Quistrachia having a more eastern range, while Westraltrachia overlaps both ranges. In the zone of sympatry, with changes proceeding from east to west, Westraltrachia diverges in feeding, and there is massive, gradual convergence in shell size, shape and colour between Amplirhagada and Westraltrachia. Shells of both genera depart markedly from their distinctive allopatric appearances to form a new morphotype in the north-west Napier Ranges. The species there are so similar in appearance that the two genera cannot be identified at a distance of 0.5 m. Thirteen species-level taxa are involved in these changes, four allopatric Amplirhagada , seven allopatric Westraltrachia, Quistrachia monogramma and Rhagada basedowana.
Westraltrachia species, the only large land snails in their main area of distribution (south-east Kimberlcy), are pre-adapted to graze on an occasionally present food resource, algal-fungal blooms on limestone seepage faces. They entered the Oscar and Napier Ranges, where this food resource is more abundant, and large land snails that are generalized feeders, Amplirhagada or Quistrachia , were already present and abundant. Hence feeding divergence occurred. Such specialization by Westraltrachia to minimize competition is readily understandable.
The selective pressures that led to the striking convergence in shell features in north-west Napier taxa are unknown.     

Genetic parallelism of protein polymorphism in nature: ecological test of the neutral theory of molecular evolution

We have conducted an ecological test of protein polymorphism in 13 unrelated genera of plants, invertebrates and vertebrates, involving 21 species, 142 populations and 5474 individuals. Each was tested, on average, for 27 enzymatic gene loci. These species varied in population size and structure, life histories and biogeographical origins, but they largely share a geographically short (260 km) and ecologically stressful gradient of increasing aridity in Israel, both eastward and (mainly) southward. We found genetic parallelism across most taxa, and most loci. Observed average heterozygosity, H , and gene diversity, He , were positively and overall significantly correlated with rainfall variation. This result corroborates the environmental theory of genetic diversity, primarily the niche-width variation hypothesis in both space and time. Our results are inconsistent with the neutral theory of molecular evolution and suggest that natural selection appears to be an important differentiating evolutionary force at the protein level.