ALTITUDINAL VARIATION AND ADAPTIVE ZONES IN ARIANTA ARBUSTORUM: A NEW LOOK AT A WIDESPREAD SPECIES

Arianta arbustorum is a European land snail, occurring in bothlowland and mountainous regions. Its shell is usually globular,which makes the species aberrant among the Campylaeinae in shellshape, habitat, and wide, not exclusively montane range. Withincreasing altitudes, the shells usually decrease in size andbecome more conical. However, in some mountain stocks thereare populations with depressed shells, provided with an openumbilicus. This form (subspecies) is only known from formerPleistocene refugia. It is hypothesized that in the Pliocene, the primary montaneA. arbustorum, with a depressed shell, entered an additional,adaptive zone, while conquering damp lowland habitats. Thisresulted in an enormous range extension. In the new habitata different (globular) shell shape evolved and the species becamepolytypic. The subsequent glaciations wiped out most of theoriginal montane A. arbustorum with depressed shells. Wherethis had occurred, post-Pleistocene re-colonizing conspecificsnails with globular shells, found vacant territory and coulddevelop the dines mentioned before. The invading populationsdid not regain the original, montane, depressed shell shape.This shape was only retained in a few refugia, where the populationssurvived amidst the glaciers. (Received 9 February 1990; accepted 31 May 1990)     

Dynamic gastropods: stable shell polymorphism despite gene flow in the land snail Arianta arbustorum

Based on a sequence fragment of mtDNA we analysed the genetic differentiation of the land snail Arianta arbustorum in the Alpine massif Gesäuse, where four morphotypes occur in close vicinity. We aimed at understanding actual and historical evolutionary processes among the morphotypes. With 135 haplotypes diverging up to 12.5%, genetic diversity was extremely high. Phylogenetic analyses indicated that A. arbustorum has colonized the Gesäuse several times in consecutive waves and that introgression of neutral markers across morphotypes and selection on shell shape have played an important role. Gene flow was more likely between locally close, different morphotypes than among distant populations of the same morphotype again indicating that cohesion of morphotypes was mainly due to selection. The actual distribution of morphotypes and haplotypes as well as the high genetic diversity also suggested that the area provided refugia during the Pleistocene glaciations.     

Mitochondrial differentiation in a polymorphic land snail: evidence for Pleistocene survival within the boundaries of permafrost

The genetic differentiation of populations having colonized formerly unsuitable habitats after the Pleistocene glaciations depends to a great extent on the speed of expansion. Slow dispersers maintain their refugial diversity whereas fast dispersal leads to a reduction of diversity in the newly colonized areas. During the Pleistocene, almost the entire current range of the land snail Arianta arbustorum has repeatedly been covered with ice or been subjected to permafrost. Owing to the low potential for dispersal of land snails, slow (re)colonization of the wide range from southern refugia can be excluded. Alternatively, fast, passive dispersal from southern refugia or survival in and expansion from multiple refugia within the area subjected to permafrost may account for the current distribution. To distinguish between these scenarios we reconstructed a phylogeography based on the sequences of a fragment of the cytochrome oxidase I from 133 individuals collected at 45 localities and analysed the molecular variance. Seventy-five haplotypes were found that diverged on average at 7.52% of positions. This high degree of diversity suggests that A. arbustorum is an old species in which the population structure, isolation and the hermaphroditic nature have reduced the probability of lineage extinction. The genetic structure was highly significant with the highest variance partition found among regions. Geographic distance and mitochondrial differentiation were not congruent. Lineages had overlapping ranges. The clear genetic differentiation and the patchy pattern of haplotype distribution suggest that colonization of formerly unsuitable habitats was mainly achieved from multiple populations from within the permafrost area.     

Thermal advantage of pale coloured morphs of the snail Arianta arbustorum (Helicidae, Pulmonata) in alpine habitats

In alpine grasslands pale colour morphs of the snail Arianta arbustorum are more frequent than dark ones As pale morphs absorb less solar radiation, they may forage for longer in sunlight and/or have a lower mortality risk from overheating This hypothesis was tested Equal numbers of pale and dark morphs were kept in enclosures in an alpine meadow During the day, more pale morphs were seen than dark ones, the latter obviously remaining hidden The difference was not significant in enclosures set up in a nearby forest, confirming that the inequality in behaviour depends on the amount of sunlight Dark snails are on average warmer than pale ones
Experiments were also conducted in grassland on a mountain slope at altitudes between 1200 and 2000 m Snails were marked with a paint which fades in light, and released When recaptured several days later, the marks were more faded in pale morphs than in dark, suggesting that they had been more exposed to sunlight, supporting our hypothesis The difference in fading was more pronounced at high than at low altitude     

