Variations in dung beetle assemblages across a gradient of hunting in a tropical forest

Populations of large mammals are severely depleted by hunting in tropical forests, with direct effects on plant regeneration. But indirect consequences on commensal taxa depending on them for food resources, like coprophagous beetles, are less documented. Cascading effects of species loss across Scarabaeinae are expected, with likely significant negative implications for ecosystem functions. We examined dung beetle assemblages using pitfall traps at three rain forest sites in French Guiana ranging from intact mammalian fauna (Nouragues) to moderate (Kaw) and heavy (Matoury) defaunation. The site with the most depauperate mammalian fauna showed significantly lower dung beetle species richness than the two other two sites, which were not different from each other. Mean abundance and biomass per trap were not different across sites whereas community composition strongly differed among sites. A positive correlation was observed between body size and the individual contribution to dissimilarity between Nouragues and Kaw. The species contributing the most to dissimilarity were large. By contrast, one medium-sized species, dominant in Matoury, contributed the most to dissimilarity between Matoury and other sites. Diurnal genera of large tunnellers showed a higher diversity and abundance in Nouragues compared to other sites, whereas a nocturnal genus showed no differences. Large rollers were more abundant in Kaw compared to other sites. None of the groups of small beetles but one were affected by defaunation. Our results suggest that loss of large mammal populations affects dung beetle assemblage structure and causes decreasing abundance or disappearance of large tunnellers species that have a major impact on several dung beetle-mediated ecological processes.     

Saproxylic beetle assemblages related to silvicultural management intensity and stand structures in a beech forest in Southern Germany

Compared to agricultural land and spruce plantations, central European beech-oak forests are often relatively close to natural conditions. However, forest management may alter these conditions. In Steigerwald, southern Germany, a large beech-dominated forest area, three management intensities were applied during the past 30–70 years. Here, we examined the influence of management intensity on saproxylic beetles in >100-year old mature stands at 69 sampling plots in 2004. We sampled beetles using flight-window traps and time standard direct searches. The community structure based on presence/absence data changed remarkably along the gradient from unmanaged to low-intensity to high-intensity management, but these differences were not evident using abundance data from flight interception traps. Saproxylic species richness decreased in intensively managed forests. Elateridae and threatened species richness peaked in unmanaged forests and in forests under low-intensity management. Saproxylic species richness was dependent on certain micro-habitat factors. These factors were (1) the amount of dead wood for Elateridae, overall and threatened saproxylic beetle richness; (2) the amount of flowering plants for Cerambycidae; (3) the richness of wood-inhabiting fungi for Staphylinidae, Melandryidae and overall saproxylic beetle richness; and (4) the frequency of Fomes fomentarius for threatened species. Species richness was better explained by plot factors, such as dead wood or fungi, than by management intensity. These results suggest that the natural variation of dead wood niches (decay stages, snag sizes, tree cavities and wood-inhabiting fungi species) must be maintained to efficiently conserve the whole saproxylic beetle fauna of beech forests. Also, intensive management may alter the specialised saproxylic beetle community even if the initial tree-species composition is maintained, which was the case in our study. For monitoring the ecological sustainability of forest management we must focus on threatened species. If structures alone are sampled then the amount of dead wood is the best indicator for a rich saproxylic beetle fauna.     

Dung beetle assemblages in forests and pastures of El Salvador: a functional comparison

This study examines the functional attributes of Neotropical dung beetles, and, based on the analysis, evaluates the role of dung beetles in pastures and tropical dry forest in El Salvador. Dung beetle diversity was lower in pastures than in forests. However, the total biomass of beetles at similarly-sized dung baits was frequently higher in pastures. Diversity loss followed structured patterns: (1) carrion and fruit feeding beetles were well represented (species number) in the forests but were largely absent from pastures; (2) large ball rollers and small fast tunnellers were present in forests but were absent from pastures; (3) large fast tunnellers and small slow tunnellers were poorly represented in pastures compared to forests but the few species that survived in pastures attained extremely high populations; and (4) each functional group in the species-poor pasture assemblages was divided almost equally into day and night active species. Substrate and habitat generalists that were present in both the forests and pastures were attracted to fruit and carrion in forested habitat but not in open pastures. In open habitats, flies and other insects may be the principal decomposers of decaying fruit and carrion.     

