Multi-scale altitudinal patterns in species richness of land snail communities in south-eastern France

Aim Species richness is an important feature of communities that varies along elevational gradients. Different patterns of distribution have been described in the literature for various taxonomic groups. This study aims to distinguish between species density and species richness and to describe, for land snails in south‐eastern France, the altitudinal patterns of both at different spatial scales. Location The study was conducted on five calcareous mountains in south‐eastern France (Etoile, Sainte Baume, Sainte Victoire, Ventoux and Queyras). Methods Stratified sampling according to vegetation and altitude was undertaken on five mountains, forming a composite altitudinal gradient ranging from 100 to 3100 m. Visual searching and analysis of turf samples were undertaken to collect land snail species. Species density is defined as the number of species found within quadrats of 25 m². Species richness is defined as the number of species found within an elevation zone. Different methods involving accumulation curves are used to describe the patterns in species richness. Elevation zones of different sizes are studied. Results Eighty‐seven species of land snails were recovered from 209 samples analysed during this study. Land snail species density, which can vary between 29 and 1 species per 25 m², decreases logarithmically with increasing altitude along the full gradient. However, on each mountain separately, only a linear decrease is observable. The climatic altitudinal gradient can explain a large part of this pattern, but the great variability suggests that other factors, such as heterogeneity of ground cover, also exert an influence on species density. The altitudinal pattern of species richness varies depending on the spatial resolution of the study. At fine resolution (altitudinal zones of 100 m) land snail species richness forms a plateau at altitudes below 1000 m, before decreasing with increasing altitude. At coarse resolution (altitudinal zones of 500 and 1000 m) the relationship becomes linear. Main conclusions This study reveals that land snail species density and land snail species richness form two different altitudinal patterns. Species density exhibits strong variability between sites of comparable altitude. A large number of samples seem necessary to study altitudinal patterns of species density. Species density decreases logarithmically with increasing altitude. Above a critical altitudinal threshold, this decrease lessens below the rate seen in the first 1500 m. Different methods exist to scale‐up species density to species richness but these often produce different patterns. In this study, the use of accumulation curves has yielded a pattern of species richness showing a plateau at low altitude, whereas simple plotting of known altitudinal ranges from single mountains would have produced stronger mid‐altitudinal peaks. This study shows that not only factors such as temperatures and habitat heterogeneity, but also an ecotone effect, are responsible for the observed patterns.     

The development of diversity in the land snail fauna of the Madeiran archipelago

The diversity and distribution of the Madeiran land snail fauna is analysed in relation to the geological and environmental history of the archipelago. High levels of single-island endemism, the varying balance of families and genera between islands and the local differentiation of populations within islands all suggest that speciation has been predominantly a within island phenomenon. Restricted patterns of distribution, nevertheless, suggest that speciation is predominantly allopatric and that it is driven by repeated and sometimes devastating environmental changes. An equilibrium model of faunal diversity is rejected; the fauna is supersaturated at archipelago level, yet individual sites are not species rich, nor are all niches full. The consequences for the success and effects of invasions and introductions are discussed, and questions still to be answered raised, in particular about the timing of colonization events and their number, and about the great range of rates of radiation apparently achieved by different stocks.     

Calcareous forest seepages acting as biodiversity hotspots and refugia for woodland snail faunas

Land-snail species richness has repeatedly been found to increase with the increasing site calcium content and humidity. These two factors, reported as the main drivers of land-snail assemblage diversity, are also among the main habitat characteristics of calcareous seepages. Here we explore local species richness and compositional variation of forest spring-fed patches (i.e. seepages), to test the hypothesis that these habitats might act as biodiversity hotspots and refugia of regional snail faunas. In contrast to treeless spring fens, only little is known about land snail faunas inhabiting forest seepages. Studying 25 isolated calcareous forest seepages, evenly distributed across the White Carpathians Protected Landscape Area (SE Czech Republic), we found that these sites, albeit spatially very limited, can harbour up to 66% of the shelled land-snail species known to occur in this well-explored protected area (in total 83 species). By comparing land snail assemblages of the studied seepages with those occurring in the woodland surroundings of each site as well as those previously sampled in 28 preserved forest sites within the study area, we found the seepages to be among the most species rich sites. Although the numbers of species did not statistically differ among these three systems, we found highly significant differences in species composition. Seepage faunas were composed of many species significantly associated with spring sites, in contrast to the assemblages of both surrounding and preserved forest sites. Our results highly support the hypothesis that calcareous forest seepages might serve as refugia and biodiversity hotspots of regional land snail faunas. Protection of these unique habitats challenges both conservation plans and forest management guidelines as they might act as sources for the recolonization and restoration of forest snail assemblages particularly in areas impoverished by harvesting and clearcutting.     

