High resilience of Mediterranean land snail communities to wildfires

In the Mediterranean region, wildfires have devastating effects on animals with limited mobility. With their poor dispersal abilities, their habitats on vegetation and in litter, and their sensitivity to humidity and shade, we expected land snails to be an interesting model to assess short, medium and long-term impact of fires on fauna biodiversity and their resilience. Stratified sampling was carried out on 12 sampling sites in garrigues and forests of Provence (southeastern France), according to fire regime (number of fires, fire intervals and age of the last fire) over the past 30 years. Data were investigated using diversity indexes, Kruskal–Wallis test, dendrogram of affinities and Correspondence Analysis (CA). We found, however, that Mediterranean land snail communities are particularly resilient to fires. Although abundance is drastically reduced in the short-term, species richness and community diversity are preserved provided that the time lapse between two successive fires is longer than the time required for recovery (i.e. around 5 years). This high community resilience in the short-term may be partly due to ecological and ethological aptitudes of land snails. However, these astonishing results, which have implications for conservation biology, are mainly due to the presence, within burned areas, of cryptic refuges that allow initial land snail survival, malacofauna persistence after successive fires and consistent biogeographical patterns in the long-term.     

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.     

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.     

The diet of the Madeiran lizard Lacerta dugesii

The gut contents of about 1700 lizards from the Madeiran Archipelago and die Salvage Islands were examined. The diet, which shows habitat and geographical variation, consists of a variety of invertebrates and a good amount of plant matter. The degree of herbivory is correlated widi lizard sex and size and widi tail autotomy. The amount of food and specialization in the diet were also correlated. Comparisons are made between die sex-size classes widiin die populations.     

Species richness in Madeiran land snails, and its causes

Aim  To unravel the reasons why a group of islands of 800 km² area should support over 200 extant land snail species.
Location  The temperate Atlantic Madeiran archipelago.
Methods  Distributional surveys and model testing.
Results  Repeated volcanic events have changed the topography of the islands and created periodic further isolates. There has been climatic fluctuation and varying sea level. Individual locations are not species-rich, but there is replacement from one subregion to another. Evidence of competitive interactions is lacking.
Main conclusions  High diversity results from the fortuitous coincidence of rates of geological and climatic change and isolation on the one hand, and migration and genetic divergence on the other. Hubbell's neutral model of biodiversity explains species richness if a factor is added describing structural instability or periodicity, here called the geodetic rate. This geodetic, biological interaction can explain why some archipelagos are species-rich.     

Effects of habitat history and extinction selectivity on species-richness patterns of an island land snail fauna

Aim Local‐scale diversity patterns are not necessarily regulated by contemporary processes, but may be the result of historical events such as habitat changes and selective extinctions that occurred in the past. We test this hypothesis by examining species‐richness patterns of the land snail fauna on an oceanic island where forest was once destroyed but subsequently recovered. Location Hahajima Island of the Ogasawara Islands in the western Pacific. Methods Species richness of land snails was examined in 217 0.25 × 0.25 km squares during 1990–91 and 2005–07. Associations of species richness with elevation, current habitat quality (proportion of habitat composed of indigenous trees and uncultivated areas), number of alien snail species, and proportion of forest loss before 1945 in each area were examined using a randomization test and simultaneous autoregressive (SAR) models. Extinctions in each area and on the entire island were detected by comparing 2005–07 records with 1990–91 records and previously published records from surveys in 1987–91 and 1901–07. The association of species extinction with snail ecotype and the above environmental factors was examined using a spatial generalized linear mixed model (GLMM). Results The level of habitat loss before 1945 explained the greatest proportion of variation in the geographical patterns of species richness. Current species richness was positively correlated with elevation in the arboreal species, whereas it was negatively correlated with elevation in the ground‐dwelling species. However, no or a positive correlation was found between elevation and richness of the ground‐dwelling species in 1987–91. The change of the association with elevation in the ground‐dwelling species was caused by greater recent extinction at higher elevation, possibly as a result of predation by malacophagous flatworms. In contrast, very minor extinction levels have occurred in arboreal species since 1987–91, and their original patterns have remained unaltered, mainly because flatworms do not climb trees. Main conclusions The species‐richness patterns of the land snails on Hahajima Island are mosaics shaped by extinction resulting from habitat loss more than 60 years ago, recent selective extinction, and original faunal patterns. The effects of habitat destruction have remained long after habitat recovery. Different factors have operated during different periods and at different time‐scales. These findings suggest that historical processes should be taken into account when considering local‐scale diversity patterns.     

