Niche variation and the maintenance of variation in body size in a burying beetle

© 2015 The Authors. Ecological Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.Online Version of Record published before inclusion in an issue.The final published version is available via DOI: 10.1111/een.122751. In burying beetles (Nicrophorinae) body size is known to provide both a fecundity advantage (in females) and successful resource defence (in males and females). Despite this, considerable variation in body sizes is observed in natural populations. 2. A possible explanation for the maintenance of this variation, even with intra- and interspecific resource competition, is that individuals might assort according to body size on different sized breeding-resources. 3. We tested the prediction that ‘bigger is always better’ in the wild, and in the laboratory, by experimentally manipulating combinations of available breeding-resource size (mouse carcasses) and competitor’s body size in Nicrophorus vespilloides (Herbst 1783). 4. In the field, large female beetles deserted small carcasses, without breeding, more often than they did larger carcasses, but small females used carcasses indiscriminately with respect to size. In the laboratory large beetles reared larger broods (with more offspring) on larger carcasses than small beetles, but on small carcasses small beetles had a reproductive advantage over large ones. Offspring size covaried with carcass size independently of parental body size. 5. Our combined results suggest breeding resource value depends on an individual’s body size, and variation in body size is environmentally induced: maintained by differences in available carcass sizes. This produces a mechanism by which individual specialisation leads to an increase in niche variation via body size in these beetles.This work was supported by a PhD studentship from the Natural Environment Research Council (NE/1528326/1) and a grant from NERC to N.J.R. and A.J.M. (NE/1025468/1)

Ageing in personal and social immunity: do immune traits senesce at the same rate?

1) How much should an individual invest in immunity as it grows older? Immunity is costly and its value is likely to change across an organism’s lifespan. A limited number of studies have focused on how personal immune investment changes with age in insects, but we do not know how social immunity, immune responses that protect kin, changes across lifespan, or how resources are divided between these two arms of the immune response. 2) In this study both personal and social immune function are considered in the burying beetle, Nicrophorus vespilloides. We show that personal immune function declines (phenoloxidase levels) or is maintained (defensin expression) across lifespan in non-breeding beetles but is maintained (phenoloxidase levels) or even upregulated (defensin expression) in breeding individuals. In contrast, social immunity increases in breeding burying beetles up to middle age, before decreasing in old age. Social immunity is not affected by a wounding challenge across lifespan, whereas personal immunity, through PO, is upregulated following wounding to a similar extent across lifespan. 3) Personal immune function may be prioritised in younger individuals in order to ensure survival until reproductive maturity. If not breeding, this may then drop off in later life as state declines. As burying beetles are ephemeral breeders, breeding opportunities in later life may be rare. When allowed to breed beetles may therefore invest heavily in ‘staying alive’ in order to complete what could potentially be their final reproductive opportunity. As parental care is important for the survival and growth of offspring in this genus, staying alive to provide care behaviours will clearly have fitness payoffs. 4) This study shows that all immune traits do not senesce at the same rate. In fact, the patterns observed depend upon the immune traits measured and the breeding status of the individual

The early-life environment and individual plasticity in life-history traits

We tested whether the early-life environment can influence the extent of individual plasticity in a life-history trait. We asked: can the early-life environment explain why, in response to the same adult environmental cue, some individuals invest more than others in current reproduction? Moreover, can it additionally explain why investment in current reproduction trades off against survival in some individuals, but is positively correlated with survival in others? We addressed these questions using the burying beetle, which breeds on small carcasses and sometimes carries phoretic mites. These mites breed alongside the beetle, on the same resource, and are a key component of the beetle's early-life environment. We exposed female beetles to mites twice during their lives: during their development as larvae and again as adults during their first reproductive event. We measured investment in current reproduction by quantifying average larval mass and recorded the female's life span after breeding to quantify survival. We found no effect of either developing or breeding alongside mites on female reproductive investment, nor on her life span, nor did developing alongside mites influence her size. In post hoc analyses, where we considered the effect of mite number (rather than their mere presence/absence) during the female's adult breeding event, we found that females invested more in current reproduction when exposed to greater mite densities during reproduction, but only if they had been exposed to mites during development as well. Otherwise, they invested less in larvae at greater mite densities. Furthermore, females that had developed with mites exhibited a trade-off between investment in current reproduction and future survival, whereas these traits were positively correlated in females that had developed without mites. The early-life environment thus generates individual variation in life-history plasticity. We discuss whether this is because mites influence the resources available to developing young or serve as important environmental cues

Asynchronous hatching provides females with a means for increasing male care but incurs a cost by reducing offspring fitness

In species with biparental care, sexual conflict occurs because the benefit of care depends on the total amount of care provided by the two parents while the cost of care depends on each parent’s own contribution. Asynchronous hatching may play a role in mediating the resolution of this conflict over parental care. The sexual conflict hypothesis for the evolution of asynchronous hatching suggests that females adjust hatching patterns in order to increase male parental effort relative to female effort. We tested this hypothesis in the burying beetle Nicrophorus vespilloides by setting up experimental broods with three different hatching patterns: synchronous, asynchronous and highly asynchronous broods. As predicted, we found that males provided care for longer in asynchronous broods whereas the opposite was true of females. However, we did not find any benefit to females of reducing their duration of care in terms of increased lifespan or reduced mass loss during breeding. We found substantial negative effects of hatching asynchrony on offspring fitness as larval mass was lower and fewer larvae survived to dispersal in highly asynchronous broods compared to synchronous or asynchronous broods. Our results suggest that, even though females can increase male parental effort by hatching their broods more asynchronously, females pay a substantial cost from doing so in terms of reducing offspring growth and survival. Thus, females should be under selection to produce a hatching pattern that provides the best possible trade-off between the benefits of increased male parental effort and the costs due to reduced offspring fitness