Assessing the resilience of Mediterranean holm oaks to disturbances using selective thinning

Climate change will increase the frequency and the intensity of droughts in the Mediterranean region, likely reducing growth and increasing mortality of holm oaks (Quercus ilex), one of the most abundant species of Mediterranean forests. In water-limited systems such as those of the Mediterranean, carbon allocation patterns strongly favour belowground accumulation, especially in large subterranean structures called lignotubers. The resilience of these forests depends largely on the replenishment rate of these carbon reserves after disturbances. An experimental thinning, with two intensities (removal of 40% and 80% of basal area), was performed in 1992 in a holm oak forest at the Prades Experimental Complex of Catchments (NE Spain). In 2002, a second thinning was carried out in subplots within the former experimental 0.5 ha plots. Samples from the lignotubers of holm oak trees were analyzed for starch, and both mobile and immobile chemical components, in order to assess the resilience of holm oaks to repeated disturbances. Our results show that after 10 years, starch stocks in the lignotubers have only recovered to half their former values. Removing 40% of the basal area instead of 80% is suggested to be the better managing option for this kind of forests.     

History, island area and habitat availability determine land snail species richness of Aegean islands

Aim We examined the species-area relation of Aegean land snails, comparing different models to describe the relation. By examining those factors other than area that may also affect species richness, we tested whether the Aegean land snail fauna was more influenced by equilibrial migration and colonization processes, or rather is conservative and relictual. Location The Aegean archipelago (Greece). Methods Sixty-five islands were examined. Data were taken from own collections and from literature sources. Multiple regression analysis was used to test the null hypothesis of no relationship between species richness and island area, elevation, distance to the next larger island, and the presence and extent of calcareous substrate. Results The single most important factor determining land snail species number was area. While colonization-extinction dynamics have frequently been cited to explain this result, this conclusion was not tenable in this study as it was contradicted by species number not being related to the islands’ distances to neighbouring larger islands, after accounting for other factors affecting species number. We also found that habitat diversity affected species richness even after accounting for the effects of area: both increased elevation and greater extent of calcareous substrate on islands resulted in higher species number. This effect was most likely due to the fact that particular ecological conditions increased the probability that particular species could survive on an island. We compared the utility of the power and extreme-value function models of the species-area relation and found that both gave substantially the same results. However, fitting the power function model using nonlinear regression was of questionable utility. Main conclusions We conclude that the snail fauna of the Aegean is relictual, not equilibrial. The unusually high number of land snail species found on Crete is consistent with this conclusion. Crete is a currently united island which was separated into at least six smaller islands for 7–9 million years during the Neogene. Our results are consistent with the hypothesis that Crete still hosts a large number of endemic species of these paleoislands, resulting in a total number of species in excess of what would be expected based on area alone.     

The role of landscape history and persistent biogeographical patterns in shaping the responses of Mediterranean land snail communities to recent fire disturbances

Aims To assess the impact of various fire regimes over the past 30 years on land snail communities and to analyse the role of recent landscape history and the influence of biogeography in shaping the response patterns of gastropod communities following disturbances by fire. Location South-eastern France (Provence) and Mediterranean region. Methods Stratified sampling within 12 sites was undertaken with regard to fire regime (i.e. number of fires, fire intervals and age of the last fire) occurring over the past 30 years. The study was complemented by a historical analysis using aerial photographs, old maps of vegetation cover and an analysis of the biogeographical composition of malacofaunas. Data were investigated using Correspondence Analysis and Sørensen coefficient of similarity. Results When a disturbance regime (land use or fire disturbances) has been maintained over decades or centuries, land snail communities appear highly modified and tend to be composed of only Mediterranean and xerophilous species. However, low fire regimes, since the 1970s, do not seem to greatly affect the composition of gastropod communities. Indeed, shade-loving, mesophilous and European range species persist even after successive fires within some sites. In addition, the malacofaunas have a higher component of European range species with increasing distance from the Mediterranean sea. Main conclusions Analysis of the response patterns of gastropod communities to fire shows a response to numerous different factors. The composition of current land snail communities is not only the result of (more or less) recent patterns of fire regimes but also of anthropogenic disturbances, of landscape changes over the last centuries and of subsequent structure of the pre-fire habitat, as well as of the influence of a biogeographical gradient. However, the response patterns observed and the persistence of pre-fire communities imply the presence of cryptic refuges located within burned areas.     

