Age,seasonal and diel variations of the uricolytic enzymatic system in the snailsXeropicta arenosa (Ziegler) andCernuella virgata (Da Costa)

Summary The nitrogen catabolism of two snail species,Xeropicta arenosa andCermuella virgata, living on coastal dunes, was studied at different seasons and times of the day and at different ages. All the nitrogenous metabolites of the uricolytic enzymatic system were found in body homogenates of the snails, although uric acid was most abundant. The highest levels of these catabolites were observed during the day when the snails were inactive, while the lowest levels occurred during the night when excretion probably took place. The levels of most of the nitrogenous catabolites and the activities of the enzymes were higher in juveniles than in adults.All the enzymes of the uricolytic pathway were shown to be present in bothX. arenosa andC. virgata.     

Dispersal and population structure of two sympatric species of the mediterranean land snail genus Mastus (Gastropoda, Pulmonata, Enidae)

We studied the population dynamics of Mastus by investigating the effects of population structure, spatial ecology and biology of reproduction on the genetic diversity of two sympatric Mastus species endemic to the island of Crete. Over a period of 27 months, we carried out both mark–recapture and random quadrat sampling techniques in order to assess the dispersal trends, the aggregation patterns, the neighbourhood size and the habitat preferences of these species. There were 154 recorded movements for M. butoti and 114 for M. cretensis . Mean monthly dispersal was estimated at d  = 0.5 m for M. butoti and d  = 1 m for M. cretensis . Both species showed a random dispersal pattern but tended to aggregate in the field. Their populations were found to be highly structured owing to their highly parsimonious dispersal behaviour and the very low population densities, estimated at D  = 2.07 ± 0.16 and D  = 0.73 ± 0.16 individuals m⁻² for M. butoti and M. cretensis , respectively. The neighbourhood size did not exceed 150 individuals for either species. The habitat occupied by each species changed during the active season, but both the immature and the adult individuals of each species seemed to prefer the same habitats throughout the active season. Partial population activation during the active season was observed in both species. We conclude that the population structure, the partial population activation and the species-specific reproductive strategies have a profound effect on maintaining the genetic diversity of the genus' populations.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 131–144.     

Population dynamics and production of the freshwater snail Chilina gibbosa Sowerby 1841 (Chilinidae,Pulmonata) in a North-Patagonian reservoir

Chilina gibbosa is an endemic snail widely distributed in Patagonia, Argentina. Due to its importance in the benthic fauna and in the diet of some fish in the oligo-mesotrophic reservoir Ezequiel Ramos Mexía (39° 30′ S, 69° 00′ W), special attention has been given to its life cycle, growth patterns and annual production. Samples were taken monthly at five littoral stations between June 1983 and July 1984. Mean abundance and biomass of C. gibbosa were much higher in vegetated stations dominated by Potamogeton berteroanus (Station 1 : 583 ind. m⁻², 5.95 g AFDM m⁻²) or by Nitella clavata (Station 5 : 275 ind. m⁻², 4.18 g AFDM m⁻²) than bare stations with low transparency or stations with other macrophytes. The snails presented a clustered spatial pattern and their abundance was significantly correlated with macrophyte wet biomass only when this was above 250 g m⁻². Analysis of size distributions showed an annual life cycle with a reproductive period in the summer. However, differences in recruitment and growth occurred probably due to differences in water temperature and food availability. Growth was maximum in summer and almost absent during winter. Hence, shell growth data fit a sigmoid curve well, and growth was somewhat higher at Station 1. Annual production at Stations 1 and 5, estimated by the ‘growth increment summation’ method (28.8 g AFDM m⁻² and 14.18 g AFDM m⁻² respectively), was among the highest recorded for pulmonate gastropods, possibly due to a low interspecific competition. The P : B ratio values were within the range for univoltine gastropods (4.84 and 3.39). The high productivity and turnover rate of these snails grant a high availability of food for the abundant molluscivore, the silverside Patagonina hatcheri.     

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.     

Characterization of 13 polymorphic microsatellite loci for two land snail species, Trochulus villosus and T. sericeus (Gastropoda: Pulmonata: Hygromiidae)

Thirteen new microsatellite loci were isolated and tested on two land snail species, Trochulus villosus and T. sericeus (Pulmonata: Hygromiidae), resulting in a set of eight polymorphic markers for each species. The expected heterozygosity was high for all loci and species (between 0.616 and 0.944). Such levels of variability will allow detailed insights into the population genetic structure of some Trochulus species.     

The annual reproductive cycle of the snail Megalobulimus abbreviatus (Bequaert, 1948) (Gastropoda, Pulmonata).

