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.     

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 .     

Molecular phylogeny, taxonomy and evolution of the land snail genus Iberus (Pulmonata: Helicidae)

Partial DNA sequences of two mitochondrial genes [cytochrome oxidase subunit I (COI) and 16S rRNA] from 59 specimens of Iberus were used to test the validity of the described morphospecies of this genus, and examine genetic divergences within and between main phylogenetic groups. Both gene fragments showed phylogenetic concordance. The COI gene was found to be faster evolving than the 16S gene and was fully protein-coding with no insertions or deletions. 16S rRNA was more informative than COI for resolving basal nodes. Both individual and combined analyses of the two gene fragments revealed five main phylogroups. These five groups are genetically unique lineages that are allopatrically distributed and considered to have full species status. Further subdivisions were also considered. Shell morphology was suitable for delimiting species boundaries, but several incongruences between morphology and mtDNA phylogeny were observed. These incongruences were considered consequence of hybridization between Iberus cobosi and Iberus marmoratus , and the result of shell shape polymorphism in Iberus rositai . According to spatial patterns of sequence divergence, life habits and shell morphology may be concluded that the keeled-flat shelled snails independently originated several times within Iberus and they could represent cases of similar shell adaptation to a karstic arid environment.     

Proximate factors influencing egg cannibalism in the land snail Arianta arbustorum (Pulmonata,Helicidae)

Summary Byttneria aculeata (Sterculiaceae), a subcanopy liane with a shrubby juvenile form has two distinct leaf color morphs in juvenile plants- a given juvenile has plain green leaves or leaves with whitish variegation. Both forms occur together in the forest and in clearings; however, the variegated morph is more common in open sites, and the plain morph predominates in the forest. Percent variegation per leaf for variegated plants increased from closed to open sites. Measurements of growth support the idea that variegation is favored in open habitats. Within a given habitat, rate of herbivory by leaf miners on a given morph increases with increasing relative frequency of that morph; however, at a given relative frequency, the variegated morph is less heavily attacked than is the plain morph when it occurs elsewhere at that same frequency.     

Radiation of the land snail genus Sagda (Pulmonata: Sagdidae): comparative morphology, biogeography and ecology of the species of north-central Jamaica

The radiation of the endemic Jamaican land snail genus Sagda is examined through analysis of the comparative shell morphology, anatomy, ecology and biogeography of the four species inhabiting north-central Jamaica: S. centralis sp. nov. , S. spei, S. bondi and S. montegoensis. The species are all rather similar in their morphology and ecology. The shells show some differences in the presence or absence of internal lamellae, the position of the lamellae, the form of the depression in the centre of the base, the regularity of sculpture (low ribs) and the pattern of microsculpture. Slight differences in the form of the radular teeth are found between some species. The structure of the secondary ureter is peculiar: in all species it is open shortly behind the mantle collar, then branches into two closed tubes which open separately on the outside of the mantle collar. Despite its great complexity, the reproductive system is rather uniform among species; differences are seen in the number of pockets that form spines on the spermatophore. The complex, three-branched spermatophore is formed within the entire epiphallar branch of the penis, whereas the appendicular branch apparently assumes the copulatory function. When inactive, all species are found associated with rocks or, less commonly, among leaf litter. Activity occurs primarily at night and mostly on leaf litter rather than rocks. The snails eat dead plant material, including a variety of plant species. All species (with the possible exception of S. montegoensis) are ovoviviparous. They overlap broadly in the range of climates occupied. The geographic ranges of the four species are overlapping, although Sagda bondi has a parapatric distribution in relation to the western species S. grandis. Fossil and subfossil evidence shows that shifts in the ranges of the species have occurred in the late Quaternary; Pleistocene-Holocene environmental changes, habitat destruction by man, and possibly other environmental changes during historical times are probably involved. Radiations of other land snail groups on tropical islands have generally produced greater morphological and ecological differentiation than in Sagda. Allopatric or parapatric species distributions appear to be more common in other land snail radiations.     

