Claudia Palestrini

Dung Beetle Conservation: Effects of Habitat and Resource Selection (Coleoptera: Scarabaeoidea)

There is increasing concern about the conservation of dung beetles which are threatened by several peculiar dangers world-wide, both at the population and community level. A noticeable threat in Europe is the dramatic reduction in the number of cattle and sheep grazing in the open, which is associated with both intensive agriculture and the progressive reforestation of previously pastured areas. We studied dung beetle habitat and resource preferences at La Mandria Park (north-west Italy) which is a mosaic of open and wooded patches where domestic (cows and horses) and wild ungulates (deer and wild boar) co-exist. Scarabaeidae were numerically dominant, accounting for 61.5% of the approximately 3000 individuals sampled (Aphodiidae accounted for 32.5% and Geotrupidae for only 6%). However, when species richness was considered, Aphodiidae were dominant, with 17 of the 27 species found (Scarabaeidae with eight and Geotrupidae with two). Assuming a null hypothesis of equal probability of colonizing any habitat or faeces, we found that most species were significantly associated with one of the four dung types or with one of the two habitats considered. On average, Scarabaeidae preferred cattle dung and open habitats whereas most Aphodiidae used deer lumps and wooded habitats. In spite of the precise ecological choices observed at La Mandria, surveys from other European areas suggest that both habitat and food selection are quite flexible. From a conservation viewpoint, the ability of coprophagous beetles to choose herbivore faeces according to their availability and to select habitats that satisfy their own microclimate requirements may certainly be useful in preserving biodiversity. Agroecology, which implies some woodland and hedgerow management practices associated with pastoralism, could be the right approach to the management of the agricultural landscape. Conversely, in reforested areas, while wild ungulate populations should be preserved, some form of human disturbance, particularly clearings used for pastoralism, should also be maintained.

The impact of forest ski-pistes on diversity of ground-dwelling arthropods and small mammals in the Alps

Forest clearing for winter sport activities is the major force driving loss and fragmentation of the alpine forests. The establishment of ski-pistes involves impacts on every ecosystem component. To assess the extent of this threat we studied ground-dwelling arthropods (namely ground beetles and spiders) and small mammals (shrews and voles) at two ski resorts in north-western Italian Alps by pitfall trapping. Diversity parameters (mean abundance, species richness and Shannon index) of spiders and macropterous carabids increased from forest interior to open habitats (i.e., ski-piste or pasture), whereas parameters of brachypterous carabids significantly decreased from forest interior to open habitats. Diversity parameters of macropterous ground beetles were higher on pastures than on ski-pistes. Small mammals were virtually absent from ski-pistes. Observed frequencies in the three adjacent habitats were significantly different from expected ones for the bank vole Myodes glareolus and the pygmy shrew Sorex minutus. Generalized linear models showed that abundance, species richness and diversity of spiders and macropterous carabids of ski-pistes were best modelled by combination of factors, including grass cover and width of the ski-piste. Indicator Species Analysis showed that species that significantly preferred ski-pistes were less than those preferring pastures, and species which were exclusive of ski-pistes were very few. To retain arthropod ground-dwelling fauna of open habitats environmentally friendly ways of constructing pistes should be developed. After tree clearing, only the roughest ground surfaces should be levelled, in order to preserve as much natural vegetation as possible. Where necessary, ski-pistes should be restored through the recovery of local vegetation.

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.

Shape - but Not Size - Codivergence between Male and Female Copulatory Structures in Onthophagus Beetles

