Relictual physiological ecology in the threatened land snail Codringtonia helenae: A cause for decline in a changing environment?

Land snails often exhibit intra-annual cycles of activity interspersed by periods of dormancy (hibernation/aestivation), accompanied by a range of behavioural and physiological adaptations to ensure their survival under adverse environmental conditions. These adaptations are useful to understand species-specific habitat requirements and to predict their response to environmental changes. We examined the seasonal physiological and biochemical composition patterns of the threatened land snail Codringtonia helenae, endemic to Greece, in relation to its behavioural ecology and climatic conditions. Fuel reserves (glycogen, lipids, proteins) and water were accumulated prior to aestivation, but subsequently were rapidly depleted. LDH exhibited substantial rise during aestivation, suggesting that anaerobic pathways may provide additional energy. The major outcome of our study was the unambiguous discrimination of the four life-cycle periods. Most remarkable was the clear distinction of the aestivation period, with hibernation, the other dormancy period, showing similarity with the two active periods but not with aestivation. We observed disassociation between behavioural and physiological responses and climatic conditions. The physiological responses of C. helenae were effective during hibernation, but only partly compensate the effect of adverse conditions during aestivation, since its aestivating behaviour is occasional and time limited. Perhaps, the behavioural ecology of Codringtonia is relictual and shaped during past environmental conditions. This constitutes an important extinction threat considering the current climatic trends and the deterioration of the habitat of that species due to human activities.     

Adjustment of metabolite composition in the haemolymph to seasonal variations in the land snail <Emphasis Type="Italic">Helix pomatia</Emphasis>

In temperate regions, land snails are subjected to subzero temperatures in winter and hot temperatures often associated to drought in summer. The response to these environmental factors is usually a state of inactivity, hibernation and aestivation, respectively, in a temperature and humidity buffered refuge, accompanied by physiological adjustments to resist cold or heat stress. We investigated how environmental factors in the microhabitat and body condition influence the metabolite composition of haemolymph of the endangered species Helix pomatia. We used UPLC and GC–MS techniques and analyzed annual biochemical variations in a multivariate model. Hibernation and activity months differed in metabolite composition. Snails used photoperiod as cue for seasonal climatic variations to initiate a physiological state and were also highly sensitive to temperature variations, therefore constantly adjusting their physiological processes. Galactose levels gave evidence for the persistence of metabolic activity with energy expenditure during hibernation and for high reproductive activity in June. Triglycerides accumulated prior to hibernation might act as cryoprotectants or energy reserves. During the last month of hibernation snails activated physiological processes related to arousal. During activity, protein metabolism was reflected by high amino acid level. An exceptional aestivation period was observed in April giving evidence for heat stress responses, like the protection of cells from dehydration by polyols and saccharides, the membrane stabilization by cholesterol and enhanced metabolism using the anaerobic succinic acid pathway to sustain costly stress responses. In conclusion, physiological adjustments to environmental variations in Helix pomatia involve water loss regulation, cryoprotectant or heatprotectant accumulation.     

Metabolic depression during aestivation does not involve remodelling of membrane fatty acids in two Australian frogs

Changes in membrane lipid composition (membrane remodelling) have been associated with metabolic depression in some aestivating snails but has not been studied in aestivating frogs. This study examined the membrane phospholipid composition of two Australian aestivating frog species Cyclorana alboguttata and Cyclorana australis. The results showed no major membrane remodelling of tissue in either frog species, or in mitochondria of C. alboguttata due to aestivation. Mitochondrial membrane remodelling was not investigated in C. australis. Where investigated in C. alboguttata, total protein and phospholipid content, and citrate synthase (CS) and cytochrome c oxidase (CCO) activities in tissues and mitochondria mostly did not change with aestivation in liver. In skeletal muscle, however, CS and CCO activities, mitochondrial and tissue phospholipids, and mitochondrial protein decreased with aestivation. These decreases in muscle indicate that skeletal muscle mitochondrial content may decrease during aestivation. Na⁺K⁺ATPase activity of both frog species showed no effect of aestivation. In C. alboguttata different fat diets had a major effect on both tissue and mitochondrial phospholipid composition indicating an ability to remodel membrane composition that is not utilised in aestivation. Therefore, changes in lipid composition associated with some aestivating snails do not occur during aestivation in these Australian frogs.     