Species-independent, geographical structuring of chloroplast DNA haplotypes in a montane herb Ipomopsis (Polemoniaceae)

Two hypotheses have been proposed to explain the occurrence of hybrid zones between red-flowered Ipomopsis aggregata and white-flowered I. tenuituba . Either local adaptation to hummingbird and hawkmoth pollinators has given rise to sympatric (or parapatric) divergence of flower colour and morphology (primary intergradation at hybrid zones), or alternatively two previously allopatric species are coming into contact at several geographical areas of secondary intergradation. We examined restriction site patterns in nuclear DNA (nrDNA), chloroplast DNA (cpDNA) and mitochondrial DNA (mtDNA) from populations of I. aggregata and I. tenuituba representing seven zones of sympatry. No variation was detected in a 350-bp fragment of mtDNA and uninformative levels of variation were observed for nrDNA. We detected 22 potentially informative restriction site polymorphisms in cpDNA, all of which united geographical areas containing populations of both species. We detected no informative species-specific markers. Studies of other species (e.g. oaks) have detected similar species-independent geographical structure of cpDNA. However, in these cases secondary interegradation could be inferred from species-specific nuclear alleles. The pattern in Ipomopsis is consistent with both primary intergradation (independent speciation in each area of sympatry) or secondary intergradation involving complete cytoplasmic replacement. Thus, additional data are needed to explain the origin of hybrid zones in Ipomopsis .     

Control of acid-base status in active and dormant land snails,Otala lactea (Pulmonata,Helicidae)

Summary Control of extracellular acid-base status was examined during activity and dormancy inOtala lactea (Pulmonata, Helicidae). Active snails showed little variation in hemolymph pH and at constant temperature. With increase of temperature, hemolymph increased from about 6 Torr at 5°C to 13 Torr at 24°C and pH decreased by about 0.017 pH units/°C, a pattern consistent with alphastat regulation of pH via ventilatory control of .During dormancy, mean hemolymph increased to about 50 Torr. Venous pH declined by about 0.4 units due to hypercapnia and fluctuated more widely than in active snails due to variability of . Hemolymph pH declined further in prolonged dormancy due to progressive metabolic acidosis; after one year of dormancy the mean hemolymph pH was about 0.8 units lower than that of active snails at similar temperature.Active snails exposed experimentally to high showed a large increase in hemolymph [HCO ₃ ^– ]. However, [HCO ₃ ^– ] declined by up to 50% during dormancy, despite the naturally occurring hypercapnia. Hemolymph osmolality and the concentrations of solutes other than [HCO ₃ ^– ] increased with increasing duration of dormancy. Concentrations of magnesium and calcium increased about 2.5 times more rapidly than those of sodium and chloride, indicating that acidosis is partially offset by the dissolution of carbonates from the shell or tissues.     

Bioacoustic variation of swimbladder disturbance sounds in Neotropical doradoid catfishes (Siluriformes: Doradidae,Auchenipteridae): Potential morphological correlates

Swimbladder disturbance sounds of doradoid catfishes (Doradidae and Auchenipteridae) demonstrated striking waveform and spectrographic variation.We surveyed sounds of 25 doradoid species in 20 genera comparing these to sounds of four vocal outgroup catfish families.Sounds were either continuous waveforms (lacking interpulses) or pulsed (groups of pulses repeated at fixed temporal intervals).This is the first evidence for swimbladder calls with fixed interpulse patterns in catfishes.Vocal mechanism components that were similar between doradids and auchenipterids included:swimbladder shape,swimbladder dimensions and sonic muscle-somatic index.Morphological traits that showed variation among taxa and were evaluated for potential correlates of call diversity are:1) diverticula (marginal outpocketings of the swimbladder with no connection to inner ear) and 2) elastic spring apparatus Millerian rami (ESA-Mr).Within the doradid subfamilies and within the Auchenipteridae most species differed significantly in dominant frequency with frequency ranges overlapping to some extent for most.Doradid swimbladder diverticula did not explain dominant frequency variation within the doradoid superfamily.Some doradids with conical ESA-Mr had the highest dominant frequency sounds.Auchenipterids included both relatively lower and higher dominant frequency sound producers but lacked diverticula and had discoidal ESA-Mr.Comparing a phylogeny of doradoid genera with outgroup taxa,we infer that complex diverticula and conical ESA-Mr are derived characters within the Doradidae.Species representing outgroup families produced either continuous lower dominant frequency sounds (aspredinids,mochokids and pseudopimelodids) or pulsed higher dominant frequency sounds (pimelodids) [Current Zoology 58 (1):171-188,2012].     