Genitalia are among the fastest evolving morphological traits in arthropods. Among the many hypotheses aimed at explaining this observation, some explicitly or implicitly predict concomitant male and female changes of genital traits that interact during copulation (i.e., lock and key, sexual conflict, cryptic female choice and pleiotropy). Testing these hypotheses requires insights into whether male and female copulatory structures that physically interact during mating also affect each others evolution and patterns of diversification. Here we compare and contrast size and shape evolution of male and female structures that are known to interact tightly during copulation using two model systems: (a) the sister species O. taurus (1 native, 3 recently established populations) and O. illyricus, and (b) the species-complex O. fracticornis-similis-opacicollis. Partial Least Squares analyses indicated very little to no correlation between size and shape of copulatory structures, both in males and females. Accordingly, comparing shape and size diversification patterns of genitalia within each sex showed that the two components diversify readily - though largely independently of each other - within and between species. Similarly, comparing patterns of divergence across sexes showed that relative sizes of male and female copulatory organs diversify largely independent of each other. However, performing this analysis for genital shape revealed a signature of parallel divergence. Our results therefore suggest that male and female copulatory structures that are linked mechanically during copulation may diverge in concert with respect to their shapes. Furthermore, our results suggest that genital divergence in general, and co-divergence of male and female genital shape in particular, can evolve over an extraordinarily short time frame. Results are discussed in the framework of the hypotheses that assume or predict concomitant evolutionary changes in male and female copulatory organs.

The impact of high-altitude ski pistes on ground-dwelling arthropods in the Alps

The development of winter sport resorts above the timberline may affect every ecosystem component. We analyzed the effect of ski-pistes on the abundance and species richness of arthropods (namely carabids, spiders, opilionids, and grasshoppers) trapped in grasslands adjacent to the ski-run, on ski-pistes and at the edge between these two habitat types. Our results showed that diversity of brachypterous carabids, spiders, and grasshoppers decreased significantly from natural grasslands to ski-pistes. This was not true for the macropterous carabid guild, which included species with contrasting ecological requirements. The analysis of indicator species (IndVal) showed that most of the species (some of them precinctive to restricted areas in the north-western Alps) had clear preferences for natural grassland and few taxa were limited to ski-pistes. Generalized linear models suggested that the local extent of grass and rock cover can significantly affect assemblages: the low grass cover of ski-pistes, in particular, was a serious hindrance to colonization by spider, grasshopper, brachypterous, and some macropterous carabid species. The results obtained, support concerns over the possible disruption of local ecosystem functionality and over the conservation of arthropod species which are endemic to restricted alpine areas. In order to retain arthropod ground-dwelling fauna we suggest that: (i) new, environmentally friendly ways of constructing pistes should be developed to preserve as much soil and grass cover as possible; (ii) existing ski-pistes should be restored through management to promote the recovery of local vegetation.

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

SUMMARY 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.

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.

The Impact of Overgrazing on Dung Beetle Diversity in the Italian Maritime Alps

ABSTRACT 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 β 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.

The effect of local anthropogenic habitat heterogeneity on assemblages of carabids (Coleoptera, Caraboidea) endemic to the Alps

We assessed the response of a guild of endemic medium and large-bodied carabid species to the local (fine-grained) habitat heterogeneity induced by man. The study was conducted by means of trapping (without killing and preservative agents) and radio-tracking methods (to collect data on the spatial ecology of the endangered species Carabus olympiae Sella). Habitat differentiation induced by man (a mosaic of beech forests, alpen rose shrubberies and pastures) affected species richness and abundance of the guild, which were significantly higher in forests and/or shrubberies than in pastures. Most species preferred forests and shrubberies, with the only exception of Carabus concolor Fabricius, which preferred open areas. Species distribution was also affected by stone density and cattle grazing intensity. In particular, petrophilous species peaked on the roadside because of the high stone density there (stones amassed during road construction), while species abundances significantly lowered in overgrazed pastures. The compactness of the paths made by C. olympiae individuals (evaluated through a tortuosity index) was significantly higher in beech forests than in alpen rose shrubberies, indicating movements in the latter habitat type were bound by the spatial distribution of shrubs, which imposes a limit on path tortuosity. Our findings suggest that local ground beetle species diversity strongly depends on small-scale anthropogenic variables (namely habitat type, stone density and grazing intensity), and that habitat modifications (namely from forest to shrubbery) may significantly affect species movement patterns. General and local conservation management suggestions are given in conformity with these results.