Integrating thermal tolerance, water balance and morphology: An experimental study on dung beetles.

The impacts of extreme and rising mean temperatures due to climate change can pose significant physiological challenges for insects. An integrated approach that focuses on mechanisms of body temperature regulation, water balance and morphology may help to unravel the functional traits underpinning thermoregulation strategies and the most relevant trade-offs between temperature and water balance regulation. Here, we focused on four species of tunneler dung beetles as important providers of ecosystem services. In this experimental research, we first quantified two traits related to desiccation resistance and tolerance via experimental tests, and subsequently defined two levels of resistance and tolerance (i.e. low and high) according to significant differences among species. Second, we identified morphological traits correlated with water balance strategies, and we found that desiccation resistance and tolerance increased with small relative size of spiracles and wings. High levels of desiccation tolerance were also correlated with small body mass. Third, by integrating thermal tolerance with functional traits based on desiccation resistance and desiccation tolerance, we found that the species with the highest survival rates under elevated temperatures (Euoniticellus fulvus) was characterized by low desiccation resistance and high desiccation tolerance. Our results suggest shared physiological and morphological responses to temperature and desiccation, with potential conflicts between the need to regulate heat and water balance. They also highlighted the sensitivity of a large species such as Geotrupes stercorarius to warm and arid conditions with potential implications for its geographic distribution and the provisioning of ecosystem services under a climate change scenario.

Ecological functions provided by dung beetles are interlinked across space and time: evidence from 15 N isotope tracing.

Maintaining multiple ecological functions ("multifunctionality") is crucial to sustain viable ecosystems. To date most studies on biodiversity-ecosystem functioning (BEF) have focused on single or few ecological functions and services. However, there is a critical need to evaluate how species and species assemblages affect multiple processes at the same time, and how these functions are interconnected. Dung beetles represent excellent model organisms because they are key contributors to several ecosystem functions. Using a novel method based on the application of 15 N-enriched dung in a mesocosm field experiment, we assessed the role of dung beetles in regulating multiple aspects of nutrient cycling in alpine pastures over appropriate spatial (up to a soil depth of 20 cm) and temporal (up to 1 yr after dung application) scales. 15 N isotope tracing allowed the evaluation of multiple interrelated ecosystem functions responsible for the cycling of dung-derived nitrogen (DDN) in the soil and vegetation. We also resolved the role of functional group identity and the importance of interactions among co-occurring species for sustaining multiple functions by focusing on two different dung beetle nesting strategies (tunnelers and dwellers). Species interactions were studied by contrasting mixed-species to single-species assemblages, and asking whether the former performed multiple functions better than the latter. Dung beetles influenced at least seven ecological functions by facilitating dung removal, transport of DDN into the soil, microbial ammonification and nitrification processes, uptake of DDN by plants, herbage growth, and changes in botanical composition. Tunnelers and dwellers were found to be similarly efficient for most functions, with differences based on the spatial and temporal scales over which the functions operated. Although mixed-species assemblages seemed to perform better than single-species, this outcome may be dependent on the context. Most importantly though, the different functions were found to be interconnected sequentially as reveled by analyzing 15 N content in dung, soil and vegetation. Taken together, our current findings offer strong support for the contention that the link between biodiversity and ecosystem functions should be examined not function by function, but in terms of understanding multiple functions and how they interact with each other.

Alpine bird distributions along elevation gradients: the consistency of climate and habitat effects across geographic regions.

