Assessing the origin of Neotropical mountain dung beetle assemblages (Scarabaeidae: Scarabaeinae): the comparative influence of vertical and horizontal colonization

Aims The fauna of mountains and their surrounding regions are likely to be influenced principally by two biological processes: horizontal colonization along similar altitudinal levels by elements originating from lineages inhabiting higher latitudes; and vertical colonization by lineages from the same latitude, but at lower altitudes. We examine whether the expected patterns derived from the latter process can be observed in mountain dung beetle assemblages. Specifically, we study the variation in species composition and richness with altitude in five regions spanning elevation gradients, analysing whether the altitudinal rates of change in the number of species and genera differ, and whether beta‐diversity scores for adjacent sites in each altitudinal gradient are different for species and genera. Location Eastern Cordillera of the Colombian Andes. Methods Field work was carried out in 1997–99 at 27 sites in five regions with elevation gradients, with 10–32 pitfall traps placed in each site. For each altitudinal level the numbers of species and genera were analysed with respect to altitude, and the slope of the linear regression between these variables was calculated. The slope of the curve of the altitude against the cumulative number of species and genera was also calculated for each altitudinal gradient to describe the compositional change between adjacent sites (beta diversity). Species and generic slopes were compared using analysis of covariance. The turnover of species along each altitudinal gradient was measured using presence/absence data and Cody's beta‐diversity index between adjacent pairs of sites. A cluster analysis was used to detect faunistically homogeneous groups of localities. Results Species richness always decreased with altitude, although the slopes did not differ significantly from zero. The number of genera also decreased with increasing altitude, but generally at a significantly slower rate than for species. Variation in the species beta‐diversity scores between altitudinal levels did not follow a homogeneous pattern in the different regions. Two main altitudinal groups of sites with a boundary c. 1500–1750 m a.s.l. can be detected with respect to faunistic similarity. Low‐ and mid‐altitude sites are inhabited by all of the genera (19) and 80% of all species collected. Eight genera and 61 species (c. 60% of the total) are unable to inhabit high‐altitude sites, and only 20 species appear to be exclusive to these high‐altitude environments (> 2000 m a.s.l.). Main conclusions The dominant processes explaining dung beetle composition in the high north‐eastern Andean mountains are probably those of vertical colonization. The limited role of horizontal colonization processes, or colonization from northern or southern lineages, could be a consequence of the isolation and recent geological origin of these mountains.     

Nesting strategies affect altitudinal distribution and habitat use in Alpine dung beetle communities

1. Dung beetles are key contributors to a suite of ecosystem services. Understanding the factors that dictate their distributions is a necessary step towards preventing negative impacts of biodiversity loss. 2. Alpine dung beetle communities were analysed along altitudinal gradients to assess how different components of the community, defined in terms of nesting strategy [dung‐ovipositing Aphodiidae (DOAs), soil‐ovipositing Aphodiidae (SOAs) and two paracoprid (PAR) groups, Geotrupidae and Scarabaeidae] and parameters relevant to dung removal rates (species richness, total biomass and functional diversity), are distributed, and to identify to which environmental factors they respond. 3. Species richness declined with altitude. There was no significant variation in functional diversity or total biomass in relation to altitude. There were significant variations when considered by nesting group: DOA species richness and biomass decreased, SOA biomass increased, and Geotrupidae biomass showed a non‐linear trend, as altitude increased. 4. Functional diversity and total species richness were positively related to vegetation cover. DOA species richness was highest in forest and scrub; SOA species richness was highest in grassland and PAR species richness was lowest in rocky areas. 5. Dung beetle species show different trends in species richness and biomass depending on nesting strategy. Management to promote the dung beetle community should include maintenance of a mosaic of habitat types. Given the likely importance of species richness and biomass to ecosystem functioning, and the complimentary effect of different dung beetle groups, such a strategy may protect and enhance the ecosystem services that Alpine dung beetles provide.     

