Effect of pony morphology and hay feeding methods on back and neck postures.

The application of hay feeding devices, such as the use of hay nets or slow feeders, can help with the management of weight in ponies; however, there is still a lack of knowledge regarding their effect on equine posture. Therefore, the aim of the study was to assess the effect of different feeding devices on the posture of ponies using morphometric analysis. Two different breed types, Shetland type (SH, n = 5) versus Welsh Cob type (WC, n = 4), were fed the same forage in four different ways: on the ground (G), using a fully filled haynet (HF), using a partially filled haynet (HL) and using a slow-feeder hay box (HB). Video recordings were obtained and then geometric morphometric analysis was applied. Breed morphology was confirmed by body morphometric measurements. Data were analysed statistically using one-way ANOVA, canonical variate analysis (CVA), principal component analysis (PCA), partial least-squares (PLS) analysis and multivariate analysis of variance (MANOVA). Moreover, a mixed model was performed to study differences in mandibular angle. SH and WC ponies were shown to have significantly different body morphometric measurements. The geometric morphometric analysis results showed that ponies arch their back and modify their neck shape differently according to the feeding method and their morphological group. For the neck, the SH and WC ponies adapted similarly to the use of small-holed hay nets, but their posture varied when feeding from the ground or hay box. The back postures consistently differed according to the breed type and feeding method. The mandibular angle for both breed types was reduced with all the feeding devices compared to feeding from the ground. Further studies are needed to evaluate the long-term effects of slow-feeding devices on posture and mandibular angle, taking into consideration animals with different morphologies.

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.

The First Morphometric Study of the Horn Morphological Pattern in a Geotrupidae: The Case of the Dor Beetle Ceratophyus rossii Jekel, 1865.

Among beetles, thousands of species develop horns, the size of which is often extraordinarily disproportionate with respect to body size. The Scarabaeidae is the family in which horned species are most predominant, but other families, such as the Geotrupidae (dor beetles), also show remarkable horns, although in a more limited number of species. Horn expression mechanisms are well documented in Scarabaeidae but, despite the wealth of studies on this family, the horn morphological pattern of the Geotrupidae, to our knowledge, has never been investigated. In this paper, we describe for the first time the horn expression pattern in a dor beetle. As a study species, we chose Ceratophyus rossii, an Italian endemic dor beetle of the protected Mediterranean maquis in Tuscany, which shows remarkable head and pronotal horns in males and a notable cephalic horn in females. We identified and modeled shape and size horn patterns combining traditional and geometric morphometric approaches. We discuss the results in the wider landscape of developmental models described for other, more well-characterized, scarab beetles.

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).

Pony feeding management: the role of morphology and hay feeding methods on intake rate, ingestive behaviors and mouth shaping.

In the last decade, haynets and slow feeders have been promoted as sustainable tools to improve the feeding management of horses and reduce forage waste, but little is known about their effects on ponies. Therefore, the aim of this study was to analyze the effects of different hay feeding methods on the ingestive behaviors, intake rate and mouth shaping of ponies belonging to two breed types, which are characterized by different head morphologies. Shetland type (SH, n = 5) and Welsh/Cob type (WC, n = 4) ponies were fed hay using four feeding methods: on the ground (G), a fully filled haynet (HF), a partially filled haynet (HL), and a slow-feeder hay box (HB). Head morphology was measured for each pony. Video recordings were then made to apply geometric morphometrics and to perform behavioral analysis. The intake rate was measured for each pony and each feeding method. Data obtained with geometric morphometrics were analyzed using principal component analysis (PCA) and canonical variate analysis (CVA). Behavioral data and intake rate measurements were analyzed using a mixed model, a post-hoc Tukey's test, a Pearson's correlation test, and a stepwise regression model. The geometric morphometrics results demonstrated that feeding method influenced mouth shaping (36% for G, 78% for HB, 77% for HF, 83% for HL, considering the total variance of shape) and affected the intake rate. Differences in mouth shaping and ingestive behaviors in SH and WC ponies also confirmed the role of morphology in feeding management. The HL proved to be the most effective tool to increase feeding consumption time when needed (5 h/kg for SH ponies and 3 h/kg for WC ponies, considering the intake time), although the HB may be the optimal choice to reduce the intake rate while maintaining a more natural posture. Future studies are suggested to fully understand how body size and morphology influence feeding in equine species.