Does vicariance shape biotas? Biogeographical tests of the vicariance model in the north‐west European land snail fauna

Aim To investigate the importance of vicariance in shaping the north‐west European land snail fauna. Location North‐west Europe. Methods We tested whether there is a non‐random congruence, i.e. a clustering of species ranges, using a Monte Carlo procedure with a null model that generates range data sets such that their range size distribution, the species richness distribution of the geographic cells and the spatial autocorrelation of the occurrences of a taxon approach the parameters in the real data set. Biotic elements, groups of species with similar ranges, were delimited with Model based Gaussian clustering. The prediction that closely related species belong to different biotic elements, has been tested with a chi‐square test. Results The distribution areas of the north‐west European land snail species are significantly clustered as predicted by the vicariance model. One widespread and seven regional biotic elements were identified. Contrary to the predictions of the vicariance model, closely related north‐west European land snail species belong significantly more often to the same biotic element than should be expected by chance. Main conclusions The clustering of closely related north‐west European land snail species within the same biotic element indicates that speciation modes other than vicariance were frequent or that the imprint of vicariance on the ranges was obscured by extensive post‐speciational dispersal. Extensive dispersal may have been caused by Pleistocene climatic fluctuations. The core areas of the regionally restricted biotic elements might indicate the positions of glacial refugia of land snails.     

The hypogean invertebrate fauna of Georgia (Caucasus)

Information about the cave invertebrates of Georgia, Caucasus, is summarised, resulting in 43 troglo- and 43 stygobiont taxa reported from 64 caves. Species distribution analyses were conducted for 61 caves harbouring 58 invertebrate taxa, with the majority of caves (39) located in Apkhazeti (north-western Georgia). In 22 caves from central-west Georgia (Samegrelo, Imereti and Racha-Lechkhumi regions of west Georgia) 31 taxa are reported. Composition of cave fauna differed strongly between the caves in Apkhazeti and the central-west of Georgia. Only two taxa of the total 86 were shared, resulting in negligible similarity (Sørensen-Dice coefficient S_s=4.8%). Rarefaction indicated an increase in number of species with additional sampling could increase species richness from 58 to 76 for caves in Apkhazeti and from 31 to 69 for caves in central-west Georgia. These findings suggest that the low invertebrate species richness observed in caves of western Georgia is the result of insufficient sampling. A pairwise approach to analysing species co-occurrence showed ten positive spatial associations in 7 out of 86 cave species, all from Kveda Shakurani and Tsebelda caves. The species co-occurring in the same microhabitat require further study to understand their relationships.     

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.     

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.     

Survey of the land snail fauna (Gastropoda: Pulmonata) of Mount Kaputar National Park in northern inland New South Wales,Australia, including a description of the listing of Australia's first legally recognised endangered land snail community

Mount Kaputar is an isolated high-elevation mesic refuge on the dry inland western slopes of eastern Australia's Great Dividing Range. A field survey in Mount Kaputar National Park in 2007–2014, and collation of existing information, identified a native land snail fauna of 24 species (seven families), including nine endemic taxa and another two occurring as disjunct populations. Additional species probably remain to be identified. Two introduced species of agriolimacid slugs were also recorded. Information from this study was the basis of the successful listing of an endangered land snail community at Mount Kaputar under state threatened species legislation in 2013; the first legal listing of an endangered land snail community in Australia. The recognition of endangered communities is a useful intermediate scale between the extremes of the single-species and broad-landscape conservation approaches, and has significant potential for multi-species conservation initiatives for terrestrial gastropods. Anthropogenic climate change is the most significant threat to this community, with a 100 m rise in environmental envelope likely to reduce the available area of habitat by over 50%. This community, and, in particular, a bright pink endemic athoracophorid slug, is contributing to raising public awareness of the threat to global biodiversity from anthropogenic climate change.     

Habitat, isolation and the evolution of Madeiran Iandsnails

The Madeiran archipelago consists of Madeira itself, Porto Santo and the Deserta islands. On Madeira, the forest arid the coastal floral associations are so different that their faunas are effectively isolated and have undergone largely independent development. There are different faunal associations on the eastern peninsula and in the SE coastal region, which may have been separated from each other in the past. On Porto Santo, western and eastern hills have different faunas. Most observations on the fauna are compatible with evolution by allopatric speciation, consequent upon isolation on different islands or mountains, as opposed to parapatric or sympatric processes following disruptive selection. Some cases where the taxonomy is difficult to unravel may, however, indicate parapatric speciation; examples belong to the genera Discula and Heterostoma (Helicidae) and Amphorella (Ferussaciidae). Most evidence relating to species composition in communities is compatible with a balance of random immigration and extinction, rather than selective interaction, allowing clusters of similar sympatric species to accumulate. However, this impression may indicate that test procedures are insufficiently sensitive to detect interactions, and detailed ecological studies are required. Questions about speciation and distribution would be clarified if dates of divergence were established.     

