Fate of mouse carcasses in a Northern Woodland

1. Small, vertebrate carrion is a significant input into terrestrial ecosystems. The relative importance of different recyclers of carrion is poorly understood because of seasonal and habitat variation and methodological differences among studies. Many studies, in particular, do not account adequately for the activity of burying beetles (Nicrophorus spp.). 2. The present work, utilising a previously unstudied woodland, addresses this concern by: placing only two carcasses in the field at one time, using microsites only once, and providing a means to easily recover successful burying beetles. 3. Burying beetles buried 77.5% of experimental mouse carcasses over their breeding season, and typically did so within three activity periods after placement (30 of 31 burials). 4. The results and a brief review suggest that burying beetles are the dominant summer competitor for small, vertebrate carrion in mid‐ and northern latitude woodlands. Information on the age of carcasses utilised by burying beetles also provides an ecological context for laboratory studies of behaviour, life history, and microbial interactions.     

Study of community assembly patterns and interspecific interactions involved in insect succession on rat carcasses

Although many forensic entomological studies have described patterns of carrion insect succession and theoretical studies have explained interspecific interactions that drive succession, empirical studies on the quantitative and ecological aspects of carrion insect succession, such as the degree of historical contingency in community assembly and interspecific interactions during succession, are limited. In this study, I investigated variability in the successional pathways of carrion insect communities in rat carcasses and their decomposition processes, and examined the interspecific interactions involved in succession, such as the effects of carcass utilization by early‐arriving species on late‐arriving species. Members of the families Calliphoridae and Formicidae and the species Eusilpha japonica (Motschulsky) and Nicrophorus concolor Kraatz were chiefly observed. In almost all carcasses, formicid species arrived first, and calliphorid species and E. japonica arrived simultaneously or immediately after. Nicrophorus concolor arrived last, with its time of colonization occurring earlier in carcasses with greater E. japonica abundance. Meanwhile, the early‐arriving species decreased when N. concolor arrived. Nicrophorus concolor tended to reproduce on carcasses with lower cumulative abundance of early‐arriving species and tended to feed on carcasses with greater cumulative abundance of early‐arriving species. These results show that the successional pathways of the chief carrion insect species are highly consistent among carcasses. In contrast, early‐arriving species seem to influence the utilization patterns of carcasses by late‐arriving species, and therefore produce variability in the decomposition process. These results also show that succession could be driven by facilitation and interspecific competition between early‐arriving and late‐arriving species.     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

Traits reveal ecological strategies driving carrion insect community assembly

1. The succession of carrion-associated (necrophilous) insects on decomposing carrion is well documented. To exploit the changing nutritious and dynamic resources available throughout the carrion decomposition process, different species colonise and consume carrion in a predictable temporal sequence. The traits of these necrophilous insects should reflect their ecological strategies. Morphological traits of these insects, such as body size and wing size, however, have not previously been examined during active and advanced decomposition. 2. We used fourth-corner multivariate generalised linear models to identify insect community morphological trait patterns and to quantify their change through time on decomposing rabbit carcasses in grassland and woodland environments. 3. We found that larger-bodied species of flies and carrion-specialist beetles were associated with the early stages of decomposition. The morphological traits of ants, in contrast, showed no changes at carcasses through time and instead showed body size differences between grassland and woodland environments. 4. Our findings indicate that specialist flies and beetles that arrive early in the decomposition process possess traits that enable rapid discovery of carrion at a large scale. Generalist beetles and ants do not share this same trait and are instead adapted to locate and consume a wider variety of resources in their preferred habitat type at their local scale. 5. Our results provide insights into the morphological adaptations linked to the ecological strategies of distinct components of carrion insect communities. Further, our results offer insights into the community assembly dynamics that structure the communities of necrophilous insect species.     

