Type materials of the species of the Planorbidae (Mollusca,Gastropoda, Hygrophila) described by Shuichi Mori

ABSTRACT

The planorbid species that were described by Shuichi (Syuiti) Mori from Japan have long been enigmatic and their taxonomy has been highly confused. This situation is also an impediment to action for the conservation of Japanese planorbids which are one of the most threatened freshwater snail groups in the country. Based on our examination of Shuichi Mori’s collection stored in the Kyoto University Museum, type materials of the taxa described by him are re-illustrated and re-examined for the first time. Our examination suggested that Gyraulus amplificatus (Mori, 1938) is not a morphological form of Gyraulus biwaensis but a distinct species. Gyraulus pulcher (Mori, 1938) can be regarded as a junior synonym of Gyraulus chinensis.     

Torsion in Patella caerulea (Mollusca, Patellogastropoda): ontogenetic process, timing, and mechanisms

Abstract. Torsion is a process in gastropod ontogenesis where the visceral body portion rotates 180° relative to the head/foot region. We investigated this process in the limpet Patella caerulea by using light microscopy of living larvae, as well as scanning electron microscopy (SEM) of larvae fixed during the torsion process. The completion of the 180° twist takes considerably less time in larvae of Patella caerulea than previously described for other basal gastropod species. At a rearing temperature of 20–22°C, individuals complete ontogenetic torsion in ?2 h. Furthermore, the whole process is monophasic, i.e., carried out at a constant speed, without any evidence of distinct ‘fast” or ‘slow” phases. Both larval shell muscles—the main and the accessory larval retractor—are already fully contractile before the onset of torsion. During the torsion process both retractors perform cramp‐like contractions at ～30 s intervals, which are followed by hydraulic movements of the foot. However, retraction into the embryonic shell occurs only after torsion is completed. The formation of the larval operculum is entirely in‐dependent from ontogenetic torsion and starts before the onset of rotation, as does the mineralization of the embryonic shell. The reported variability regarding the timing (mono‐ versus biphasic; duration) of torsion in basal gastropod species precludes any attempt to interpret these data phylogenetically. The present findings indicate that the torsion process in Patella caerulea, and probably generally in basal gastropods, is primarily caused by contraction of the larval shell muscles in combination with hydraulic activities. In contrast, the adult shell musculature, which is independently formed after torsion is completed, does not contribute to ontogenetic torsion in any way. Thus, fossil data relying on muscle scars of adult shell muscles alone appear inappropriate to prove torted or untorted conditions in early Paleozoic univalved molluses. Therefore, we argue that paleontological studies dealing with gastropod phylogeny require data other than those based on fossilized attachment sites of adult shell muscles.     

Climate-driven range dynamics of the freshwater limpet, Ancylus fluviatilis (Pulmonata, Basommatophora)

Aim Our aim was to understand the processes that have shaped the present‐day distribution of the freshwater limpet Ancylus fluviatilis sensu stricto in order to predict the consequences of global climate change for the geographical range of this species. Location North‐western Europe. Methods We sampled populations of A. fluviatilis sensu stricto over the entire range of the species (north‐western Europe) and sequenced 16S ribosomal RNA (16S) and cytochrome oxidase subunit I (COI) mitochondrial fragments to perform phylogenetic and phylogeographical analyses. Climatic niche modelling allowed us to infer the climatic preferences of the species. A principal components analysis identified the most important climatic factors explaining the actual range of A. fluviatilis. We also identified which climatic factor was the most limiting at range margins, and predicted the species’ geographical range under a climate change scenario [Community Climate Model 3 (CCM3)]. Results By means of the phylogeographical analysis, we infer that A. fluviatilis sensu stricto occupied northern refuges during the Last Glacial Maximum. We show that the climatic preferences of Baltic populations are significantly different from those of Central European populations. The projection of the occupied area under the CCM3 climate model predicts a moderate poleward shift of the northern range limits, but a dramatic loss of areas currently occupied, for instance in northern Germany and in southern Great Britain. Main conclusions The post‐glacial range dynamics of A. fluviatilis are not governed by niche conservatism. Therefore, we must be cautious about bioclimatic model predictions: the expected impact of climate change could be tempered by the adaptive potential this species has already shown in its evolutionary history. Thus, modelling approaches should rather be seen as conservative forecasts of altered species ranges as long as the adaptive potential of the organisms in question cannot be predicted.     

