Silphidae Research Articles

Ecological drivers of carrion beetle (Staphylinidae: Silphinae) diversity on small to large mammals.

Silphinae (Staphylinidae; carrion beetles) are important contributors to the efficient decomposition and recycling of carrion necromass. Their community composition is important for the provision of this ecosystem function and can be affected by abiotic and biotic factors. However, investigations are lacking on the effects of carrion characteristics on Silphinae diversity. Carrion body mass may affect Silphinae diversity following the more individuals hypothesis (MIH). The MIH predicts a higher number of species at larger carrion because higher numbers of individuals can be supported on the resource patch. Additionally, biotic factors like carrion species identity or decomposition stage, and the abiotic factors elevation, season and temperature could affect Silphinae diversity. To test the hypotheses, we collected Silphinae throughout the decomposition of 100 carcasses representing 10 mammal species ranging from 0.04 to 124 kg. Experimental carcasses were exposed in a mountain forest landscape in Germany during spring and summer of 2021. We analysed Silphinae diversity using recently developed transformation models that considered the difficult data distribution we obtained. We found no consistent effect of carrion body mass on Silphinae species richness and, therefore, rejected the MIH. Carrion decomposition stage, in contrast, strongly influenced Silphinae diversity. Abundance and species richness increased with the decomposition process. Silphinae abundance increased with temperature and decreased with elevation. Furthermore, Silphinae abundance was lower in summer compared to spring, likely due to increased co-occurrence and competition with dipteran larvae in summer. Neither carrion species identity nor any abiotic factor affected Silphinae species richness following a pattern consistent throughout the seasons. Our approach combining a broad study design with an improved method for data analysis, transformation models, revealed new insights into mechanisms driving carrion beetle diversity during carrion decomposition. Overall, our study illustrates the complexity and multifactorial nature of biotic and abiotic factors affecting diversity.

Intraspecific trait variation of carrion beetle species and communities across elevations

Abstract Filtering processes across environmental gradients can structure patterns of trait variation within communities. The community‐weighted mean (CWM) is a metric that is commonly used to indicate the directionality of such filtering processes and the optimal adaptive strategy of taxa within community. Proximity to the CWM indicates higher fitness, and deviations from this optimal value result in changes in the relative abundances of coexisting species. We investigated patterns of intraspecific trait variation in four coexisting carrion beetle species (Coleoptera: Silphidae) across elevational gradients. The study was conducted in temperate forest ecosystems with distinct natural vegetation zones ranging from 950 m to 1700 m above sea level. Of the 12 traits measured, we found that intraspecific variation ranged from 34% (body length) to 100% (ratio of elytra length, head length and head width to body length) and accounted for a larger proportion of variation than interspecific variation in 7 traits. For most traits, trait range, which indicates the niche breadth of species at a given elevation, was positively correlated with relative abundance. The CWMs of traits associated with long‐distance dispersal decreased with elevation, whereas those associated with microhabitat use showed the opposite trend. Soil temperature influenced tibia length after controlling for the effects of species identity, but soil water content had no effect on trait variation. Patterns of variation in body width and thorax width of two beetle species supported the CWM‐optimality hypothesis; however, patterns of trait variation in body width and thorax length of another species showed an opposite trend. Our study identifies several traits that can be highly variable within carrion beetle species. Such high levels of intraspecific trait variation may enable populations to adapt across a wide range of elevations and vegetation types.

Large carrion and burying beetles evolved from Staphylinidae (Coleoptera, Staphylinidae, Silphinae): a review of the evidence.