The use of carrion as breeding sites by the blowfly Lucilia sericata and other Calliphoridae

Abstract. To examine the species composition of flies breeding in carrion in the field, the carcasses of mice and quail were exposed on sheep farms in the South West of England. Calliphora vicina was the dominant species of Diptera; 19,294 individuals emerged from 175 of the 241 infested carcasses recovered. Lucilia sericata emerged from only 39 of the infested carcasses, at a median of 10 adults per infested carcass. Other species of Lucilia present were L.ampullacea, L.caesar and L.illustris. The highest number of L.sericata emerged from carcasses placed in open pasture, the highest number of C.vicina emerged from carcasses in hedgerow, whereas the highest numbers of L.caesar, L.ampullacea and L.illustris emerged from carcasses in woodland. The duration of exposure of carcasses in the field was negatively related to the size of the adult L.sericata which emerged and, in woodland and hedgerow habitats, to the number of L.sericata which emerged. These data indicate that the larvae of L.sericata in carcasses experience significant levels of competition and that the intensity of this competition may be sufficient to reduce the numbers of L.sericata able to emerge successfully. The size distributions of female L. sericata which emerged from carcasses or which were caught as adults in the field showed only a small degree of overlap, suggesting that only a relatively small proportion of the wild L.sericata population emerge from carcass breeding sites. The results are discussed in relation to the development of new approaches to the control of blowfly strike of sheep.     

Habitat Fragmentation and Burying Beetle Abundance and Success

Four species of burying beetle (Nicrophorus marginatus F., N. tomentosus Weber, N. orbicollis Say and N. defodiens Mannerheim) are attracted to small, fresh mouse carcasses in northern Michigan. The number of burying beetles and their success (burial of a carcass) were greater in woodlands than in edge or field habitats. Species diversity was least in open fields as assessed by two different indices of diversity. Nicrophorus marginatus was the only species captured in large fields (<25ha). This species was never trapped in small fields (>5ha) suggesting that a minimum habitat size might be necessary to maintain local populations. In contrast to previous studies which employed pitfall traps baited with a large quantity of carrion, N. tomentosus was caught exclusively in woodlands at single mouse carcasses. In Connecticut woodlands, burying beetle success, assayed as the proportion of carcasses buried and held for 7 days, was significantly greater in larger as compared to smaller woodlands. The limited success of burying beetles in smaller woodlands was due, in part, to a higher rate of scavenging by vertebrates.     

Patch size drives colonization by aquatic insects,with minor priority effects of a cohabitant

Patch size is one of the most important factors affecting the distribution and abundance of species, and recent research has shown that patch size is an important niche dimension affecting community structure in aquatic insects. Building on this result, we examined the impact of patch size in conjunction with presence of larval anurans on colonization by aquatic insects. Hyla chrysoscelis (Cope''s gray treefrog) larvae are abundant and early colonists in fishless lentic habitats, and these larvae can fill multiple ecological roles. By establishing larvae in mesocosms prior to colonization, we were able to assess whether H. chrysoscelis larvae have priority effects on aquatic insect assemblages. We conducted a series of three experiments in naturally colonized experimental landscapes to test whether (1) H. chrysoscelis larval density affects insect colonization, (2) variation in patch size affects insect colonization, and (3) the presence and larval density of H. chrysoscelis shift colonization of insects between patches of different size. Larval density independently had almost no effect on colonization, while patch size had species‐specific effects consistent with prior work. When larvae and patch size were tested in conjunction, patch size had numerous, often strong, species‐specific effects on colonization; larval density had effects largely limited to the assemblages of colonizing beetles and water bugs, with few effects on individual species. Higher larval densities in large mesocosms shifted some insect colonization to smaller patches, resulting in higher beta diversity among small patches in proximity to high density large mesocosms. This indicates establishing H. chrysoscelis larvae prior to insect colonization can likely create priority effects that slightly shape insect communities. Our results support the importance of patch size in studying species abundances and distributions and also indicate that colonization order plays an important role in determining the communities found within habitat patches.     

Contrasting diversity dynamics of phoretic mites and beetles associated with vertebrate carrion

Carrion is an ephemeral and nutrient-rich resource that attracts a diverse array of arthropods as it decomposes. Carrion-associated mites often disperse between animal carcasses using phoresy, the transport of one species by another. Yet few studies have contrasted the dynamics of mite assemblages with other insect taxa present at carrion. We examined and compared the changes in abundance, species richness and composition of mite and beetle assemblages sampled at kangaroo carcasses in a grassy eucalypt woodland at four different times over a 6-month period. We found that the majority of mites were phoretic, with the mesostigmatid genera Uroseius (Uropodidae), Macrocheles (Macrochelidae) and Parasitus (Parasitidae) the most abundant taxa (excluding astigmatid mites). Abundance and richness patterns of mites and beetles were very different, with mites reaching peak abundance and richness at weeks 6 and 12, and beetles at weeks 1 and 6. Both mites and beetles showed clear successional patterns via changes in species presence and relative abundance. Our study shows that mesostigmatid mite assemblages have a delay in peak abundance and richness relative to beetle assemblages. This suggests that differences in dispersal and reproductive traits of arthropods may contribute to the contrasting diversity dynamics of carrion arthropod communities, and further highlights the role of carrion as a driver of diversity and heterogeneity in ecosystems.     