The volatile organic compounds of introduced and native dung and carrion and their role in dung beetle foraging behaviour

1. The decomposition of biological material produces a plethora of volatile organic compounds (VOCs), which are implicated in the foraging behaviour of coprophagous and necrophagous insects. Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) have an acute olfactory system used to locate food resources. Accordingly, identification of food resource VOCs potentially used in food location is integral to understanding dung beetle foraging ecology. 2. In this study, volatile emissions from dung and carrion of native and introduced animals in New Zealand were analysed using solid‐phase microextraction (SPME) and gas chromatography‐mass spectrometry (GC‐MS). Volatile profiles were compared via principal component analyses (PCAs) and cluster solutions based on attractiveness using canonical discriminant analysis (CDA). 3. A total of 115 compounds were detected from 21 food types. Statistical analyses showed that dung and carrion volatile profiles clustered according to attractiveness to the dung beetle Saphobius edwardsi, and that different dung types formed distinct clusters and grouped separately from carrion. 4. This study suggests that volatile profiles emitted by food resources used by dung beetles are complex, producing distinct odours, which potentially mediate foraging decisions.     

The importance of streamside sandbars to ground beetle (Coleoptera, Carabidae) communities in a deciduous forest

We used pitfall traps to sample ground beetles on sandbars along a small woodland stream and in the adjacent floodplain forest (Oglethorpe Co., GA, USA). We captured a total of 1,477 ground beetles representing 41 species. Twenty-two species were exclusive to sandbars, while eight were found only in the forested habitat. Ground beetles were captured in significantly greater numbers from sandbars, especially Brachinus janthinipennis and Omophron americanum. The B. janthinipennis record represents a new state record for the species. This study demonstrates that many unique species can be found in specialized microhabitats and emphasizes the need for biodiversity assessment surveys to include a wide range of these microhabitats within a survey area. In addition, it appears that many generalist forest species might use sandbars seasonally to exploit available resources. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Assessing tillage disturbance on assemblages of ground beetles (Coleoptera: Carabidae) by using a range of ecological indices

Many ecological studies have used diversity indices to assess the impact of environmental disturbance. In particular, ground beetles have been advocated as a good group for assessing disturbance. Most studies on various organisms have used only one or two indices. For our study of the impact of tillage disturbance on carabid beetles in farm fields in southern Ontario, Canada, we used seven different diversity indices (richness, Shannon–Wiener, Berger–Parker, Q-statistic, Margalef, and evenness). Few studies have used deviation from diversity abundance models as a measure of disturbance; however, we use three that are applicable to our data (geometric, log-normal and log-series). The indices and models were used to test the null hypothesis that there is no change in diversity with increasing tillage disturbance, and that there is no difference in diversity with different crops or years. We were not able to reject the null hypothesis that there is any diversity difference among farms. We also found that there was no single diversity index or model that was better than any other at detecting disturbance. These results are supplemented by a meta-analysis of 45 published data sets for the same taxon but in different habitats. The meta-analysis supports the conclusions from our field research that diversity indices and models are not useful for detecting the possible effect of disturbance on assemblages of carabid beetles.     

Mechanical limits to maximum weapon size in a giant rhinoceros beetle

The horns of giant rhinoceros beetles are a classic example of the elaborate morphologies that can result from sexual selection. Theory predicts that sexual traits will evolve to be increasingly exaggerated until survival costs balance the reproductive benefits of further trait elaboration. In Trypoxylus dichotomus, long horns confer a competitive advantage to males, yet previous studies have found that they do not incur survival costs. It is therefore unlikely that horn size is limited by the theoretical cost–benefit equilibrium. However, males sometimes fight vigorously enough to break their horns, so mechanical limits may set an upper bound on horn size. Here, I tested this mechanical limit hypothesis by measuring safety factors across the full range of horn sizes. Safety factors were calculated as the ratio between the force required to break a horn and the maximum force exerted on a horn during a typical fight. I found that safety factors decrease with increasing horn length, indicating that the risk of breakage is indeed highest for the longest horns. Structural failure of oversized horns may therefore oppose the continued exaggeration of horn length driven by male–male competition and set a mechanical limit on the maximum size of rhinoceros beetle horns.     

