Male burying beetles extend, not reduce, parental care duration when reproductive competition is high

This is the author accepted manuscript. The final version is available on open access from Wiley via the DOI in this recordThe published version is also available in ORE: http://hdl.handle.net/10871/22254Male parents spend less time caring than females in many species with biparental care. The traditional explanation for this pattern is that males have lower confidence of parentage, so they desert earlier in favour of pursuing other mating opportunities. However, one recent alternative hypothesis is that prolonged male parental care might also evolve if staying to care actively improves paternity. If this is the case, an increase in reproductive competition should be associated with increased paternal care. To test this prediction, we manipulated the level of reproductive competition experienced by burying beetles, Nicrophorus vespilloides (Herbst, 1783). We found that caregiving males stayed for longer and mated more frequently with their partner when reproductive competition was greater. Reproductive productivity did not increase when males extended care. Our findings provide support for the increased paternity hypothesis. Extended duration of parental care may be a male tactic both protecting investment (in the current brood) and maximizing paternity (in subsequent brood(s) via female stored sperm) even if this fails to maximize current reproductive productivity and creates conflict of interest with their mate via costs associated with increased mating frequency

Selection on an antagonistic behavioral trait can drive rapid genital coevolution in the burying beetle, Nicrophorus vespilloides

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Male and female genital morphology varies widely across many taxa, and even among populations. Disentangling potential sources of selection on genital morphology is problematic because each sex is predicted to respond to adaptations in the other due to reproductive conflicts of interest. To test how variation in this sexual conflict trait relates to variation in genital morphology we used our previously developed artificial selection lines for high and low repeated mating rates. We selected for high and low repeated mating rates using monogamous pairings to eliminate contemporaneous female choice and male-male competition. Male and female genital shape responded rapidly to selection on repeated mating rate. High and low mating rate lines diverged from control lines after only 10 generations of selection. We also detected significant patterns of male and female genital shape coevolution among selection regimes. We argue that because our selection lines differ in sexual conflict, these results support the hypothesis that sexually antagonistic coevolution can drive the rapid divergence of genital morphology. The greatest divergence in morphology corresponded with lines in which the resolution of intrasexual conflict over mating rate was biased in favor of male interests.Funding was provided by Natural Environment Research Council grants NE/I025468/1 to N.J.R. and A.J.M., and NE/H003738/1 to A.J.M

The effect of size and sex ratio experiences on reproductive competition in Nicrophorus vespilloides burying beetles in the wild.

PublishedJournal ArticleThis is an open access article.Male parents face a choice: should they invest more in caring for offspring or in attempting to mate with other females? The most profitable course depends on the intensity of competition for mates, which is likely to vary with the population sex ratio. However, the balance of pay-offs may vary among individual males depending on their competitive prowess or attractiveness. We tested the prediction that sex ratio and size of the resource holding male provide cues regarding the level of mating competition prior to breeding and therefore influence the duration of a male's biparental caring in association with a female. Male burying beetles, Nicrophorus vespilloides were reared, post-eclosion, in groups that differed in sex ratio. Experimental males were subsequently translocated to the wild, provided with a breeding resource (carcass) and filmed. We found no evidence that sex ratio cues prior to breeding affected future parental care behaviour but males that experienced male-biased sex ratios took longer to attract wild mating partners. Smaller males attracted a higher proportion of females than did larger males, securing significantly more monogamous breeding associations as a result. Smaller males thus avoided competitive male-male encounters more often than larger males. This has potential benefits for their female partners who avoid both intrasexual competition and direct costs of higher mating frequency associated with competing males.Natural Environment Research Council. NE/1528326/1, NE/1025468/

Do female Nicrophorus vespilloides reduce direct costs by choosing males that mate less frequently?

PublishedJournal ArticleSexual conflict occurs when selection to maximize fitness in one sex does so at the expense of the other sex. In the burying beetle Nicrophorus vespilloides, repeated mating provides assurance of paternity at a direct cost to female reproductive productivity. To reduce this cost, females could choose males with low repeated mating rates or smaller, servile males. We tested this by offering females a dichotomous choice between males from lines selected for high or low mating rate. Each female was then allocated her preferred or non-preferred male to breed. Females showed no preference for males based on whether they came from lines selected for high or low mating rates. Pairs containing males from high mating rate lines copulated more often than those with low line males but there was a negative relationship between female size and number of times she mated with a non-preferred male. When females bred with their preferred male the number of offspring reared increased with female size but there was no such increase when breeding with non-preferred males. Females thus benefited from being choosy, but this was not directly attributable to avoidance of costly male repeated mating.Funding was provided by Natural Environment Research Council grant no. NE/I025468 to N.J.R. and A.J.M

Comparison of the fracture resistance of endodontically treated teeth restored with prefabricated posts and composite resin cores with different post lenghts