From forest to pasture: an evaluation of the influence of environment and biogeography on the structure of beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region

The objective of this study is to evaluate the effect of environmental (associated with the expansion of cattle ranching) and biogeographical factors on the diversity of dung beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region. One gradient is located in the Mexican Transition Zone, on the Cofre de Perote mountain, the other two are in the northern Andes (the Chiles Volcano and the Río Cusiana Basin). For the three gradients, the number of species and of individuals was similar in both forest and pasture, while species composition was different between habitats. On this mountain, species turnover in pastures was characterized by the addition of new species as altitude increased. In the northern Andes, species diversity was always greater in the forest than in the pasture, and species turnover between habitats was notably influenced by species loss with increasing altitude. As such the pasture fauna of the northern Andes was an impoverished derivative of the fauna present in the forests at the same altitude characterized by species of Neotropical affinity with a limited capacity for colonizing open, sunnier habitats. The opposite occurs in the areas used by cattle on the Cofre de Perote. This habitat has its own fauna, which is mainly comprised of Holarctic and Afrotropical species adapted to the prevailing environmental conditions of areas lacking arboreal vegetation. These results suggest that the impact on beetle communities caused by human activities can differ depending on the geographic position of each mountain and, particularly, the biogeographical history of the species assemblage that lives there.     

Space-for-time substitution reveals a hump-shaped distribution of dung beetles

Unraveling how climate change impacts the diversity and distribution patterns of organisms is a major concern in ecology, especially with climate-sensitive species, such as dung beetles. Often found in warmer weather conditions, beetles are used as bio-indicators of environmental conditions. By using an altitudinal gradient as a proxy for climate change (i.e., space-for-time substitution), we assessed how changes in climatic variables, such as temperature and precipitation, impact patterns of dung beetle diversity and distribution in the Peruvian Andes. We recorded dung beetle diversity using three different types of baits, feces, carrion, and fruits, distributed in 18 pitfall traps in five different altitudinal sites (from 900 to 2500 m, 400 m apart from each other) in the rainy and dry season. We found that (i) dung beetle richness and abundance were influenced by the climate gradient, (ii) seasonality influenced beetle richness, which was high in the wet season, but did not influence abundance, (iii) dung beetle richness and abundance fit to a hump-shaped distribution pattern along the altitudinal gradient, and (iv) species richness is the beta-diversity component that best describes the composition of dung beetle species along the altitudinal gradient. Our data show that the distribution and diversity of dung beetles are different at larger scales, with different patterns resulting from the response of species to both abiotic and biotic factors.     

Regional and local influence of grazing activity on the diversity of a semi-arid dung beetle community

This study analyses the effect of resource availability (i.e. sheep dung) on dung beetle communities in an arid region of Central Spain, both at regional and at local scales. A total of 18 sites within 600 km² were sampled for the regional analysis and 16 sites within the 30 km² of an Iberian municipality were sampled for the local analysis. Spatial and environmental characteristics of sampling sites were also compiled at both scales, including measures of grazing activity (livestock density at regional scale, and two counts of rabbit and sheep dung at local scale). At a regional scale, any environmental or spatial variable can help to explain the variation in abundance. However, species richness was related to summer precipitation and composition was related to elevation. At local scale, abundance is not significantly related to any of the environmental variables, but species richness was related to the local amount of sheep dung (27% of variance). The amount of dung in a 2‐km buffer around the site accounts for 27–32% of variance in abundance and 60–65% of variance in species richness. The presence of the flock with the highest sheep density explains 53% of abundance variability and 73% of species richness variance. A cluster analysis of localities identified two main groups, one characterized by a lower abundance and species richness that can be considered a nested subsample of the species‐rich group. The mean and maximum amount of sheep dung in the sites separated by less than 2 km are the only significant explanatory variables able to discriminate both groups. These results suggest that grazing intensity (and the associated increase in the amount of trophic resources) is a key factor in determining local variation in the diversity and composition of dung beetle assemblages. However, dung beetle assemblages are not spatially independent at the analysed resolution, and the amount of dung in the surroundings seems to be more important for locally collected species than the dung effectively found in the site. Although differences in the availability and quantity of trophic resources among nearby sites could be affecting the population dynamics and dispersion of dung beetles within a locality, sites with larger populations, and greater species numbers would not be able to exercise enough influence as to bring about a complete local faunistic homogenization.     