Many species have shown recent shifts in their distributions in response to climate change. Patterns in species occurrence or abundance along altitudinal gradients often serve as the basis for detecting such changes and assessing future sensitivity. Quantifying the distribution of species along altitudinal gradients acts as a fundamental basis for future studies on environmental change impacts, but in order for models of altitudinal distribution to have wide applicability, it is necessary to know the extent to which altitudinal trends in occurrence are consistent across geographically separated areas. This was assessed by fitting models of bird species occurrence across altitudinal gradients in relation to habitat and climate variables in two geographically separated alpine regions, Piedmont and Trentino. The ten species studied showed non-random altitudinal distributions which in most cases were consistent across regions in terms of pattern. Trends in relation to altitude and differences between regions could be explained mostly by habitat or a combination of habitat and climate variables. Variation partitioning showed that most variation explained by the models was attributable to habitat, or habitat and climate together, rather than climate alone or geographic region. The shape and position of the altitudinal distribution curve is important as it can be related to vulnerability where the available space is limited, i.e. where mountains are not of sufficient altitude for expansion. This study therefore suggests that incorporating habitat and climate variables should be sufficient to construct models with high transferability for many alpine species.

Coccidioidomycosis in Biopsies with Presumptive Diagnosis of Malignancy in Dogs: Report of Three Cases and Comparative Discussion of Published Reports.

Coccidioidomycosis is a respiratory fungal infection with occasional systemic dissemination. The disseminated coccidioidomycosis is considered a multifaceted disease. In medicine, disseminated coccidioidomycosis is included within a group of infectious diseases that have been referred as the great imitators. In many cases, malignancies are included in the presumptive diagnosis. In veterinary medicine, disseminated coccidioidomycosis is common in dogs. Nonetheless, despite of being a diagnostic dilemma, disseminated coccidioidomycosis is underestimated and frequently not included into differentials, even in endemic zones. Herein, we describe three cases of granulomatous inflammation caused by Coccidioides spp. which were masquerading malignancies in dogs (0.39 %). The presumptive diagnoses in these cases were osteosarcoma, lymphoma and neurofibroma, respectively. A PCR assay employing tissues in paraffin blocks resulted positive for C. posadasii in one of these cases. A comparative discussion on the ambiguous clinic-pathological presentation of disseminated coccidioidomycosis in dogs and humans is included.

Color polymorphism and mating trends in a population of the alpine leaf beetle Oreina gloriosa.

The bright colors of Alpine leaf beetles (Coleoptera, Chrysomelidae) are thought to act as aposematic signals against predation. Within the European Alps, at least six species display a basal color of either blue or green, likely configuring a classic case of müllerian mimicry. In this context, intra-population color polymorphism is paradoxical as the existence of numerous color morphs might hamper the establishment of a search image in visual predators. Assortative mating may be one of the main factors contributing to the maintenance of polymorphic populations. Due to the marked iridescence of these leaf beetles, the perceived color may change as the viewing or illumination angle changes. The present study, conducted over three years, involved intensive sampling of a population of Oreina gloriosa from the Italian Alps and applied colorimetry and a decision tree method to identify the color morphs in an objective manner. The tertiary sex ratio of the population was biased in favor of males, suggesting that viviparous females hide to give birth. Seven color morphs were identified, and their frequencies varied significantly over the course of the study. Three different analyses of mating (JMating, QInfomating, and Montecarlo simulations) recognized a general trend for random mating which coexists with some instances of positive and negative assortative mating. This could help explain the pre-eminence of one morph (which would be favored because of positive selection due to positive assortative mating) in parallel with the persistence of six other morphs (maintained due to negative assortative mating).

Behavioural Repeatability and Behavioural Syndrome in the Dung Beetle Copris umbilicatus (Coleoptera, Scarabaeidae).

Although personality studies have primarily focused on vertebrates, the evidence showing invertebrates to be capable of displaying personalities has been steadily growing in recent years. In this study, we investigated the behavioural repeatability (repetition of a behaviour over time) and behavioural syndromes (a set of correlated behaviours) in Copris umbilicatus, which is a dung beetle species showing complex sub-social behaviour. We analysed three behaviours (activity, thanatosis and distress call emission) by measuring seven distinct behavioural traits (i.e., three activity-, one thanatosis- and three distress call-related traits). We found moderate to high levels of individual repeatability in all behavioural traits considered. The duration of thanatosis was inversely correlated with two activity traits, hinting a behavioural syndrome for thanatosis and activity, with bolder individuals exhibiting shorter thanatosis and higher locomotor activity in contrast with fearful individuals, which display longer thanatosis and poor locomotor activity. No relationships were found between the behavioural traits and body size or sex. Results of the principal component analysis (PCA) suggested personality differences among individuals. Dung beetles provide an impressive variety of ecosystem services. Since the provision of these services may depend on the personalities represented in local populations and communities, studies on the ecology of personality in dung beetles should be encouraged in future research.