OBJECTIVE: This study evaluated the fracture strengths of endodontically treated teeth restored with prefabricated posts with different post lengths. MATERIAL AND METHODS: Thirty freshly extracted canines were endodontically treated. They were randomly divided into groups of 10 teeth and prepared according to 3 experimental protocols, as follows; Group 1/3 PP: teeth restored with prefabricated post and composite resin core (Z250) with post length of 5.0mm; Group 1/2 PP and Group 2/3 PP: teeth restored with prefabricated post and composite resin core (Z250) with different combinations of post length of 7.5mm and 10mm, respectively. All teeth were restored with full metal crowns. The fracture resistance (N) was measured in a universal testing machine (crosshead speed 0.5mm/min) at 45 degrees to the tooth long axis until failure. Data were analyzed by one-way analysis of variance (alpha=.05). RESULTS: The one-way analysis of variance demonstrated no significant difference among the different post lengths (P>;.05) (Groups 1/3 PP = 405.4 N, 1/2 PP = 395.6 N, 2/3 PP = 393.8 N). Failures occurred mainly due to core fracture. CONCLUSIONS: The results of this study showed that an increased post length in teeth restored with prefabricated posts did not significantly increase the fracture resistance of endodontically treated teeth

Alpha shapes: Determining 3D shape complexity across morphologically diverse structures

Background. Following recent advances in bioimaging, high-resolution 3D models of biological structures are now generated rapidly and at low-cost. To utilise this data to address evolutionary and ecological questions, an array of tools has been developed to conduct 3D shape analysis and quantify topographic complexity. Here we focus particularly on shape techniques applied to irregular-shaped objects lacking clear homologous landmarks, and propose the new ‘alpha-shapes’ method for quantifying 3D shape complexity. Methods. We apply alpha-shapes to quantify shape complexity in the mammalian baculum as an example of a morphologically disparate structure. Micro- computed-tomography (μCT) scans of bacula were conducted. Bacula were binarised and converted into point clouds. Following application of a scaling factor to account for absolute differences in size, a suite of alpha-shapes was fitted to each specimen. An alpha shape is a formed from a subcomplex of the Delaunay triangulation of a given set of points, and ranges in refinement from a very coarse mesh (approximating convex hulls) to a very fine fit. ‘Optimal’ alpha was defined as the degree of refinement necessary in order for alpha-shape volume to equal CT voxel volume, and was taken as a metric of overall shape ‘complexity’. Results Our results show that alpha-shapes can be used to quantify interspecific variation in shape ‘complexity’ within biological structures of disparate geometry. The ‘stepped’ nature of alpha curves is informative with regards to the contribution of specific morphological features to overall shape ‘complexity’. Alpha-shapes agrees with other measures of topographic complexity (dissection index, Dirichlet normal energy) in identifying ursid bacula as having low shape complexity. However, alpha-shapes estimates mustelid bacula as possessing the highest topographic complexity, contrasting with other shape metrics. 3D fractal dimension is found to be an inappropriate metric of complexity when applied to bacula. Conclusions. The alpha-shapes methodology can be used to calculate ‘optimal’ alpha refinement as a proxy for shape ‘complexity’ without identifying landmarks. The implementation of alpha-shapes is straightforward, and is automated to process large datasets quickly. Beyond genital shape, we consider the alpha-shapes technique to hold considerable promise for new applications across evolutionary, ecological and palaeoecological disciplines

Covetable Corpses and Plastic Beetles - The Socioecological Behavior of Burying Beetles

Among-individual variation in behavioral plasticity-the modification of behavior in response to changes in environment experienced by individuals-is increasingly recognized as an important, but relatively poorly understood, feature of organisms that facilitates adaptation to environmental change. It is expected to evolve when there is rapidly fluctuating or directional environmental change during the lifetime of individuals. This is particularly likely to occur in the context of reproductive behaviors, when the outcomes of unpredictable social interactions with other individuals during mating and parental care determine how selection acts on males and females and mating systems evolve. To better understand patterns of mating and parental care and organismal adaptation to environmental change, we need to know why there is so much variation in behavioral plasticity between and within species. Here we address this question using burying beetles as a model. Burying beetles have unusually variable, facultatively expressed, modes of parental care and variation between the sexes and among individuals in the plasticity of reproductive behaviors. We present evidence to show that variation in male plasticity of mating behavior is a key driver of the evolution of patterns of parental care in Nicrophorus vespilloides burying beetles. More generally, we conclude that behavioral plasticity in burying beetles, and likely other taxa, has evolved as a consequence of a resource requirement bottle-neck (niche specialization) in combination with highly unpredictable availability of such suitable resources and the social unpredictability that arises as a result: constraint is the mother of plastic invention.The burying beetle research has been supported by funding from the Natural Environment Research Council (grants NE/I025468/1 and NE/H003738/1 and studentship NE/1528326/1). We are most grateful to all the colleagues that we have worked with on the beetles, including: Kyle Benowitz, Lisa Berry, Mauricio Carter, Emma Davey, Megan Head, Camilla Hinde, Eleanor Jordan, Victoria Lee, Geoffrey Mazué, Allen Moore, Tom Tregenza, and Cam Williams. In addition, we particularly thank Suzanne Alonzo, Allen Moore, Andy Russell, and Alastair Wilson for discussions and collaborations that have shaped some of the ideas on plasticity presented in this review. Thanks also to Cosawes Park in Cornwall for permission and logistical support to collect and study beetles on their land

Burying beetles.

PublishedJournal ArticleWhat are they? Burying beetles are members of the coleopteran family Silphidae (the carrion beetles) of the genus Nicrophorus. There are approximately 75 species in this Northern hemisphere genus. As for most other silphids, the use of vertebrate carrion is an essential part of a burying beetles life. But unlike other silphids, which use carrion primarily as an adult food source or somewhere to lay eggs, Nicrophorus beetles bury the carcasses. This ‘grave-digging’ behaviour gives them their common English name: Sexton beetles