Panmixia across elevation in thermally sensitive Andean dung beetles

Janzen's seasonality hypothesis predicts that organisms inhabiting environments with limited climatic variability will evolve a reduced thermal tolerance breadth compared with organisms experiencing greater climatic variability. In turn, narrow tolerance breadth may select against dispersal across strong temperature gradients, such as those found across elevation. This can result in narrow elevational ranges and generate a pattern of isolation by environment or neutral genetic differentiation correlated with environmental variables that are independent of geographic distance. We tested for signatures of isolation by environment across elevation using genome‐wide SNP data from five species of Andean dung beetles (subfamily Scarabaeinae) with well‐characterized, narrow thermal physiologies, and narrow elevational distributions. Contrary to our expectations, we found no evidence of population genetic structure associated with elevation and little signal of isolation by environment. Further, elevational ranges for four of five species appear to be at equilibrium and show no decay of genetic diversity at range limits. Taken together, these results suggest physiological constraints on dispersal may primarily operate outside of a stable realized niche and point to a lower bound on the spatial scale of local adaptation.     

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

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

Effects of Degradation and Replacement of Southern Brazilian Coastal Sandy Vegetation on the Dung Beetles (Coleoptera: Scarabaeidae)

In this paper we address the effects of anthropogenic disturbance and replacement of Brazilian Coastal sandy vegetation (restingas) on dung beetles communities. We sampled dung beetles in the four main vegetative physiognomies of Guriri Island, Espírito Santo State: forest restinga, restinga Clusia, disturbed restinga (from burning events), and pastures. We placed four sets of two pitfall traps (baited with horse and human dung) in four independent areas of each vegetation type, and collected 14,534 individuals of 13 dung beetle species. Neither log₁₀ of individuals nor log₁₀ of species richness were good predictors of restinga disturbance. However, a significant amount of variation in dung beetle abundance and richness could be explained by bait type. Ordination of these sites using hybrid multidimensional scaling revealed a gradient of habitat disturbance from undisturbed restinga samples to pasture. Dung beetle communities along this gradient demonstrated a complete turnover in species composition, from restinga‐specialists to invasive and generalists species respectively. This complete turnover signals the local extirpation of forest‐adapted species in disturbed and converted areas. Only a single dung beetle species in preserved restingas is protected by Brazilian law (Dichotomius schiffleri). Given the extent of the clearing of restinga habitat, the conservation status of dung beetles associated with restinga forest gives cause for concern.     