Large carrion beetles (Silphidae) are the focus of ongoing behavioral ecology, forensic, ecological, conservation, evolutionary, systematic, and other research, and were recently reclassified as a subfamily of Staphylinidae. Twenty-three analyses in 21 publications spanning the years 1927-2023 that are relevant to the question of the evolutionary origin and taxonomic classification of Silphidae are reviewed. Most of these analyses (20) found Silphidae nested inside Staphylinidae (an average of 4.38 branches deep), two found Silphidae in an ambiguous position, and one found Silphidae outside Staphylinidae, as sister to Hydrophilidae. There is strong evidence supporting the hypothesis that large carrion beetles evolved from within Staphylinidae and good justification for their classification as the subfamily Silphinae of the megadiverse, and apparently now monophyletic, Staphylinidae. Considerable uncertainty remains regarding the interrelationships and monophyly of many staphylinid subfamilies. Nonetheless, the subfamily Tachyporinae was found to be the sister of Silphinae in more analyses (7) than any other subfamily.

Detection of extra‐pair maternity in a carrion beetle under natural conditions

Abstract The provision of care to offspring is a costly endeavour that can be hijacked by others—known as brood parasitism. Females can choose to lay eggs in the nest of a conspecific female, resulting in extra‐pair maternity (EPM). Burying beetles of the genus Nicrophorus use small vertebrate carcasses for reproduction and provide biparental care to their offspring. Carcasses are a rare resource; thus, competition often occurs among adults of the same sex. A previous study examined the outcomes of competition and subsequent parental care of Nicrophorus vespilloides using laboratory experiments. Larvae of losing females were sometimes cared for by the winning females, which is generally an evidence of EPM. However, the presence of EPM in burying beetles under natural conditions has not yet been well established. Here, we focused on the reproduction of N. quadripunctatus in the field. To examine the presence of EPM, maternity analysis was performed on 149 offspring and their caring females in 10 broods, using 8 microsatellite DNA loci. Seven larvae originating from three broods were unrelated to their caring females. There were one to three unrelated larvae in each of the three broods. Thus, EPM occurs in N. quadripunctatus reproduction under natural conditions.

Competition, cooperation, and parental effects in larval aggregations formed on carrion by communally breeding beetles Necrodes littoralis (Staphylinidae: Silphinae).

Aggregations of juveniles are dominant forms of social life in some insect groups. Larval societies are shaped by competitive and cooperative interactions of the larvae, in parallel with parental effects. Colonies of necrophagous larvae are excellent systems to study these relationships. Necrodes littoralis (Staphylinidae: Silphinae), a carrion beetle that colonizes cadavers of large vertebrates, forms massive juvenile aggregations. By spreading over carrion anal and oral exudates, the beetles form the feeding matrix, in which the heat is produced and by which adults presumably affect the fitness of the larvae. We predict that exploitative competition shapes the behavior of N. littoralis larvae in their aggregations. However, cooperative interactions may also operate in these systems due mainly to the benefits of collective exodigestion. Moreover, indirect parental effects (i.e., formation of the feeding matrix) probably modulate larval interactions within the aggregations. By manipulating parental effects (present/absent) and larval density (0.02-1.9 larvae/g of meat), we found a strong negative group-size effect on fitness components of N. littoralis, in colonies with parental effects over almost the entire density range, and in colonies without parental effects for densities larger than 0.5 larva/g. This was accompanied by positive group-size effects in terms of development time (it shortened with larval density) and thermogenesis (it increased with larval density). A pronounced positive group-size effect on juvenile fitness was found only in colonies without parental effects and only in the low-density range. These results support the hypothesis that larval societies of N. littoralis are shaped by exploitation competition.

Friction-reduction effect of the hierarchical surface microstructure of carrion beetle by controlling the real contact area.

The discovery and elucidation of the surface microstructure functions of living organisms are crucial to resolving issues, such as friction. We newly discovered that Necrophila japonica, a type of carrion beetle that lives on the ground surface, exhibited a hierarchical surface microstructure comprising a submicron-sized wrinkle structure on top of a micron-sized microstructure. The surface microstructure of this beetle improved wettability but did not exhibit superhydrophobicity, a well-known function of hierarchical structures, so it was expected to have a different function. By combining the insights in the field of structural mechanics that avoidance of stress concentration by the structural geometry affects deformation with the basic principles of friction, the frictional properties and mechanisms of the hierarchical surface microstructure of carrion beetle were investigated. The measurements of frictional force indicated that the mimicked structure exerted lower frictional forces than flat and single-layer microstructure surfaces. Analysis of finite element method simulations showed that even though the mimicked structure was prone to pressure concentration due to small contact points, the surface contact pressure was reduced more than that of the single-layer structure by hierarchical load dispersion like that of metamaterials. As a result, the suppression of the increase in the real contact area due to deformation suppression contributed to effective friction-reduction. The effective friction-reduction by the hierarchical structure provides not only new insight into the surface microstructure function of various organisms, but also new design for the lubricant-free friction reduction that has been the focus of attention in carbon neutrality and other fields.