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 Pleistocene glaciations and the evolutionary history of the polytypic snail species Arianta arbustorum (Gastropoda, Pulmonata, Helicidae)

The evolutionary history of the snail Arianta arbustorum is controversial. This diverse, polytypic species has two distinct forms: one, with a globular shell and closed umbilicus, is found from lowland to high altitudes; the other, with a depressed shell and open umbilicus, is found at a few scattered, high altitude localities. What is the origin of these two forms? Some believe that the depressed shell is a recent, local, ecotypic adaptation to alpine environments. Others believe that this form is a relic of an ancestral condition that may have survived the Pleistocene glaciations on nunatak-like montane refugia, while the globular shell is a derived condition and its presence at high altitudes follows post-Pleistocene recolonisation. We analysed a portion of the mitochondrial gene cytochrome oxidase I for 100 snails of the species A. arbustorum, three additional Arianta species, and nine outgroup taxa from five genera, in order to understand the phylogeographic history of the species. Despite some confounding artefacts that are likely due to introgression among the morphological forms, the resulting phylogeny shows that the depressed shell is plesiomorphic, while the globular shell is derived. Moreover, their disparate histories suggest that the depressed shell variety survived the glaciations in pockets of alpine refugia, while the globular shell variety recolonised the alpine environment post-glacially.     

Pallial systems in the Athoracophoridae (Gastropoda: Pulmonata)

Abstract

Pallial systems are described for 10 athoracophorid species: Athoracophorus bitentaculatus, Pseudaneitea dendyi, P. gigantea, P. papillata, P. pallida, P. aspera, P. schauinslandi, P. multistriata, and Palliopodex verrucosus from the New Zealand and sub antarctic area; and Triboniophorus graeffei from Australia. In the New Zealand and subantarctic species the kidney is invariably bilobed, the ureter is extensively folded and connected to the intestine by an elongate tubule running initially through the lung, and the pneumostome, anus, and excretory pores are widely sepamted. Triboniophorus graeffei has the pneumostome, anus, and excretory pore closely associated, a compact kidney, and a more simply folded ureter. The New Zealand and subantarctic representatives are considered to be more advanced. Reasons for the development of the features they display are discussed, as is the function and evolution of the entire pallial system. An account of the variation in the pallial complex of Athoracophorus bitentaculatus is included.     

Mitochondrial phylogeny and biogeographic history of the Greek endemic land-snail genus Codringtonia Kobelt 1898 (Gastropoda, Pulmonata, Helicidae)

The aim of this work was to infer the phylogeny of the Greek endemic land-snail genus Codringtonia Kobelt 1898, estimate the time frame of the radiation of the genus, and propose a biogeographic scenario that could explain the contemporary distribution of Codringtonia lineages. The study took place in the districts of Peloponnese, Central Greece and Epirus of mainland Greece. Sequence data originating from three mtDNA genes (COI, COII, and 16S rDNA) were used to infer the phylogeny of the eight nominal Codringtonia species. Furthermore, the radiation time-frame of extant Codringtonia species was estimated using a relaxed molecular clock analysis and mtDNA substitution rates of land snails. The phylogenetic analysis supported the existence of six Codringtonia lineages in Greece and indicated that one nominal species (Codringtonia neocrassa) might belong to a separate genus distantly related to Codringtonia. The time frame of differentiation of Codringtonia species was placed in the Late Miocene-Pleistocene epoch. The dispersal-vicariance analysis performed indicated that most probably Codringtonia exhibited a north-to-south spread with the ancestral area being that of central Greek mainland, accompanied with duplication (speciation) and vicariance events.     