Morphological changes in the sexual organs of the pulmonates were observed throughout a year and correlated with reproductive-cycle periods. Reproductive-organ weights of the snail Megalobulimus abbreviatus were recorded seasonally and gonad sections were analyzed morphologically. The weights were used to obtain the organosomatic index. Mean oocytic diameter and oocytic maturation index were based on gonad sections. It was concluded that M. abbreviatus is an iteroparous snail whose annual reproductive cycle is characterized by mating and egg laying throughout spring and early summer, and also by reproductive system preparation, occurring over the remainder of the summer until the end of winter, for a new breeding season.     

Phenotypic diversity,population structure and stress protein-based capacitoring in populations of Xeropicta derbentina,a heat-tolerant land snail species

The shell colour of many pulmonate land snail species is highly diverse. Besides a genetic basis, environmentally triggered epigenetic mechanisms including stress proteins as evolutionary capacitors are thought to influence such phenotypic diversity. In this study, we investigated the relationship of stress protein (Hsp70) levels with temperature stress tolerance, population structure and phenotypic diversity within and among different populations of a xerophilic Mediterranean snail species (Xeropicta derbentina). Hsp70 levels varied considerably among populations, and were significantly associated with shell colour diversity: individuals in populations exhibiting low diversity expressed higher Hsp70 levels both constitutively and under heat stress than those of phenotypically diverse populations. In contrast, population structure (cytochrome c oxidase subunit I gene) did not correlate with phenotypic diversity. However, genetic parameters (both within and among population differences) were able to explain variation in Hsp70 induction at elevated but non-pathologic temperatures. Our observation that (1) population structure had a high explanatory potential for Hsp70 induction and that (2) Hsp70 levels, in turn, correlated with phenotypic diversity while (3) population structure and phenotypic diversity failed to correlate provides empirical evidence for Hsp70 to act as a mediator between genotypic variation and phenotype and thus for chaperone-driven evolutionary capacitance in natural populations.     

Preliminary studies on the reproduction and population dynamics of Monodonta lineata and Gibbula umbilicalis (Mollusca,Gastropoda) on the central coast of Asturias (N. Spain)

Summary Populations of Monodonta lineata and Gibbula umbilicalis were studied at Aramar beach (Luanco, Asturias) from October 1982 to November 1983. Both species show a sexual ratio of 1: 1 but differ in their gonadal cycle patterns. M. lineata gonadal development stages are found from November to May–June in a significant part of the population. Breeding stages occur from June–July to September, and in some individuals to November, with complete discharge of all spawn at the end of the reproductive season. Some degree of inter-annual variation is detected. Spawning stages of mature G. umbilicalis are found through the year but the higher frequencies occur in summer and autumn samples, with development phases predominating in January to May. Animals over 10 mm in shell diameter maintain their gonads close to spawning stages, completely spawned individuals being less than 10 percent. In both species variability in gonad stages for a given month reduces as size increases, indicating a certain synchrony.The new recruits of M. lineata are detected on the bare rock by September to November and are recognizable through their first year. Due to the amplitude of the breeding period and differing growth rates size distribution within an age class is wide. Size class frequencies rapidly decrease after the second year of life. The data for G. umbilicalis are difficult to interpret given the scarce number of juveniles and size overlapping.     

Neurosecretion in the basommatophoran snail Bulinus truncatus (Gastropoda,Pulmonata)

Summary The neurosecretory system of the freshwater snail Bulinus truncatus was investigated. With the Alcian blue-Alcian yellow (AB/AY) staining method at least 10 different types of neurosecretory cells (NSC) were distinguished in the ganglia of the central nervous system. The differences in staining properties of the NSC — with AB/AY the cells take on different shades of green and yellow — are borne out at the ultrastructural level: the NSC types contain different types of neurosecretory elementary granules.The neurosecretory system of B. truncatus is compared to that of Lymnaea stagnalis, the species which has received the most attention among the pulmonates. It appears from the comparison that the systems of both species show many similarities, although some differences are also apparent.     

Seasonal and inter-stream variations in the population dynamics, growth and secondary production of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) in monsoonal Hong Kong