The foraging ecology of yellow-billed and red- billed choughs changed between two climatically different years.

Climate change is affecting the alpine ecosystem at an unprecedented rate, with marked changes in spring phenology and the elevation distribution of birds. Changes in the European Alps are happening rapidly, and it is possible behaviours stand to change from one year to the next. The year 2022 was characterised by climatic extremes: Italy experienced its hottest year ever, and it was the driest since 1800. Here, we assessed whether the foraging ecology of two coexisting upland bird species, the yellow-billed and the red-billed chough, changed from 2021 to 2022. We assessed foraging stay times, flock size, propensity to mixed flocking, foraging home ranges and altitudinal distribution. Stay times of both species when foraging in monospecific flocks significantly shortened in 2022, especially in the case of the red-billed chough. The two corvids are known to influence each other when foraging together. In 2021, as expected, the stay times of the red-billed chough decreased when in the presence of the congener, but this did not occur in 2022. Instead, the yellow-billed chough increased its altitudinal foraging distribution in 2022. The results are in line with the hypothesis that large climate variations may disrupt the foraging ecology of mountain birds. However, as it is not possible to draw solid conclusions from just two years of observations, further field research will have to be planned in the future.

Dung removal increases under higher dung beetle functional diversity regardless of grazing intensification.

Dung removal by macrofauna such as dung beetles is an important process for nutrient cycling in pasturelands. Intensification of farming practices generally reduces species and functional diversity of terrestrial invertebrates, which may negatively affect ecosystem services. Here, we investigate the effects of cattle-grazing intensification on dung removal by dung beetles in field experiments replicated in 38 pastures around the world. Within each study site, we measured dung removal in pastures managed with low- and high-intensity regimes to assess between-regime differences in dung beetle diversity and dung removal, whilst also considering climate and regional variations. The impacts of intensification were heterogeneous, either diminishing or increasing dung beetle species richness, functional diversity, and dung removal rates. The effects of beetle diversity on dung removal were more variable across sites than within sites. Dung removal increased with species richness across sites, while functional diversity consistently enhanced dung removal within sites, independently of cattle grazing intensity or climate. Our findings indicate that, despite intensified cattle stocking rates, ecosystem services related to decomposition and nutrient cycling can be maintained when a functionally diverse dung beetle community inhabits the human-modified landscape.

Continuous phenotypic modulation explains male horn allometry in three dung beetle species.

Many dung beetle species show male horn polyphenism, the ability of males to develop into distinct phenotypes without intermediate forms as a response to the larval growth environment. While males with long (majors) and rudimentary (minor) horn have been widely reported in literature, little is known about the existence of individuals with intermediate horn length. Here we investigate the occurrence of intermediates in natural populations of three dung beetle species (Onthophagus furcatus, Copris lunaris and C. hispanus). We analysed the body size-horn length relationship using linear, exponential, and sigmoidal models with different error structures. We inferred the number of individuals in the minor, intermediate, and major groups by combining changepoint analysis and simulation from fitted allometric models. The sigmoidal equation was a better descriptor of the body size-horn length relationship than linear or exponential equations in all the three studied species. Our results indicated that the number of intermediates equals or exceeds the number of minor and major males. This work provides evidence that, at least in the studied species, males with intermediate horn length exist in natural populations. For similar cases we therefore suggest that continuous phenotypic modulation rather than discrete polyphenism can explain variation in male horn allometry.

Physiological, morphological and ecological traits drive desiccation resistance in north temperate dung beetles.