Water balance and resistance to desiccation in rock-dwelling snails

We have examined the resistance to desiccation among rock-dwelling land snails of various phylogenetic groups:Cristataria genezarethana (Clausiliidae),Rupestrella rhodia (Chondrinidae) andLevantina caesareana (Helicidae), all from the same location in Israel.L. caesareana was the most resistant andR. rhodia the least resistant to desiccation andC. genezarethana was of intermediate resistance. Differences in the rates of water loss during desiccation were determined mainly by rate of water loss during the first 2 days of desiccation. The high rates of water loss in rock-dwelling species exceed those of other snails in the Mediterranean habitat of Israel. However, snails collected in the field at the end of aestivation were in only a mild state of dehydration, suggesting that the rocky habitat protects its occupants against desiccation. We also suggest that among the rock-dwelling species, the protective role of the rock is more important in the more evolutionarily primitive genera (the chondrinidRupestrella and the clausiliidCristataria) and that physiological capacities are more effective in the more highly evolved helicidLevantina.     

Enzyme activity in the aestivating Green-striped burrowing frog (Cyclorana alboguttata)

Green-striped burrowing frogs (Cyclorana alboguttata) can depress their resting metabolism by more than 80% during aestivation. Previous studies have shown that this species is able to withstand long periods of immobilisation during aestivation while apparently maintaining whole muscle mass and contractile performance. The aim of this study was to determine the effect of prolonged aestivation on the levels of metabolic enzymes (CCO, LDH and CS) in functionally distinct skeletal muscles (cruralis, gastrocnemius, sartorius, iliofibularis and rectus abdominus) and liver of C. alboguttata. CS activity was significantly reduced in all tissues except for the cruralis, gastrocnemius and the liver. LDH activity was significantly reduced in the sartorius and rectus abdominus, but remained at control (active) levels in the other tissues. CCO activity was significantly reduced in the gastrocnemius and rectus abdominus, and unchanged in the remaining tissues. Muscle protein was significantly reduced in the sartorius and iliofibularis during aestivation, and unchanged in the remaining muscles. The results suggest that the energy pathways involved in the production and consumption of ATP are remodelled during prolonged aestivation but selective. Remodelling and subsequent down-regulation of metabolic activity seem to target the smaller non-jumping muscles, while the jumping muscles retain enzyme activities at control levels during aestivation. These results suggest a mechanism by which aestivating C. alboguttata are able to maintain metabolic depression while ensuring that the functional capacity of critical muscles is not compromised upon emergence from aestivation.     

Physiology of drought tolerance and cold hardiness of the Mediterranean tiger moth Cymbalophora pudica during summer diapause

Prepupae of the arctiid moth Cymbalophora pudica spend spring and summer months in a summer diapause (aestivation), the duration of which is photoperiodically controlled. Cold hardiness, drought tolerance and some physiological and biochemical parameters were measured in aestivating prepupae. Large amounts of metabolic reserves, in the form of lipids and glycogen, accumulated prior to aestivation. Glycogen served as the main metabolic fuel for aestivating prepupae. Metabolic rate decreased rapidly after the onset of the inactive prepupal stage and remained low (5-15% of the level in active larva) during aestivation. A spontaneous increase of the respiration rate occurred before pupation. Neither low mol. wt sugars or alcohols (polyols) accumulated nor the haemolymph osmotic pressure changed during aestivation. Drought tolerance of aestivating prepupae was high (no decrease in survival after exposure to r.h.<10% at a temperature of 23 degrees C for a substantial part of diapause) owing to their extensive capacity to stabilize the relative body water content irrespective of the r.h. of surrounding air. Cold hardiness was low (>90% decrease in survival after exposure to -7 degrees C for 24h). Cold and drought acclimations did not lead to significant changes in the measured physiological and biochemical parameters but cold (not drought) acclimation caused a significant increase in cold hardiness. Neither drought tolerance nor the increase in cold hardiness after cold acclimation appear to be related to presence/accumulation of polyols in aestivating C. pudica prepupae.     