Diversity of surface dwelling beetle assemblages in open-cast lignite mines in Central Germany

We investigated species richness of ground dwelling beetle assemblages in two non-reclaimed lignite mines and a dump in Central Germany by means of pitfall trapping. During a period of five months, a total of 203 beetle species within 27 families represented by 4099 individuals were trapped. This included 75 species of ground beetles represented in a sample of 957 individuals from which 10 species are regionally endangered. The number of individuals, species richness, as well as the proportions of endangered species did not differ between successional stages whereas species composition of sites could be related well to a set of environmental variables. High values of beta-diversity between sites indicated that the total number of species recorded is caused by habitat diversity. From the viewpoint of nature conservation, we conclude that postmining areas can play a key role in conservation of beetle diversity in agricultural areas since they harbour threatened species whose original habitats are now rare due to human impact. An important task for future management of postmining areas is to maintain successional processes and to prevent loss of habitat diversity through afforestation. Areas with extreme soil conditions should also be preserved for long-term availability of bare soil and pioneer vegetation and associated fauna.     

Environmental effects on sexual size dimorphism of a seed-feeding beetle

Sexual size dimorphism is widespread in animals but varies considerably among species and among populations within species. Much of this variation is assumed to be due to variance in selection on males versus females. However, environmental variables could affect the development of females and males differently, generating variation in dimorphism. Here we use a factorial experimental design to simultaneously examine the effects of rearing host and temperature on sexual dimorphism of the seed beetle, Callosobruchus maculatus. We found that the sexes differed in phenotypic plasticity of body size in response to rearing temperature but not rearing host, creating substantial temperature-induced variation in sexual dimorphism; females were larger than males at all temperatures, but the degree of this dimorphism was smallest at the lowest temperature. This change in dimorphism was due to a gender difference in the effect of temperature on growth rate and not due to sexual differences in plasticity of development time. Furthermore, the sex ratio (proportion males) decreased with decreasing temperature and became female-biased at the lowest temperature. This suggests that the temperature-induced change in dimorphism is potentially due to a change in non-random larval mortality of males versus females. This most important implication of this study is that rearing temperature can generate considerable intraspecific variation in the degree of sexual size dimorphism, though most studies assume that dimorphism varies little within species. Future studies should focus on whether sexual differences in phenotypic plasticity of body size are a consequence of adaptive canalization of one sex against environmental variation in temperature or whether they simply reflect a consequence of non-adaptive developmental differences between males and females.     

Introduced and native ground beetle assemblages (Coleoptera: Carabidae) along a successional gradient in an urban landscape

According to the intermediate disturbance hypothesis (IDH), species diversity should be higher at sites with intermediate levels of disturbance. We tested this hypothesis using ground beetles (Coleoptera: Carabidae) collected in pitfall traps from sites that varied in time since last disturbance. This successional gradient was embedded in an urban landscape near Montreal, Quebec. We predicted that diversity in young forests and old fields would be higher than in agricultural fields and old forests. Fifty-five species (2932 individuals) were found in 2003 and 46 species (2207 individuals) in 2004. In both years, species richness was highest from traps placed in agricultural fields. We collected nine introduced species; these had higher catch rates than the native species in both years (64.8% of total catch). When introduced species were removed from the Nonmetric Multidimensional Scaling ordination analysis, the assemblages from agricultural fields were less distinct compared to those of the other habitats, suggesting the introduced fauna is important in structuring carabid assemblages from the agricultural fields. Introduced species may play a significant role in the community composition of ground beetles in urban landscapes, and their influence may be the cause of the lack of support found for the IDH.     