A chemically triggered transition from conflict to cooperation in burying beetles

Although interspecific competition has long been recognised as a major driver of trait divergence and adaptive evolution, relatively little effort has focused on how it influences the evolution of intraspecific cooperation. Here we identify the mechanism by which the perceived pressure of interspecific competition influences the transition from intraspecific conflict to cooperation in a facultative cooperatively breeding species, the Asian burying beetle Nicrophorus nepalensis. We not only found that beetles are more cooperative at carcasses when blowfly maggots have begun to digest the tissue, but that this social cooperation appears to be triggered by a single chemical cue – dimethyl disulfide (DMDS) – emitted from carcasses consumed by blowflies, but not from control carcasses lacking blowflies. Our results provide experimental evidence that interspecific competition promotes the transition from intraspecific conflict to cooperation in N. nepalensis via a surprisingly simple social chemical cue that is a reliable indicator of resource competition between species.     

Gut microbiota in the burying beetle,Nicrophorus vespilloides,provide colonization resistance against larval bacterial pathogens

Carrion beetles, Nicrophorus vespilloides, are reared on decomposing carrion where larvae are exposed to high populations of carcass‐derived bacteria. Larvae do not become colonized with these bacteria but instead are colonized with the gut microbiome of their parents, suggesting that bacteria in the beetle microbiome outcompete the carcass‐derived species for larval colonization. Here, we test this hypothesis and quantify the fitness consequences of colonization with different bacterial symbionts. First, we show that beetles colonized by their endogenous microbiome produce heavier broods than those colonized with carcass‐bacteria. Next, we show that bacteria from the endogenous microbiome, including Providencia rettgeri and Morganella morganii, are better colonizers of the beetle gut and can outcompete nonendogenous species, including Serratia marcescens and Escherichia coli, during in vivo competition. Finally, we find that Providencia and Morganella provide beetles with colonization resistance against Serratia and thereby reduce Serratia‐induced larval mortality. This effect is eliminated in larvae first colonized by Serratia, suggesting that while competition within the larval gut is determined by priority effects, these effects are less important for Serratia‐induced mortality. Our work suggests that an unappreciated benefit of parental care in N. vespilloides is the social transmission of the microbiome from parents to offspring.     

Starving the competition: a proximate cause of reproductive skew in burying beetles (Nicrophorus vespilloides)

Proximate mechanisms underlying reproductive skew are obscure in many animals that breed communally. Here, we address causes of reproductive skew in brood-parasitic associations of burying beetles (Nicrophorus vespilloides). Male and female burying beetles feed and defend their larvae on buried carcasses. When several females locate the same small carcass, they engage in violent physical altercations. The subordinate then acts as an intraspecific brood parasite, laying eggs, but not providing care. The dominant female largely monopolizes access to the carcass; she alone provides parental care and her share of the brood is much larger than the subordinate's. On larger carcasses, subordinates have greater access to the carcass than on small ones, and reproductive skew is reduced. Differential fecundity, ovicide and larvicide have been suggested as causes of skew on small carcasses. Here, we report the results of the experiments pertaining to the first two of these potential mechanisms. Ovicide did not significantly contribute to reproductive skew on small carcasses, but differential fecundity did. Fecundity differences were due to dominance status, not body size per se. Fecundity differences disappeared when supplemental food was available, suggesting that reduced access to the carcass limits fecundity by causing nutritional deficiencies. Supplemental food prevented such nutritional deficiencies and allowed subordinates to produce as many eggs as dominants. Apparently, aggressive behaviour by dominants functions in the context of reproductive competition, limiting subordinate reproduction by preventing food intake on the carcass.     