Histochemical changes of carbohydrate and protein contents in the digestive gland cells of the land snail Monacha cartusiana following starvation

The present study was designed to investigate histochemically the detection of carbohydrate and protein in the normally feeding snails and after 15 and 30 days of starvation. Generally, abundant carbohydrate and protein materials were detected in the component cells of the digestive gland of normally feeding snails. The results of this investigation revealed a pronounced decline of carbohydrates in the digestive gland cells of Monacha cartusiana snails after starvation. Severe decline in carbohydrate content was observed especially after 30 days of starvation. Moreover, protein inclusions have exhibited a week stainability in the digestive gland cells of these snails as a consequence of starvation.     

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.     

Differential responses of zooplankton assemblages to environmental variation in temporary and permanent ponds

Permanent and temporary wetlands in Mediterranean shrublands represent unique repositories of biodiversity, which are increasingly threatened by human-induced habitat loss. The zooplankton of a permanent (P1) and a temporary pond (T35) in the Natural Reserve of Castelporziano, a rare residual stretch of such a shrubland in Central Italy (Latium), was investigated to: (1) expand and deepen knowledge of these endangered freshwater habitats, which represent a crucial component of Mediterranean biodiversity; (2) identify environmental controls regulating the development of zooplankton communities of each environment; and (3) highlight differences in the adaptive responses of the zooplankton community in relation to the different ecological conditions experienced by permanent and temporary habitats. Despite summer desiccation in T35, the two ponds exhibited a relative homogeneity in hydrological and physico-chemical dynamics. Zooplankton assemblages contained 41 total taxa, of which 32 were found in P1 and 28 in T35. Out of the 41 taxa identified, 22 (>50%) were exclusively present in one of the two ponds. On a yearly basis, the community dynamics of P1 seemed to be conditioned by physical and chemical factors and by hydrological cycle characteristics, while the community of T35 responded to algal blooms, food competition and predator/prey equilibria rather than correlating to abiotic factors. The main differences amongst zooplankton assemblages were observed over short time scales and occurred both within and between seasons, highlighting the role of some structural taxa that dominated the average composition of the community throughout the year, and the importance of “quick-response” taxa in determining the short-term composition and structure variation of pond zooplankton. A year-round cyclic community succession peculiar to each pond is described.     

The determinants of land snail diversity along a tropical elevational gradient: insularity,geometry and niches

Aim We investigated the patterns of species richness in land snails and slugs along a tropical elevational gradient and whether these patterns correlate with area, elevation, geographic constraints, and productivity. We did so both at the scale at which land snail population processes take place and at the coarser scale of elevational zones. Location Mount Kinabalu (4096 m) and the adjacent Mount Tambuyukon (2588 m) in Kinabalu Park, Sabah, Malaysian Borneo. Methods We used an effort‐controlled sampling protocol to determine land snail and slug species richness in 142 plots of 0.04 ha at elevations ranging from 570 to 4096 m. Extents of elevational ranges were determined by interpolation, extended where appropriate at the lower end with data from lowlands outside the study area. We used regression analysis to study the relationships between species density and richness on the one hand and elevation and area on the other. This was done for point data as well as for data combined into 300‐m elevational intervals. Results Species density (based on the individual samples) showed a decline with elevation. Elevational range length profiles revealed that range lengths are reduced at greater elevations and that a Rapoport effect is absent. Diversity showed a mild mid‐domain effect on Kinabalu, but not on Tambuyukon. When the data were combined into 300‐m elevational intervals, richness correlated more strongly with elevation than with area. Ecomorphospace was seen to shrink with increasing elevation. Main conclusions The elevational species richness patterns show the combined effects of (1) reduced niche diversity at elevations with lower productivity and (2) historical events in which the upward migration of lowland species as well as the speciation of highland endemics took place.     