The effect of temperature on the activity of three species of Helicid snail (Mollusca: Gastropoda)

The activity of the land snails Cepaea nemoralis (L.), C. hortensis (Müll.) and Arianta arbustorum (L.) was measured over 24 hours at the constant temperatures of 0, 3, 8, 17, 22 and 26°C at 100% relative humidity and with 16 hours light and 8 hours dark. The activity of all three species varies significantly with temperature and light, and there are differences in the activities of the species at each temperature. A. arbustorum is relatively more active than Cepaea at low rather than high temperatures, as is C. hortensis with respect to C. nemoralis , though to a lesser extent. There is little diurnal rhythm in activity at 0°C, but all species become more nocturnal as temperature increases; C. nemoralis is most, and A. arbustorum least nocturnal. The results obtained assist in explaining the observed geographical and local distributions of the species concerned.     

Do the risks of egg cannibalism and desiccation influence the choice of oviposition sites in the land snail Arianta arbustorum?

Laboratory tests were conducted to examine whether egg-laying Arianta arbustorum prevents egg cannibalism in its offspring by the choice of oviposition sites. Arianta arbustorum did not react to the presence of conspecific eggs when starting to oviposit, thus exposing its eggs to the risk of between-batch cannibalism. Snails preferred to oviposit in soil of higher moisture content even at high air humidity, reducing the risks of later egg desiccation and within-batch cannibalism. Both the types of oviposition sites chosen and their spatial distribution, as observed in 11 natural populations of A. arbustorum near Uppsala, Sweden, suggest that abiotic factors, such as a permanently moist microhabitat, may exert a stronger selection pressure on the oviposition-site choice than does between-batch egg cannibalism. As a result, the dispersion of egg batches observed in the field mainly reflects the spatial heterogeneity of habitats.     

Influence of shell colour on the mortality rate of the land snail Arianta arbustorum under different microclimatic regimes

The mortality of phenotypic shell colour morphs and age classes of the snail Arianta arbustorum in several microclimatic conditions was recorded. An analysis of variance was performed on five factors: adaptation temperature, relative humidity, shell colour, age class and test temperature. There were no significant differences in the mortality between different adaptation temperatures or relative humidities, but the interaction of these two factors was highly significant. There were significant differences in mortality rate between test temperature and age class. The mortality of the brown morph was higher than the yellow one at all adaptation temperatures (overall, P < 0.05). There were no significant differences between the mortality rates of the two morphs at different relative humidities. The mortality of the brown morph was higher than that of the yellow at six out of seven test temperatures. Juvenile snails survived significantly better than adults at all test temperatures.     

Self-fertilization and cross-fertilization in the land snail Arianta arbustorum (Mollusca, Pulmonata: Helicidae)

The simultaneously hermaphroditic land snail Arianta arbustorum (L.) (Pulmonata: Helicidae) was long believed to be self-incompatible. In the present study, however, uniparental reproduction was observed under laboratory conditions. Snails from two natural populations and from laboratory culture reared in isolation from the subadult stage produced fertile eggs. Over a period of three years after maturation, 38.5% of the unmated snails produced fertile eggs, mostly in the second and third years instead of the first year. Egg production and hatching success (the proportion of eggs that hatched) were lower in unmated snails than in mated snails. The number of hatchlings produced by unmated snails was 1–2% of that produced by mated snails. Hatchlings from unmated snails survived as well as those from mated snails under laboratory conditions. The result indicate that A. arbustorum can self-fertilize but with a great fitness reduction.     

The survival, weight-loss and behaviour of three species of land snail in conditions of low humidity

Cepaea nemoralis L., C. hortensis Mull, and Arianta arbustorum L. show differences in survival, rate of water-loss and behaviour under conditions of low humidity in laboratory studies. C. nemoralis survives better, ceases activity more rapidly, and loses proportionally less weight in low humidity than C. hortensis , which in turn shows the same properties with respect to A. arbustorum . The differences between these three species are reflected in differences in distribution in parts of Britain.     