Fitness costs of phoretic nematodes in the burying beetle,Nicrophorus vespilloides

Nicrophorusvespilloides is a social beetle that rears its offspring on decomposing carrion. Wild beetles are frequently associated with two types of macrobial symbionts, mites, and nematodes. Although these organisms are believed to be phoretic commensals that harmlessly use beetles as a means of transfer between carcasses, the role of these symbionts on N. vespilloides fitness is poorly understood. Here, we show that nematodes have significant negative effects on beetle fitness across a range of worm densities and also quantify the density‐dependent transmission of worms between mating individuals and from parents to offspring. Using field‐caught beetles, we provide the first report of a new nematode symbiont in N. vespilloides, most closely related to Rhabditoides regina, and show that worm densities are highly variable across individuals isolated from nature but do not differ between males and females. Next, by inoculating mating females with increasing densities of nematodes, we show that worm infections significantly reduce brood size, larval survival, and larval mass, and also eliminate the trade‐off between brood size and larval mass. Finally, we show that nematodes are efficiently transmitted between mating individuals and from mothers to larvae, directly and indirectly via the carcass, and that worms persist through pupation. These results show that the phoretic nematode R. regina can be highly parasitic to burying beetles but can nevertheless persist because of efficient mechanisms of intersexual and intergenerational transmission. Phoretic species are exceptionally common and may cause significant harm to their hosts, even though they rely on these larger species for transmission to new resources. However, this harm may be inevitable and unavoidable if transmission of phoretic symbionts requires nematode proliferation. It will be important to determine the generality of our results for other phoretic associates of animals. It will equally be important to assess the fitness effects of phoretic species under changing resource conditions and in the field where diverse interspecific interactions may exacerbate or reduce the negative effects of phoresy.     

Burying beetle Nicrophorus investigator reproduction on Pacific salmon carcasses

Abstract.  1. In many undisturbed watersheds along the Pacific Rim, anadromous salmon ( Oncorhynchus spp.) provide a predictable source of carrion to the riparian zone, largely due to horizontal transfer of salmon carcasses by bears ( Ursus spp.) and other vertebrates.
2. Burying beetles are important members of the north-temperate carrion fauna, and may utilise salmon carcasses and remnants for breeding. In this study, isotopic and observational data are reported that demonstrate previously unrecognised Nicrophorus investigator (Zetterstedt) reproduction on large salmon carcasses from five watersheds in coastal British Columbia.
3. Stable isotope signatures (δ¹⁵N and δ¹³C) of adult beetles collected in autumn indicate a diet of salmon origin in all but one individual from all watersheds, suggesting that this beetle–salmon association is widespread. Comparison of autumn isotope signatures to individuals collected randomly in summer suggests that isotope signatures represent the larval carrion source from the previous autumn rather than immediate adult diet.
4. In a survey of N. investigator use of salmon carcasses from two watersheds, 35 broods were observed on chum and pink salmon carcasses, including 16 natural brood complexes containing over 100 larvae, and five ranging from 250 to 750 larvae.
5. Overall, north-coastal populations of N. investigator breed on the rich and reliable salmon resource and may exhibit a system of communal breeding on these carcasses. This is most relevant when the dramatic reduction in salmon spawning biomass over the last century is considered.     

Effects of diet on female fecundity and larval development in the carrion beetle Necrophila japonica

The carrion beetle subfamily Silphinae (Coleoptera: Silphidae) contains dominant macroinvertebrates of soil ecosystems in temperate zones. However, their feeding habits, which determine the role of each species in the ecosystem, have not been sufficiently studied. Moreover, although a diet shift from necrophagy on vertebrate carcasses to predatory feeding on invertebrates is known to occur in this subfamily, the processes and mechanisms of this shift have also been inadequately addressed. We examined female fecundity and larval development on various diets in a Silphinae species, Necrophila (Eusilpha) japonica (Motschulsky). The experimental diets included a meat diet and various invertebrate diets, which reflect the ‘ancestral’ feeding habit in Silphinae, necrophagy, and the ‘derivative’ feeding habit, predatory feeding. Female fecundity was significantly higher on the meat diet (minced beef) than on an insect larvae diet (mealworms and dipteran larvae) but did not significantly differ from that on an earthworm diet. Larval developmental performance was significantly higher on the earthworm diet than on the meat and insect larvae diets. Our results for larval development were consistent with those of previous stable isotope analyses of the same species, in which isotopic values of larval samples agree with those of hypothetical consumers that utilize earthworms. The consistency of results among different methods indicates that N. japonica larvae are most likely earthworm feeders. In contrast, our results for the female fecundity experiment differed from those of previous stable isotope analyses, in which vertebrate carcasses unlikely serve as the staple diet of adults in the field; thus, the feeding habits of N. japonica adults remain unresolved. Our observations that females and larvae performed best on the meat and earthworm diets, respectively, may indicate that, in Silphinae, the diet shift from necrophagy to predatory habits occurs earlier in larvae than in adults.     