Morphological and ecological shifts in a land snail caused by the impact of an introduced predator

Introduced predators have become major threats to native animal species in oceanic islands. A number of studies have shown that alien predators have caused serious extinctions of island endemics. However, little attention has been paid to the evolutionary impacts of alien predators on native species. The present study shows that predation by black rats, Rattus rattus, has resulted in ecological and morphological changes in the land snail Mandarina anijimana from the island of Anijima in the Ogasawara archipelago. The frequency of empty predated shells has increased over the past 17–19 years in southern areas of the island. The shells of these snails were found to be significantly higher, smaller and darker in the survey in 2006 than in the survey in 1987–1989 performed in central and southern parts of Anijima, where predation by Rattus was serious. M. anijimana were formerly restricted to shallow broad-leaved litter, whereas they are currently found in deep palm litter, where predation pressure from Rattus may be lower. This suggests that increased predation pressure by Rattus has changed the habitat use of M. anijimana. The close association between shell morphology and habitat use of Mandarina species suggests that the habitat shift induced by the predation of Rattus has caused these changes in the shell morphology of M. anijimana over a period of 17–19 years.     

Geographic variation in thermal tolerance and strategies of heat shock protein expression in the land snail <Emphasis Type="Italic">Theba pisana</Emphasis> in relation to genetic structure

Land snails are exposed to conditions of high ambient temperature and low humidity, and their survival depends on a suite of morphological, behavioral, physiological, and molecular adaptations to the specific microhabitat. We tested in six populations of the land snail Theba pisana whether adaptations to different habitats affect their ability to cope with thermal stress and their strategies of heat shock protein (HSP) expression. Levels of Hsp70 and Hsp90 in the foot tissue were measured in field-collected snails and after acclimation to laboratory conditions. Snails were also exposed to various temperatures (32 up to 54 °C) for 2 h and HSP messenger RNA (mRNA) levels were measured in the foot tissue and survival was determined. To test whether the physiological and molecular data are related to genetic parameters, we analyzed T. pisana populations using partial sequences of nuclear and mitochondrial DNA ribosomal RNA genes. We show that populations collected from warmer habitats were more thermotolerant and had higher constitutive levels of Hsp70 isoforms in the foot tissue. Quantitative real-time polymerase chain reaction (PCR) analysis indicated that hsp70 and hsp90 mRNA levels increased significantly in response to thermal stress, although the increase in hsp70 mRNA was larger compared to hsp90 and its induction continued up to higher temperatures. Generally, warm-adapted populations had higher temperatures of maximal induction of hsp70 mRNA synthesis and higher upper thermal limits to HSP mRNA synthesis. Our study suggests that Hsp70 in the foot tissue of T. pisana snails may have important roles in determining stress resistance, while Hsp90 is more likely implicated in signal transduction processes that are activated by stress. In the phylogenetic analysis, T. pisana haplotypes were principally divided into two major clades largely corresponding to the physiological ability to withstand stress, thus pointing to genetically fixed tolerance.     

Vegetation dynamics in north-western Mediterranean regions: Instability of the Mediterranean bioclimate

Abstract

Pollen and plant macrofossil records from a selection of sites in Spain, France and Italy are used to explore the origin and the development of the Mediterranean vegetation. The role of evergreen and summergreen taxa is discussed. The expansion of sclerophyllous trees during the Pleistocene interglacials is well correlated with the orbital forcing. During the Holocene, the large number of data illustrates latitudinal and longitudinal differences in vegetation dynamics. Multidisciplinary studies identify short terms climatic fluctuations. The debate is still open between those who attribute an increase of sclerophyllous forests during the late Holocene to a trend toward aridity and those who consider this dynamic as human-induced.     