Extreme mtDNA divergences in a terrestrial slug (Gastropoda, Pulmonata, Arionidae): accelerated evolution, allopatric divergence and secondary contact

Extremely high levels of intraspecific mtDNA differences in pulmonate gastropods have been reported repeatedly and several hypotheses to explain them have been postulated. We studied the phylogeny and phylogeography of 51 populations (n = 843) of the highly polymorphic terrestrial slug Arion subfuscus (Draparnaud, 1805) across its native distribution range in Western Europe. By combining the analysis of single stranded conformation polymorphisms (SSCP) and nucleotide sequencing, we obtained individual sequence data for a fragment of the mitochondrial 16S rDNA and a fragment of the nuclear ITS1. Additionally, five polymorphic allozyme loci were scored. Based on the 16S rDNA phylogeny, five monophyletic haplotype groups with sequence divergences of 9-21% were found. Despite this deep mitochondrial divergence, the haplotype groups were not monophyletic for the nuclear ITS1 fragment and haplotype group-specific allozyme alleles were not found. Although there is evidence for an accelerated mtDNA clock, the divergence among the haplotype groups is older than the Pleistocene and their current allopatric ranges probably reflect allopatric divergence and glacial survival in separate refugia from which different post-glacial colonization routes were established. A range-overlap of two mtDNA groups (S1 and S2, 21% sequence divergence) stretched from Central France and Belgium up to the North of the British Isles. The nuclear data suggest that this secondary contact resulted in hybridization between the allopatrically diverged groups. Therefore, it seems that, at least for two of the groups, the deep mtDNA divergence was only partially accompanied by the formation of reproductive isolation.     

Shell polymorphism in a southern peripheral population of Cepaea nemoralis (L.) (Pulmonata: Helicidae) in Spain

Populations of the land snail Cepaea nemoralis have been investigated at the southern limits of its range in Spain, the Upper Tajo region, in order to analyse the distribution of the species and the variation of polymorphism in relation to environmental conditions. For this purpose two natural subareas, which differ in topography, soil nature, climate and vegetation have been considered. Cepaea nemoralis is confined to the river banks of the area; it is found in open and shaded biotopes of subarea 1 where it is relatively common, and it is almost restricted to shaded biotopes of subarea 2 where it becomes progressively rarer towards the south. Relative abundance of the species may be related to the amount of calcareous soils and climate, the restriction in its southern distribution being probably the result of extreme solar radiation, high temperature and low humidity. The two subareas differ in the frequency of the main polymorphic characters. These differences may be related to the kind of biotope. Subarea 1 richer in more open and exposed biotopes have higher frequencies of yellow, 00000, 00300, and unfused bands, whereas the restriction to poplar groves because of the lack of suitable exposed biotopes in subarea 2 may account for the higher proportion of pink five-bandeds with the bands fused together. This kind of variation with biotope is consistent with an explanation in terms of differences in the temperature absorbance of different morphs and a metabolic response to differences in temperature amplitude of the biotopes. Other characters (lip colour, punctate bands, hyalozonate bands, etc.) are not polymorphic in this region, thus, suggesting that at the periphery environmental conditions are important not only in determining the distribution of the species but exerting a marked effect on the genetic structure of the populations.     

Evolution of distribution and habitat patterns in endemic millipedes of the genus Dolichoiulus (Diplopoda: Julidae) on the Canary Islands, with notes on distribution patterns of other Ganarian species swarms

Forty-six species of the genus Dolichoiulus , all endemic, occur on the Canary Islands The highest number of species occur on the largest, highest island (Tenerife); fewest occur on Lanzarote, Fuerteventura (low, xeric), El Hierro and La Palma (small, remote). Most of the Dolichoiulus species live on one island only, as in other endemic Canarian species swarms. The scarcity of pluri-insular Dolichoiulus species, in connection with information on phylogeny, suggests that speciation has mainly taken place within individual islands. Distribution patterns are partly governed by habitat differences between species, but vicariance patterns between species living in the same kind of habitat are evident on La Gomera and Tenerife. Dolichoiulus species occur in all kinds of natural habitats. Laurisilva and cave species are generally paler than other species. In the laurisilva of eastern Tenerife, microhabitat differentiation between species is pronounced. In some, but far from all, cases, species coexisting in the same microhabitat are of different sizes. The ancestral colonizing species of Dolichoiulus is/are hypothesized to have been small and to have lived in coastal habitats. Colonization of higher altitudes was usually accompanied by an increase in body size. Invasion of the laurisilva was usually accompanied by a habitat shift from the ground layer to logs.     

