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In contrast to other necromass, such as leaves, deadwood, or dung, the drivers of insect biodiversity on carcasses are still incompletely understood. For vertebrate scavengers, a richer community was shown for randomly placed carcasses, due to lower competition. Here we tested if scavenging beetles similarly show a higher diversity at randomly placed carcasses compared to easily manageable fixed places. We sampled 12,879 individuals and 92 species of scavenging beetles attracted to 17 randomly and 12 at fixed places exposed and decomposing carcasses of red deer, roe deer, and red foxes compared to control sites in a low range mountain forest. We used rarefaction-extrapolation curves along the Hill-series to weight diversity from rare to dominant species and indicator species analysis to identify differences between placement types, the decay stage, and carrion species. Beetle diversity decreased from fixed to random locations, becoming increasingly pronounced with weighting of dominant species. In addition, we found only two indicator species for exposure location type, both representative of fixed placement locations and both red listed species, namely Omosita depressa and Necrobia violacea. Furthermore, we identified three indicator species of Staphylinidae (Philonthus marginatus and Oxytelus laqueatus) and Scarabaeidae (Melinopterus prodromus) for larger carrion and one geotrupid species Anoplotrupes stercorosus for advanced decomposition stages. Our study shows that necrophilous insect diversity patterns on carcasses over decomposition follow different mechanisms than those of vertebrate scavengers with permanently established carrion islands as important habitats for a diverse and threatened insect fauna.

Rove beetles (Coleoptera: Staphylinidae) are one of the most important colonizers of vertebrate cadavers. We have previously investigated carrion-associated rove beetle communities across various forests and demonstrated that the study regions are the main drivers of the local rove beetle species pool that is, in turn, able to colonize a carcass. Nevertheless, little is known about their temporal variation in community composition during the decomposition process. The aim of our study has been to analyze the temporal changes of the composition of the rove beetle community and to identify new, potentially useful candidate species for forensic entomological evaluations. We determined 80 rove beetle species that were attracted to 60 piglet cadavers across various forest stands in Germany. Both the abundance and the community composition changed over the decomposition process, independently of the species-specific geographical variation across study regions. In the region Schorfheide-Chorin, species from the genus Philonthus proved to be a suitable group for future forensic entomological investigations. They appeared in markedly high numbers at piglet cadavers from the bloated stage until the advanced decay stage. For the study region Schwäbische Alb, we newly describe the species Omalium septentrionis as a member of the carrion-associated rove beetle fauna. Based on the geographical variation in rove beetle community compositions, we have filtered out several species that might contribute to advances in postmortem interval estimations or other applied fields in forensic entomological contexts. [Display omitted] •Rove beetles (Staphylinidae) are underexplored in a forensic entomological context.•Aim to test temporal change over decay and to identify species useful for forensics.•80 rove beetle species were determined on 60 piglet cadavers in various forests.•Genus Philonthus was a post-bloating indicator group for small vertebrate carrion.•Omalium septentrionis was a new member of the carrion-associated rove beetle fauna.

Carrion of wild ungulates is an important resource for microbes, invertebrates and vertebrates, and affects local plant communities and soils. Most wild ungulates in central Europe, however, are intensely hunted and removed from the ecosystem, thus not available as carrion. This study explores the use of evisceration residues as a resource by necrophagous insects in a temperate mountainous forest in Germany. We experimentally compared the relative abundance of necrophagous insects between sites of caged roe deer carcasses and evisceration residues of roe deer using pitfall‐traps. While evisceration residues were used by necrophagous insects, significantly more individuals were sampled at entire deer carcasses; when corrected for biomass, however, no significant difference in abundance for Coleoptera groups was found. Overall, evisceration residues were used earlier and for a shorter period than entire deer carcasses. Leaving evisceration residues on site where game animals are obtained might help maintain specialized necrophagous insect communities, although their ecological role cannot completely replace that of whole carcasses of wild ruminants.