Effects of vulture exclusion on carrion consumption by facultative scavengers

Vultures provide an essential ecosystem service through removal of carrion, but globally, many populations are collapsing and several species are threatened with extinction. Widespread declines in vulture populations could increase the availability of carrion to other organisms, but the ways facultative scavengers might respond to this increase have not been thoroughly explored. We aimed to determine whether facultative scavengers increase carrion consumption in the absence of vulture competition and whether they are capable of functionally replacing vultures in the removal of carrion biomass from the landscape. We experimentally excluded 65 rabbit carcasses from vultures during daylight hours and placed an additional 65 carcasses that were accessible to vultures in forested habitat in South Carolina, USA during summer (June–August). We used motion‐activated cameras to compare carrion use by facultative scavenging species between the experimental and control carcasses. Scavenging by facultative scavengers did not increase in the absence of competition with vultures. We found no difference in scavenger presence between control carcasses and those from which vultures were excluded. Eighty percent of carcasses from which vultures were excluded were not scavenged by vertebrates, compared to 5% of carcasses that were accessible to vultures. At the end of the 7‐day trials, there was a 10.1‐fold increase in the number of experimental carcasses that were not fully scavenged compared to controls. Facultative scavengers did not functionally replace vultures during summer in our study. This finding may have been influenced by the time of the year in which the study took place, the duration of the trials, and the spacing of carcass sites. Our results suggest that under the warm and humid conditions of our study, facultative scavengers would not compensate for loss of vultures. Carcasses would persist longer in the environment and consumption of carrion would likely shift from vertebrates to decomposers. Such changes could have substantial implications for disease transmission, nutrient cycling, and ecosystem functioning.     

Review. When to leave the brood chamber? Routes of dispersal in mites associated with burying beetles

Most nests of brood-caring insects are colonized by a rich community of mite species. Since these nests are ephemeral and scattered in space, phoresy is the principal mode of dispersal in mites specializing on insect nests. Often the mites will arrive on the nest-founding insect, reproduce in the nest and their offspring will disperse on the insect's offspring. A literature review shows that mites reproducing in the underground brood chambers of burying beetles use alternative routes for dispersal. For example, the phoretic instars of Poecilochirus spp. (Mesostigmata: Parasitidae) disperse early by attaching to the parent beetles. Outside the brood chamber, the mites switch host at carcasses and pheromone-emitting male beetles, where juvenile and mature burying beetles of several species congregate. Because they preferably switch to beetles that are reproductively active and use all species of burying beetles within their ranges, they have a good chance of arriving in a new brood chamber. Other mite associates of burying beetles (Alliphis necrophilus and Uropodina) disperse from the brood chamber on the beetle offspring. We suggest that these mites forgo the possible time gain of dispersing early on the parent beetles because their mode of attachment precludes host switching. Their phoretic instars, once attached, have to stay on their host and so only dispersing on the beetle offspring guarantees that they are present on reproducing burying beetles of the next season. The mites associated with burying beetles providean example of multiple solutions to one life history problem – how to find a new brood chamber for reproduction. Mites that have mobile phoretic instars disperse on the parent beetles and try to arrive in the next brood chamber by host switching. They are independent of the generation cycle of a single host and several generations of mites per host generation are possible. Mites that are constrained by their mode of attachment disperse on the beetle offspring and wait until their host becomes mature and reproduces. By doing this they synchronize their generation time with the generation time of their host species. Exp Appl Acarol 22: 621–631 © 1998 Kluwer Academic Publishers     

Species Traits Predict Assemblage Dynamics at Ephemeral Resource Patches Created by Carrion