Does natural reforestation represent a potential threat to dung beetle diversity in the Alps?

Traditional agro-pastoral practices are in decline over much of the Alps, resulting in the complete elimination of livestock grazing in some areas. Natural reforestation following pastoral abandonment may represent a significant threat to alpine biodiversity, especially that associated with open habitats. This study presents the first assessment of the potential effects of natural reforestation on dung beetles by exploring the relationships between the beetle community (abundance, diversity, species turnover and assemblage structure) and the vegetation stages of ecological succession following pastoral abandonment. A hierarchical sampling design was used in the montane belt of the Sessera Valley (north-western Italian Alps). Dung beetles were sampled across 16 sampling sites set in four habitat types corresponding to four different successional stages (pasture, shrub, pioneer forest and beech forest) at two altitudinal levels. The two habitats at the extremes of the ecological succession, i.e. pasture and beech forest, had the greatest effect on the structure of local dung beetle assemblages. Overall, dung beetle abundance was greater in beech forest, whereas species richness, Shannon diversity and taxonomic diversity were significantly higher in pasture, hence suggesting this latter habitat can be considered as a key conservation habitat. Forests and pastures shared a lower number of species than the other pairs of habitats (i.e. species turnover between these two habitats was the highest). The two intermediate seral stages, i.e. shrub and pioneer forest, showed low dung beetle abundance and diversity values. Local dung beetle assemblages were also dependent on season and altitude; early-arriving species were typical of pastures of high elevation, whereas late-arriving species were typical of beech forests. It is likely that grazing in the Alps will continue to decrease in the future leading to replacement of open habitats by forest. This study suggests therefore that, at least in the montane belt, reforestation may have potentially profound and negative effects on dung beetle diversity. Maintaining traditional pastoral activities appears to be the most promising approach to preserve open habitats and adjacent beech forests, resulting in the conservation of species of both habitats.     

Species richness and trait composition of butterfly assemblages change along an altitudinal gradient

Species richness patterns along altitudinal gradients are well-documented ecological phenomena, yet very little data are available on how environmental filtering processes influence the composition and traits of butterfly assemblages at high altitudes. We have studied the diversity patterns of butterfly species at 34 sites along an altitudinal gradient ranging from 600 to 2,000 m a.s.l. in the National Park Berchtesgaden (Germany) and analysed traits of butterfly assemblages associated with dispersal capacity, reproductive strategies and developmental time from lowlands to highlands, including phylogenetic analyses. We found a linear decline in butterfly species richness along the altitudinal gradient, but the phylogenetic relatedness of the butterfly assemblages did not increase with altitude. Compared to butterfly assemblages at lower altitudes, those at higher altitudes were composed of species with larger wings (on average 9 %) which laid an average of 68 % more eggs. In contrast, egg maturation time in butterfly assemblages decreased by about 22 % along the altitudinal gradient. Further, butterfly assemblages at higher altitudes were increasingly dominated by less widespread species. Based on our abundance data, but not on data in the literature, population density increased with altitude, suggesting a reversed density–distribution relationship, with higher population densities of habitat specialists in harsh environments. In conclusion, our data provide evidence for significant shifts in the composition of butterfly assemblages and for the dominance of different traits along the altitudinal gradient. In our study, these changes were mainly driven by environmental factors, whereas phylogenetic filtering played a minor role along the studied altitudinal range.     