Influence of Molting and Starvation on Digestive Enzyme Activities and Energy Storage in Gammarus fossarum

Among the many biological responses studied in ecotoxicology, energy-based biomarkers such as digestive enzyme activities and energy reserves appear to be useful predictive tools for detecting physiological disturbances in organisms. However, the use of these biological responses as biomarkers could be limited by the effects of confounding factors (biotic and abiotic) and physiological processes, such as the reproductive cycle. Thus, the optimal use of these biomarkers will be facilitated by understanding the effects of these factors on the energy metabolism of the sentinel species being studied. We considered abiotic factors (temperature and conductivity) in a previous study, whereas the present study investigated the effects of gender, the female reproductive stage, and food availability on the digestive enzyme activities and energy storage of Gammarus fossarum. The results indicated that, during the female reproductive cycle, the activities of digestive enzymes (amylase, cellulase, and trypsin) decreased significantly, whereas the levels of reserves (proteins, lipids, and sugar) increased until the last premolt stage. Restricted food diets only led to decreased amylase activities in both sexes. Food starvation also induced a decrease in the energy outcomes in females, whereas there were no effects in males. In general, the biochemical (digestive enzyme activities) and physiological (energy reserves) responses were more stable in males than in females. These results support the use of males fed ad libitum to limit the effects of confounding factors when using these energy biomarkers in Gammarus fossarum during biomonitoring programs.     

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.     

Validation of Fecal Glucocorticoid Metabolite Assays for South African Herbivores

ABSTRACT Fecal glucocorticoid metabolite (FGM) assays are a popular means of monitoring adrenocortical activity (i.e., physiological stress response) in wildlife. Species-specific differences in glucocorticoid metabolism and excretion require assay validation, including both laboratory and biological components, before assay use in new species. We validated a commercially available radioimmunoassay (MP ¹²⁵I corticosterone RIA kit [MP Biomedicals, Solon, OH]) for measuring FGMs of several South African herbivores, including giraffe (Giraffa camelopardalis), impala (Aepyceros melampus), nyala (Tragelaphus buxtoni), kudu (Tragelaphus strepsiceros), wildebeest (Connochaetes taurinus), and zebra (Equus burchelli). These herbivores are important in South African parks and reserves for ecotourism and as a prey base for predators and serve an integral role in ecosystem processes. Standard biochemical validations (e.g., recovery of exogenous corticosterone, intra- and interassay variation, and parallelism) demonstrated that the assay accurately and precisely measured FGMs of all 6 herbivore species. Our biological validations demonstrated that the assay was sensitive enough to detect changes in FGM production associated with season. Samples collected during the dry season (Jun-Aug) contained higher FGM concentrations than those from the wet season (Dec-Feb) in all species. We established optimal sample dilutions and reference FGM levels for these 6 herbivores, which can now be used to monitor the effects of management and ecotourism activities on the stress responses of these herbivores.     

Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms

Human alteration of habitat has increased the proportion of forest edge in areas of previously continuous forest. This edge habitat facilitates invasion of exotic species into remaining fragments. The ability of native species to resist invasion varies and may depend on intrinsic variables such as dispersal and reproductive rates as well as external factors such as rate of habitat change and the density of populations of introduced species in edge habitat. We examined the distributional and competitive relationships of two members of the class Chilopoda, Scolopocryptops sexspinosus, a centipede native to the eastern US, and Lithobius forficatus, an exotic centipede introduced from Europe. We found that L. forficatus was most abundant in edge habitat and S. sexspinosus was most abundant in the interior habitat at our field sites. Although L. forficatus was present in habitat interiors at 11 of 12 sites, there was no correlation between fragment size and numbers of L. forficatus in interior habitat. The native centipede was rarely found occupying fragment edges. We used laboratory microcosms to examine potential competitive interactions and to indirectly assess prey preferences of the two species. In microcosms both species consumed similar prey, but the native centipede, S. sexspinosus, acted as an intraguild predator on the introduced centipede. Native centipedes were competitively superior in both intraspecific and interspecific pairings. Our results suggest that intraguild predation may aid native centipedes in resisting invasion of introduced centipedes from edge habitat.     

Is Aquatic Life Correlated with an Increased Hematocrit in Snakes?

Background

Physiological adaptations that allow air-breathing vertebrates to remain underwater for long periods mainly involve modifications of the respiratory system, essentially through increased oxygen reserves. Physiological constraints on dive duration tend to be less critical for ectotherms than for endotherms because the former have lower mass-specific metabolic rates. Moreover, comparative studies between marine and terrestrial ectotherms have yet to show overall distinct physiological differences specifically associated with oxygen reserves.

Methodology/Principal Findings

We used phylogenetically informed statistical models to test if habitat affects hematocrit (an indicator of blood oxygen stores) in snakes, a lineage that varies widely in habitat use. Our results indicate that both phylogenetic position (clade) and especially habitat are significant predictors of hematocrit. Our analysis also confirms the peculiar respiratory physiology of the marine Acrochordus granulatus.

Conclusion/Significance

Contrary to previous findings, marine snakes have significantly–albeit slightly–elevated hematocrit, which should facilitate increased aerobic dive times. Longer dives could have consequences for foraging, mate searching, and predation risks. Alternatively, but not exclusively, increased Hct in marine species might also help to fuel other oxygen-demanding physiological adaptations, such as those involved in osmoregulation.     

Are invasive species drivers of native species decline or passengers of habitat modification? A case study of the impact of the common myna (Acridotheres tristis) on Australian bird species

Habitat modification and invasive species are significant drivers of biodiversity decline. However, distinguishing between the impacts of these two drivers on native species can be difficult. For example, habitat modification may reduce native species abundance, while an invasive species may take advantage of the new environment. This scenario has been described as the driver‐passenger model, with ‘passengers’ taking advantage of habitat modification and ‘drivers’ causing native species decline. Therefore, research must incorporate both habitat modification and invasive species impact to successfully investigate native species decline. In this paper, we used the common myna (Acridotheres tristis) as a case study to investigate the driver‐passenger model. We investigated changes in bird abundance, over 2 years, in relation to different habitat types and common myna abundance. We hypothesized that the common myna is both a passenger of habitat change and a driver of some bird species decline. Our results indicated that the abundance of many native species is greater in high tree density nature reserves, while the common myna was uncommon in these areas. Common myna abundance was almost three times higher in urban areas than nature reserves and declined rapidly as tree density in nature reserves increased. Our findings indicated that the common myna is primarily a passenger of habitat change. However, we also observed negative associations between common myna abundance and some bird species. We stress the importance of simultaneously investigating both invasive species impact and habitat modification. We suggest habitat restoration could be a useful tool for both native species recovery and invasive species control. Understanding the drivers of native species decline will help inform impact mitigation and direct further research.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Environmental variables associated with the distribution and occupancy of habitat specialist tadpoles in naturally acidic,oligotrophic waterbodies