High diversity beetle assemblages attracted to carrion and dung in threatened tropical oak forests in Southern Mexico

Despite high diversity levels of beetles inhabiting dung and carcasses, very few studies have attempted a comparative assessment of copro-necrophile beetle communities in relation to spatio-temporal variations, particularly in the tropics where the vast majority of beetles occur. We compared beetle assemblages attracted to pads of cattle dung and rat carcasses in four contrasting vegetation types associated with oak forest. In a total of 52 transects including 3,952 dung pad days and 2,600 carcass-trap days we recorded 14,989 beetles representing 406 species and 33 families. Necrophiles (323 species and 33 families) were considerably more diverse than coprophiles (172 species and 16 families). Staphylinidae were taxonomically and numerically dominant, comprising 45% of species and 66% of individuals, respectively. Species estimators (Chao 2) suggested that the observed beetle richness represented 68% of coprophile and 56% of necrophile species richness, with rare species constituting the majority of the species: singletons and doubletons?=?89 species (52%) of coprophiles and 166 species (51%) of necrophiles. Beetle assemblages varied in diversity and composition as regards to vegetation type and season: samples from less disturbed vegetation types (continuous oak forest and ravines) had higher beetle diversity, and a strong seasonality effect was recorded for necrophiles, but not for coprophiles. In addition, an indicator value analysis (IndVal), showed that relatively preserved vegetation types recorded more indicator species as compared to disturbed sites. Our studies clearly demonstrates that the least fragmented oak forest and ravine are the most valuable areas for necrophile and coprophile beetles in Neotropical Mexico, especially for specialist beetles.     

Diversity of dung and carrion beetles in a disturbed Mexican tropical montane cloud forest and on shade coffee plantations

This paper analyzes the diversity of dung and carrion beetles (Scarabaeinae and Silphidae) in four human-induced habitats of a disturbed tropical montane cloud forest: polyspecific shade coffee plantations, monospecific shade coffee plantations, tropical montane cloud forest fragments, and clear cuts. The four habitats had similar richness, species composition, and assemblage structure of dung and carrion beetles. Differences were found in abundance and biomass levels for the four dominant species in the landscape. Dung beetles were more abundant than carrion beetles, but the biomass was higher for the latter. Carrion beetles were seasonal, while dung beetles were clearly not. When forest fragments and shade coffee plantations were compared to other similar habitats in the region, the same general pattern was observed. However, forests with high disturbance and monospecific shade coffee plantations had lower species richness than forests with low and medium disturbance and polyspecific shade coffee plantations. Thus shade coffee plantations maintain connectivity between patches of cloud forest in a landscape that is strongly affected by human activities. Protecting landscape diversity appears to ensure high species richness.     

Dung beetle assemblages in primary forest and disturbed habitats in a tropical dry forest landscape in western Mexico

Dung beetles (Scarabaeinae) are conspicuous components in most terrestrial ecosystems, performing important ecological functions and services. Being sensitive to several types of disturbance, they have been successfully used as indicators of habitat change. Dung beetle communities in tropical rainforests have been well studied, but considerably less information is available for tropical dry forests. In this study I sampled dung beetles in two undisturbed habitats, deciduous forest and semideciduous forest, and two disturbed habitats, secondary forest and open area habitat, in the Chamela-Cuixmala region of western Mexico. Dung beetle species with high indicator value for each habitat were identified. Beetle abundance, observed species richness and estimated species richness were similar in the three forest habitats, but significantly lower in the open area habitat. A more detailed analysis of species-specific abundances in the three forest habitats revealed some differences. Transects of one of the undisturbed habitats, the deciduous forest, were more similar to the non-adjacent transects of disturbed secondary forest, than to the adjacent undisturbed semideciduous forest transects. Unlike studies in other tropical sites that have found a decrease in equitability in Scarabaeinae assemblages between undisturbed forest and disturbed habitat (particularly open habitats), in the Chamela-Cuixmala region all four habitats showed similar low equitability in community structure, with two or three very dominant species.     

