Intraspecific variation in resistance to desiccation and climatic gradients in the distribution of the land snail Xeropicta vestalis

Land snails are subject to desiccating conditions in their terrestrial habitat. Our previous studies have revealed significant differences in resistance to desiccation among closely related species and among genera that share a similar life style, suggesting that the distribution pattern is correlated with the abiotic regime in its specific habitat. Our present study extends the scope of comparison to the intraspecific level, by examining the resistance to desiccation in five populations of Xeropicta vestalis , a Mediterranean-dwelling species that ranges from the 1000 mm to the 200 mm isohyet.
The resistance to desiccation varied among populations in correlation with the specific habitat of each population and with the north-to-south and the west-to-easl climatic gradients. Furthermore, in one case it exceeded the resistance of other, desert-dwelling species ( Trochoidea simulata, Sphincterochila zonata). We suggest that, in spite of its physiological capacity to invade deserts, X. vestalis is prevented from doing so because it is an annual, semelparous species. The Negev Desert is an unpredictable environment, susceptible to year-to-year fluctuations to such an extent that one rainless year would wipe out all its populations.     

The heat shock response in congeneric land snails (Sphincterochila) from different habitats

Land snails are subject to daily and seasonal variations in temperature and in water availability, and use heat shock proteins (HSPs) as part of their survival strategy. We used experimental heat stress to test whether adaptation to different habitats affects HSP expression in two closely related Sphincterochila snail species, a desert species, Sphincterochila zonata, and a Mediterranean-type species, Sphincterochila cariosa. Our findings show that in S. cariosa, heat stress caused rapid induction of Hsp70 proteins and Hsp90 in the foot and kidney tissues, whereas the desert-inhabiting species S. zonata displayed delayed induction of Hsp70 proteins in the foot and upregulation of Hsp90 alone in the kidney. Our study suggests that Sphincterochila species use HSPs as part of their survival strategy following heat stress and that adaptation to different habitats results in the development of distinct strategies of HSP expression in response to heat, namely the reduced induction of HSPs in the desert-dwelling species. We suggest that the desert species S. zonata relies on mechanisms and adaptations other than HSP induction, thus avoiding the fitness consequences of continuous HSP upregulation.     

Heat shock proteins (Hsp70) and water content in the estivating Mediterranean Grunt Snail (Cantareus apertus)

Pulmonate land snails often are able to estivate to survive dry hot seasons were water and food are scarce. The aperture of the shell is closed with an epiphragm, and metabolism is depressed to approximately one fourth of basal metabolism. We investigated a molecular aspect of estivation focussing on the heat shock protein 70 (Hsp70) stress response during estivation in the Mediterranean Grunt Snail Cantareus apertus. Sequences of a new inducible hsp70 and of actin are presented and expression of the hsp70 gene as well as Hsp70 protein content was measured in estivating animals. Both Hsp70 protein and mRNA do not show a significant change from the control, although there is a trend that hsp70 mRNA is less abundant in estivating specimens. After heat shock, the expression of hsp70 increased and a higher Hsp70 protein content was detected. Water relations were also investigated. After a period of 6 months in the dormant state, the snails contained 14% less water than active ones, implying a constricted protection against desiccation, compared to the desert snail Sphincterochila zonata, and a Mediterranean-type water economy.     

Heat shock proteins and survival strategies in congeneric land snails (Sphincterochila) from different habitats

Polmunate land snails are subject to stress conditions in their terrestrial habitat, and depend on a range of behavioural, physiological and biochemical adaptations for coping with problems of maintaining water, ionic and thermal balance. The involvement of the heat shock protein (HSP) machinery in land snails was demonstrated following short-term experimental aestivation and heat stress, suggesting that land snails use HSPs as part of their survival strategy. As climatic variation was found to be associated with HSP expression, we tested whether adaptation of land snails to different habitats affects HSP expression in two closely related Sphincterochila snail species, a desert species Sphincterochila zonata and a Mediterranean-type species Sphincterochila cariosa. Our study suggests that Sphincterochila species use HSPs as part of their survival strategy following desiccation and heat stress, and as part of the natural annual cycle of activity and aestivation. Our studies also indicate that adaptation to different habitats results in the development of distinct strategies of HSP expression in response to stress, namely the reduced expression of HSPs in the desert-inhabiting species. We suggest that these different strategies reflect the difference in heat and aridity encountered in the natural habitats, and that the desert species S. zonata relies on mechanisms and adaptations other than HSP induction thus avoiding the fitness consequences of continuous HSP upregulation.     

