Seasonal variation in parental care, offspring development, and reproductive success in the burying beetle, Nicrophorus vespillo

1. Beetles of the genus Nicrophorus reproduce on small vertebrate carcasses that they bury in the soil to provide the larvae with food. Usually, both parents cooperate in brood care by feeding and guarding their progeny. 2. In pairs of the common European species N. vespillo, the duration of care depended on the time of year when the beetles reproduced. Both in 1990 and in 1991, male and female parents stayed longer with their broods when reproduction started in spring than when reproduction started in early or late summer. This was probably due to the longer development time of the larvae caused by lower temperatures in spring, because laboratory experiments suggested a strong influence of temperature on both the duration of brood care and offspring development. 3. The number of adult offspring produced by a beetle pair did not vary among different times of the year. 4. The median time required for offspring development, measured as time from burial of the carcass to emergence of young adults, was between 62 and 84 days. When the beetles reproduced in late summer, only about three-quarters of the offspring left the soil and hibernated as adults. The remaining offspring stayed underground and adults appeared on the soil surface the following spring. They still showed the flexible cuticle typical of newly-hatched beetles, suggesting that they may have overwintered in a pre-adult stage.     

Species packing and temperature dependent competition among burying beetles (Silphidae, Nicrophorus)

Abstract. 1. Burying beetles are carrion feeders that specialize on small carcasses. We investigated interactions among congeners at two sites in Michigan, U.S.A., that differ in both number and relative proportion of species.
2. Intra- and interspecific competition for carcasses is intense. The majority of carcasses are found within 24 h of their placement.
3. Competition between N.arbicollis and N.defodiens is temperature dependent. N.orbicollis can displace N.defodiens on single carcasses, but requires warm temperatures to find the carcasses. Cool weather therefore serves as a refuge for N.defadiens .
4. The southern edge of N.defodien's geographical range is probably determined by competition with N.orbicollis .
5. N.sayi and N.tomenrosus are spring and autumn breeders respectively, and rarely interact with each other or the other two species. Nevertheless, interspecific competition is the most likely evolutionary force leading to seasonal segregation.     

Timing of oviposition and reproductive skew in cobreeding female burying beetles (Nicrophorus vespilloides)

Burying beetles (genus Nicrophorus) are known for their elaborate parental care. Two or more conspecific females may reproduceon the same carcass, especially when the carcass is large.Here we present the results of experiments in which we observedpatterns of larval hatching and parental care in unmanipulatedcobreeders, manipulated hatching synchrony between cobreeders,and compared patterns of oviposition in cobreeding and single females. Our results show that in these cobreeding associations,one of the females may or may not monopolize the carcass duringthe period of larval hatching. We present evidence that ineither case, infanticide based on temporal cues constitutesan important proximate mechanism underlying the observed reductionin average reproductive success in cobreeding females. Femaleswith higher synchrony (i.e., greater overlap between their oviposition patterns) produce larger broods with lower reproductive skew.Cobreeding females oviposit later and less synchronously thansingle breeders. Such delayed oviposition may reduce the riskthat a female's larvae fall victim to cannibalistic acts committedby her cobreeder or maximize her own opportunity to selectivelykill her cobreeder's larvae.     

Regulation of brood size in a burying beetle,Nicrophorus tomentosus (Silphidae)

Regulation of brood size in a biparental burying beetle, Nicrophorus tomentosusWeber, was studied by providing pairs with one of two sizes of mouse carcasses in the laboratory. For a given carcass size, there was an inverse relationship between number and mass offspring in a brood. The requirement for regulation was that brood size was adjusted such that mean mass of individual larvae was constant for carcasses of different size. Brood size was regulated if parents were present but regulation did not occur if parents were removed prior to hatching of larvae. Pairs bred in quick succession on two carcasses raised fewer than the regulated number of young in the second reproductive attempt. Reasons for regulation of brood size in this genus are discussed.     

Social stimuli affect juvenile hormone during breeding in biparental burying beetles (Silphidae: Nicrophorus)

Extended biparental care is rare in insects but provides an excellent opportunity to investigate the interaction between the endocrine system and the physical and social environment in the regulation of this behavior. Burying beetles (Nicrophorus spp.) have facultative biparental care and depend on locating a small vertebrate carcass that they bury and prepare as food for their young. Commonly, both male and female Nicrophorus orbicollis remain in the burial chamber after eggs hatch to feed and guard the larvae. In both sexes, juvenile hormone (JH) rises rapidly in response to the discovery and assessment of the carcass; it returns to near baseline in 24 h; then in females it reaches very high titers at the onset of maternal care. In this paper, we investigate some social (presence of a mate, mating history, larval age) and environmental (carcass size) factors that may affect this endocrine profile. For females, neither the presence of a mate nor mating status (i.e., virginity) affected the initial rise of JH. However, the absence of a mate significantly depressed the JH rise in males. Eighty-seven percent of the single males buried the carcass like paired males but 87% also released pheromones to attract a mate. JH hemolymph titers in females whose broods were replaced every 24 h with newly hatch larvae were significantly higher than those of females rearing aging broods. Lastly, even though larger carcasses took longer to bury and prepare and oviposition was delayed, neither JH titers nor speed of ovarian development was affected by carcass size.     