Molecular evidence for Pleistocene refugia at the eastern edge of the Tibetan Plateau

The role of the Quaternary ice ages in forming the contemporary genetic structure of populations has been well studied in a number of global regions. However, due to the different nature of glaciations and complex topography, their role in shaping eastern Eurasian genetic diversity, particular in areas surrounding the Tibetan Plateau have remained largely unstudied. We aimed to address this question by examining the genetic structure of an alpine forest-associated taxon, the blood pheasant (Ithaginis cruentus) to infer its phylogeographic history. We detected three phylogenetic lineages and four current population groups. By comparing molecular and palaeovegetation data, we found that major glaciations during the Pleistocene have had a major impact upon the current genetic diversity of this species. Coalescent simulations indicate that the populations retreated to different refugia during some glacial periods in the Pleistocene, but persisted through the last glacial maximum (LGM). The most significant recent population expansion was found to have occurred before the LGM, during which palaeoclimatic data indicate that the climate was both warmer and wetter than today. In contrast, during the LGM populations may have adopted an altitudinal shift strategy in order to track changes in alpine glaciers, exemplifying a general response for montane species in the region where alpine glaciations were not large enough to cause qualitative changes in vegetation. Although analysis based on a plumage related gene showed that divergent selection may have contributed to current patterns of intra-specific diversity, demographic isolation is inferred to have played a more dominant role.     

Phylogeography of the high alpine plant Senecio halleri (Asteraceae) in the European Alps: in situ glacial survival with postglacial stepwise dispersal into peripheral areas

Whether alpine plant species survived Pleistocene glaciations in situ on high alpine nunatak mountains is still under debate. To test this hypothesis, Senecio halleri, a high alpine and endemic species with a narrow distribution range in the European Alps, was chosen as a model organism. Polymerase chain reaction-restriction fragment length polymorphisms of chloroplast DNA (cpDNA PCR-RFLPs) were used in a phylogeographic analysis of 14 populations of S. halleri, covering its total distribution area. The results of haplotype diversity and distribution gave evidence of in situ glacial survival on siliceous central-alpine nunatak mountains in two areas, southwest and northeast of the Aosta valley. According to the absence of genetic differentiation between these two nunatak areas (based on amova), nested clade analysis implied a history of preglacial gene flow, in situ survival and extinction of intermediate populations during glaciation and postglacial stepwise recolonization of peripheral and intermediate areas.     

Altitudinal variation in Arianta arbustorum (Mollusca,Pulmonata) in the Swiss Alps

Arianta arbustorum is a highly variable helieid snail found in Europe. We have studied biometrically its transition from low altitude where large dark phenotypes are typical, to higher altitudes in the Alps, where small light ones are found. Seven measurements and 10 colour characters were considered. Size varies with altitude while colours rather vary with habitats. Our observations on 5 alpine transects agree with those of previous authors who sampled other alpine areas. It indicates that the species' reaction to alpine conditions is general to large mountain regions. This provides a striking example of consistent variation. Previous authors claim that midland and alpine Arianta stem from different colonizing stocks. However, part of the variation found in the present study appears to be adaptive. Both interpretations can be reconciled.     

Is the incidence of survival in interior Pleistocene refugia (nunataks) underestimated? Phylogeography of the high mountain plant Androsace alpina (Primulaceae) in the European Alps revisited

Temperate mountain ranges such as the European Alps have been strongly affected by the Pleistocene glaciations. Glacial advances forced biota into refugia, which were situated either at the periphery of mountain ranges or in their interior. Whereas in the Alps peripheral refugia have been repeatedly and congruently identified, support for the latter scenario, termed “nunatak hypothesis,” is still limited and no general pattern is recognizable yet. Here, we test the hypothesis of nunatak survival for species growing in the high alpine to subnival zones on siliceous substrate using the cushion plant Androsace alpina (Primulaceae), endemic to the European Alps, as our model species. To this end, we analyzed AFLP and plastid DNA sequence data obtained from a dense and range‐wide sampling. Both AFLPs and plastid sequence data identified the southwestern‐most population as the most divergent one. AFLP data did not allow for discrimination of interior and peripheral populations, but rather identified two to three longitudinally separated major gene pools. In contrast, in the eastern half of the Alps several plastid haplotypes of regional or local distribution in interior ranges—the Alpine periphery mostly harbored a widespread haplotype—were indicative for the presence of interior refugia. Together with evidence from other Alpine plant species, this study shows that in the eastern Alps silicicolous species of open habitats in the alpine and subnival zone survived, also or exclusively so, in interior refugia. As the corresponding genetic structure may be lost in mostly nuclear‐derived, rapidly homogenizing marker systems such as AFLPs or RAD sequencing tags, markers not prone to homogenization, as is the case for plastid sequences (Sanger‐sequenced or extracted from an NGS data set) will continue to be important for detecting older, biogeographically relevant patterns.     