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.

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.

Studying the Shape Variations of the Back, the Neck, and the Mandibular Angle of Horses Depending on Specific Feeding Postures Using Geometric Morphometrics.

The effects of specific feeding positions upon the horse's shape variations of the back and neck postures as well as the variations of the mandibular angle have never been objectively studied. For this reason, geometric morphometrics was applied. Six horses, aged 14 ± 8 years (mean ± standard deviation, SD), were video-recorded while using three different feeding positions: on the ground-control position (CP); neck held 15 ± 3° below withers height with low hay net position (LP); neck held 15 ± 3° above withers height with high hay net position (HP). Data were analyzed using principal component analysis (PCA), multivariate analysis of variance (MANOVA), and partial least squares (PLS). A mixed model procedure was applied to evaluate differences in the magnitude of the mandibular angle. Whilst differences between individual horses were confirmed by canonical variate analysis (CVA), PCA analysis showed that a characteristic feeding position could also be identified on a group level. During the HP hay net position, the back and neck postures as well the mandibular angle were different compared to those exhibited by horses feeding from CP. In LP hay net position, the back posture more closely resembled those exhibited while feeding from CP; however, no similarity between LP and CP was found for neck posture and mandibular angle. Since only a few degrees of variation of the feeding position can influence back and neck postures, this aspect should be further investigated. The right compromise between horse welfare, horse safety, and management practices need to be further explored and long-term effects should be investigated.

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.

Male Horn Lack of Allometry May be Tied to Food Relocation Behaviour in Lifting Dung Beetles (Coleoptera, Scarabaeidae, Eucraniini).

The small dung beetle tribe Eucraniini includes extremely specialized species that have been defined as "lifters" according to their food relocation behaviour. They are characterized by the presence of well-developed expansions on the head and pronotum, which can be included in the large and varied group of horns, whose presence is usually related to complex reproductive tactics. In this study, two closely related species, Anomiopsoides cavifrons and A. heteroclyta, were examined employing traditional and geometric morphometrics to test whether the Eucraniini has polymorphic males that might exhibit different reproductive tactics, as in the sister tribe Phanaeini, for which a male trimorphism was demonstrated. If also present in Eucraniini polyphenism could be considered a plesiomorphy common to the two clades. The inter- and intraspecific shape variation and object symmetry of the head and the scaling relationships between body size and traits were evaluated. Marked interspecific and small intraspecific differences in shape variation, high symmetry, and similar isometric growth patterns were shown in both species. The hypothesis of male polymorphism in Anomiopsoides was thus rejected. Instead, the results supported the alternative hypothesis that Eucraniini lacks male polymorphism, perhaps due to functional constraints affecting the shape of the structures involved in their peculiar food relocating behaviour.

The Phylogenetic Relationships of Tiaronthophagus n.gen. (Coleoptera, Scarabaeidae, Onthophagini) Evaluated by Phenotypic Characters.

A necro-coprophagous new genus tha is widespread in the whole Sub-Saharan Africa was identified within the tribe Onthophagini and named Tiaronthophagus n.gen. The new genus, which is well characterized by an exclusive set of characters, comprises, at present, 26 species. Twenty species were formerly included in the genus Onthophagus and six were identified and here described as new species: Tiaronthophagus angolensis n.sp., T. jossoi n.sp., T. katanganus n.sp., T. rolandoi n.sp., T. saadaniensis n.sp., and T. zambesianus n.sp. A phylogenetic analysis that is based on a combined matrix, including discrete and landmark characters, was done. The landmark characters were tested using the geometric morphometrics techniques before their inclusion in the matrix. One single, fully resolved tree was obtained, with Tiaronthophagus constituting a distinct, monophyletic clade within Onthophagini, which was clearly separated from the other genera examined here. The biogeographical analysis identified the Central Africa as the ancestral area of the new genus and it mainly accounted for dispersal events leading to the present distribution. The generic rank that is assigned to the taxon is supported by the results of the morphological, phylogenetic, and biogeographical analyses, and by the comparison to the outgroups.

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.

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.