The effect of pioneer carrion beetles on the emission of volatile organic compounds and carrion insect community assembly.

Mechanisms of carrion insect succession have been interpreted separately from interspecific interactions between early and later colonists or from changes in volatile organic compounds perceived by insects resulting from the progression of decomposition. To link these perspectives, we examined through laboratory and field experiments whether the modification of volatile organic compounds by early colonists could be a mechanism of succession. In the laboratory experiment, we used Necrophila japonica (Coleoptera, Staphylinidae) as an early colonist and examined its effects on the emissions of important volatile attractants for carrion insects, dimethyl disulfide (DMDS) and dimethyl trisulfide (DMTS) from carcasses. We collected DMDS and DMTS, using the static headspace method, under the following conditions: (i) rat carcass, (ii) rat carcass with artificial damage to the abdomen, (iii) rat carcass fed on by 10 Ne. japonica individuals, and (iv) 10 Ne. japonica individuals, and analyzed the collected gases using a gas chromatograph-mass spectrometer. After 12 and 30 h, carcasses fed on by Ne. japonica emitted higher concentrations of DMDS and DMTS than in other conditions. In the field experiment, we examined the effects of DMDS and DMTS on the attraction of carrion insects using traps baited with a mixture of DMDS and DMTS, hexane (odors unrelated to carcasses), or an empty microtube. Traps baited with DMDS and DMTS attracted more necrophagous species and individuals than traps not baited with this combination. These results showed that accelerated emissions of DMDS and DMTS from carcasses caused by early colonists may contribute to community assembly during carrion insect succession.

Culture-independent analyses of carrion beetle (Coleoptera: Silphidae) secretion bacterial communities.

In the central plains of North America, the beetle family Silphidae comprised two subfamilies, Silphinae and Nicrophorinae, differentiated by reproductive behaviors. Silphinae, known as carrion beetles, feed on carrion and fly larvae and produce free-living larvae that receive no parental care. Adult Nicrophorinae, known as burying beetles, prepare a vertebrate carcass into a brood ball and provide biparental care to their offspring. Preparation of a brood ball involves coating the carcass in antimicrobial oral and anal secretions. These secretions contain a community of microbes, referred to as the secretion microbiome, which inhibit carcass microbe succession, preventing normal decomposition. Here, the secretion microbiomes of five species of Nicrophorinae and two species of Silphinae, both sampled from Oklahoma, with additional Nicrophorinae from Nebraska, were characterized using culture-independent analyses to understand and decipher factors shaping diversity and community structure. We identify the core secretion microbiome across Silphidae and show that, while the host subfamily, secretion type, and collection locality had no significant effect on the bacterial community alpha diversity, these factors significantly influenced bacterial community structure. Global and local tests of phylogenetic associations identified 14 genera with phylogenetic signals to the host subfamily and species. Description of the bacterial communities present in silphid secretions furthers our understanding of how these beetles interact with microbes for carcass nutrient processing. Future culture-dependent studies from silphid secretions may identify novel antimicrobials and nontoxic compounds that can act as meat preservatives or sources for antimicrobials.IMPORTANCEThe manuscript explores the secretion bacterial community of carrion and burying beetles of the central plains of North America. A core secretion microbiome of 11 genera is identified. The host subfamily, secretion type, and collection locality significantly affects the secretion microbiome. Future culture-dependent studies from silphid secretions may identify novel antimicrobials and nontoxic compounds that can act as meat preservatives or sources for antimicrobials.