The distribution and variation of three species of land snail near Rickmansworth, Hertfordshire

The distribution of Arianta arbustorum, Cepaea nemoralis and C. hortensis , and the variations in the shell polymorphism of the Cepaea species have been studied in an area of low chalk downland and alluvial river valleys near Rickmansworth, Hertfordshire. The distribution of A. arbustorum is restricted, the species being largely confined to the bottoms of river valleys, especially in hedgerows. Variations in dampness seem the most likely limiting factor. C. hortensis is widespread and found in many habitats, but C. nemoralis is largely confined to woods, and is especially common in beechwoods. The morph-frequencies of C. nemoralis and C. hortensis vary in such a way as to suggest the operation of climatic and visual selection. In C. hortensis there are area effects for the presence or absence of pink five-banded shells, and there is an overall linkage disequilibrium resulting in an excess of unbandeds in non-yellow shells. Reasonable evidence for the operation of random factors in determining the morph-frequencies of populations of Cepaea in the area has been found only for three very disturbed populations. It is suggested that the large number of selective forces known to act on Cepaea populations in this area will account for the variation in morph-frequencies in populations from the same area or habitat.     

High levels of population subdivision in a morphologically conserved Mediterranean toad (Alytes cisternasii) result from recent,multiple refugia: evidence from mtDNA,microsatellites and nuclear genealogies

Pleistocene glaciations often resulted in differentiation of taxa in southern European peninsulas, producing the high levels of endemism characteristic of these regions (e.g. the Iberian Peninsula). Despite their small ranges, endemic species often exhibit high levels of intraspecific differentiation as a result of a complex evolutionary history dominated by successive cycles of fragmentation, expansion and subsequent admixture of populations. Most evidence so far has come from the study of species with an Atlantic distribution in northwestern Iberia, and taxa restricted to Mediterranean‐type habitats remain poorly studied. The Iberian Midwife toad (Alytes cisternasii) is a morphologically conserved species endemic to southwestern and central Iberia and a typical inhabitant of Mediterranean habitats. Applying highly variable genetic markers from both mitochondrial and nuclear genomes to samples collected across the species’ range, we found evidence of high population subdivision within A. cisternasii. Mitochondrial haplotypes and microsatellites show geographically concordant patterns of genetic diversity, suggesting population fragmentation into several refugia during Pleistocene glaciations followed by subsequent events of geographical and demographic expansions with secondary contact. In addition, the absence of variation at the nuclear β‐fibint7 and Ppp3caint4 gene fragments suggests that populations of A. cisternasii have been recurrently affected by episodes of extinction and recolonization, and that documented patterns of population subdivision are the outcome of recent and multiple refugia. We discuss the evolutionary history of the species with particular interest in the increasing relevance of Mediterranean refugia for the survival of genetically differentiated populations during the Pleistocene glaciations as revealed by studies in co‐distributed taxa.     

A comparative phylogeographic study reveals discordant evolutionary histories of alpine ground beetles (Coleoptera,Carabidae)

Taiwan, an island with three major mountain ranges, provides an ideal topography to study mountain–island effect on organisms that would be diversified in the isolation areas. Glaciations, however, might drive these organisms to lower elevations, causing gene flow among previously isolated populations. Two hypotheses have been proposed to depict the possible refugia for alpine organisms during glaciations. Nunatak hypothesis suggests that alpine species might have stayed in situ in high mountain areas during glaciations. Massif de refuge, on the other hand, proposes that alpine species might have migrated to lower ice‐free areas. By sampling five sympatric carabid species of Nebria and Leistus, and using two mitochondrial genes and two nuclear genes, we evaluated the mountain–island effect on alpine carabids and tested the two proposed hypotheses with comparative phylogeographic method. Results from the phylogenetic relationships, network analysis, lineage calibration, and genetic structure indicate that the deep divergence among populations in all L. smetanai, N. formosana, and N. niitakana was subjected to long‐term isolation, a phenomenon in agreement with the nunatak hypothesis. However, genetic admixture among populations of N. uenoiana and some populations of L. nokoensis complex suggests that gene flow occurred during glaciations, as a massif de refuge depicts. The speciation event in N. niitakana is estimated to have occurred before 1.89 million years ago (Mya), while differentiation among isolated populations in N. niitakana, N. formosana, L. smetanai, and L. nokoensis complex might have taken place during 0.65–1.65 Mya. While each of the alpine carabids arriving in Taiwan during different glaciation events acquired its evolutionary history, all of them had confronted the existing mountain ranges.     