Anthropogenic land use causes global declines in biodiversity. Despite the knowledge that animal carrion is the most nutrient-rich form of dead organic matter, studies on landscape and local scales determining whether and the means by which land use intensity influences the diversity of the carrion-associated insect fauna are globally scarce. We investigated the effects of land use intensity and abiotic and biotic environmental factors on the abundance, species richness, and diversity of the important ecosystem-service-providing silphid beetle taxon (carrion beetles) in three regions of Germany. In 61 forest stands distributed over three geographically distinct regions in Central Europe, we trapped silphid beetles on exposed piglet cadavers during late summer. In all three regions, higher ambient temperatures and higher fine sand contents were associated with the abundance of the silphid beetle taxa. The carrion community silphid diversity was negatively affected by an increase in mean ambient temperature in all three regions. Although management intensity in forests did not affect the overall abundance of Silphidae, the abundance of Nicrophorus humator decreased significantly with higher forest management intensity across all three regions. Unmanaged and age-class forests showed a higher abundance of N. humator compared with extensively managed forest stands. These findings indicate that N. humator has potential as an indicator species for anthropogenic disturbances in forests. Overall, the direct responses of the silphid beetle community to diverse soil characteristics underline soil as an important factor determining the abundance and diversity of necrophagous carrion beetles in Central Europe. To protect these valuable ecosystem-service providers, forest-management-induced soil modifications need to be paid close attention.

Carrion plays an essential role in shaping the structure and functioning of ecosystems and has far‐reaching implications for biodiversity conservation. The change in availability and type of carcasses throughout ecosystems can involve negative effects for scavenging communities. To address this issue, there have been recent conservation management measures of carrion provision in natural systems. However, the optimal conditions under which exposing carcasses to optimize conservation outcomes are still limited. Here, we used camera traps throughout elevational and vegetational gradients to monitor the consumption of 48 deer carcasses over a study period of six years by evaluating 270,279 photographs resulting out of 15,373 trap nights. We detected 17 species visiting carcass deployments, including five endangered species. Our results show that large carcasses, the winter season, and a heterogeneous surrounding habitat enhanced the frequency of carcass visits and the species richness of scavenger assemblages. Contrary to our expectations, carcass species, condition (fresh/frozen), and provision schedule (continuous vs single exposure) did not influence scavenging frequency or diversity. The carcass visitation frequency increased with carcass mass and lower temperatures. The effect of large carcasses was especially pronounced for mesopredators and the Eurasian lynx (Lynx lynx). Lynx were not too influenced in its carrion acquisition by the season, but exclusively preferred remote habitats containing higher forest cover. Birds of prey, mesopredators, and top predators were also positively influenced by the visiting rate of ravens (Corvus corax), whereas no biotic or abiotic preferences were found for wild boars (Sus scrofa). This study provides evidence that any ungulate species of carrion, either in a fresh or in previously frozen condition, attracts a high diversity of scavengers especially during winter, thereby supporting earlier work that carcass provisions may support scavenger communities and endangered species.

Infectious diseases in wildlife threaten not only those species but also domestic animals and human health, necessitating strategies to prevent pathogen spread. The natural decomposition of carcasses may lead to pathogen inactivation due to associated increases in temperature and changes in the pH of the carcass and in the surrounding soil. In this study, the internal temperatures of 64 decomposing wild boar carcasses, the pH in the topsoil beneath 74 carcasses, and the pH of muscle and rectal tissue from 12 carcasses were monitored throughout the decomposition process. Carcass temperatures increased during decomposition, frequently exceeding 30 °C during aerobic decomposition in summer (maximum 58 °C). The pH in the carcasses increased until skeletonization, ranging from pH 4 to above pH 8. Soil pH also continuously increased during the decomposition, reaching a pH above 9 and remaining stable for at least 30 days post-skeletonization. The information on natural carcass decomposition processes provided by our study can serve as a basis for future studies to assess if elevated carcass temperatures and pH changes are sufficient for pathogen inactivation. However, our results suggest that, in most cases, neither the increase in carcass temperatures nor the changes in pH exceed the thresholds required to inactivate African swine fever virus.

During decomposition, vertebrate carrion emits volatile organic compounds to which insects and other scavengers are attracted. We have previously found that the dung beetle, Anoplotrupes stercorosus, is the most common dung beetle found on vertebrate cadavers. Our aim in this study was to identify volatile key compounds emitted from carrion and used by A. stercorosus to locate this nutritive resource. By collecting cadaveric volatiles and performing electroantennographic detection, we tested which compounds A. stercorosus perceived in the post-bloating decomposition stage. Receptors in the antennae of A. stercorosus responded to 24 volatiles in odor bouquets from post-bloating decay. Subsequently, we produced a synthetic cadaver odor bouquet consisting of six compounds (benzaldehyde, DMTS, 3-octanone, 6-methyl-5-hepten-2-ol, nonanal, dodecane) perceived by the beetles and used various blends to attract A. stercorosus in German forests. In field assays, these beetles were attracted to a blend of DMTS, 3-octanone, and benzaldehyde. Generalist feeding behavior might lead to the super-dominant occurrence of A. stercorosus in temperate European forests and have a potentially large impact on the exploitation and rapid turnover of temporally limited resources such as vertebrate cadavers.