Bioaccumulation of contaminants in Scarabaeidae and Silphidae beetles at sites polluted by coal combustion residuals and radiocesium

Anthropogenic contamination from coal-fired power plants and nuclear reactors is a pervasive issue impacting ecosystems across the globe. As a result, it is critical that studies continue to assess the accumulation and effects of trace elements and radionuclides in a diversity of biota. In particular, bioindicator species are a powerful tool for risk assessment of chemically contaminated habitats. Using inductively coupled plasma mass spectrometry (ICP-MS) and auto-gamma counting, we analyzed trace element and radiocesium contaminant concentrations in Scarabaeidae and Silphidae beetles (Order: Coleoptera), important taxa in decomposition and nutrient cycling, at contaminated and reference sites on the Savannah River Site, South Carolina, U.S. Our results revealed variability in trace element concentrations between Scarabaeidae and Silphidae beetles at uncontaminated and contaminated sites. Compared to Scarabaeidae, Silphidae had higher levels of chromium (Cr), copper (Cu), iron (Fe), manganese (Mn), nickel (Ni), and zinc (Zn). Unexpectedly, concentrations of Cr, Cu, and Ni were higher in both taxa at the uncontaminated sites. Scarabaeidae and Silphidae beetles at the coal combustion waste site consistently had high concentrations of arsenic (As), and Scarabaeidae had high concentrations of selenium (Se). Of the 50 beetles analyzed for radiocesium levels, two had elevated radioactivity concentrations, both of which were from a site contaminated with radionuclides. Our results suggest carrion beetles may be particularly sensitive to bioaccumulation of contaminants due to their trophic position and role in decomposition, and thus are useful sentinels of trace element and radionuclide contamination.

Calibrating insect age at eclosion by size in a gregarious carrion beetle Thanatophilus sinuatus (Staphylinidae: Silphinae).

Recent discoveries have shown that the physiological age at eclosion of forensically useful beetles differs between males and females and between beetles of various sizes. Accordingly, it was postulated that the size and sex of the beetles at eclosion may be used to calibrate their age, which may improve the accuracy of age (and post-mortem interval) estimates in forensic entomology. In this study, we derived thermal summation models for the eclosion for the Central European population of carrion beetles Thanatophilus sinuatus (Fabricius, 1775), (Staphylinidae: Silphinae), and tested the usefulness of sex and size for the calibration of beetle age at eclosion. Although in previous developmental studies, the beetles were reared individually, we reared them in larval aggregations, since in natural conditions T. sinuatus beetles are gregarious. Weak (r2 between 5% and 13%) negative correlations were observed between the size and age of T. sinuatus males or females at eclosion, demonstrating that calibration of age by beetle size and sex may bring only minimal benefits regarding the accuracy of age estimation in this species. However, it may still be worthwhile in the case of extremely large or small beetles. Moreover, the total development times recorded in this study were much shorter than in the previous T. sinuatus study, at 14°C by about 15 days and at 26°C by about 2 days. These differences emphasise the importance of gregariousness for the development of carrion beetles, and at the same time highlight the need for the ecologically-relevant protocols of development studies in forensic entomology.

A beetle for prosecution: exogenous DNA detection from larval and adult gut of a Neotropical carrion beetle.

The use of recovered DNA ingested by necrophagous or hematophagous insects has increased in forensic sciences, mainly with representatives of flies. However, some beetles are also important for medico-legal forensic entomology because they feed on carcasses until advanced decomposition. This study evaluated whether the Neotropical carrion beetle Oxelytrum discicolle (Silphidae) has the potential for the detection of exogenous DNA into the gut. The whole gut or the gut contents were extracted from O. discicolle larvae and adult previously fed on pig carcass. The pig DNA recovery rate was 33.3% in larvae and 25% in adults, indicating that the carrion beetle's gut may be useful for DNA identification of ingested food. Samples with the whole gut or only gut contents showed the same DNA recovery rate. Exogenous DNA from the whole gut was recovered from samples stored in ethanol at -20 ºC for 11 days, showing that samples of O. discicolle can be stored in the forensic laboratory without loss in DNA recovery rate.