Effects of intermittent starvation on mature Biomphalaria glabrata (Gastropoda: Planorbidae)

Mature Biomphalaria glabrata, submitted to four weeks of varied starvation times (0, 1, 3, 5, 6 & 7 d.week^–1, were thereafter refed during four weeks. The different intermittent starvation times had no significant effect on snails survival. As weekly starvation increased, the rate of change in body weight and fecundity decreased. In snails fed one or two d.week^–1, the rate of change in body weight was negative, while fecundity remained at a low level. Continuous hunger stopped oviposition.Starvation had no further effect on body growth after the first week of refeeding; however, its effect on fecundity remained significant over the two first weeks.     

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.     

Comparative reproductive biology of Ariolimax californicus and A. dolichophallus (Gastropoda; Stylommiatophora)

Summary

Traits which differ among species in a genus are considered to be the product of relatively rapid evolution. Laboratory studies indicate that copulation differs between two species of the banana slug Ariolimax with A. dolichophallus having a pattern of a single, long duration (typically greater than 1 h) intromission which is simultaneously reciprocal, and A. californicus having mating encounters typically involving a bout of brief (typically 10–20 min) unilateral intromissions. Reciprocal apophallation was observed for both species. Field observations confirm simultaneously reciprocal copulation and apophallation in A. dolichophallus. Mating was more frequent in the spring and summer months. In laboratory studies the two species also differed in life history parameters; mean growth rate was greater in A. californicus than A. dolichophallus, whereas eggs are larger in A. dolichophallus (range = 0.33–0.5 g; n=59) than in A. californicus (range= 0.08–0.27 g; n=164).     

Effects of starvation on growth and reproductive apparatus of two immature freshwater snails Biomphalaria pfeifferi and Biomphalaria glabrata (Gastropoda: Planorbidae)

Starvation of immature snails of Biomphalaria pfeifferi and B. glabrata results in an arrest of growth and animals remain immature. Spermatogenesis is limited to spermatogonia (B.g.) or spermatocytes 1 (B.p.). The number and the size of oocytes remain inferior to that of controls. Animals show reduced genital tracts.Once feeding is restored, growth is resumed but wet weight and shell diameter do not reach the same level as in controls. Fecundity and gametogenesis do not differ from that in controls. Genital tracts weight is proportional to body weight.     

Microsatellites in the freshwater snail Bulinus globosus (Gastropoda: Planorbidae) from Zanzibar

Six microsatellite loci were isolated and characterized in Bulinus globosus, a freshwater snail with a wide distribution throughout sub‐Saharan Africa. Bulinus globosus is an intermediate host of Schistosoma haematobium, the causative agent of human urinary schistosomiasis. Microsatellites were tested using 32 snails from four populations collected from Pemba and Unguja islands of Zanzibar. The microsatellite loci displayed relatively low levels of variation, with between two and five alleles per locus. F_ST estimates indicate that gene flow is low, as has previously been suggested for other species of Bulinus.     

The cephalic sensory organ in veliger larvae of pulmonates (Gastropoda: Mollusca).