BACKGROUND: Increasing temperatures and changes in precipitation patterns threaten the existence of many organisms. It is therefore informative to identify the functional traits that underlie differences in desiccation resistance to understand the response of different species to changes in water availability resulting from climate change. We used adult dung beetles as model species due to their importance to ecosystem services. We investigated: (i) the effect of physiological (water loss rate, water loss tolerance, body water content), morphological (body mass) and ecological (nesting behaviour) traits on desiccation resistance; (ii) the role of phylogenetic relatedness in the above associations; and, (iii) whether relatively large or small individuals within a species have similar desiccation resistance and whether these responses are consistent across species. RESULTS: Desiccation resistance decreased with increasing water loss rate and increased with increasing water loss tolerance (i.e. proportion of initial water content lost at the time of death). A lack of consistent correlation between these traits due to phylogenetic relatedness suggests that the relationship is not determined by a shared evolutionary history. The advantage of a large body size in favouring desiccation resistance depended on the nesting behaviour of the dung beetles. In rollers (one species), large body sizes increased desiccation resistance, while in tunnelers and dwellers, desiccation resistance seemed not to be dependent on body mass. The phylogenetic correlation between desiccation resistance and nesting strategies was significant. Within each species, large individuals showed greater resistance to desiccation, and these responses were consistent across species. CONCLUSIONS: Resistance to desiccation was explained mainly by the dung beetles' ability to reduce water loss rate (avoidance) and to tolerate water loss (tolerance). A reduction in water availability may impose a selection pressure on body size that varies based on nesting strategies, even though these responses may be phylogenetically constrained. Changes in water availability are more likely to affect dweller species, and hence the ecosystem services they provide.

Rapid shape divergences between natural and introduced populations of a horned beetle partly mirror divergences between species.

Onthophagus taurus is a polyphenic beetle in which males express alternative major (horned) and minor (hornless) morphologies largely dependent on larval nutrition. O. taurus was originally limited to a Turanic-European-Mediterranean distribution, but became introduced to several exotic regions in the late 1960s. Using geometric morphometrics, we investigate the present-day morphological shape differentiation patterns among native (Italian) and introduced (Western Australian and Eastern US) populations. We then contrast these divergences to those observed between native O. taurus and its sympatric sister species O. illyricus. Our analysis failed to find significant divergences between O. taurus populations in external morphological traits (head, pronotum) when analyses were conducted separately for each sex. However, when sexes and male morphs were analyzed together, three important differences among populations emerged. First, relative warp analyses showed that native and introduced populations diverged in certain shape components that normally distinguish major and minor male morphs. Second, comparison of covariation of body regions (head vs. pronotum) in the three populations showed that populations diverged in the nature of this covariation, suggesting that different body regions are not totally constrained to evolve in concert. Lastly, and most importantly, the analysis of genitalic shape revealed little to no divergence of female genitalia, but unexpected substantial differentiation of male genitalia among the three O. taurus populations. This suggests that genitalic shape divergence can occur extremely rapidly even in the absence of sympatry and possible reinforcement, and that the genitalia of males and females may diverge independent of one another, at least during the early stage of interpopulational divergence. Interpopulation divergences in O. taurus mirrored aspects of interspecific divergences between O. taurus and O. illyricus in some cases but not others.

Movement and demographics of Libellula fulva (Odonata, Libellulidae).

Many capture-recapture studies on adult dragonflies have found male-biased sex ratios. However, few have estimated survivorship of males and females separately from data on frequency of recaptures in the field. Even when daily survival and capture probabilities are estimated separately, controversies can arise on whether sex biases in local survival are to be attributed to mortality or permanent emigration from the study site. The knowledge of male and female movements, assessed on an appropriate scale (i.e., within and outside the breeding site), can help address this issue. In this paper, we performed a 4-yr capture-recapture study of two Libellula fulva Müller populations in northwest Italy. Cormack-Jolly-Seber models were used to get unbiased estimates of demographic parameters (daily survival and capture probabilities, sex ratio, mean life span, and population size). Movement parameters were measured directly by georeferencing encounters. Moderate differences in survival, with males surviving better than females, were found in one population and not in the other, suggesting that these differences are not an inherent characteristic of the species. In the population with lower female survival, females were not more vagile than males, thus indicating their lower survival was caused by actual mortality rather than to emigration. In the population with no survival differences between males and females, marked males outnumbered females, but estimated sex ratios were approximately 1:1 or female biased. Therefore, raw field data were misleading because they led to underestimates of the more elusive sex and overestimates of the more detectable one (males). Survival and movement differences detected in the two populations are discussed in the framework of local environmental and demographic factors.