Substantial niche overlap in carrion beetle habitat and vegetation use

Abstract The habitat affinities of carrion beetles (Coleoptera: Silphidae), a speciose group with wide cooccurrence, are only coarsely described for well‐studied species, particularly in the western United States. We aim to identify if the 15 species of montane carrion beetle in Colorado differ substantially in their use of habitats and across more uniquely defined, fine‐scale vegetation characteristics. Habitat and vegetation data as well as carrion beetle abundance were collected along four elevational transects in the Front Range and San Juan Mountains from 2010 to 2012 across 32 sites. Multiple habitat types were sampled, including forest, meadow, riparian, rocky outcrops and tundra. The fine‐scale vegetation characteristics included percent coverage of ground vegetation (grass, forb, shrub, cacti, bare ground), understory vegetation biomass and height, canopy cover and tree species, number and size. Canonical correspondence analysis models examined whether vegetation characteristics strongly segregated species using abundances and presence‐absences. Habitat and vegetation models explained a maximum of 18.93% of the variation in species' abundances and 2.48% in species' presence–absences. Only one likely habitat specialist was identified by the models (Heterosilpha ramosa) and the remaining species had substantial overlap in habitat and vegetation use. The arid, low productivity and generally open understory habitats in Colorado mountains likely play a large role in the substantial vegetation overlap. Other mechanisms of partitioning likely exist in this system to reduce niche overlap, which could include differences in activity time and seasonality, physiological traits, other life history strategies and body size.

Evidence of ancestral nocturnality, locomotor clock regression, and cave zone-adjusted sleep duration modes in a cave beetle

The small carrion beetle Ptomaphagus hirtus is an abundant inhabitant of the exceptionally biodiverse Mammoth Cave system. Previous studies revealed negative phototaxis and the expression of biological clock genes in this microphthalmic cave beetle. Here we present results from probing P. hirtus for the entrainment of locomotor rhythms using the TriKinetics activity monitor setup. Although curtailed by low adjustment frequency of animals to the test environment, the data obtained from successfully monitoring two animals in constant darkness (DD) and six animals exposed to 12 hour light-dark cycles (LD) revealed a strong effect of light on locomotor activity in P. hirtus. In LD, activity was prevalent during the artificial night phases while close to absent during the presumptive day phases, suggesting conserved nocturnality. Upon transitioning LD animals to constant darkness, none displayed detectable evidence of free-running activity rhythms, suggesting complete regression of the central circadian clock. Equally notable, overall locomotor activity of the two DD-monitored animals was about three-fold lower compared to LD animals due to longer rest durations in the former. We, therefore, propose the existence of cave zone-specific energy expenditure modes that are mediated through light schedule responsive modification of sleep duration in P. hirtus.

The Impact of Diet and Photoperiodism on the Life History of Thanatophilus sinuatus (Coleoptera: Silphidae).

The forensically important genus Thanatophilus Leach, 1815 (Coleoptera: Silphidae) is a widespread group of carrion beetles that occupies the Holarctic and Afrotropical realms. It has recently received more recognition, as its species are frequently detected on large carrion, including humans, and can be useful in estimating the time of colonization (TOC) and the post-mortem interval (PMI). The immature stages of Thanatophilus sinuatus (Fabricius, 1775) were reared on three different meat diets (pork liver, pork muscle, and chicken muscle). Diet influence on some of the essential life history traits of the species was examined for different populations. Our findings indicated an accelerated development on pork muscle followed closely by pork liver and a prolonged development on chicken muscle. The diet significantly affected the size, and the body condition (weight/ size) as both pork muscle and liver produced individuals in better physical condition (larger and heavier). The impact of two cyclic photoperiod regimes (16 h Light (L): 8 h Dark (D) and 12 h L: 12 h D) was further evaluated and indicated no significant variability.