The apical area of larvae of four primitive pulmonate species was investigated by means of serial ultrathin and light microscope sections. Cephalic sensory organs (CSOs) were found in the larvae of Onchidium cf. branchiferum (Onchidiidae) and Laemodonta octanfracta (Ellobiidae), while no trace of the organ was present in the larvae of Ovatella myosotis (Ellobiidae) or Williamia radiata (Siphonariidae). TEM investigation revealed very similar CSOs in O. cf. branchiferum and L. octanfracta, with characteristic putative sensory cell types: ampullary cells with an internal ampulla containing densely packed cilia, para-ampullary cells with external cilia parallel to the surface, and ciliary tuft cells, bearing short ciliary tufts. The epithelium covering the organ has a thick microvillar border with microvilli laterally bearing a pair of electron-dense accumulations and a glycocalyx with interspersed flat plaque-like elements. While homologues of all major elements of the CSO can be found in other gastropod taxa, for example caenogastropods and opisthobranchs, the homology of the ampullary cell with similar cells in nongastropods appears unlikely. The CSO of L. octanfracta is associated with an additional structure, an epithelial external protrusion, lying ventral to the CSO. The absence of the organ in W. radiata weakens hypotheses on the organ's function of examining settlement conditions and velar control.     

Neotropical schistosomiasis: African affinities of the host snail Biomphalaria glabrata (Gastropoda: Planorbidae)

Schistosoma mansoni , the blood fluke responsible for human intestinal schistosomiasis in the Neotropics, was imported repeatedly with African slaves during the period 1500–1800. This trematode, and its intermediate host snails of the genus Biomphalaria , are widely distributed across Africa and the disease is thought to have quickly become established in South America and the West Indies because of the presence of an endemic susceptible congener, B. glabrata. We compared B. glabrata with four other Neotropical and three African species of Biomphalaria using 20 allozyme loci and found that it is phenetically and phylogenetically more like the African species; both parasite and American host snail are apparently of historically or geologically recent African origin. Furthermore, genetic distances, cladistic analyses and fossil data suggest the African Biomphalaria species may themselves have evolved from Neotropical founders following an initial trans-Atlantic dispersal in the reverse direction 2.3–4.5 Mya. Interpretation of existing patterns remains problematic as few African snails have been characterized genetically and both B. glabrata and African B. pfeifferi appear to comprise several cryptic species.     

A peculiar new species in the genus Landouria Godwin-Austen, 1918 from China (Gastropoda: Heterobranchia: Stylommatophora: Camaenidae)

Landouria omphalostoma n. sp. is described from northern Yunnan, China. The new species is placed in the genus Landouria Godwin-Austen, 1918 based on the presence of tuberculated flagellum and swollen basal part in the bursa copulatrix, and on the absence of dart sacs and mucous glands. L. omphalostoma n. sp. is characterized by a horizontal aperture of the shell which is unique in the genus. This feature resembles those of only two other camaenid/bradybaenid taxa, species of the Chinese genus Pseudaspasita Möllendorff, 1902 and species of the Japanese genus Coelorus Pilsbry, 1900. This is the first verified record of the genus in China and the easternmost record of Landouria. This paper provides the taxonomic ground work for further studies of the repeated evolution of a horizontal aperture.     

The life history of Brotia hainanensis (Brot, 1872) (Gastropoda: Prosobranchia: Thiaridae) in a tropical forest stream

An investigation of the prosobranch snail Brotia hainanensis (Brot, 1872) (Gastropoda: Thiaridae) in Tai Po Kau Forest Stream, New Territories, Hong Kong, has been undertaken with population samples collected at approximately 14-day intervals between August, 1977 and March, 1979. This species shows two peaks of recruitment per annum, one in spring and a second in the latter part of summer. Growth to sexual maturity is rapid, taking approximately 8.5 months, and in this respect, as well as its dual peak of recruitment, B. hainanensis displayed some variation from the iteroparous pattern of life cycles considered to be characteristic of freshwater prosobranch snails. Mortality after the young snails reach maturity is high due to severe seasonal spates in the stream and it is suggested that the modification of the life cycle of B. hainanensis from the type more usually encountered in freshwater prosobranchs reflects that this is an r-selected species adapted to a habitat where high density-independent mortality occurs.     