Heat shock proteins and resistance to desiccation in congeneric land snails

Land snails are subject to daily and seasonal variations in temperature and in water availability and depend on a range of behavioral and physiological adaptations for coping with problems of maintaining water, ionic, and thermal balance. Heat shock proteins (HSPs) are a multigene family of proteins whose expression is induced by a variety of stress agents. We used experimental desiccation to test whether adaptation to different habitats affects HSP expression in two closely related Sphincterochila snail species, a desiccation-resistant, desert species Sphincterochila zonata, and a Mediterranean-type, desiccation-sensitive species Sphincterochila cariosa. We examined the HSP response in the foot, hepatopancreas, and kidney tissues of snails exposed to normothermic desiccation. Our findings show variations in the HSP response in both timing and magnitude between the two species. The levels of endogenous Hsp72 in S. cariosa were higher in all the examined tissues, and the induction of Hsp72, Hsp74, and Hsp90 developed earlier than in S. zonata. In contrary, the induction of sHSPs (Hsp25 and Hsp30) was more pronounced in S. zonata compared to S. cariosa. Our results suggest that land snails use HSPs as part of their survival strategy during desiccation and as important components of the aestivation mechanism in the transition from activity to dormancy. Our study underscores the distinct strategy of HSP expression in response to desiccation, namely the delayed induction of Hsp70 and Hsp90 together with enhanced induction of sHSPs in the desert-dwelling species, and suggests that evolution in harsh environments will result in selection for reduced Hsp70 expression.     

Resistance to desiccation and distribution patterns in bush-dwelling land snails

Bush-dwelling land snails are exposed to desiccating conditions that are more severe than those of snails that seek the shelter of rock crevices, litter or the upper layers of the soil. We studied the resistance to desiccation in four bush-dwelling species of Israeli snails to evaluate a possible relation between their water economy and their distribution pattern. The resistance to desiccation decreased in the following order: Xeropicta vestalis (a widely distributed Mediterranean species), Trochoidea simulata (a desert species), Theba pisana (a Mediterranean species from the sand dunes of the coastal plain), and Monacha haifaensis (a Mediterranean species). Xeropicta vestalis also had the quickest response to the desiccating conditions. It is probably prevented from establishing itself in the desert, in spite of its superior water economy, because it is an annual, semelparous species, and the desert is a highly unpredictable environment. An immediate response to desiccating conditions may be a key factor in the success of desert-inhabiting land snails. Snails from more humid regions take several days to recruit their water preserving mechanisms, a delay which may be crucial to their water balance. The conchiometrics of X. vestalis and T. simulata suggest that the main water preserving mechanisms of these species are located in the mantle, rather than in the shell and epiphragm.     

Heat shock protein expression in relation to reproductive cycle in land snails: Implications for survival