When the red-lined carrion beetle disrupts successional dynamics on large vertebrate carcasses

In a field study of insect colonization on domestic pig carcasses, the typical colonization sequence took a different turn midway through decomposition when Necrodes surinamensis (Fabricius) (Coleoptera: Silphidae) dramatically increased in abundance on some carcasses. This provided an opportunity to test the prediction that N. surinamensis would aggregate at clumped carcasses but not as much at dispersed carcasses, thus leaving enough resources in less heavily colonized habitat patches to support weaker competitors, as predicted by the aggregation model of coexistence. Larger aggregations of this silphine beetle resulted in the collapse of blow fly (Diptera: Calliphoridae) maggot populations due to intraguild predation, and a decline or exclusion of late-successional insect species. This phenomenon has been reported in a European Necrodes species but it is the first time it is documented with N. surinamensis, a widespread species in North America. Substantial increases in adult and larval populations of N. surinamensis were first documented on clumped carcasses and were not as pronounced on dispersed carcasses. The specifics of N. surinamensis aggregations are discussed with reference to ecological and forensic implications, including the fact that feeding by N. surinamensis larvae makes the remains appear to have been altered by scavengers and decomposing for longer than they have.

Identification of Forensically Important Carrion Beetles (Coleoptera: Staphilinidae) in China Based on COI and COII.

Unambiguous and speedy necrophagous insect species identification is common task in forensic entomological study. Carrion beetles (Staphilinidae: Silphinae) belong to a small group of Coleoptera with less than 200 species worldwide. Some species are commonly found on dead body during forensic entomological investigation. Despite some species are hard to be categorized morphologically, present DNA-based technologies offer a potential identification strategy. Here, 37 carrion beetle specimens were collected from 15 locations throughout Chinese mainland. The cytochrome oxidase subunit I (COI) and II (COII) genes among all specimens were successfully sequenced, which provided reliable markers for precise identification. Mostly, the interspecific distance could clarify the capability of these genes for identifying included carrion beetle species. Exceptions existed between close species in Nicrophorus genus (Fabricius 1775). The sequenced gene's phylogenetic analysis revealed that all carrion beetle specimens were correctly classified into eight genera, and most have relatively high supporting values (>90%). Our data gives genetic diversity and a reference for global forensically important carrion beetle species identification, as well as a conductive significance for future application of Chinese carrion beetles in forensic entomology.

Larval and adult digestive tract of the carrion beetle Oxelytrum discicolle (Brullé, 1840) (Coleoptera: Silphidae)

Oxelytrum discicolle is a necrophagous beetle occurring in Central and South America, with potential use in forensic entomology for investigations in the context of legal medicine. The present work aimed to describe the morphology of the digestive tract of larvae and adults and contributes to the knowledge about the digestion associated with necrophagy. In the three larval instars, the foregut is short and narrow, the midgut is a dilated tube, elongated with a smooth surface, and the hindgut is narrow and long, with small lateral projections and a dilated terminal region. The gut epithelium in the second and third instar larvae is vacuolated in the mid- and hindgut, with high production of apocrine secretions in the midgut lumen. In adults, the foregut is short, with small spines in the cuticular intima that covers the flattened epithelium. The midgut is dilated, with many short gastric caeca with regenerative cells in the blind portion, which differentiate in digestive columnar cells towards the midgut lumen. The hindgut is long and narrow, with an enlarged distal portion with folded epithelium lined by a thin cuticle. The histochemical tests reveal the absence of protein storage granules in the gut epithelium of larvae and adults. The gut characteristics of the carrion beetle O. discicolle are similar to those of other predatory and phytophagous Coleoptera, which may indicate that ancestry may influence the alimentary canal morphology more than the feeding habits.

Gene expression underlying parenting and being parented shows limited plasticity in response to different ambient temperatures.