Delayed spermatophore removal in the land snail Euhadra peliomphala

Although many studies have demonstrated that variation in male genital morphology can influence male fertilization success, it remains unclear in many cases which mechanism(s) of sexual selection and sexual conflict produce the morphological divergence of genitalia. Distinguishing between such mechanisms requires, at the very least, information on the process by which genital morphology influences fertilization success. The length of the spermatophore‐producing organ (SPO) of the stylommatophoran land snails is a case in point. The length of the spermatophore may have an important influence on the transfer of the spermatophore to the recipient's genital tract. To test this hypothesis, we investigated in Euhadra peliomphala (Pfeiffer, 1850) the relationship between the length of the SPO in sperm donors and the position of their spermatophore in sperm recipients at 72, 84, and 96 h after mating. The results showed that longer spermatophores required longer times to reach a gametolytic organ and to begin to be digested. These findings suggest that this delay in spermatophore digestion explains the mechanism for the high fertilization success shown by snails with a long SPO. © 2013 The Linnean Society of London     

Difference in habitat selection by different morphs of the land snail Arianta arbustorum

The behaviour of morphs of Arianta arbustorum has been investigated using a new technique involving a paint which fades in daylight. The technique was used to study the response of different morphs to exposure to sunlight at a homogeneous woodland habitat and a heterogeneous open habitat. The yellow morph showed a significantly higher rate of fading than the brown morph. A mark, release and recapture programme was used to analyse the association between frequencies of different morphs recaptured and not recaptured. In both habitats, there was a general tendency for the frequency of recapture to increase with increasing age in both morphs. Habitat selection resulting from different behaviour of different morphs of A. arbustorum is discussed. Morph frequencies can be related to habitat, the brown morphs of A. arbustorum are significantly less frequent in the open habitat. The behavioural responses of Cepaea hortensis , present with A. arbustorum in two other populations, to the exposure of sunshine were also studied. Climatic selection by solar energy associated with different morphs, species and habitat are also discussed.     

The linyphiid fauna of eastern Africa (Araneae: Linyphiidae)—distribution patterns, diversity and endemism

Distributions, endemism and diversity among East African linyphiids are analysed and discussed in relation to other forest organisms and the environmental history of eastern African. A total of 231 species are reported from eastern Africa, of which 14 are confined to the Afroalpine region and 114 species to moist forests. Only 12 of the latter are widely distributed. The rest are only known from one or two localities. Information on habitats and distributions of all species is tabulated. Few species are shared between East African mountains and there are no detectable gradients of species diversity between mountains. There is, however, a gradient of decreasing species diversity from high latitudes to the Equator. Vicariance patterns are demonstrated for Elgonia, Ophrynia and Callitrichia in the Eastern Arc Mountains, Tanzania—areas that also hold the highest degree of endemism (> 80% on individual mountains) among linyphiids. The many endemic species on nearby mountains suggest that intermontane dispersal (ballooning) is rare or non-existent. There is no evidence for a distinction between highland and lowland linyphiid faunas, but altitudinal segregation of single species is demonstrated. The question of the reality of highland and lowland faunas cannot be solved by studying the altitudinal distribution of single species. Phylogenetic relationships must be taken into consideration to determine where sister-groups/species are located (lowland or highland).     

Spatial ecology, habitat and speciation in the Porto Santan land snail genus Heterostoma

The Madeiran land snail genus Heterostoma expresses two forms of genital anatomy. One form (hemiphallic) lacks the distal male organs while the other (euphallic) has fully developed hermaphroditic genitalia. Recent evidence suggests this genital variation characterizes two sibling or incipient species. The spatial ecology and habitat associations of these taxa are described in the light of recent ideas on the role of ecology in speciation. Hemiphallic snails are found at a lower density and show a tendency to be more separated from each other compared to euphallic snails. Both taxa are more likely to be closer to other members of the same taxon. The genus as a whole is not found in the presence of sand or pale rock. However, when only areas actually containing snails are examined, hemiphallics show a positive association with sand and rock while euphallics continue to show a negative association. The differences in population density and spacing may suggest an adaptive explanation for reduction of male organs in hemiphallics since this taxon would be expected to have significantly fewer mating encounters. The observed differences in habitat association may suggest a mechanism for sympatric or parapatric divergence in keeping with current models of habitat-driven adaptive speciation. Both of these factors (selection on the mating system in a marginal environment and small scale separation based on habitat) may have been significant in the divergence of hemiphallic and euphallic taxa.     