Environmental factors play an integral role, either directly or indirectly, in structuring faunal assemblages. Water chemistry, predation, hydroperiod and competition influence tadpole assemblages within waterbodies. We surveyed aquatic predators, habitat refugia, water height and water chemistry variables (pH, salinity and turbidity) at 37 waterbodies over an intensive 22‐day field survey to determine which environmental factors influence the relative abundance and occupancy of two habitat specialist anuran tadpole species in naturally acidic, oligotrophic waterbodies within eastern Australian wallum communities. The majority of tadpoles found were of Litoria olongburensis (wallum sedge frog) and Crinia tinnula (wallum froglet) species, both habitat specialists that are associated with wallum waterbodies and listed as Vulnerable under the IUCN Red List. Tadpoles of two other species (Litoria fallax (eastern sedge frog), and Litoria cooloolensis (cooloola sedge frog)) were recorded from two waterbodies. Tadpoles of Litoria gracilenta (graceful treefrog) were recorded from one waterbody. Relative abundance and occupancy of L. olongburensis tadpoles were associated with pH and water depth. Additionally, L. olongburensis tadpole relative abundance was negatively associated with turbidity. Waterbody occupancy by C. tinnula tadpoles was negatively associated with predatory fish and water depth and positively associated with turbidity. Variables associated with relative abundance of C. tinnula tadpoles were inconclusive and further survey work is required to identify these environmental factors. Our results show that the ecology of specialist and non‐specialist tadpole species associated with ‘unique’ (e.g. wallum) waterbodies is complex and species specific, with specialist species likely dominating unique habitats.     

Differential effect of wave stress on the physiology and behaviour of native versus non-native benthic invertebrates

In fresh waters, non-native invertebrate species preferentially spread via navigation waterways, where they often dominate assemblages. Littoral habitats in navigation waterways are regularly exposed to ship-induced waves. We conducted experiments to test the effects of artificial wave exposure on the relative performance of wide-spread native and non-native species. We compared physiological and behavioural response variables (i.e. growth rate, glycogen content, feeding and swimming activity) of two amphipods (native Gammarus roeselii and non-native Dikerogammarus villosus) and two gastropods (native Bithynia tentaculata and non-native Physella acuta) subject to wave and control (i.e. no wave) treatment flumes across a 6-week period. Growth, and in part glycogen content (as a measure of energy storage), were significantly reduced after exposure to waves in native invertebrates, but not in non-native invertebrates. The reduction in growth may be associated with the disturbance effects of waves, such as the higher swimming activity of G. roeselii and lower food uptake of B. tentaculata. In comparison, the effective hiding behaviour observed for D. villosus and good swimming ability of P. acuta, were identified as important traits facilitating the successful colonisation of the harsh habitat conditions of littoral waterways. Our study demonstrates that artificial wave regimes may contribute significant selective pressure, thus explaining the observed dominance of non-native species in navigational waterways. The success of non-native species under the harsh hydraulic habitat conditions of these socio-economically driven ecosystems may consequently be traced directly to behavioural and/or physiological traits.     

Carbohydrate and amino acid metabolism in fasting and aestivating African lungfish (Protopterus dolloi)

The potential importance of carbohydrates and amino acids as fuels during periods of fasting and aestivation in the African lungfish, Protopterus dolloi, were examined. No significant decreases in tissue glycogen levels were observed following 60 days of fasting or aestivation, suggesting lungfish may undergo 'glycogen sparing'. Yet glycogenolysis may be important during aestivation based on the differing responses of two flux-generating enzymes of the glycolytic pathway, hexokinase (HK) and pyruvate kinase (PK). PK is required for glycogen breakdown whereas HK is not. HK activity is significantly down-regulated in the heart and gill tissues during aestivation, while PK activity is sustained. The significant negative correlation between the activity of HK and glucose levels in the heart of aestivating lungfish suggests HK may be regulated by glucose concentrations. There was no indication of anaerobic glycolytic flux during aestivation as lactate did not accumulate in any of the tissues examined, and no significant induction of lactate dehydrogenase (LDH)activity was observed. The increase in glutamate dehydrogenase (GDH) and aspartate aminotransferase (Asp-AT) activities in the liver of aestivating P. dolloi suggests some energy may be obtained via increased aminoacid catabolism, leading to the generation of tricarboxylic acid (TCA) cycle intermediates. These findings indicate the importance of both carbohydrate and amino acid fuel stores during aestivation in aphylogenetically ancient, air-breathing fish.     