1. Balitorid loaches are widespread and highly diverse in Asian streams, yet their life history and ecology have received little attention. We investigated seasonal (wet versus dry season) and spatial variation in populations of algivorous Pseudogastromyzon myersi in Hong Kong, and estimated the magnitude of secondary production by this fish in pools in four streams (two shaded and two unshaded) over a 15‐month period. 2. Mean population densities of P. myersi ranged from 6.0 to 23.2 individuals m⁻², constituting more than half (and typically >70%) of benthic fishes censused. Abundance was c. 25% greater in the wet season, when recruitment occurred. Significant density differences among streams were not related to shading conditions and were evident despite small‐scale variations in P. myersi abundance among pools. Mean biomass varied among streams from 0.85 to 3.87 g ash‐free dry weight (AFDW) m⁻². Spatial and seasonal patterns in biomass and density were similar, apart from some minor disparities attributable to differences in mean body size among populations. 3. All four P. myersi populations bred once a year in June and July, and life spans varied from 24 to 26 months. Populations consisted of three cohorts immediately after recruitment but, for most of the study period, only two cohorts were evident. Cohort‐specific growth rates did not differ significantly among streams but, in all streams, younger cohorts had higher cohort‐specific growth rates. 4. Secondary production of P. myersi estimated by the size‐frequency (SF) method was 2.7–11.5 g AFDW m⁻² year⁻¹ and almost twice that calculated by the increment‐summation (IS) method (1.2–6.6 g AFDW m⁻² year⁻¹). Annual P/B ratios were 1.17 – 2.16 year⁻¹ (IS) and 2.73 – 3.22 year⁻¹ (SF). Highest production was recorded in an unshaded stream and the lowest in a shaded stream, but site rankings by production did not otherwise match shading conditions. Wet‐season production was six times greater than dry‐season production, and daily production fell to almost zero during January and February. Cool temperatures (<17 °C) may have limited fish activity and influenced detectability during some dry‐season censuses. Estimates of abundance and annual production by P. myersi are therefore conservative. 5. Comparisons with the literature indicate that the abundance and production of P. myersi in Hong Kong was high relative to other benthic fishes in tropical Asia, or their temperate counterparts in small streams. Manipulative experiments are needed to determine the influence of P. myersi, and algivorous balitorids in general, on periphyton dynamics and energy flow in Asian streams.     

Reproductive cycle of the Azorean endemic land snail Oxychilus (Drouetia) brincki Riedel, 1964 (Gastropoda: Pulmonata)

The life history of Oxychilus (Drouetia) brincki (Riedel, 1964), an endemic species from Santa Maria island (Azores), is addressed here for the first time. One population was analyzed for 1 year, to study the reproductive cycle and to assess the validity of three morphometric parameters as maturation diagnostic characters. The positive correlation between morphometric parameters (maximum diameter and number of whorls) can be useful to define the reproductive maturation status particularly during the active reproductive season. Oxychilus brincki is reproductively active between May and October/November; the residual values of mature oocytes between February and April allow reproduction to occur throughout the year. Our observations on gonadal maturation show that there is a functional protandric tendency and that photoperiod probably triggers the maturation process, whereas temperature acts mainly as a regulatory factor. Contrary to other terrestrial pulmonate snails that have a well demarcated reproductive cycle, Azorean species are gametogenically active throughout almost all the year.     

Biology of Latia neritoides Gray 1850 (Gastropoda,Pulmonata, Basommatophora): the Only Light-producing Freshwater Snail in the World

This report on the freshwater pulmonate basommatophoran Latia neritoides of the North Island of New Zealand is based on field and laboratory studies. It deals with aspects of the habitat, describes biological adaptations of organs and organ systems in Latia, reviews the chemical basis of bioluminescence in this species, discusses the possible roles of light production in Latia, and addresses zoogeographic and phylogenetic questions.     

Polymorphic microsatellites in the African freshwater snail,Bulinus forskalii (Gastropoda,Pulmonata)

Eleven microsatellites were isolated in the freshwater snail Bulinus forskalii, intermediate host for the medically important trematode Schistosoma intercalatum. Characterization in 60 snails from three populations of B. forskalii from Cameroon revealed 4 to 18 alleles per locus. Low observed heterozygosity but higher expected heterozygosity, high F_IS estimates, significant departures from Hardy–Weinberg equilibrium and genotypic linkage disequilibria all indicate that B. forskalii is a preferential selfer. High F_ST estimates suggest that effective dispersal is limited and genetic drift is an important determinant of genetic structure. The potential utility of the microsatellite primers in other closely related Bulinus species was explored.     

Population dynamics and population structure ofFerrissia wautieri (Mirolli, 1960) (Gastropoda,Ancylidae) in a pond near Nijmegen (The Netherlands)

Ferrissia wautieri, a freshwater limpet, is a widely distributed species in The Netherlands. In a pond near Nijmegen samples were taken twice a month over the year to study the population size and structure of this species in relation to the water temperature. Only ancyloids were found. Production of juveniles is temperature-dependent; peak numbers occurred in July and August. Just-hatched juveniles (shell length 0.6–1.0 mm) occurred over a lengthy period in the year, but were absent in March and April. During these months the collected numbers of specimens were very low. The largest specimens were collected during March, April and May.     