Burying beetles: intraspecific interactions and reproductive success in the field

Abstract. 1. The discovery and utilization of small carcasses by burying beetles (Silphidae, Nicrophorus ) was studied by placing dead mice at random points on large grids at two Iocations in Michigan, U.S.A.
2. The majority of mice are found within 24 h by more beetles than ultimately will utilize the carcass. If a carcass is likely to be usurped by a larger species of beetle or by a vertebrate, then intraspecific competition may be postponed until the carcass is concealed and buried.
3. Both males and females practice parental care. Maturing broods are tended by no adults, a single female, a single male, or a male—female pair. No differences in brood success were observed among these categories.
4. The female lays a larger clutch than ultimately will survive. Brood size is regulated after the egg stage, such that offspring number varies, but individual offspring size does not.
5. A large amount of unexplained variation exists in brood size, in both the laboratory and the field. This variation is probably caused by the environment, and not the reproductive physiology of the beetles. Competition with microbes is a likely candidate.
6. Differences exist not only between Nicrophorus species, but also between localities for a single species, suggesting adaptation to local environments.     

Juvenile hormone-mediated reproduction in burying beetles: From behavior to physiology

The burying beetle, Nicrophorus orbicollis, is the first beetle exhibiting parental care for which endocrinological studies have been initiated. Burying beetles bury and prepare small vertebrate carcasses as a breeding resource for their offspring. After emergence as an adult, hemolymph titers of juvenile hormone and ovarian size increase in concert for 2–3 weeks, and both plateau until an appropriate resource is discovered. Upon finding a suitable carcass, titers of juvenile hormone increase extremely rapidly (<20 min), and within 18 h ovarian mass increases threefold and oviposition begins. This rapid reproductive development is hypothesized to be selected by the intense competition for these protein-rich but quickly deteriorating resources. Burying beetle females exhibit an additional juvenile hormone surge at the time young hatch. This peak in juvenile hormone is hypothesized to be associated with either the considerable behavioral demands which accompany care of young larvae or with a female's willingness to oviposit a replacement clutch should brood failure occur early in the care-giving phase. Parental care has evolved in at least 16 additional families within the Coleoptera. Comparative studies of nonparental and parental groups will be important for understanding how the physiology of ancestral groups affects the evolution of complex social behavior. Arch. Insect Biochem. Physiol. 35:479–490, 1997. © 1997 Wiley-Liss, Inc.     

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.     

Dormancy and the influence of photoperiod and temperature on sexual maturity in Nicrophorus nepalensis (Coleoptera: Silphidae)

Abstract Carrion beetles (Nicrophorus spp.) use small vertebrate carcasses for food and reproduction. Their ecology and behaviors are highly affected by the availability of carcasses and the surrounding environmental conditions. Our results revealed that in subtropical Fushan, northern Taiwan, N. nepalensis was mainly active in spring (February to May), and could also be found in autumn (October and November); but there was no capture record in summer (June to September) and winter (December and January). A laboratory temperature tolerance study indicated that N. nepalensis adults become inactive at temperatures above 26°C, and had the highest mortality when the temperature was raised from 27°C to 28°C. Furthermore, N. nepalensis became sexually mature at 20°C, depending on the photoperiod: the longer the day, the lower the percentage of sexually mature 2‐week‐old females after emergence. In another experiment, N. nepalensis virgins were paired under three possible conditions at Fushan. At 15°C and 20°C, if carcasses were presented to the pairs within 3 days after emergence, all laid eggs in the second week after emergence. If carcasses were presented 1 week after emergence, most began to reproduce at 20°C with 12.5 h of daylight. However, at 15°C with 11 h of daylight, the carrion beetles hibernated first, and reproduced in the ninth week after emergence. At 25°C with 14 h of daylight, carrion beetles did not bury the mouse carcasses, the females did not lay eggs, and the adult lifespan was only one‐third of that at 20°C. This study revealed that both photoperiod and temperature influence the time needed to reach the sexual maturity of N. nepalensis; and also implied that the narrow temperature tolerance range and dormancy behavior of carrion beetles are highly regulated by those environmental factors.     

Effect of forest removal on the abundance of the endangered American burying beetle, Nicrophorus americanus (Coleoptera: Silphidae)

We test the hypothesis that the decline of the endangered American burying beetle (Nicrophorus americanus) from over 90% of its original range is the result of habitat loss and fragmentation of eastern North America. Forest removal at a site in southeastern Oklahoma known to have a significant population of N. americanus gave us a unique opportunity to test this hypothesis. At the local scale of this experiment, N. americanus declined significantly after forest removal while beetle numbers at adjacent forested plots did not change. Our results indicate that local disturbances such as forest removal, if occurring across relatively broad spatial scales, can cause wholesale geographic range collapse in this species.     