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.     

Spire index and preferred surface orientation in some land snails

Fourteen species of land snails have been tested for their preference for surfaces at 0, 90 or 180 degrees under laboratory conditions. They range from high-spired (height/breadth = 4.1) to discoidal forms (height/breadth = 0.4). There is a positive association of spire height with tendency to adopt the 90 degree surface. Species of intermediate (globular) shape show less specificity for a particular surface than high- or low-spired species. The exception is Helix aspersa , which behaves more like one of the high-spired species than like one of its similarly shaped relatives. The differences in preference will help to reduce interaction between co-existing species in the field.     

铜尾矿废弃地土壤动物多样性特征

土壤动物的恢复、群落演替及其多样性对于铜尾矿废弃地的生态重建具有重要意义.为了解铜尾矿废弃地土壤动物群落多样性特征,在铜陵市铜尾矿区设置4个样地23采样点,共捕获土壤动物4622只个体,隶属5门10纲18目29类.结果表明,铜尾矿自然废弃地与其对照组土壤动物多样性差异明显,自然废弃地土壤动物丰富度指数d、大型土壤动物密度、中小型土壤动物密度、多样性指数H'和D·G指数均小于其对照组且存在显著差异.而复垦废弃地中小型土壤动物密度和D·G指数小于其对照组且存在显著差异,多样性指数H'大于其对照组且差异极显著,其他土壤动物群落指标与其对照组均无显著差异.相似系数q表明,外围林地间土壤动物群落相似性最大,其次是复垦废弃地与外围林地间,自然废弃地与其它3个样地间相似性最小.从垂直分层来看,复垦废弃地土壤动物表聚性强于自然废弃地,尾矿废弃地表聚性强于外围林地.灰色关联分析表明,铜尾矿区土壤理化性质与土壤动物群落结构指标关系密切,其中最重要的因素是土壤含水量和总钾含量,重金属全镉含量也不容忽视,植被因素和土壤全铜含量对土壤动物的影响相对较小.由此可见,土壤含水量、土壤基质的优劣、土壤有机质与营养元素的含量等因素限制了铜尾矿废弃地土壤动物的恢复与重建,而这些因素的改善主要归功于尾矿废弃地的土地复垦和作物种植.所以,尾矿废弃地的复垦与利用有利于土壤动物的恢复与重建.     

High diversity and local endemism in Aotearoa New Zealand's groundwater crustacean fauna

We used DNA barcoding to assess the diversity and distribution of New Zealand''s groundwater amphipods and isopods (Crustacea) and to determine whether biodiversity and endemism within tectonically active New Zealand are similar to those of more tectonically stable continents. Sixty‐five wells were sampled in seven aquifers across four regions within the North and South islands of New Zealand, and resident invertebrates were morphologically identified and then assessed using sequencing of the mitochondrial DNA cytochrome c oxidase subunit one (COI) gene. Invertebrates were found in 54 wells. Of the 228 individual amphipods and isopods found in 36 of the wells, 154 individuals were successfully sequenced for COI (68% success rate) from 25 wells, with at least one well in each aquifer containing sequenced individuals. Of the 45 putative species identified using Barcode Index Numbers (BINs), 30 BINs (78% of all taxa and 83% of amphipods) were previously unrecorded. Substantial morphologically cryptic, species‐level diversity was revealed, particularly within the amphipod Family Paraleptamphopidae. Similarly, one isopod taxon morphologically identified as Cruregens fontanus was assigned to five well‐separated BINs based on COI sequences. Endemism appeared high, with all taxa regionally endemic; 87% of species were restricted to one aquifer and more than 50% restricted to one well. Non‐saturated species accumulation curves indicated that, while additional sampling may increase the range of some currently identified taxa, additional range‐restricted taxa are also likely to be discovered. Patterns of diversity and short‐range endemism were similar to those found elsewhere, including locations which are more tectonically stable. The predominance of local endemism within New Zealand''s groundwater fauna suggests that land‐use activities and groundwater extraction require careful evaluation to minimize threats to groundwater biodiversity.