Evidence for Male Horn Dimorphism and Related Pronotal Shape Variation in Copris lunaris (Linnaeus, 1758) (Coleoptera: Scarabaeidae, Coprini).

Male horn dimorphism is a rather common phenomenon in dung beetles, where some adult individuals have well-developed head horns (i.e., major males), while others exhibit diminished horn length (i.e., minor males). We focused on horn dimorphism and associated head and pronotum shape variations in Copris lunaris. We examined the allometric relationship between horn length (i.e., cephalic and pronotal horns) and maximum pronotum width (as index of body size) by fitting linear and sigmoidal models for both sexes. We then asked whether head and pronotum shape variations, quantified using the geometric morphometric approach, contributed to this allometric pattern. We found that female cephalic and pronotal horn growth showed a typical isometric scaling with body size. Horn length in males, however, exhibited sigmoidal allometry, where a certain threshold in body size separated males into two distinct morphs as majors and minors. Interestingly, we highlighted the same allometric patterns (i.e., isometric vs. sigmoidal models) by scaling horn lengths with pronotum shape, making evident that male horn dimorphism is not only a matter of body size. Furthermore, the analysis of shape showed that the three morphs had similar heads, but different pronota, major males showing a more expanded, rounded pronotum than minor males and females. These morphological differences in C. lunaris can ultimately have important functional consequences in the ecology of this species, which should be explored in future work.

Dung beetles as drivers of ecosystem multifunctionality: Are response and effect traits interwoven?

Rapid biodiversity loss has emphasized the need to understand how biodiversity affects the provisioning of ecological functions. Of particular interest are species and communities with versatile impacts on multiple parts of the environment, linking processes in the biosphere, lithosphere, and atmosphere to human interests in the anthroposphere (in this case, cattle farming). In this study, we examine the role of a specific group of insects - beetles feeding on cattle dung - on multiple ecological functions spanning these spheres (dung removal, soil nutrient content and greenhouse gas emissions). We ask whether the same traits which make species prone to extinction (i.e. response traits) may also affect their functional efficiency (as effect traits). To establish the link between response and effect traits, we first evaluated whether two traits (body mass and nesting strategy, the latter categorized as tunnelers or dwellers) affected the probability of a species being threatened. We then tested for a relationship between these traits and ecosystem functioning. Across Scandinavian dung beetle species, 75% of tunnelers and 30% of dwellers are classified as threatened. Hence, nesting strategy significantly affects the probability of a species being threatened, and constitutes a response trait. Effect traits varied with the ecological function investigated: density-specific dung removal was influenced by both nesting strategy and body mass, whereas methane emissions varied with body mass and nutrient recycling with nesting strategy. Our findings suggest that among Scandinavian dung beetles, nesting strategy is both a response and an effect trait, with tunnelers being more efficient in providing several ecological functions and also being more sensitive to extinction. Consequently, functionally important tunneler species have suffered disproportionate declines, and species not threatened today may be at risk of becoming so in the near future. This linkage between effect and response traits aggravates the consequences of ongoing biodiversity loss.

Greenhouse gas emissions from dung pats vary with dung beetle species and with assemblage composition.