Flexible interactions between parents and offspring are essential for buffering families against variable, unpredictable, and challenging environmental conditions. In the subsocial carrion beetle, Nicrophorus orbicollis, mid-summer temperatures impose steep fitness costs on parents and offspring but do not elicit behavioural plasticity in parents. Here, we ask if plasticity of gene expression underpins this behavioural stability or facilitates independent compensation by larvae. To test this, we characterized gene expression of parents and offspring before and during active parenting under benign (20°C) and stressful (24°C) temperatures to identify genes of parents and offspring associated with thermal response, parenting/being parented, and gene expression plasticity associated with behavioural stability of parental care. The main effects of thermal and social condition each shaped patterns of gene expression in females, males, and larvae. In addition, we implicated 79 genes in females as "buffering" parental behaviour across environments. The majority of these underwent significant changes in expression in actively parenting mothers at the benign temperature, but not at the stressful temperature. Our results suggest that neither genetic programmes for parenting nor their effects on offspring gene expression are fundamentally different under stressful conditions, and that behavioural stability is associated primarily with the maintenance of existing genetic programmes rather than replacement or supplementation. Thus, while selection for compensatory gene expression could expand the range of thermal conditions parents will tolerate, without expanding the toolkit of genes involved selection is unlikely to lead to adaptive changes of function.

Entomological identification of the post-mortem colonization of wolf cadavers in different decomposition stages

Human-driven biodiversity loss is progressively becoming a problem with dramatic consequences for the conservation of vital ecosystems. The increasing number of illegal killings of the grey wolf (Canis lupus, Linnaeus, 1758), a threatened species, displays the need for investigation and prosecution of such offences. Forensic entomology makes use of the knowledge about necrophagous insects to estimate a minimum time-since-death interval of the deceased person or animal, which can give important information on a possible perpetrator. The cadaver fauna along five decomposition stages of wolves in Germany was investigated in the period 2014–2021. The insects from 70 wolf cadavers, originating from all over Germany, were provided by the Leibniz Institute for Zoo and Wildlife Research Berlin. The accumulated degree day (ADD) model was applied for the post-mortem interval estimation on wolf cadavers for the first time. A total of 20 coleopteran species and 14 different dipteran species were discovered and identified. Almost 99 % of all insect specimens were from the order of Diptera, and beetles (Coleoptera) accounted for only 1 % of the cadaver fauna. The blowflies (Calliphoridae) are of particular importance for forensic issues, accounting for about 66 % of all families. Carrion beetles (Silphidae) were found as the second most abundant family (about 21 %). In addition, combining all cases, a steadily increasing insect species richness S was detected from early decay to advanced decay (fresh S = 8; bloated S = 12; active decay S = 21; advanced decay S = 34). In the following remains stage, the species number decreased again (S = 24). However, no significant difference in the number of species was found between the stages of decay when the cases were considered individually. The temporal pattern of insect appearance was found to be congruent with those of previous studies. Furthermore, a time of death was determined for each case and compared to the pathologist's estimates. This study provides insights into the arthropod fauna of wolf remains for the first time, applies the ADD-Model for post-mortem interval estimation, and discusses the suitability of forensic entomology for wildlife death investigations.

Temperature models of development for Necrodes littoralis L. (Coleoptera: Silphidae), a carrion beetle of forensic importance in the Palearctic region

Analysis of insects can provide evidence in death cases, for example, by answering the question about the time of death. Apart from flies, beetles are the second most useful insect group in forensic entomology. To elucidate the time of death based on insect evidence, developmental models of a given species are necessary. In this study, we developed such models for Necrodes littoralis, a necrophagous beetle, which is common in the Palearctic region and has great potential in forensic entomology. We monitored the development at 10 constant temperatures (14–30 °C). Larvae were reared in aggregations. Thermal summation models, isomorphen and isomegalen diagrams and growth curves were derived using the data. Depending on the temperature, development lasted between about 23 and 89 days. Mortality was high at the extremes of the temperature range. The thermal summation constant for the total development was 434.7 ± 28.86 accumulated degree-days above a developmental threshold of 9.04 ± 0.55 °C. This is the first comprehensive dataset on the development of N. littoralis. Implications for its use in forensic casework are discussed.