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.
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An important resource partitioning strategy allowing dung beetles to coexist in the same habitat, while utilising the same food, is species’ separation of activity times. After establishing the diel ...activity period of three closely related, co-occurring dung beetles, we examined their eye and wing morphology. Absolute and relative eye size, and facet size were greater in the nocturnal
Escarabaeus satyrus
, followed by the crepuscular
Scarabaeus zambesianus
and then the diurnal
Kheper lamarcki
. The diurnal
K. lamarcki
had the highest wing aspect ratio (long, narrow wings), followed by the crepuscular
S. zambesianus
and the nocturnal
E. satyrus
(short, broad wings), suggesting that dim-light active species fly slower than diurnal species. In addition, the two species active in dim light had a lower wing loading than the diurnal species, indicating the need for greater manoeuvrability in the dark. Analyses of wing shape revealed that the diurnal
K. lamarcki
wing had a proportionally larger jugal and anal region than both dim light species. Our results show that different species of dung beetles have a combination of optical and morphological wing adaptations to support their foraging activities in diverse light conditions.
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EMUNI, FIS, FZAB, GEOZS, GIS, IJS, IMTLJ, KILJ, KISLJ, MFDPS, NLZOH, NUK, OBVAL, OILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
In nature, nothing is wasted, not even waste. Dung, composed of metabolic trash and leftovers of food, is a high‐quality resource and the object of fierce competition. Over 800 dung beetle species ...(Scarabaeinae) compete in the South African dung habitat and more than 100 species can colonize a single dung pat. To coexist in the same space, using the same food, beetles divide the day between them. However, detailed diel activity periods and associated morphological adaptations have been largely overlooked in these dung‐loving insects. To address this, we used a high‐frequency trapping design to establish the diel activity period of 44 dung beetle species in their South Africa communities. This allowed us to conclude that the dung beetles show a highly refined temporal partitioning strategy, with differences in peak of activity even within the diurnal, crepuscular, and nocturnal guilds, independent of nesting behavior and taxonomic classification. We further analyzed differences in eye and body size of our 44 model species and describe their variability in external eye morphology. In general, nocturnal species are bigger than crepuscular and diurnal species, and as expected, the absolute and relative eye size is greatest in nocturnal species, followed by crepuscular and then diurnal species. A more surprising finding was that corneal structure (smooth or facetted) is influenced by the activity period of the species, appearing flat in the nocturnal species and highly curved in the diurnal species. The role of the canthus—a cuticular structure that partially or completely divides the dung beetle eye into dorsal and ventral parts—remains a mystery, but the large number of species investigated in this study nevertheless allowed us to reject any correlation between its presence and the nesting behavior or time of activity of the beetles.
Dung is the site of fierce competition. To coexist in exactly the same space, using precisely the same food, over 40 species of dung beetles are shown in our paper to finely divide the day between them. This is driven by changes in light, which in turn reflect in morphological adaptations among these species which in many cases override their taxonomic heritage.
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To guide their characteristic straight-line orientation away from the dung pile, ball-rolling dung beetles steer according to directional information provided by celestial cues, which, among the most ...relevant are the sun and polarised skylight. Most studies regarding the use of celestial cues and their influence on the orientation system of the diurnal ball-rolling beetle have been performed on beetles of the tribe Scarabaeini living in open habitats. These beetles steer primarily according to the directional information provided by the sun. In contrast, Sisyphus fasciculatus, a species from a different dung-beetle tribe (the Sisyphini) that lives in habitats with closely spaced trees and tall grass, relies predominantly on directional information from the celestial pattern of polarised light. To investigate the influence of visual ecology on the relative weight of these cues, we studied the orientation strategy of three different tribes of dung beetles (Scarabaeini, Sisyphini and Gymnopleurini) living within the same biome, but in different habitat types. We found that species within a tribe share the same orientation strategy, but that this strategy differs across the tribes; Scarabaeini, living in open habitats, attribute the greatest relative weight to the directional information from the sun; Sisyphini, living in closed habitats, mainly relies on directional information from polarised skylight; and Gymnopleurini, also living in open habitats, appear to weight both cues equally. We conclude that, despite exhibiting different body size, eye size and morphology, dung beetles nevertheless manage to solve the challenge of straight-line orientation by weighting visual cues that are particular to the habitat in which they are found. This system is however dynamic, allowing them to operate equally well even in the absence of the cue given the greatest relative weight by the particular species.
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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.
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Animals commonly integrate multiple sources of information to guide their behavior. Among insects, previous studies have suggested that the relative reliability of cues affects their weighting in ...behavior, but have not systematically explored how well alternative integration strategies can account for the observed directional choices. Here, we characterize the directional reliability of an ersatz sun at different elevations and wind at different speeds as guiding cues for a species of ball-rolling dung beetle. The relative reliability is then shown to determine which cue dominates when the cues are put in conflict. We further show through modeling that the results are best explained by continuous integration of the cues as a vector-sum (rather than switching between them) but with non-optimal weighting and small individual biases. The neural circuitry in the insect central complex appears to provide an ideal substrate for this type of vector-sum-based integration mechanism.