Because animal carcasses often serve as reservoirs for pathogens, their location and removal are crucial in controlling the spread of diseases. During carcass decomposition, heat is emitted due to microbial activity and the development of maggots. Recent studies have shown that infrared sensors can be used to locate animal carcasses, but little is known about the factors influencing detection success. In this study, we investigated the potential of infrared technology to locate wild boar carcasses, as they play an important role in the spread of African swine fever. Specifically, we tested the effects of environmental and carcass conditions on the detection probability. A drone-based thermal camera was used to collect data during 379 flyovers of 42 wild boar carcasses in different stages of decomposition between September 2020 and July 2021. Generalized mixed-effect models and conditional inference trees were used to identify the environmental and carcass conditions that influenced the detection probability. Our results showed that the thermal camera accurately measured carcass temperature (R2 = 0.75, RMSE = 5.89°C). The probability of finding carcasses was higher in open habitats with air temperatures >3.0°C and thus conducive to maggot development (detection rate ≤80%). A forest canopy openness >29.3% and cloudy conditions or flights at dawn increased the detection rate. Moreover, carcasses infested with large amounts of maggots could be detected even in habitats with a more extensive canopy cover, whereas in dense forests, the detection probability was limited (<25%). Carcasses in an advanced stage of decomposition could still be detected as long as the difference between the carcass temperature and the air temperature was >6.4°C (≤62%). Our study demonstrates the utility of thermal imaging in searching for wild boar carcasses under specific environmental and carcass conditions and thus its use in supporting ground searches.

Abstract There is growing evidence that a wide range of insect sex pheromones are condition dependent and play a fundamental role in mate choice. However, the effectiveness of pheromonal communication might not only depend on internal factors of the sender, but also on attributes of the microhabitat, in which the signaler chooses to emit its chemical signal. For example, the degree of anthropogenic land use might affect how successful the signal is transmitted, as land use has been shown to affect animal communities and the complexity of biotic interactions. To test the hypothesis that parameters of the microenvironment determine males’ ability to attract females via their sex pheromone, we used the burying beetle Nicrophorus vespilloides Herbst (Coleoptera: Silphidae) as our model system. We exposed 144 males across differently managed forest stands and analyzed the impact of 29 environmental parameters. Our data revealed that human land use intensity had no effect on a male’s attractiveness. However, the harvested tree biomass positively affected the proportion of competitors attracted. Furthermore, we found that soil characteristics were important factors determining the amount and body size of females a male was able to attract. Consequently, we present evidence that the environmental context of a signaling male influences the effectiveness of chemical signaling either because it affects the transmission process or the prevailing abundance of potential signal receivers. Thus, our results demonstrate that males need to make careful decisions about the location where they emit their pheromone, as this choice of microhabitat has an impact on their fitness.

The necrophagous burying beetle Nicrophorus vespilloides reproduces on small carcasses that are buried underground to serve as food for their offspring. Cadavers that are too large to bury have previously been postulated to be important food sources for newly emerged beetles; however, the attractiveness of distinct successive stages of decomposition were not further specified. Therefore, we investigated the potential preference of newly emerged N. vespilloides females for odour bouquets of piglet cadavers at specific stages of decomposition. Analyses of walking tracks on a Kramer sphere revealed a significantly higher mean walking speed and, consequently, a higher mean total track length when beetles were confronted with odour plumes of the decomposition stages 'post-bloating', 'advanced decay' or 'dry remains' in comparison with the solvent control. Such a change of the walking speed of newly emerged N. vespilloides females indicates a higher motivation to locate such food sources. In contrast to less discriminating individuals this behaviour provides the advantage of not wasting time at unsuitable food sources. Furthermore, in the advanced decay stage, we registered a significantly higher preference of beetles for upwind directions to its specific odour plume when compared with the solvent control. Such a change to upwind walking behaviour increases the likelihood that a large cadaver will be quickly located. Our findings are of general importance for applied forensic entomology: newly emerged N. vespilloides females on large cadavers can and should be regarded as potential indicators of prolonged post mortem intervals as our results clearly show that they prefer emitted odour bouquets of later decomposition stages.