Historical biogeography of scarabaeine dung beetles

Abstract Aim (1) To review briefly global biogeographical patterns in dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), a group whose evolutionary history has been dominated by ecological specialization to vertebrate dung in warmer climates. (2) To develop hypotheses accounting for the evolution of these patterns. Location Six principal biogeographical regions: Palaearctic, Oriental, Afrotropical, Australasia, Neotropical, Nearctic and five outlying islands or island groups harbouring endemic genera: Caribbean, Madagascar, Mauritius, New Caledonia, New Zealand. Methods Major patterns of tribal, generic and species distribution are investigated using cluster analysis, ordination, parsimony analysis of endemism and track analysis. Attempts are made to resolve biogeographical patterns with findings in the fields of plate tectonics, fossil and evolutionary history, plus phylogeny of both mammals and dung beetles. Results Because of conflict between published findings, it is uncertain at what point in time density of dinosaur dung, mammal dung or both became sufficiently great to select for specialized habits in dung beetles. However, biogeographical evidence would suggest a Mesozoic origin followed by further taxonomic radiation during the Cenozoic, possibly in response to the increasing size and diversity of mammalian dung types in South America and Afro‐Eurasia. Proportional generic distribution in fourteen tribes and subtribes showed four principal biogeographical patterns: (1) southerly biased Gondwanaland distribution, (2) Americas or (3) Madagascar endemism, and (4) northerly biased, Afro‐Eurasian‐centred distribution with limited numbers of genera also widespread in other regions. Proportional composition of faunas in eleven geographical regions indicated three principal distributional centres, East Gondwanaland fragments, Afro‐Eurasia and the Americas. These patterns probably result from three principal long‐term range expansion and vicariance events (Mesozoic: Gondwanaland interchange and fragmentation, Cenozoic: Afro‐Eurasian/Nearctic interchange and the Great American interchange). It is suggested that old vicariance caused by the Mesozoic fragmentation of Gondwanaland leads to a high degree of regional endemism at generic or tribal level across one or more Gondwanaland tracks. In contrast, it is suggested that the more recent Cenozoic range expansions occurred primarily towards northern regions leading to endemism primarily at species level. These Cenozoic radiations were facilitated by the re‐linking of continents, either because of tectonic plate movements (Africa to Eurasia in Miocene), climatically induced sea‐level change (Afro‐Eurasia to Nearctic in Miocene and Pleistocene), or similar coupled with orogenics (Nearctic to Neotropical in Pliocene). Speciation has followed vicariance either because of climatic change or physical barrier development. These recent range expansions probably occurred principally along an Afro‐Eurasian land track to the Nearctic and Neotropical and an Americas land track northwards from the Neotropics to the Nearctic, with limited dispersal from Eurasia to Australia, probably across a sea barrier. This accounts for the overall, spatially constrained, biogeographical pattern comprising large numbers of species‐poor genera endemic to a single biogeographical region and fewer more species‐rich genera, many of which show wider biogeographical distributions. In most southerly regions (Australasia, Madagascar, Neotropical), faunal composition and generic endemism is primarily dominated by elements with Gondwanaland ancestry, which is consistent with the Gondwanaland origin claimed for Scarabaeinae. In Afro‐Eurasia (Palaearctic, Oriental, Afrotropical), generic endemism of monophyletically derived Afro‐Eurasian and widespread lineages is centred in the Afrotropical region and faunal composition is numerically dominated by Afro‐Eurasian and widespread elements. In the Nearctic region, the fauna is jointly dominated by widespread elements, derived from Afro‐Eurasia, and Gondwanaland and Americas elements derived from the Neotropical region. Main conclusions Global biogeographical patterns in scarabaeine dung beetles primarily result from Mesozoic and Cenozoic range expansion events followed by vicariance, although recent dispersal to Australia may have occurred across sea barriers. Detailed phylogenetics research is required to provide data to support dispersal/vicariance hypotheses.     

Tropical dung beetle morphological traits predict functional traits and show intraspecific differences across land uses

Functional traits and functional diversity measures are increasingly being used to examine land use effects on biodiversity and community assembly rules. Morphological traits are often used directly as functional traits. However, behavioral characteristics are more difficult to measure. Establishing methods to derive behavioral traits from morphological measurements is necessary to facilitate their inclusion in functional diversity analyses. We collected morphometric data from over 1,700 individuals of 12 species of dung beetle to establish whether morphological measurements can be used as predictors of behavioral traits. We also compared morphology among individuals collected from different land uses (primary forest, logged forest, and oil palm plantation) to identify whether intraspecific differences in morphology vary among land use types. We show that leg and eye measurements can be used to predict dung beetle nesting behavior and period of activity and we used this information to confirm the previously unresolved nesting behavior for Synapsis ritsemae. We found intraspecific differences in morphological traits across different land use types. Phenotypic plasticity was found for traits associated with dispersal (wing aspect ratio and wing loading) and reproductive capacity (abdomen size). The ability to predict behavioral functional traits from morphology is useful where the behavior of individuals cannot be directly observed, especially in tropical environments where the ecology of many species is poorly understood. In addition, we provide evidence that land use change can cause phenotypic plasticity in tropical dung beetle species. Our results reinforce recent calls for intraspecific variation in traits to receive more attention within community ecology.     

How localized are the distributions of Australian scarabs (Coleoptera,Scarabaeoidea)?