A morphological study and hybridization analysis of Caloglossa leprieurii (Ceramiales, Rhodophyta) from Japan, Singapore and Australia

Morphological and hybridization experiments were performed on Caloglossa leprieurii (Montagne) J. Agardh collected from Japan, Singapore and Australia in order to evaluate taxonomic characters of this species. Within C. leprieurii at least four mating groups were recognized from the Indo-Pacific region. These mating groups could be characterized by the blade width at the inter-node and the cell-row numbers on the opposite side derived from the first axial cell at the main axis, though these properties showed a certain variability even in the same plant under both field and culture conditions. The phylogenetic relationship and geographic distribution of the four mating groups are discussed.     

The chromosomes of Cepaea vindobonensis (Pulmonata: Helicidae) and the relationship between this species and the rest of the genus

The karyotype of Cepaea vindobonensis (Pfeiffer) (2 n = 50) is defined and compared with those of C. sylvatica (Draparnaud), C. nemoralis (L.) and C. hortensis (Müller). Cepaea vindobonesis shares its chromosome number with C. sylvatica. The genus Cepaea thus consists of two pairs of species, one with 2 n = 50 the other with 2 n = 44. The karyotype of C. vindobonensis is distinct from that of C. sylvatica as it lacks any discontinuity in its range of chromosome length and has all three of its longest chromosome pairs nearly identical. The two 2 n = 50 species both have an unusual meiotic bivalent showing a 'satellite-type' structure. It is apparent that the 2 n = 50 species are more closely related to each other than to the 2 n = 44 species. No precise definition of the evolutionary history can be given but a working hypothesis is suggested.     

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.     

The life cycle, population dynamics, growth and secondary production of the snail Xeropicta arenosa Ziegler (Gastropoda: Pulmonata) in northern Greece

The life cycle, population dynamics and secondary production of the xerothermophilic land snail Xeropicta arenosa were studied in northern Greece. The study of biology and the demographic analysis of the populations of X. arenosa revealed that it is a semelparous annual species. Reproduction takes place in autumn and the majority of the adults die after egglaying. Growth is continuous throughout the year and no seasonal fluctuations occur. Estimation of annual secondary production with the Hynes' size frequency method revealed a mean standing crop ( aa ) of 0.075 g m^-2 year^-1 and a productivity ( P ) of 0.267+ 0.043 gm^-2 year^-1. The annual turnover ratio ( P/ ) was equal to 3.56.     

A new species of Caracolus (Pulmonata: Camaenidae) from the Oligocene of Nebraska and the biotic history of the American camaenid land snails

A fossil land snail from the Oligocene White River Group of Nebraska is described as Caracolus aquilonaris sp nov. Living members of the genus Caracolus occur in tropical forest on the islands of the Greater Antilles. It is likely that C. aquilonaris occupied a similar habitat and that the present restricted occurrence of the genus Caracolus is a fragment of a previously wider range. The American carnaenid genera Pleurodonte, Polydontes and Zachrysia form a well defined monophyletic group, but there is an unresolved trichotomy involving this group, Labyrinthus and Caracolus . Well characterized fossil camaenids assigned to Pleurodonte (Jamaica, Carriacou) and the related Pleurodontites (Florida) occur in the Miocene. Antecedents to Labyrinthus were isolated in South America and to Caracolus and Pleurodonte in North America, possibly in the late Cretaceous. The latter genera reached the Antilles where Caracolus has persisted relatively unchanged. Ancestral Pleurodonte has diversified producing the distinctive subgenera Eurycratera and Thelidomus on Jamaica and the genus Polydontes on the other Greater Antilles. Drier conditions on Cuba have resulted in the divergence of Zachysia from Polydontes .