Pan-European phylogeography of the aquatic snail Theodoxus fluviatilis (Gastropoda: Neritidae)

Investigating the geographical distribution of genetic lineages within species is critical to our understanding of how species evolve. As many species inhabit large and complex ranges, it is important that phylogeographical research take into account the entire range of widespread species to clarify how myriad extrinsic variables have affected their evolutionary history. Using phylogenetic, nested clade, and mismatch distribution analyses on a portion of the mitochondrial COI gene, I demonstrate that the wide-ranging freshwater snail Theodoxus fluviatilis possesses in parallel many of the phylogeographical patterns seen in less widespread freshwater species of Europe. Fragmentary forces play a major part in structuring the range of this species, with 12 of 14 geographically structured nested clades displaying a distribution consistent with fragmentation or restricted dispersal. Certain regions of southern Europe harbour the majority of genetic diversity (total haplotype diversity, H = 0.87), particularly Italy (H = 0.87) and areas surrounding the Black Sea (H = 0.81). Post-Pleistocene range expansion is pronounced, with the majority of northern European populations (95% of sample sites) having arisen from northern Italian individuals that initially colonized northern Germany. Additionally, two highly divergent haplotype lineages present in northern Germany imply that there were at least two postglacial recolonization routes. Estuaries may also provide a means of dispersal given that no genetic differentiation was found between estuarine populations and neighbouring freshwater populations. Taken together, these data reveal a species with a complex genetic history resulting from the fragmentary effects of European geology as well as continuous and discrete range expansion related to their aquatic biology.     

Degradation of the radula in the snails Biomphalaria glabrata say and Limnaea stagnalis L. (Gastropoda,Pulmonata)

Summary The radula of snails is formed at the posterior end of the radular gland or pocket, and degraded at the same rate at its anterior end. Degradation is due to different secretory activities of the inferior epithelium of the radular gland. Its secretions seem to degrade enzymatically the matrix of the radular membrane and basal plates of teeth, leaving only chitin containing microfibres and degradation products. The sclerotized parts of the teeth remain unchanged, but as they are now only loosely connected with the radular membrane. they are torn off easily during feeding movements. The rest of the degraded and frayed radular membrane and the subradular membrane are also lost by abrasion during feeding. The cells of the inferior epithelium are connected with each other by septate desmosomes and an elaborate system of deep lateral interdigitation which may provide tensile strength. Extrusion of degraded cells of the inferior epithelium into the subradular membrane takes place, although the thick basal lamina forms a continuous sheath which is closely adjoined to the basal parts of the inferior epithelium. Nerve fibres containing vesicles with electron dense neurosecretory material (deduced from the diameter of 200–250 nm) are attached to this sheath or penetrate into it; they may be involved in the regulation of production and degradation processes during radula replacement. Problems of the forward transport of radula and inferior epithelium are discussed.     

Heritability of male outcrossing ability in the simultaneous hermaphrodite, Bulinus truncatus (Gastropoda: Planorbidae)

The heritability of phally, a dichotomous trait defined by the functional state of the male reproductive tract, was measured in a laboratory population of the simultaneous hermaphrodite snail Bulinus truncatus by means of a breeding experiment and a selection experiment. Euphallic individuals develop a fully functional male and female tract and are capable of receiving and donating sperm. In aphallic individuals the male tract does not develop fully, preventing sperm donation. There was no evidence of a heritable component to phally in the breeding experiment, but the selection experiment demonstrated a slight heritable effect. In both experiments there was more variation in the observed proportion of euphallics than expected by chance alone and no evidence of line or family effects, implicating environmental determination of male outcrossing ability even under controlled laboratory conditions. Previous studies of populations of B. truncatus reported that the proportion of euphallics was under strong genetic control. We suggest that there may be population differences in the extent of environmental control over phally, analogous to that reported for sex determination.     