The degree of localization in the distribution of Australian scarabaeine (320 spp.), aphodiine (160 spp.), geotrupid (166 spp.) and dynastine (182 spp.) scarabs is analysed. Localization differs among these groups, appearing to increase marginally from the Dynastinae, to the Aphodiinae, to the Scarabaeinae, to the Geotrupidae, but with significant differences within groups, e.g. the scarabaeine tribes Onthophagini and Scarabaeini. However, the Australian fauna lacks the high proportion of widespread species characteristic of the West Palaearctic Scarabaeinae, Aphodiinae and Geotrupidae. This difference reflects the different ecological histories of the two regions.     

Dung beetle diversity and functions suggest no major impacts of cattle grazing in the Brazilian Pantanal wetlands

1. Dung beetles perform relevant ecological functions in pastures, such as dung removal and parasite control. Livestock farming is the main economic activity in the Brazilian Pantanal. However, the impact of cattle grazing on the Pantanal's native dung beetle community, and functions performed by them, is still unknown. 2. This study evaluated the effects of cattle activity on dung beetle community attributes (richness, abundance, biomass, composition, and functional group) as well as their ecological functions (dung removal and soil bioturbation) in the Pantanal. In January/February 2016, dung beetles were sampled and their ecological functions measured in 16 sites of native grasslands in Aquidauana, Mato Grosso do Sul, Brazil, 10 areas regularly grazed by cattle and six control ungrazed areas (> 20 years of abandonment). 3. In all, 1169 individuals from 30 species of dung beetles were collected. Although abundance, species richness, and biomass did not differ between grasslands with and without cattle activity, species composition and functional groups differed among systems. Large roller beetles were absent from non‐cattle grasslands, and the abundance, richness, and biomass of medium roller beetles was higher in those systems. 4. Despite causing changes in species/functional group composition, the results of this study show that a density compensation of functional groups in cattle‐grazed natural grasslands seems to have conserved the ecological functions (dung removal and soil bioturbation), with no significant differences between systems. 5. Therefore, these results provide evidence that cattle breeding in natural grasslands of the Brazilian Pantanal can integrate livestock production with the conservation of the dung beetle community and its ecological functions.     

Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes

Aim We describe the changes in species richness, rarity and composition with altitude, and explore whether the differences in Scarabaeinae dung beetle composition along five altitudinal transects of the same mountain range are related to altitude or if there are interregional differences in these altitudinal gradients. Location Field work was carried out on the eastern slope of the eastern Cordillera, Colombian Andes, between Tamá Peak to the north, in the Tamá National Park (07°23′ N, 72°23′ W) and the San Miguel River (00°28′ N, 77°17′ W) to the south. Methods Sampling was carried out between February 1997 and November 1999 in five regions spanning elevation gradients. In each gradient, six sites were chosen at 250 m intervals between 1000 and 2250 m a.s.l. Results We found a curvilinear relationship between altitude and mean species richness, with a peak in richness at middle elevations. However, the diversity of dung beetle assemblages does not seem to be related to the interregional differences in environmental conditions. The number of geographically restricted species is negatively and significantly related to altitude, with geographically restricted species more frequent at low altitude sites. Ordination delimited the two main groups according to altitude: one with all the highest sites (1750–2250 m a.s.l.) and a second group with the remaining sites (< 1750 m a.s.l.). Analysis of species co‐occurrence shows that these dung beetle assemblages seem to be spatially structured when all sites have the same probability of being chosen. In contrast, the spatial structure of species assemblages seems to be random when the probability of choosing any site is proportional to its altitude. Main conclusions The altitude of sites is the main factor that influences the diversity of these dung beetle assemblages. The peak in species richness at middle elevations, the higher number of geographically restricted species at lower altitudinal levels, and the compositional differences along these mountain gradients seem to result from the mixing at these altitudes of dung beetle assemblages that have different environmental adaptations and, probably, different origins. The relevance of altitude in these assemblages is related to the limited role of these Neotropical high altitude environments as centres of refuge and vicariance for a monophyletic group of warm‐adapted species, for which the vertical colonization of these high mountain environments by lineages distributed at lower altitudes would have been very difficult.