Metabolic depression during aestivation in Cyclorana alboguttata

The green striped burrowing frog, Cyclorana alboguttata, spends, on average, nine to ten months of every year in aestivation. Recently, C. alboguttata has been the focus of much investigation regarding the physiological processes involved in aestivation, yet our understanding of this frog's capacity to metabolically depress remains limited. This study aimed to extend our current knowledge of metabolic depression during aestivation in C. alboguttata. C. alboguttata reduced whole animal metabolism by 82% within 5 weeks of aestivation. The effects of aestivation on mass specific in vitro tissue metabolic rate (VO₂) varied among individual organs, with muscle and liver slices showing significant reductions in metabolism; kidney VO₂ was elevated and there was no change in the VO₂ of small intestine tissue slices. Organ size was also affected by aestivation, with significant reductions in the mass of all tissues, except the gastrocnemius. These reductions in organ size, combined with changes in mass specific VO₂ of tissue slices, resulted in further energy savings to aestivating animals. This study shows that C. alboguttata is capable of selectively down- or up-regulating individual tissues, using both changes in metabolic rate and morphology. This strategy allows maximal energy savings during aestivation without compromising organ functionality and survival at arousal.     

ALCOHOL TOLERANCE,ADH ACTIVITY,AND ECOLOGICAL NICHE OF DROSOPHILA SPECIES

In vitro alcohol dehydrogenase (ADH) activity was measured in adults of species belonging to Drosophila and to the related genus Zaprionus. Data were analyzed according to the known breeding sites and the level of ethanol tolerance of these species. Alcohol dehydrogenase activity was assayed with both ethanol (E) and isopropanol (I). Our results show a very broad range of activities among the 71 species investigated, the ratio of the highest value observed (D. melanogaster) to the lowest (D. pruinosa) being 65:1. A general positive correlation was found between the level of ADH activity and the capacity to detoxify ethanol. Nevertheless, many species show exceptions to this rule. Contrary to a logical expectation, adaptation to high alcoholic resources, which has been a recurrent evolutionary event, was not mediated by a more efficient use of ethanol, that is, an increase of the E/I ratio. This ratio seems to be quite variable according to the phylogeny and is especially low in the subgenus Sophophora as well as in Zaprionus. Alcohol tolerance clearly is related to the larval habitat of the species and shows that adaptation to alcoholic resources has been a major evolutionary challenge in drosophilids. This adaptation is not related to phylogeny, having occurred independently several times during the evolution of the group. Finally, it should be borne in mind that, besides metabolization and detoxification, other physiological processes such as nervous-system tolerance or ethanol excretion may be involved in ethanol tolerance, and such functions also should be investigated. Environmental ethanol, which is certainly a major ecological parameter for many drosophilids, has selected a diversity of physiological adaptations, all related to the Adh locus, but presumably much more complicated than was previously believed.     

Patterns of daily behaviour in two lizard species Lacerta agilis L. and Lacerta vivipara Jacquin

Summary During the period May to August 1978, individuals of both Lacerta agilis and Lacerta vivipara were observed in a specially constructed out-door vivarium. The daily patterns of behaviour of the two species were recorded along with surface and air temperatures. Daily activity of both species followed a similar basic pattern but was modified by variations in the local weather. Adults of L. vivipara reached a thermal preferendum more quickly and had a longer daily activity period than adults of L. agilis. The larger L. agilis can increase its period of activity by making use of different thermal conditions in a diverse habitat with varied vegetation structure under conditions of high solar radiation. The status of viable populations of L. agilis could be maintained or enhanced by management aimed at modifying the microenvironment of the lizard's habitat.     

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.