Land snails are subject to daily and seasonal variations in temperature and in water availability and use heat shock proteins (HSPs) as part of their survival strategy. We tested whether the reproductive cycle of land snails affects the endogenous levels of HSPs, and their involvement in the reproductive process. We examined HSP levels in the foot tissue of two Sphincterochila species, S. cariosa and S. zonata, before and after laying eggs, and analyzed the albumen gland (reproductive organ) of both species and eggs of S. cariosa for the presence and quantity of various HSPs. Our study shows reduction in the expression level of Hsp70 isoforms and Hsp90 in S. zonata foot and of Hsp74 in S. cariosa foot during the period preceding egg laying compared to the post-reproductive stage. Hsp70 isoforms and Hsp25 were highly expressed in both large albumen glands and in freshly laid eggs of S. cariosa, whereas large albumen glands of S. zonata expressed mainly Hsp70 isoforms. We conclude that a trade-off between survival and fertility is responsible for the expression level of HSPs in the foot tissue of Sphincterochila snails. Our study shows that HSPs are involved in the reproductive process. We propose that parental provision of HSPs may be part of a "be prepared" strategy of Sphincterochila snails, and that HSPs may play important roles in the survival strategy of land snails during the early life stages. Our observations also highlight the importance of the reproductive status in study of whole organisms, especially when assessing the HSP response to stress.     

Climatic niche partitioning following successive invasions by fruit flies in La Réunion

1. Biological invasions have profound effects on community structure. The community composition following invasions can be influenced by the habitat diversity and the species' responses to abiotic factors. 2. We evaluated the tolerance to climatic factors and analysed the field distribution of four polyphagous fruit flies (Diptera: Tephritidae) of La Réunion Island (three exotic species that successively invaded the island and the endemic species Ceratitis catoirii) in order to evaluate the opportunities of coexistence by niche differentiation. 3. Atmospheric humidity and immersion in water in the laboratory greatly influence the survival of fruit fly pupae. While C. catoirii and C. rosa are very sensitive to desiccation, C. capitata and especially Bactrocera zonata are relatively tolerant. B. zonata also tolerated immersion in water much longer than did C. rosa and C. catoirii, that in turn were more resistant than C. capitata. Overall, field distributions agree with the predictions based on this study of humidity combined with previous data on the effects of temperature. 4. Climatic niche partitioning promotes coexistence between some but not all pairs of invasive species. Thus, C. rosa can coexist with both C. capitata and B. zonata at the regional scale, while climatic niches are not different enough to promote coexistence of the latter two species. The endemic species has no private climatic niche either and this now very rare species could be in the process of extinction. 5. By promoting coexistence or not, climatic diversity in invaded areas can directly affect the community composition following invasions.     

Intraspecific variation in resistance to desiccation and climatic gradients in the distribution of the bush-dwelling land snail Trochoidea simulata

Resistance to desiccation was examined in six populations of the Israeli bush-dwelling snail Trochoidea simulata , a desert species distributed mainly between the 100–200 mm isohyets.
The present study revealed significant intraspecific differences in resistance to desiccation which are correlated with habitat and climatic gradients within the distribution range of the species. Populations from more arid sites were more resistant to desiccation and heat exposure than those from more mesic areas. However, the population from the Rift Valley (an extremely arid region) was surprisingly poorly resistant. Rates of population water loss under the controlled experimental conditions in the laboratory generally matched the calculated water losses during natural summer aestivation.     

Including climate change in pest risk assessment: the peach fruit fly, Bactrocera zonata (Diptera: Tephritidae)

Bactrocera zonata (Saunders) is one of the most harmful species of Tephritidae. It causes extensive damage in Asia and threatens many countries located along or near the Mediterranean Sea. The climate mapping program, CLIMEX 3.0, and the GIS software, ArcGIS 9.3, were used to model the current and future potential geographical distribution of B. zonata. The model predicts that, under current climatic conditions, B. zonata will be able to establish itself throughout much of the tropics and subtropics, including some parts of the USA, southern China, southeastern Australia and northern New Zealand. Climate change scenarios for the 2070s indicate that the potential distribution of B. zonata will expand poleward into areas which are currently too cold. The main factors limiting the pest's range expansion are cold, hot and dry stress. The model's predictions of the numbers of generations produced annually by B. zonata were consistent with values previously recorded for the pest's occurrence in Egypt. The ROC curve and the AUC (an AUC of 0.912) were obtained to evaluate the performance of the CLIMEX model in this study. The analysis of this information indicated a high degree of accuracy for the CLIMEX model. The significant increases in the potential distribution of B. zonata projected under the climate change scenarios considered in this study suggest that biosecurity authorities should consider the effects of climate change when undertaking pest risk assessments. To prevent the introduction and spread of B. zonata, enhanced quarantine and monitoring measures should be implemented in areas that are projected to be suitable for the establishment of the pest under current and future climatic conditions.     