Pattern of ant diversity in Korea: An empirical test of Rapoport's altitudinal rule

Two diversity patterns (hump-shaped and monotonic decrease) frequently occur along altitude or latitude gradients. We examined whether patterns of ant species richness along altitudes in South Korea can be described by these patterns and whether ranges of ant species follow Rapoport's altitudinal rule. Ants on 12 high mountains (> 1100 m) throughout South Korea (from 33° N to 38° N) were surveyed using pitfall traps at intervals of 200–300 m altitude. The temperatures at the sampling sites were determined from digital climate maps. Ant species richness decreased monotonically along the altitudinal gradient and increased along the temperature gradient. However, species richness of cold-adapted species (highland species) showed a hump-shaped pattern along altitude and temperature gradients. The altitude and temperature ranges of ant species followed Rapoport's rule. Sampling site temperature ranges were significantly correlated with coldness. Therefore, Rapoport's rule can be explained by high cold-tolerance of species inhabiting high altitudes or latitudes.     

Spatial and temporal variation in species composition of dung beetle assemblages in southern Ireland

Abstract. 1. This study attempts to identify the main community characteristics that contribute to variability in dung beetle assemblage composition and structure across a range of spatial and temporal scales.
2. Dung beetle assemblages ( Aphodius , Sphaeridium, and Geotrupes species) were monitored by dung-baited pitfall trapping at 10-day intervals during the seasonally active period at eleven sites in southern Ireland. Three of the sites were monitored over at least 2 years between 1991 and 1996.
3. Although the species composition of the above taxonomic groups was comparable among sites and years, relative abundances of component species varied considerably. Detrended correspondence analysis ordinations indicated a similar level of variability in dung beetle assemblage structure among years, and among sites ≈1–180 km apart.
4. Processes that may contribute to spatio-temporal variability in dung beetle assemblages are discussed, and strategies for future research are suggested.     

Does an aridity and trophic resource gradient drive patterns of dung beetle food selection across the Botswana Kalahari?

1. Aridity gradients are paralleled by both reductions in resources and decreased species richness of animals. Across the aridity gradient of the Botswana Kalahari, a reduction in mammal species richness leads to reduced density and diversity of dung types, accompanied by reduced dung beetle species richness. We investigated whether this gradient also drives changes in dung beetle food type association and specialisation owing to a loss of some dung types to the arid southwest. 2. Dung beetles were sampled from three study sites in each of the six study areas using 2 × 10 grids of pitfall traps baited with dung (pig, elephant, cattle, and sheep) or carrion (chicken livers). 3. Canonical correspondence analysis (CCA) showed that distributions of dung beetle species between bait types deviated significantly from random associations. 4. Central Kalahari assemblages were more specialist than those at the mesic and arid extremes of the gradient. 5. Patterns of selection and specialisation to bait types differed between mesic northeast and arid southwest study areas. There were specialist faunas on carrion and more generalist faunas on ruminant herbivore dung (cattle and sheep) in each region. However, specialist species associated with elephant dung in the northeast were replaced by a more generalist fauna in the southwest with an opposite trend on pig dung. 6. Reduced species richness and high species turnover from the mesic northeast to the arid southwest is paralleled by a shift in patterns of food association that may reflect changes in the diversity of food types, particularly the absence of elephant dung from the southwest.     

Effects of dung-pad conditions and density on coprophagous beetle assemblages in a Mediterranean rangeland

Dung beetles highly depend on the ephemeral microhabitat dung which is food resource and larval habitat at the same time. Environmental conditions surrounding a dung pad, such as vegetation structure, have an impact on dung beetle assemblages. We investigated the influence of dung conditions and surrounding habitat characteristics on Mediterranean dung beetle assemblages in a permanently grazed landscape in northern Sardinia. We sampled the dung beetle assemblages of donkey and horse dung in three different vegetation types and assessed species richness and abundance of dung beetles. Species richness was determined by dung and surrounding habitat conditions, whereas abundance was solely affected by dung conditions. However, species richness and abundance decreased with increasing dung density. The effect of dung density on species richness varied depending on vegetation type, with dry grassland exhibiting the highest number of dung beetles species at high dung density. Species composition in dung pads was influenced by abiotic factors with dwellers being negatively affected by increasing dung-pad temperature. Our results underline the importance of diverse vegetation, particularly with respect to the complexity of vegetation which interrelates with the microclimate. Furthermore, our findings illustrate the negative effect of high dung densities on dung beetle assemblages, suggesting that the degree of the intensity of use by grazing animals is important when considering measures for the conservation of dung beetles.     