Difference in habitat selection by different morphs of the land snail Arianta arbustorum

The behaviour of morphs of Arianta arbustorum has been investigated using a new technique involving a paint which fades in daylight. The technique was used to study the response of different morphs to exposure to sunlight at a homogeneous woodland habitat and a heterogeneous open habitat. The yellow morph showed a significantly higher rate of fading than the brown morph. A mark, release and recapture programme was used to analyse the association between frequencies of different morphs recaptured and not recaptured. In both habitats, there was a general tendency for the frequency of recapture to increase with increasing age in both morphs. Habitat selection resulting from different behaviour of different morphs of A. arbustorum is discussed. Morph frequencies can be related to habitat, the brown morphs of A. arbustorum are significantly less frequent in the open habitat. The behavioural responses of Cepaea hortensis , present with A. arbustorum in two other populations, to the exposure of sunshine were also studied. Climatic selection by solar energy associated with different morphs, species and habitat are also discussed.     

Parasitic mites influence fitness components of their host, the land snail Arianta arbustorum

Abstract. Parasites can influence the population dynamics of their hosts by affecting life-history strategies and behavior. The hematophageous mite Riccardoella limacum lives in the lung cavity of terrestrial gastropods. We used correlational and experimental approaches to investigate the influence of parasite infection on the behavior and life-history traits of the simultaneously hermaphroditic land snail Arianta arbustorum , a common host of R. limacum. Naturally infected individuals of A. arbustorum , collected in the wild, showed a decreased activity compared with uninfected snails. The reproductive output, expressed as the number of eggs deposited in a reproductive season, was reduced in mite-infected hosts. However, the hatching success of the eggs laid by parasitized snails was slightly higher than that of uninfected individuals. We also examined winter survival in 361 adults of A. arbustorum collected from four natural populations. The prevalence of mite infection ranged from 44.8% to 70.1% in three populations (snails in the fourth population were not infected). Winter survival was reduced in infected snails in two out of three populations. Furthermore, experimentally infected snails from an uninfected population showed a reduced winter survival compared with control snails. Our results indicate that parasite pressure imposed by members of R. limacum may influence life history in A. arbustorum.     

Patterns of dispersion, density and dispersal in alpine populations of the land snail Arianta arbustorum (L.) (Helicidae)

In studying populations of the land snail Arianta arbustorum in the eastern Swiss Alps, juvenile and subadult snails were found to be aggregated all the time, while adults were mostly aggregated during summer, the main mating period. The mean size of aggregations was 25 cm × 25 cm but varied, according to the area a grass tuft covered, while the intensity of aggregations was influenced by weather conditions. Densities of 10–15 subadults and 3–6 adults m⁻² were found along ditches and streams, 2–25 subadults and 2–20 adults on uncultivated alpine meadows, while on scree-covered alpine grassland the density of subadults and adults was 0.3–0.5 m⁻². The extent of daily movements varied with microclimatic factors and season and depended on the structure of the habitat. The dispersal rate was greatest in June (2.6 m month⁻¹) and least in August (1.1 m month⁻¹). In summer and autumn, active uphill movements were noted, a process which compensates for the loss of altitude by passive downhill displacements. Roiling down on snowfields and being carried away by avalanches and streams were observed.     

Responses of plants to changes in Qinghai–Tibetan Plateau and glaciations: Evidence from phylogeography of a Sibiraea (Rosaceae) complex

The genetic structure and phylogeographical history of the alpine shrubs Sibiraea angustata (Rosaceae) and Sibiraea laevigata from the Qinghai–Tibetan Plateau (QTP) were investigated to identify alpine plant responses to changes in the QTP and glaciations. Fifty-five populations were analyzed using four chloroplast DNA (cpDNA) regions and (nuclear ribosomal internal transcribed spacer) nrITS sequence data. In all, 21 cpDNA haplotypes and 13 nrITS sequence types were detected. Analyses of the genetic diversity and phylogenetic relationships detected two rarely reported glacial refugia. One was the Yushu–Nangqian area, and the other consisted of the area from the Songpan Plateau to the southeastern margin of the QTP. Sibiraea species populations experienced divergent evolution and founder effects when they recolonized the QTP platform and adjacent high-altitude regions following glaciations. The divergence times of the main lineages and haplotypes were in the range of 1.60–2.58 Ma. The population size of Sibiraea species in the QTP decreased approximately 23-fold during the last 0.12 Ma, indicating that Sibiraea species were significantly affected by environmental changes in the QTP. Therefore, the rapid uplift of the QTP and subsequent glaciations likely played an important role in driving genetic divergence and population size changes of Sibiraea species in the QTP.