Eco-evolutionary dynamics: disentangling phenotypic,environmental and population fluctuations

Decomposing variation in population growth into contributions from both ecological and evolutionary processes is of fundamental concern, particularly in a world characterized by rapid responses to anthropogenic threats. Although the impact of ecological change on evolutionary response has long been acknowledged, the converse has predominantly been neglected, especially empirically. By applying a recently published conceptual framework, we assess and contrast the relative importance of phenotypic and environmental variability on annual population growth in five ungulate populations. In four of the five populations, the contribution of phenotypic variability was greater than the contribution of environmental variability, although not significantly so. The similarity in the contributions of environment and phenotype suggests that neither is worthy of neglect. Population growth is a consequence of multiple processes, which strengthens arguments advocating integrated approaches to assess how populations respond to their environments.     

Elements of cold hardiness in a littoral population of the land snail Helix aspersa (Gastropoda: Pulmonata)

The land snail Helix aspersa can be considered partially tolerant to freezing, in the sense it can survive some ice formation within its body for a limited time, and possesses a limited ability to supercool. This study aimed at understanding what factors are responsible for the variation of the temperature of crystallization ( Tc) in a littoral temperate population. The ability to supercool was maximal (ca. -5 degrees C) during dormancy periods (hibernation and aestivation) and minimal (ca. -3 degrees C) during spring and autumn, in relation with the decrease of water mass and the increase of osmolality. Tc decreased in October to remain stable through late autumn and winter; it increased quickly with the awakening of animals in April. Snails with an epiphragm had a significantly higher ability to supercool (ca. -4.8 degrees C) than snails which did not form an epiphragm (ca. -4.2 degrees C). The animals' size had a weak but significant influence on the realization of the Tc. It appeared that there was not a real cold-hardiness strategy in this population; rather a sum of parameters, varying in consequences of the external conditions and of the activity cycle, which are responsible for the enhancement of the supercooling ability during winter.     

Small effective population sizes in a widespread selfing species, Lymnaea truncatula (Gastropoda: Pulmonata)

We present here a spatial and temporal population genetic survey of a common freshwater snail, also a predominantly selfing species, Lymnaea truncatula. The rate of genetic diversity loss was quantified by estimating the effective size (Ne) of the snail populations, using two different methods. A temporal survey allowed estimation of a variance effective size of the populations, and a spatial survey allowed the estimation of an inbreeding effective size, from two-locus identity disequilibria estimates. Both methods were consistent and provided low Ne values. Drift due to (i) high amounts of selfing and (ii) fluctuations in population sizes because of temporary habitats, and also selection coupled to genome-wide linkage disequilibria, could explain such reductions in Ne. The loss of genetic diversity appears to be counterbalanced only very partially by low apparent rates of gene flow.     

Characterization of 15 polymorphic microsatellite markers in the freshwater snail Aplexa marmorata (Mollusca, Gastropoda)

We describe 15 new variable microsatellites in the freshwater snail Aplexa (Physa) marmorata, as well as conditions for multiplexing and simultaneous genotyping sets of loci. Two to 11 alleles were detected per locus over the five populations studied, and up to eight alleles per locus and population. The observed heterozygosity was extremely low, most probably as the consequence of high selfing rates. The genetic differentiation among populations was large. These markers will constitute efficient tools for investigating the population structure in fragmented habitats. Cross-species amplification in three Physa species (P. acuta, P. fontinalis and P. gyrina) was rather unsuccessful.     

Solar radiation stress in climbing snails: behavioural and intrinsic features define the Hsp70 level in natural populations of Xeropicta derbentina (Pulmonata)

Ectotherms from sunny and hot environments need to cope with solar radiation. Mediterranean land snails of the superfamily Helicoidea feature a behavioural strategy to escape from solar radiation-induced excessive soil heating by climbing up vertical objects. The height of climbing, and also other parameters like shell colouration pattern, shell orientation, shell size, body mass, actual internal and shell surface temperature, and the interactions between those factors may be expected to modulate proteotoxic effects in snails exposed to solar radiation and, thus, their stress response. Focussing on natural populations of Xeropicta derbentina, we conducted a ‘snapshot’ field study using the individual Hsp70 level as a proxy for proteotoxic stress. In addition to correlation analyses, an IT-model selection approach based on Akaike’s Information Criterion was applied to evaluate a set of models with respect to their explanatory power and to assess the relevance of each of the above-mentioned parameters for individual stress, by model averaging and parameter estimation. The analysis revealed particular importance of the individuals’ shell size, height above ground, the shell colouration pattern and the interaction height × orientation. Our study showed that a distinct set of behavioural traits and intrinsic characters define the Hsp70 level and that environmental factors and individual features strongly interact.