Environmental features influencing Carabid beetle (Coleoptera) assemblages along a recently deglaciated area in the Alpine region

Abstract.  1. The spatio-temporal approach was used to evaluate the environmental features influencing carabid beetle assemblages along a chronosequence of an Italian Alpine glacier foreland. The influence of environmental variables on species richness, morphology (wing and body length), and distribution along the chronosequence was tested.
2. Species richness was found to be a poor indicator of habitat due to weak influences by environmental variables. It seems that the neighbouring habitats of a glacier foreland are not able to determine significant changes in carabid species richness.
3. Instead it appears that history (age since deglaciation) and habitat architecture of a glacier foreland are strongly correlated to species adaptive morphological traits, such as wing morphology and body length. Assemblages characterised by species with reduced wing size are linked to the older stages of the chronosequence, where habitat is more structured. Assemblages characterised by the largest species are linked to the younger sites near the glacier. These morphological differentiations are explained in detail.
4. Habitat age can therefore be considered the main force determining assemblage composition. On the basis of the relationship between morphological traits and environmental variables, it seems likely that age since deglaciation is the main variable influencing habitat structure (primary effect) on the Forni foreland. The strong relationship between carabid assemblages and habitat type indicates that site age has but a secondary effect on carabid assemblages. This may be utilised to interpret potential changes in assemblages linked to future glacier retreat.     

Sampling saproxylic beetle assemblages in dead wood logs: comparing window and eclector traps to traditional bark sieving and a refinement

The use of saproxylic beetle community as a metric to evaluate nature conservation measures in forests requires efficient methods. We first compare traditional bark sieving to a potential improvement (extracting beetles from whole bark with Tullgren funnels) to determine the most efficient. Secondly we compare this most efficient bark sampling to eclector and window traps. At the species, family, and functional group levels, we consider species richness, abundance and practical aspects. Traditional bark sieving missed >50% of the individual beetles compared to whole bark sampling so we recommend the latter. Window traps caught large numbers of mobile saproxylic beetles, but a high proportion of non-saproxylics results in high sorting cost; bark sampling and eclector traps had a high proportion of saproxylics and obligate saproxylics. Compared to bark sampling, eclector traps are non-destructive, and monitor the whole saproxylic assemblage (i.e. also beetles inside the wood). Overall, window traps are useful because they capture saproxylic beetles attracted to dead wood and sample the local species pool, whereas eclector traps capture the saproxylics that actually emerge from a particular piece of dead wood, and thus are suited to detailed studies. Overall, we suggest that a combination of these two best methods is highly complementary.     

Local and regional ecological morphology of dung beetle assemblages across four biogeographic regions

Aim Niche partitioning within species assemblages is thought to influence species packing and/or total niche space occupied. The evolution of dung beetles (Scarabaeinae) is likely to have been strongly influenced by inter‐specific competition, leading to niche partitioning. We consider whether local‐scale processes leave a signature in regional patterns of functional diversity in dung beetle assemblages, and investigate the correlation between total exploited ecomorphological space and density of species packing with increased species richness. We test whether ecomorphological space occupied by local assemblages reflects that of their regional species pool, and the extent to which ecomorphological space is convergent or divergent within functional groups across regional pools. Location Neotropics, Africa, Australia and Madagascar. Methods Dung beetle assemblages were collected in a standardized manner from four biogeographic regions. Ecomorphological similarity among the assemblages was assessed by multivariate analysis of 19 linear measurements for 300 species and three functional nesting types (roller, tunneller or dweller), firstly on a local level within the Neotropics and Afrotropics, and then between the regional species pools. Results Key body measurements, in particular the hind tibia, separated rollers and tunnellers into largely non‐overlapping entities along the first three axes of the shape analysis. Three Neotropical assemblages, which vary widely in species numbers, each harboured a similar amount of morphometric variation, resulting in increasingly dense species packing with greater species richness. Similar findings were obtained in two South African assemblages. Assemblages in the four biogeographic regions showed largely similar distributions of ecomorphological variation, including the separation of rollers and tunnellers, despite their distant phylogenetic relationships. Ecomorphological similarity among regions was particularly high in tunnellers, whilst the rollers exhibited greater regional differentiation. Main conclusions Local assemblages evidently represent the full diversity of functional groups available in the regional pool, even in species‐poor assemblages. There is a strong trend towards convergence in morphology separating tunnellers and rollers in phylogenetically independent lineages. The ecomorphological similarity of regional assemblages suggests that morphological convergence is the result of common selective forces active within the assemblages themselves. This lends support to the widely hypothesized effect of inter‐specific interactions and niche partitioning in determining assemblage composition and lineage evolution in the Scarabaeinae.     