Cattle farming is a major source of greenhouse gases (GHGs). Recent research suggests that GHG fluxes from dung pats could be affected by biotic interactions involving dung beetles. Whether and how these effects vary among beetle species and with assemblage composition is yet to be established. To examine the link between GHGs and different dung beetle species assemblages, we used a closed chamber system to measure fluxes of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) from cattle dung pats. Targeting a total of four dung beetle species (a pat-dwelling species, a roller of dung balls, a large and a small tunnelling species), we ran six experimental treatments (four monospecific and two mixed) and two controls (one with dung but without beetles, and one with neither dung nor beetles). In this setting, the overall presence of beetles significantly affected the gas fluxes, but different species contributed unequally to GHG emissions. When compared to the control with dung, we detected an overall reduction in the total cumulative CO2 flux from all treatments with beetles and a reduction in N2O flux from the treatments with the three most abundant dung beetle species. These reductions can be seen as beneficial ecosystem services. Nonetheless, we also observed a disservice provided by the large tunneler, Copris lunaris, which significantly increased the CH4 flux-an effect potentially traceable to the species' nesting strategy involving the construction of large brood balls. When fluxes were summed into CO2-equivalents across individual GHG compounds, dung with beetles proved to emit less GHGs than did beetle-free dung, with the mix of the three most abundant species providing the highest reduction (-32%). As the mix of multiple species proved the most effective in reducing CO2-equivalents, the conservation of diverse assemblages of dung beetles emerges as a priority in agro-pastoral ecosystems.

Identifying key conservation threats to Alpine birds through expert knowledge.

Alpine biodiversity is subject to a range of increasing threats, but the scarcity of data for many taxa means that it is difficult to assess the level and likely future impact of a given threat. Expert opinion can be a useful tool to address knowledge gaps in the absence of adequate data. Experts with experience in Alpine ecology were approached to rank threat levels for 69 Alpine bird species over the next 50 years for the whole European Alps in relation to ten categories: land abandonment, climate change, renewable energy, fire, forestry practices, grazing practices, hunting, leisure, mining and urbanization. There was a high degree of concordance in ranking of perceived threats among experts for most threat categories. The major overall perceived threats to Alpine birds identified through expert knowledge were land abandonment, urbanization, leisure and forestry, although other perceived threats were ranked highly for particular species groups (renewable energy and hunting for raptors, hunting for gamebirds). For groups of species defined according to their breeding habitat, open habitat species and treeline species were perceived as the most threatened. A spatial risk assessment tool based on summed scores for the whole community showed threat levels were highest for bird communities of the northern and western Alps. Development of the approaches given in this paper, including addressing biases in the selection of experts and adopting a more detailed ranking procedure, could prove useful in the future in identifying future threats, and in carrying out risk assessments based on levels of threat to the whole bird community.

The effects of body mass on dung removal efficiency in dung beetles.

Understanding of the role of body mass in structural-functional relationships is pressing, particularly because species losses often occur non-randomly with respect to body size. Our study examined the effects of dung beetle body mass on dung removal at two levels. First, we used the lab experiment to evaluate the efficiency of eight dung beetle species belonging to two functional groups (tunnelers, dwellers) on dung removal. Second, the same species employed in the lab were used in field mesocosms to examine the effects of the two functional groups on dung removal maintaining realistic differences in the total body mass between tunneler and dweller assemblages. Furthermore, the experimental assemblages contained one and four species within each functional group, so the effect of body mass heterogeneity was examined. We used a statistical approach (offset method) which took into account a priori constraints due to the study design allowing us to analyse the effect of larger species in mesocosm style experiments. Body size played a crucial role in dung removal: large beetles were more efficient than small ones and the percentage of removed dung increased with higher body mass heterogeneity. Tunnelers were more efficient than dwellers over both short and long time periods (one month and one year). Significant effects of dwellers were found only after one year. Moreover, our study showed that not including the body mass as an offset in the model resulted in sometimes different results, as the offset expresses dung removal independently of the body mass. This approach confirmed that body size is likely a pivotal factor controlling dung removal efficiency at multiple levels, from single species to overall dung beetle assemblages. Even though other specific traits should be examined, this study has begun to address the consequences of losing individuals with specific traits that are especially sensitive to perturbations.