Carrion is an ephemeral and spatially patchy resource that supports a diverse subset of species linked to nutrient cycling and the decomposition process. A number of studies have separately documented changes in the diversity of plants, arthropods and vertebrates at individual carcasses, but there are few studies that have examined how functional traits of different groups of organisms underpin their responses to carrion patches. We used a carrion addition experiment to compare changes in composition and functional traits of insect and plant assemblages at carcasses compared with control sites. We found that significant changes in insect assemblage evenness and heterogeneity was associated with species’ dispersal traits, and that plant assemblage responses to subsequent soil nitrogen changes was most apparent among graminoids and exotic species. Beetles at carcasses were twice as large as their counterparts at control sites during the first week of carrion decomposition, and also had higher wing loadings. Plants with high specific leaf area responded faster to the carcass addition, and twice as many species recolonised the centre of carcasses in exotic-dominated grassland compared with carcasses in native-dominated grassland. These results provide an example of how traits of opportunist species enable them to exploit patchy and dynamic resources. This increases our understanding of how carcasses can drive biodiversity dynamics, and has implications for the way carrion might be managed in ecosystems, such as appropriate consideration of spatial and temporal continuity in carrion resources to promote heterogeneity in nutrient cycling and species diversity within landscapes.     

Nestmate recognition in burying beetles: the "breeder's badge" as a cue used by females to distinguish their mates from male intruders

Burying beetles use small vertebrate carcasses as food for theirlarvae and defend these carcasses against intra- and interspecificcompetitors. Breeding associations on carcasses can consistof single females, heterosexual pairs, or various combinationsof males and females. When a heterosexual pair collaborate ina breeding attempt, they do not typically exhibit aggressivebehavior toward each other, but do attack newly arrived conspecificsthat attempt to usurp the carcass. We investigated the cuesinvolved in discrimination between breeding partners and intrudersby female burying beetles. We found that resident females toleratemales that have cared for a brood, as well as males that havenot cared for a brood but have been on a carcass for a day ortwo. Males that have had no prior contact with a carcass areattacked. Females appear to use a chemical cue, the "breeder'sbadge," an apolar substance on the male's cuticle that can beremoved by washing with pentane. This cue is reliably correlatedwith recent male experience with a carcass that is suitablefor reproduction. The breeder's badge develops as a result ofprolonged contact with such a carcass, and disappears on removalfrom the carcass; its presence does not require contact witha female or with larvae. Female recognition of their male partnersin burying beetles thus does not involve individual recognition,but rather recognition of reproductive condition.     

Flexible parents: joint effects of handicapping and brood size manipulation on female parental care in Nicrophorus vespilloides

Parental care is highly variable, reflecting that parents make flexible decisions in response to variation in the cost of care to themselves and the benefit to their offspring. Much of the evidence that parents respond to such variation derives from handicapping and brood size manipulations, the separate effects of which are well understood. However, little is known about their joint effects. Here, we fill this gap by conducting a joint handicapping and brood size manipulation in the burying beetle Nicrophorus vespilloides. We handicapped half of the females by attaching a lead weight to their pronotum, leaving the remaining females as controls. We also manipulated brood size by providing each female with 5, 20 or 40 larvae. In contrast to what we predicted, handicapped females spent more time provisioning food than controls. We also found that handicapped females spent more time consuming carrion. Furthermore, handicapped females spent a similar amount of time consuming carrion regardless of brood size, whereas controls spent more time consuming carrion as brood increased. Females spent more time provisioning food towards larger broods, and females were more likely to engage in carrion consumption when caring for larger broods. We conclude that females respond to both handicapping and brood size manipulations, but these responses are largely independent of each other. Overall, our results suggest that handicapping might lead to a higher investment into current reproduction and that it might be associated with compensatory responses that negate the detrimental impact of higher cost of care in handicapped parents.     

Effects of Carrion-Independent Pheromone Emission by Male Burying Beetles (Silphidae: Necrophorus)

All former publications on the reproductive biology of burying beetles (genus Necrophorus) state that in these species copulations take place on carcasses only, this being the only place for the sexes to meet. Our laboratory investigations have shown that males emit pheromones if no carrion is around. We now present data from a field investigation showing the effects of pheromone emission by males that have not secured a carcass. Such males are successful in attracting conspecific females. But conspecific males and individuals of other species are also attracted by the scent. Possible benefits for attracted individuals of different species and sex are discussed. Pheromone emission and attraction of conspecifics are restricted to a distinct species-specific period of the day in at least two of the common species.