Population trends and flight behavior of the American burying beetle, Nicrophorus americanus (Coleoptera: Silphidae), on Block Island, RI

The endangered American burying beetle, Nicrophorus americanus, was monitored on Block Island, RI, USA, from 1991–2003 using mark-recapture population estimates of adults collected in pitfall traps. Populations increased through time, especially after 1994 when a program was initiated that provided carrion for beetle production. Beetle captures increased with increasing temperature and dew point, and decreased with increasing wind speed. Short distance movement was not related to wind direction, while longer distance flights tended to be downwind. Although many individuals flew considerable distances along transects, most recaptures were in traps near the point of release. These behaviors probably have counterbalancing effects on population estimates.The U.S. Goverment's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Satellite DNA repeat sequence variation is low in three species of burying beetles in the genus Nicrophorus (Coleoptera: Silphidae)

Three satellite DNA families were identified in three species of burying beetles, Nicrophorus orbicollis, N. marginatus, and N. americanus. Southern hybridization and nucleotide sequence analysis of individual randomly cloned repeats shows that these satellite DNA families are highly abundant in the genome, are composed of unique repeats, and are species-specific. The repeats do not have identifiable core elements or substructures that are similar in all three families, and most interspecific sequence similarity is confined to homopolymeric runs of A and T. Satellite DNA from N. marginatus and N. americanus show single-base-pair indels among repeats, but single-nucleotide substitutions characterize most of the repeat variability. Although the repeat units are of similar lengths (342, 350, and 354 bp) and A + T composition (65%, 71%, and 71%, respectively), the average nucleotide divergence among sequenced repeats is very low (0.18%, 1.22%, and 0.71%, respectively). Transition/transversion ratios from the consensus sequence are 0.20, 0.69, and 0.70, respectively.      

Factors affecting overwinter survival of the American burying beetle, <Emphasis Type="Italic">Nicrophorus americanus</Emphasis> (Coleoptera: Silphidae)

The endangered American burying beetle (Nicrophorus americanus) is relatively abundant at Fort Chaffee Maneuver Training Center in northwestern Arkansas. There is a paucity of basic life-history information available, particularly with respect to factors affecting overwintering success. In a field experiment we: (1) captured beetles at Fort Chaffee; (2) bred them in captivity; (3) in the fall on Fort Chaffee placed offspring individually in well-ventilated, lidded 21.1-l buckets containing original soil plugs in grassland or woodland, either provisioned or not with a rat carcass as potential food; (4) overwintered the beetles; (5) checked in the spring to determine survival; and (6) released surviving beetles. Overall, 59.6% of 104 beetles survived the winter, with 77.1% and 44.6% survival in provisioned and nonprovisioned buckets, respectively. No differences were evident between habitats. Beetle age was an important survival predictor, with older beetles having a higher survival probability, but only if nonprovisioned. Gender and body size were not predictive of survival. Many surviving beetles were at or near the surface; depth averaged 6.0 cm, with some as deep as 20 cm. Our findings suggest that American burying beetles will have a higher probability of overwinter survival if carcasses are readily available as winter approaches.     

Silvicultural practices,forest age,forest type,and large-scale regional factors influencing numbers of American burying beetles,Nicrophorus americanus (Coleoptera: Silphidae)

Journal of Insect Conservation - The federally threatened American burying beetle, Nicrophorus americanus previously occurred throughout much of eastern North America but now is self-sustaining...     

Isolation of nine polymorphic microsatellite loci from the burying beetle,Nicrophorus quadripunctatus (Coleoptera: Silphidae)

The burying beetle, Nicrophorus quadripunctatus Kraatz, is a common species in Japan, and its biparental care has been well studied. It exploits small vertebrate carrion as food for its young. Therefore, its reproductive success is restricted to large habitats in which carrion is abundant and available. Hence, the abundance of Nicrophorus species can be used as an indicator of forest fragmentation. Forests have recently become fragmented due to anthropogenic activities, and thus the population sizes of Nicrophorus species have decreased. To investigate the population genetic structure of N. quadripunctatus, we developed polymorphic microsatellite markers using magnetic particles. Nine microsatellites were polymorphic, with two to 12 alleles observed in the samples collected from our study sites, Matsunoyama and Nagaoka A and B. Deviation from the Hardy–Weinberg equilibrium was detected at five to seven loci in each population examined, which is indicative of significant heterozygous deficiencies. Higher genetic diversity was found in the Matsunoyama and Nagaoka B populations, the collection plots of which were covered by continuous woodlands, compared to Nagaoka A. The isolated microsatellite markers will be used to determine the genetic structures of the fragmented populations of N. quadripunctatus.