Asymmetrical competition between Neotropical dung beetles and its consequences for assemblage structure

Abstract.  1. This study combines the results of laboratory experiments using representative assemblage components and pitfall trapping over a large geographical area to examine the hypothesis that ongoing interspecific competition structures Neotropical dung beetle assemblages.
2. From Guatemala to Panama assemblages of large to medium-sized, fast-tunnelling dung beetles include a single large, nocturnal dichotomiine species, Dichotomius annae (Kohlmann & Solís, 1997). In competition experiments, this species out-competed the medium-sized coprine species, Copris lugubris Boheman and Phanaeus demon Laporte-Castelnau, for dung and nesting space, in spite of earlier colonisation by the diurnal species, P. demon .
3. Differences in the abundance of D. annae at Central American sites did not affect total fast-tunnelling dung beetle assemblage richness over the rainy season. However, D. annae rank order was directly related to the probability of interspecific encounters (Hurlbert's Δ ₁) among species. These trends were also observed when species lists from published and unpublished studies of other large allopatric dichotomiine species, with a more northerly distribution, were included in the analyses.
4. The results obtained suggest that where large dichotomiine species are abundant, their efficient pre-emption of a considerable proportion of available resources drives all, or most, other fast-tunnelling species to a lower population density, thereby decreasing assemblage diversity.     

Species-richness patterns of vascular plants along seven altitudinal transects in Norway

Altitudinal richness patterns were investigated along altitudinal gradients located in northern Norway (two transects) and along a west–east gradient in southern Norway (five transects). The transects were sampled for vascular plant species richness using a uniform sampling method. Each transect consisted of 38–48 5×5 m sample plots regularly spaced from sea level or valley bottom to a local mountain top. In five transects species richness peaked at mid-altitudes, whereas in the two northern transects species richness decreased with altitude. The observations were qualitatively evaluated in relation to the influence of the area of the species pool, hard boundaries, temperature and precipitation, and mass effect. The observed patterns cannot be fully accounted for by any of these factors. However, the altitude of the peak in species richness was above the forest-limit for all the humped relationships, which may suggest that species richness above the forest-limit might be enhanced by a mass effect from forest taxa. The two monotonic relationships found in the north may be caused by the relatively low number of alpine species at these sites. The monotonic pattern may result from a decrease in "forest species" towards the mountain tops.     

Altitudinal distribution of alien plant species in the Swiss Alps

In summer 2003 we recorded the presence and abundance of alien plant species at 232 sites (107 railway stations and 125 road sites) along mountain passes in the Swiss Alps. The altitudinal distribution of species was related to the current abundance of the species in Switzerland and time since introduction. A total of 155 alien taxa were recorded. Numbers of species per site declined exponentially with altitude, and only a few species were found in the alpine zone (>2000 m). In contrast, species richness among comparable native taxa appeared to be nearly independent of altitude over the range investigated. Maximum altitude reached by alien species was related positively to both total area occupied in Switzerland and to time since introduction. A comparison of the results with earlier records suggests that many species, particularly those previously restricted to low or intermediate altitudes, have advanced their altitudinal limits over the past few decades. Various hypotheses are presented to explain the declining abundance of alien species with altitude: low-altitude filter effects, low propagule pressure, and genetic swamping of peripheral populations at higher altitudes. However, at present we do not have sufficient evidence to determine the relative importance of these effects. We conclude that invasion into mountain areas such as the Swiss Alps tends to proceed rather slowly, though the process may be accelerated by climatic warming. For this reason, further research to investigate the processes determining how plants invade mountain areas is urgently needed. And more generally, investigations into the distribution of alien species along strong altitudinal gradients may provide valuable insights into the mechanisms driving the spread of alien organisms.     