Effects of repeated burning on Mediterranean communities of the northeastern Iberian Peninsula

Question: In relation to a single fire, do repeated wildfires in Mediterranean type ecosystems (1) reduce plant species richness or diversity; (2) modify patterns of abundance or dominance of plant species or (3) alter plant composition? Location: Pinus halepensis dominated communities of Catalonia, northeastern Iberian Peninsula, western Mediterranean Basin. Methods: Regional, paired design with 14 study sites, each consisting of a once burnt area (1994) and a twice burnt area (1975–1993 and 1994). Ten years after the last fire, we recorded all vascular plant species present in nested plots and quantified their relative abundances on transects. We compared species richness, diversity, dominance and relative abundance and species‐area correlations between paired once and twice burnt areas and assessed their floristic composition similarity. Results: No statistically significant differences were found in species richness or diversity. Slopes of species‐area correlations were higher in once burnt areas. In twice burnt areas, dominance by one or two species was higher. P. halepensis showed lower relative abundance and nanophanerophytes showed higher relative abundance. No differences were found for resprouter, seeder or resprouter‐seeder species. Floristic composition similarity between paired areas tended to be higher in less productive sites. Conclusions: Fire recurrence had contrasting effects on species richness at different spatial scales. Repeated burning reduced the relative abundance of the dominant tree species, which resulted in a higher relative abundance of shrubs. It also promoted the dominance of herbs, particularly Brachypodium retusum. However, it did not change the relative abundance of regenerative groups. Paired areas were more similar as they were more Mediterranean in terms of climatic conditions.     

Effects of the exotic invader Musculista senhousia on benthic communities of two Mediterranean lagoons

The impact that the alien mussel Musculista senhousia has on benthic biodiversity and community structure was investigated in two Italian transitional environments, where the species was particularly abundant: the Sacca di Goro (Adriatic Sea), and the Rio Padrongiano Deltaic area (Sardinia, Tyrrhenian Sea). Sampling campaigns were conducted by sampling mussel beds from each of two different patch size classes (small: <1.0 m², and large: >10.0 m²), and the surrounding bare sediments. At both study sites, mussel density varied greatly with time within and between patches. Despite the huge difference in benthic species complement at Goro and Padrongiano (33 and 115 taxa, respectively), at both sites the presence of mussel patches had a gross positive effect on the benthic community. At Goro, many trophic guilds showed a positive relationship with mussel abundance. At Padrongiano, a continuous and directional process of patches colonization (both large and small) through time seemed evident, while at Goro the process was interrupted by high-summer mortalities. At both study sites, M. senhousia played a double role: (1) as secondary substratum, enhancing the environmental structural complexity, and (2) as densely aggregated, living organisms, thus giving rise to a number of possible interactions with other invertebrates. The outcome, however, was strictly dependent by the level of anthropogenic stress, in this case water oxygen deficiency, which was different at the two study sites. Guest editors: A. Razinkovas, Z. R. Gasiūnaitė, J. M. Zaldivar & P. Viaroli European Lagoons and their Watersheds: Function and Biodiversity     

Measures of resilience: the response of coastal sage scrub to fire

Measures of four components of resilience are developed and used to quantify the response of coastal sage scrub to fire in southern California: (1) elasticity (rate of recovery following disturbance), (2) amplitude (threshold of disturbance beyond which recovery to the original state no longer occurs), (3) malleability (extent of alteration of the new stable-state from the original) and (4) damping (extent and duration of oscillation in an ecosystem parameter following disturbance). Vegetation and soil properties measured before fire, and for the first 5–6 yr after fire on four coastal (Venturan association) and four inland (Riversidian association) sites of coastal sage were used to follow changes. In addition, results from a simulation model of post-fire succession in Venturan coastal sage scrub (the FINICS model of Malanson) were used to examine resilience behavior over a 200 yr period. Resilience behavior of coastal sage scrub is critically influenced by the presence of a competitive mix of inherently strongly and weakly resprouting species. Sites dominated by weak resprouters exhibit lower elasticity and less damping of year-to-year fluctuations in composition in the early post-fire years. Sites with a mixture of weak and strong resprouters have a lower threshold of disturbance (amplitude) before species extirpation occurs, a result intensified by a higher frequency of disturbance. Malleability is also greater in these systems under higher disturbance frequency.Nomenclature follows. P.A. Munz, 1974. A Flora of Southern California. Univ. California Press.     

Comparison of ecosystems in the Mediterranean Basin and Western Australia

A comparison between fire-prone ecosystems in the Mediterranean basin and Western Australia shows convergent adaptations in structure, life forms and flora evolution. Both areas have in common summer drought and winter rainfall. Reproductive biology is mainly correlated with fire ecology; the different length of human impact and history seem to be responsible for floristic divergency in both areas. The ecological significance of pollination by insects, birds and mammals and problems of the seed banks are discussed.