Pastoral practices to reverse shrub encroachment of sub-alpine grasslands: dung beetles (coleoptera, scarabaeoidea) respond more quickly than vegetation.

In recent decades, pastoral abandonment has produced profound ecological changes in the Alps. In particular, the reduction in grazing has led to extensive shrub encroachment of semi-natural grasslands, which may represent a threat to open habitat biodiversity. To reverse shrub encroachment, we assessed short-term effects of two different pastoral practices on vegetation and dung beetles (Coleoptera, Scarabaeoidea). Strategic placement of mineral mix supplements (MMS) and arrangement of temporary night camp areas (TNCA) for cattle were carried out during summer 2011 in the Val Troncea Natural Park, north-western Italian Alps. In 2012, one year after treatment, a reduction in shrub cover and an increase in bare ground cover around MMS sites was detected. A more intense effect was detected within TNCA through increases in forage pastoral value, and in the cover and height of the herbaceous layer. Immediately after treatment, changes in dung beetle diversity (total abundance, species richness, Shannon diversity, taxonomic and functional diversity) showed a limited disturbance effect caused by high cattle density. In contrast, dung beetle diversity significantly increased one year later both at MMS and TNCA sites, with a stronger effect within TNCA. Multivariate Regression Trees and associated Indicator Value analyses showed that some ecologically relevant dung beetle species preferred areas deprived of shrub vegetation. Our main conclusions are: i) TNCA are more effective than MMS in terms of changes to vegetation and dung beetles, ii) dung beetles respond more quickly than vegetation to pastoral practices, and iii) the main driver of the rapid response by dung beetles is the removal of shrubs. The resulting increase in dung beetle abundance and diversity, which are largely responsible for grassland ecosystem functioning, may have a positive effect on meso-eutrophic grassland restoration. Shrub encroachment in the Alps may therefore be reversed, and restoration of grassland enhanced, by using appropriate pastoral practices.

The impact of overgrazing on dung beetle diversity in the Italian Maritime Alps.

Dung beetles (Coleoptera: Scarabaeidae) are undoubtedly the most typical and ecologically relevant insects of grazed alpine habitats because they provide valuable ecological services such as biological pest control and soil fertilization. Despite the great ecological contribution of these insects to pasture ecosystem functioning, little is known about their direct or indirect relationships with pastoral activities. The main aim of the study was to assess whether dung beetle diversity was influenced by different intensities of cattle grazing. Dung beetle communities of two adjacent alpine valleys within the Maritime Alps Natural Park (north-western Italian Alps), representing overgrazed and ungrazed pastures, were studied by pitfall trapping. A hierarchical design (three levels: valleys, transects, and replicates) was established for additive partitioning of γ-diversity and Indicator Species Analysis. Evenness and Shannon diversity were significantly higher at the ungrazed than at the overgrazed site because abundances were much more evenly distributed at the former than at the latter site (where one species was dominant over all the others). Dung beetle abundance and species richness of the overgrazed graminaceous pasture vegetation types were in most cases significantly lower than those of the ungrazed nongraminaceous vegetation type. In the additive partitioning of γ -diversity analysis relative to the whole study area, the randomization procedure indicated that the contribution of ß to γ-diversity was significantly different from that expected by chance, suggesting that one or more environmental factors has intervened to change the partition of total diversity in the system considered. The analysis of the preferences and fidelity of species (Indicator Species Analysis) showed that only one species chose overgrazed pastures; all the others positively selected the ungrazed site, or the only ungrazed pasture vegetation type (Rumicetum alpini Beger) occurring at the overgrazed site. Results conformed to evidences that overgrazing represents a serious threat to the conservation of alpine dung beetles. To conserve local dung beetle assemblages, especially in protected areas, cattle overgrazing should be avoided. This does not mean, however, that pastoral activities are incompatible with biodiversity conservation. The contemporaneous presence of wild ungulates and low intensity extensive pastoral activities may be useful to preserve both local dung beetle assemblages and alpine pasture ecosystems.