Early development and pathogenesis of Lobatostoma manteri Rohde (Trematoda: Aspidogastrea)

The prosobranch Planaxis sulcatus is reported as a new natural host of Lobatostoma manteri at Heron Island, Great Barrier Reef. Planaxis sulcatus and Cerithium moniliferum were experimentally infected with large numbers of eggs. The larvae hatch in the stomach and migrate immediately along the ducts of the digestive gland into the digestive follicles. The larvae feed on the secretion and probably epithelial cells of the follicles. The acetabulum is used for adhesion to the epithelium and contributes to its erosion. In heavily infected snails, the digestive follicles disappear gradually and the larvae live in cavities lined by a flattened epithelium, parts of which show secretory activity. In snails dissected 47–49 and 65–66 days after infection, the cavities are fused, forming several large spaces which communicate with each other; only small parts of the epithelium are still secretory. Concentrations of amoebocytes occur in the walls of the digestive gland and in the wall between digestive gland and stomach of infected Planaxis. Some young worms were found in the stomach of Planaxis. No tissue reactions were seen around the stomach except in the wall between digestive gland and stomach. In Cerithium with 65–67 days old infection, the cavities contain much detritus and disintegrating cells, the epithelium is practically non-secretory and surrounded by loose connective tissue. In larvae with a body length of approximately 0·5–0·6 mm, the acetabulum begins to divide into alveoli and its anterior end grows forward; the anterior alveoli gradually increase in size and new alveoli are formed in the posterior undivided zone. In two specimens of approximately 1·3 mm body length, the whole adhesive disk was divided into half the number of alveoli usually found in adults. Allometric shifts during growth of the worms are analysed.     

Biogeography of the Helicoidea (Mollusca: Gastropoda: Pulmonata): land snails with a Pangean distribution

The biogeography of helicoid land snails was investigated using cladistic methods. Parsimony analysis under Assumption 0 yielded twelve area cladograms (length=25, c.i.=0.76, r.i.=0.86). The pattern of vicariance for the Helicoidea indicated that families originated with the break up of eastern Gondwana and Laurasia between the late Mesozoic and mid-Tertiary, and possible vicariance events are identified. It is proposed that Asian terranes, located between India and Australia, maintained contact with northern Australia until the late Cretaceous, which is later than is suggested in current palaeogeographical hypotheses.     

Effects of environmental calcium deprivation on the egg masses of Physa marmorata Guilding (Gastropoda: Physidae) and Biomphalaria glabrata Say (Gastropoda: Planorbidae)

Eggs of the basommatophoran snails Physa marmorata and Biomphalaria glabrata were cultured in low concentrations of calcium to determine effects on growth and development. In both species there was some development in media with 0.12 mg/l Ca²⁺ but embryos were unable to hatch. 61.04% of embryos of P. marmorata could develop to hatching in 0.22 mg/l Ca²⁺ but those of B. glabrata required a level of 0.42 mg/l Ca²⁺, to attain even a 31.07% hatch. Marked effects on growth rate, embryo size and on time taken to achieve hatching were noted in both species at very low calcium levels. The possibility of cation-controlling mechanisms in the egg membrane is discussed.     

Comparative ultrastructure of the terminal portions of the male copulatory apparatus in Planorbidae (Gastropoda: Pulmonata)

Terminal portions of the male copulatory apparatus of Planorbis planorbis, Segmentina oelandica, and Anisus vortex were studied using whole-mount preparations, serial semi-thin sections, and transmission electron microscopy. In the latter species, stylet formation was investigated at several stages of postembryonic development. Organization of the penial distal portion in the species studied varies greatly. In P. planorbis, the distal end of the penis lacks developed papillae and is armed with a stylet built up of the covering epithelial cells of the penis proper. In A. vortex, the stylet is formed by the secretory activity of the middle cells of the distal portion of the penis. To the time of maturation, the cells encompassing the stylet are broken down exposing its solid chitinous structure and characteristic shape. In S. oelandica, the distal end of the penis bears the long probably flexible papilla with the characteristics of an internal ‘skeleton,’ organized as a line of connective tissue cells and a system of hydrocoelic cavities.