Altitudinal effects on habitat selection of dung beetles (Scarabaeoidea: Aphodiidae) in the northern Iberian peninsula

The effects or vegetation cover on the abundance and species richness of dung beetles (Aphodiidae) were examined in three altitudinal zones: lowland (520–600 m), mid-altitude (950–963 m) and highland (1230–1275 m). Pitfall traps baited with fresh cattle dung were placed in three vegetation types at each altitudinal zone: forest, heathland and pasture. Species richness was lower in forests than in the other vegetation types. The total abundance of beetles also varied with vegetation type, but the effect was dependent on altitude. Number of individuals was higher in pasture at highlands and in heathland and forest at mid-altitude and lowlands. An altitudinal dependence of distribution of individuals between vegetation types was also found for the more abundant species in the study area. These results could be explained if the local distribution of individuals between vegetation types was determined by microcli-matic factors, mainly temperature. Differences in altitude between sites are associated with temperature changes, determining the habitat selection of species at each altitude.     

Plant species and growth form richness along altitudinal gradients in the southwest Ethiopian highlands

Questions: Do growth forms and vascular plant richness follow similar patterns along an altitudinal gradient? What are the driving mechanisms that structure richness patterns at the landscape scale? Location: Southwest Ethiopian highlands. Methods: Floristic and environmental data were collected from 74 plots, each covering 400 m². The plots were distributed along altitudinal gradients. Boosted regression trees were used to derive the patterns of richness distribution along altitudinal gradients. Results: Total vascular plant richness did not show any strong response to altitude. Contrasting patterns of richness were observed for several growth forms. Woody, graminoid and climber species richness showed a unimodal structure. However, each of these morphological groups had a peak of richness at different altitudes: graminoid species attained maximum importance at a lower elevations, followed by climbers and finally woody species at higher elevations. Fern species richness increased monotonically towards higher altitudes, but herbaceous richness had a dented structure at mid‐altitudes. Soil sand fraction, silt, slope and organic matter were found to contribute a considerable amount of the predicted variance of richness for total vascular plants and growth forms. Main Conclusions: Hump‐shaped species richness patterns were observed for several growth forms. A mid‐altitudinal richness peak was the result of a combination of climate‐related water–energy dynamics, species–area relationships and local environmental factors, which have direct effects on plant physiological performance. However, altitude represents the composite gradient of several environmental variables that were interrelated. Thus, considering multiple gradients would provide a better picture of richness and the potential mechanisms responsible for the distribution of biodiversity in high‐mountain regions of the tropics.     

Nematode distributions as spatial null models for macroinvertebrate species richness across environmental gradients: A case from mountain lakes

Nematode species are widely tolerant of environmental conditions and disperse passively. Therefore, the species richness distribution in this group might largely depend on the topological distribution of the habitats and main aerial and aquatic dispersal pathways connecting them. If so, the nematode species richness distributions may serve as null models for evaluating that of other groups more affected by environmental gradients. We investigated this hypothesis in lakes across an altitudinal gradient in the Pyrenees. We compared the altitudinal distribution, environmental tolerance, and species richness, of nematodes with that of three other invertebrate groups collected during the same sampling: oligochaetes, chironomids, and nonchironomid insects. We tested the altitudinal bias in distributions with t‐tests and the significance of narrow‐ranging altitudinal distributions with randomizations. We compared results between groups with Fisher's exact tests. We then explored the influence of environmental factors on species assemblages in all groups with redundancy analysis (RDA), using 28 environmental variables. And, finally, we analyzed species richness patterns across altitude with simple linear and quadratic regressions. Nematode species were rarely biased from random distributions (5% of species) in contrast with other groups (35%, 47%, and 50%, respectively). The altitudinal bias most often shifted toward low altitudes (85% of biased species). Nematodes showed a lower portion of narrow‐ranging species than any other group, and differed significantly from nonchironomid insects (10% and 43%, respectively). Environmental variables barely explained nematode assemblages (RDA adjusted R² = 0.02), in contrast with other groups (0.13, 0.19 and 0.24). Despite these substantial differences in the response to environmental factors, species richness across altitude was unimodal, peaking at mid elevations, in all groups. This similarity indicates that the spatial distribution of lakes across altitude is a primary driver of invertebrate richness. Provided that nematodes are ubiquitous, their distribution offers potential null models to investigate species richness across environmental gradients in other ecosystem types and biogeographic regions.     