Structural adaptation for reducing water-loss in three species of terrestrial snail

Laboratory measurements indicate that observed differences in the rate of water-loss from inactive Helix aspersa, Otala lactea and Sphincterochila boissieri are due to morphological adaptations such as a thicker shell, reduced aperture and thicker epiphragm. These modifications enable terrestrial snails to inhabit dryer environments by increasing the length of time the snail can remain inactive.     

Where and why? Bees,snail shells and climate: Distribution of Rhodanthidium (Hymenoptera: Megachilidae) in the Iberian Peninsula

Species distribution patterns are widely studied through species distribution models (SDMs), focusing mostly on climatic variables. Joint species distribution models (JSDMs) allow inferring if other factors (biotic interactions, shared phylogenetic history or other unmeasured variables) can also have an influence on species distribution. We identified current distributional areas and optimal suitability areas of three species of the solitary snail‐shell bee Rhodanthidium (Hymenoptera: Megachilidae), and their host gastropod species in the Iberian Peninsula. We undertook SDMs using Maxent software, based on presence points and climatic variables. We also undertook JSDMs for the bees and the snails to infer if co‐occurrence could be a result of biotic interactions. We found that the three bee species: (1) use at least five different species of Mediterranean snails; (2) use empty shells not only for nesting but also for sheltering when there is adverse weather and during the night; (3) have their most suitable areas in the eastern and southern Iberian Peninsula, mostly on limestone areas; and (4) have their optimal range under Mediterranean climatic values for the studied variables. There is positive co‐occurrence of Rhodanthidium with the gastropod species, especially with the snail Sphincterochila candidissima. The contribution of the environmental component to the co‐occurrence is less than that of the residual component in those cases, suggesting that: (i) the use of biotic resources (between Rhodanthidium and the gastropod species); (ii) shared phylogenetic history (between R. septemdentatum and R. sticticum); or (iii) unmeasured variables are largely responsible for co‐occurrence.     

Patterns,origin and possible effects of sediment pollution in a Mediterranean lake

The influence of water permanence and high intra- and inter-annual hydrological variability on macrobenthos (organisms >1 mm) was studied using a taxonomical and a functional approach. The study was carried out in a Mediterranean salt marsh. Monthly samples of macrobenthic fauna were collected during two consecutive hydroperiods from six ponds with different water permanence (temporary, semi-permanent and permanent waters). Organisms were assigned to five functional response groups based on life-strategies according to their capacity to survive desiccation events, their dispersion capability and the necessity of water for their reproduction. Results from both approaches showed that the benthic community was more related to pond type than to intra- and inter-annual variability. The second aim was to analyse to which extent patterns in functional groups were determined by the existence of succession patterns or to environmental variability. In this sense, a clear succession pattern was not observed. In contrast, in most of the functional groups (4 out of 5), species within each functional group showed similar responses to water fluctuations. However, species of the fifth functional group, which comprised species without any particular adaptation to desiccation survival or avoidance, showed different responses to water level fluctuations. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Desiccation resistance and water balance in southern African keratin beetles (Coleoptera, Trogidae): the influence of body size and habitat