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•The weight of a sun cue for orientation decreases with increasing elevation•The weight of wind as an orientation cue increases with increasing speed•Behaviorally, dung beetles integrate cues using a vector summation strategy•The insect brain is well suited to encode vector summation
Biological sciences; Zoology; Ethology
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NLZOH, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UILJ, UL, UM, UPCLJ, UPUK, ZAGLJ, ZRSKP
Traditional agro-pastoral practices are in decline over much of the Alps (MacDonald et al. 2000), leading to shrub and tree encroachment, and this represents one of the main threats for the ...conservation of alpine biodiversity, as many plant and animal species are dependent on the presence of semi-natural open habitats. However, quantifying this environmental change and assessing its impact on biodiversity may be difficult, especially in the context of sparse historical survey data. The accessibility of contemporary data about local biodiversity surveys in general, and indicator taxa in particular, is an essential consideration for planning future evaluations of conservation status in the Alps and for conservation plans that use ecological indicators to monitor temporal changes in biodiversity. Dung beetles are important ecosystem service providers (Nichols et al. 2008) that have been assessed as a good ecological indicator taxon in several studies (reviewed by Nichols and Gardner 2011), and although the Alps is perhaps one of the best-studied regions in respect of dung beetles, there are still only eight readily-accessible publications. We have augmented and comprehensively reviewed the data from these publications.
We first provide data about changes on a temporal scale of seasons in a dung beetle community in the western Italian Alps, an issue that has to be addressed in the local assemblages because it would affect regional biomonitoring and conservation research. This survey of 12 099 individuals belonging to 22 species illustrates a distinct seasonal pattern at a single site. Second, we collate the results of 13 published surveys of the presence of 46 species of dung beetles in 11 valleys in the western Italian Alps in the period from 2005 to 2012, a period of accelerated change in land use that started around 1945 (MacDonald et al. 2000). Because ten of the surveys used baited pitfall traps and four more used manual collection of specimens, the abundance data were not strictly comparable and they were therefore transformed to binary data (presence-or-absence records) with measures of sampling effort. The results illustrate both spatial variation and temporal variation at the scale of years.Because of the importance of dung beetles in agro-pastoral ecosystems and the high sensitivity of montane ecosystems to climate change, these spatially and temporally explicit data sets provide important baseline information about western Italian Alpine dung beetles for investigations of the effects of land use change under ongoing climate change scenarios.
Cooperative transport allows for the transportation of items too large for the capacity of a single individual. Beyond humans, it is regularly employed by ants and social spiders where two or more ...individuals, with more or less coordinated movements, transport food to a known destination. In contrast to this, pairs of male and female dung beetles successfully transport brood balls to a location unknown to either party at the start of their common journey. We found that, when forced to overcome a series of obstacles in their path, transport efficiency of pairs of beetles was higher than of solo males. To climb tall obstacles with their common ball of dung, the female assisted the leading male in lifting the ball by steadying and pushing it upwards in a 'headstand' position during the climb initiation. Finally, we show that pairs were faster than single beetles in climbing obstacles of different heights. Our results suggest that pairs of
beetles cooperate in the transportation of brood balls with coordinated movements, where the male steers and the female primarily assists in lifting the ball. Taken together, this is to our knowledge, the first quantitative study of cooperative food transport without a known goal to aim for.
Insect declines have been attributed to several drivers such as habitat loss, climate change, invasive alien species and insecticides. However, in the global context, these effects remain patchy, ...whereas insect losses appear to be consistent worldwide. Increases in atmospheric CO2 concentrations are known to have indirect effects on herbivorous insects, but the effects on other insects are largely unexplored. We wondered if elevated atmospheric CO2 (eCO2) could influence the growth and survival of insects, not via rising temperature, nor through their changes in food quality, but by other means. Rearing tunnelling dung beetle Euoniticellus intermedius (Reiche, 1848) at pre‐industrial (250 parts per million ppm), current (400 ppm) and eCO2 levels (600 and 800 ppm), we found that exposure to eCO2 resulted in longer developmental times and increased mortality. Elevated CO2 also caused reduction of adult size and mass which is detrimental to dung beetle fitness. Additional results showed associated increases in CO2 levels inside dung brood balls, dung pH and respiration rates of the soil surrounding the developing dung beetles (CO2 flux). We thus hypothesize that elevated CO2 increases competition for O2 and nutrients between soil microbiota and subterranean insects. Given that many insect orders spend at least part of their life underground, our findings indicate the possibility of a negative ubiquitous effect of eCO2 on a large portion of the earth's insect biota. These findings therefore suggest an important area for future research on the soil community in the context of atmospheric change.
In the context of global insect declines, we wondered if elevated atmospheric CO2 (eCO2) could affect insects, not via rising temperature, nor changes in food quality, but other means. A tunnelling dung beetle reared under eCO2 showed reduction in adult size and survival, along with longer developmental time. In addition, CO2 levels inside beetle brood balls, brood ball pH and CO2 flux in the surrounding soil increased under eCO2. Given that many insect orders spend part of their lifecycle underground, this negative effect of eCO2 on a tunnelling insect might be a ubiquitous explanation for the decline in insect numbers.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
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