From lowlands to highlands: searching for elevational patterns of species richness and distribution of scarab beetles in Costa Rica

Aim Understanding the heterogeneous distribution of species on mountains is an important aim in ecology. Altitudinal gradients have enormous potential for improving our knowledge of trends in biodiversity and conservation. In this study, we investigated the variation in scarab beetle diversity (Dynastinae, Rutelinae and Melolonthinae) along an elevational tropical forest gradient. Location The Atlantic slope of the Guanacaste mountain range in Costa Rica. Methods Ultraviolet light traps placed in six forests situated from 100 to 1510 m were used. Changes in species composition and richness among elevations were investigated. Differences in the altitudinal patterns using different groups of species were examined: the whole assemblage, each separate subfamily and two different trophic habits (phytophagous or saproxylic). The effects of temperature, humidity and elevation on scarab distribution were tested using canonical correspondence analyses. The relationship between the community similarity of the studied forests and the altitudinal distance among them was also analysed. Results Species composition and richness changed along the gradient. The peak in species richness varied depending on the species group considered and in all cases occurred 500 or 800 m. Forests at these altitudes were also the richest in exclusive species. Species composition turnover among elevations appeared with a clear separation between lowland and highland fauna. The latter was lower in richness but also had exclusive species. Temperature, humidity and altitude affected species distribution, with altitude being the most important factor for all the subfamilies studied. Main conclusions Our results showed that species distribution fits a hump‐shaped pattern. The peak of this pattern varied depending on the taxonomic group and mountain analysed, highlighting the importance of evolutionary processes as species distribution drivers. The fact that species richness peaked at elevations where human impact is currently important underlines the value of the development of conservation strategies for these areas.     

Species Richness of Nonvolant Small Mammals Along Elevational Gradients in Northwestern Argentina

The Nevados del Aconquija (5500 m) and Cumbres Calchaquíes (4600 m) are isolated mountain ranges that contain at least three physiognomic units in their eastern slopes: Neotropical rainforests, Andean grasslands and High Andean Steppes. Despite phytogeographical similarities, the two ranges differ in the amount and spatial distribution of rainfall over the elevation gradient. We studied terrestrial small mammals by direct trapping in two altitudinal transects on the eastern slope of the two mountain ranges. We recorded the changes in richness and species composition, as well as the relationships between species and microhabitats at each altitudinal level. The results show a similar structure of the small mammal assemblage in the two ranges. The largest differences, in terms of species composition, were registered at lower elevation forests, and faunal affinities increased with elevation to the point of finding identical species composition at the top of the mountains. Species richness showed a clear curvilinear pattern with a peak at the upper limit of the forests. Our findings suggest that total rainfall has an important influence on the composition and abundance of small mammal species but apparently not on the species richness along the elevation gradient. The highest values of species richness were observed at the sites where a contact between two different physiognomic units exists. These results indicate that habitat heterogeneity plays an important role in allowing the juxtaposition of small terrestrial mammal assemblages of the highlands and lowlands at a given point, contributing significantly to the considerable diversity of species observed in intermediate altitudinal sites.
     

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.