Desiccation resistance and water balance were examined in the adults of seven trogid species, which differed both in body size and in the habitats from which they were collected. Body water contents (51–58% fresh mass) and desiccation rates at 27 °C (0.00026–0.00093 g h⁻¹) in these species were very similar to those of unrelated, similar-sized beetles from arid habitats. The keratin beetles differed markedly from many other adult Coleoptera by virtue of their very high haemolymph osmolality and inability to regulate haemolymph osmolality, and to catabolise lipids for water production, during desiccation. Like most other insects, the xeric trogid species had lower rates of water loss and longer survival times than trogids from mesic areas. This was due both to lower rates of water loss and to the larger body size of species from the more arid areas. Because absolute body water content was higher in large beetles than in small ones, larger body size conferred higher desiccation resistance on the very large Kalahari desert species. This suggests that there may be strong selection for large body size in such insects from arid areas. Most ecological and ecophysiological investigations of geographical variation in body size, and the species-body size distribution, have focused on temperature and metabolic rate as explanatory variables. This study suggests that attention should also be given to desiccation resistance. Accepted: 29 September 1997     

Geographic variation for climatic stress resistance traits in the springtail Orchesella cincta

Multiple traits of stress resistance were investigated in the epedaphic springtail Orchesella cincta. Second generation adults from five laboratory populations were compared with respect to resistance to extreme temperatures and desiccation, and traits relevant to climatic adaptation. Populations were collected along a 2000-km latitudinal gradient ranging from Denmark to southern Italy and reared under the same standard laboratory conditions. Traits investigated were resistance to high and low temperature, desiccation resistance, body size and water loss rate (WLR). Results showed genetically based differences in resistance to high and low temperature, desiccation, WLR, water pool and body size between populations. Individuals from the most northern population had the highest desiccation-and cold shock resistance, and the lowest heat shock resistance. Females were significantly more desiccation resistant than males. The results of cold shock resistance showed a positive increase with lowest environmental temperature recorded at the sites of population origin, whereas heat shock resistance showed a positive increase with highest recorded temperature at the sites of population origin. Desiccation resistance increased towards the most southern and northern population, suggesting that both low and high temperature extremes affect desiccation resistance. Body mass, water pool and WLR showed interpopulation as well as sex specific variation. This provides evidence for geographical variation in stress resistance of springtails related to climatic conditions.     

The importance of cuticular permeability, osmolyte production and body size for the desiccation resistance of nine species of Collembola

Euedaphic collembolans have recently been shown to actively regulate internal osmotic pressure by means of sugars and polyols in response to desiccation. In contrast, studies of cuticular permeability have shown that some, especially epedaphic, species of collembolans may primarily rely on a low cuticular permeability to survive desiccation. To elucidate to what extent these strategies are important for desiccation resistance, the survival of 7-day acute desiccation stress (LRH(50)), the cuticular water conductance constant and osmolyte production were investigated in nine species of collembolans, covering euedaphic, hemiedaphic and epedaphic species. The LRH(50) values ranging from 98.8% to 95.2% RH showed no correlation with the vertical distribution of species, since both the highest and lowest values were found in epedaphic species. The water conductance varied from 698+/-141 to 41+/-13 microg h(-1) cm(-2) mmHg(-1) and showed good agreement with the vertical distribution of species in their natural habitats. Modelling the drying curves showed that, in addition to cuticular permeability and osmolyte production, body size also plays an important role in the survival of short-term severe desiccation stress. Furthermore, the model pointed to the need for behavioural responses to desiccation, particularly in epedaphic species. Thus, in keeping with expected humidity regimes in their respective microhabitats, euedaphic species rely on small body size, high cuticular permeability and the ability to actively regulate the osmotic pressure of their body fluids, hemiedaphic species have similar strategies but with reduced cuticular permeability, whereas in epedaphic species, active regulation of osmotic pressure is replaced by combinations of greatly reduced cuticular permeability or greatly reduced surface area to volume ratio combined with behavioural responses to desiccation.     