Influence of carrion smell and rebaiting time on the efficiency of pitfall traps to dung beetle sampling

Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) are very useful insects, as they improve the chemo-physical properties of soil, clean pastures from dung pads, and help control symbovine flies associated with bovine cattle. Their importance makes it fundamental to sample and survey them adequately. The objectives of the present study were to determine the influence of decaying insects trapped in pitfalls on the attractiveness of Moura pig Sus scrofa L. (Suidae) and collared peccary Tayassu tajacu (L.) (Tayassuidae) dung used as baits to lure dung beetles, and to establish how long these baits remain attractive to dung beetles when used in these traps. Some dung beetle species seemed to be able to discriminate against foul smell from decaying insects within the first 24 h, hence decreasing trap efficiency. This was more evident in peccary dung-baited traps, which proved to be the least attractive bait. Attractiveness lasted only 24 h for peccary dung, after which it became unattractive, whereas the pig dung bait was highly attractive for 48 h, after which its attractiveness diminished but was not completely lost.     

Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Mechanisms and fitness effects of antibacterial defences in a carrion beetle

Parents of many species care for their offspring by protecting them from a wide range of environmental hazards, including desiccation, food shortages, predators, competitors, and parasites and pathogens. Currently, little is known about the mechanisms and fitness consequences of parental defences against bacterial pathogens and competitors. Here, we combine approaches from microbiology and behavioural ecology to investigate the role and mechanistic basis of antibacterial secretions applied to carcasses by parents of the burying beetle Nicrophorus vespilloides. This species rears its larvae on vertebrate carcasses, where larvae suffer significant fitness costs due to competition with bacterial decomposers. We first confirm that anal secretions produced by parents are potently bactericidal and that their effects are specific to gram-positive bacteria. Next, we identify the source of bacterial killing as a secreted lysozyme and show that its concentration changes throughout the breeding cycle. Finally, we show that secreted lysozyme is crucial for larval development, increasing survival by nearly two-fold compared to offspring reared in its absence. These results demonstrate for the first time that anal secretions applied to carrion is a form of parental care and expand the mechanistic repertoire of defences used by parent insects to protect dependent offspring from microbial threats.     

Selection does not favor larger body size at lower temperature in a seed-feeding beetle

Body size of many animals increases with increasing latitude, a phenomenon known as Bergmann's rule (Bergmann clines). Latitudinal gradients in mean temperature are frequently assumed to be the underlying cause of this pattern because temperature covaries systematically with latitude, but whether and how temperature mediates selection on body size is unclear. To test the hypothesis that the "relative" advantage of being larger is greatest at cooler temperatures we compare the fitness of replicate lines of the seed beetle, Stator limbatus, for which body size was manipulated via artificial selection ("Large,"Control," and "Small" lines), when raised at low (22 degrees C) and high (34 degrees C) temperatures. Large-bodied beetles (Large lines) took the longest to develop but had the highest lifetime fecundity, and highest fitness (r(C)), at both low and high temperatures. However, the relative difference between the Large and Small lines did not change with temperature (replicate 2) or was greatest at high temperature (replicate 1), contrary to the prediction that the fitness advantage of being large relative to being small will decline with increasing temperature. Our results are consistent with two previous studies of this seed beetle, but inconsistent with prior studies that suggest that temperature-mediated selection on body size is a major contributor to the production of Bergmann clines. We conclude that other environmental and ecological variables that covary with latitude are more likely to produce the gradient in natural selection responsible for generating Bergmann clines.