Traits underpinning desiccation resistance explain distribution patterns of terrestrial isopods

Predicted changes in soil water availability regimes with climate and land-use change will impact the community of functionally important soil organisms, such as macro-detritivores. Identifying and quantifying the functional traits that underlie interspecific differences in desiccation resistance will enhance our ability to predict both macro-detritivore community responses to changing water regimes and the consequences of the associated species shifts for organic matter turnover. Using path analysis, we tested (1) how interspecific differences in desiccation resistance among 22 northwestern European terrestrial isopod species could be explained by three underlying traits measured under standard laboratory conditions, namely, body ventral surface area, water loss rate and fatal water loss; (2) whether these relationships were robust to contrasting experimental conditions and to the phylogenetic relatedness effects being excluded; (3) whether desiccation resistance and hypothesized underlying traits could explain species distribution patterns in relation to site water availability. Water loss rate and (secondarily) fatal water loss together explained 90 % of the interspecific variation in desiccation resistance. Our path model indicated that body surface area affects desiccation resistance only indirectly via changes in water loss rate. Our results also show that soil moisture determines isopod species distributions by filtering them according to traits underpinning desiccation resistance. These findings reveal that it is possible to use functional traits measured under standard conditions to predict soil biota responses to water availability in the field over broad spatial scales. Taken together, our results demonstrate an increasing need to generate mechanistic models to predict the effect of global changes on functionally important organisms.     

Impact of body melanisation on contrasting levels of desiccation resistance in a circumtropical and a generalist <Emphasis Type="Italic">Drosophila</Emphasis> species

We investigated the role of cuticular lipids, body melanisation and body size in conferring contrasting levels of desiccation resistance in latitudinal populations of Drosophila melanogaster and Drosophila ananassae on the Indian subcontinent. Contrary to the well known role of cuticular lipids in water proofing in diverse insect taxa, there is lack of geographical variations in the amount of cuticular lipids per fly in both the species. In D. ananassae, quite low levels of body melanisation are correlated with lower desiccation resistance. By contrast, increased levels of desiccation resistance are correlated with quite high melanisation in D. melanogaster. Thus, species specific cuticular melanisation patterns are significantly correlated with varying levels of desiccation resistance within as well as between populations and across species. Role of body melanisation in desiccation resistance is further supported by the fact that assorted dark and light flies differ significantly in cuticular water loss, hemolymph and dehydration tolerance. However, similar patterns of body size variation do not account for contrasting levels of desiccation resistance in these two Drosophila species. Climatic selection is evidenced by multiple regression analysis with seasonal amplitude of thermal and humidity changes (Tcv and RHcv) along latitude on the Indian subcontinent. Finally, the contrasting levels of species specific distribution patterns are negatively correlated with RHcv of sites of origin of populations i.e. a steeper negative slope for D. ananassae corresponds with its desiccation sensitivity as compared with D. melanogaster. Thus, evolutionary changes in body melanisation impact desiccation resistance potential as well as distribution patterns of these two Drosophila species on the Indian subcontinent.     

桃儿七光合生理特性的地理差异研究

对中国由南向北5个种源地桃儿七(Sinopodophyllum hexandrum)的光合生理生态特性进行了研究.结果表明:(1)北部宁夏六盘山地区植株的光饱和点(LSP)和光补偿点(LCP)最高,表观光量子效率(AQY)、瞬时光能利用效率(ILUE)和最大光合速率(Pmax)最低,光合能力最差;最南部云南纳帕海植株的LSP和LCP很低,但因其AQY和ILUE最高,故其Pmax最大,强光下的光合能力最高.(2)最南部云南纳帕海和最北部甘肃兴隆山个体的羧化效率(CE)最高,且CO2补偿点(CCP)较低,因此CO2利用率较高;而较南部四川刷经寺个体的CE最低,且CCP最高,故CO2利用效率最低.(3)各种源植株叶片的叶绿素总量及叶绿素a含量无显著差异,而不同地区个体的Chla/Chlb值差异显著,最北部甘肃兴隆山植株的Chla/Chlb值最大,而南部四川刷经寺的Chla/Chlb值最小.(4)各种源地植株的水分利用效率(WUE)和蒸腾速率(Tr)对光量子通量密度(PFD)与CO2浓度的响应过程也表现出显著的地理分异.总的来看,桃儿七是一种喜光但又较耐阴植物,但不同分布区桃儿七的光合生理生态特征已经出现了较显著的地理分化.