Exaggerated allometric structures in relation to demographic and ecological parameters in Lucanus cervus (Coleoptera: Lucanidae)

Enlarged weapons and ornamental traits under sexual selection often show a positive allometric relationship with the overall body size. The present study explores the allometry of mandibles and their supporting structure, the head, in males of the European stag beetle, Lucanus cervus. This species shows a remarkable dimorphism in mandible shape and size that are used by males in intraspecific combats. Stag beetles were captured, measured, weighed, and released in the framework of a capture‐mark‐recapture study. The relationship of mandible length (ML) and head width in respect to the overall body size was described by a segmented regression model. A linear relationship was detected between ML and head width. The scaling relationships for both ML and head width identified the same switchpoint, highlighting the advantages of using combined results of weapons and their supporting structures in such analysis. These results led to a more consistent distinction of males in two morphologies: minor and major. The survival probability of individuals was dependent on the morphological class and was higher for minor males than for major. Elytron length and body mass of the individuals did not show any significant variation during the season. Differences in predatory pressure were detected between morphs by the collection and analysis of body fragments due to the predatory activity of corvids. Morphological differences and shift in demographic and ecological parameters between the two classes suggested that selection continues to favor intrasexual dimorphism in this species throughout a trade‐off mechanism between costs and benefits of carrying exaggerated traits. J. Morphol. 276:1193–1204, 2015. © 2015 Wiley Periodicals, Inc.     

Aspects of variation in the stag beetle Lucanus cervus (L.) (Coleoptera: Lucanidae)

Abstract. Variation in external dimensions of Lucanus cervus (L.) has been studied using 573 specimens of both sexes from one area over a period of 3 years. Allometric relationships between various parts of the body are calculated. The taxonomy of this very variable population is discussed.     

Monitoring presence,abundance and survival probability of the stag beetle,Lucanus cervus,using visual and odour-based capture methods: implications for conservation

A capture-mark-recapture study on the threatened saproxylic beetle Lucanus cervus was carried out in a chestnut (Castanea sativa) woodland located in northern Italy, using visual encounter surveys (VES) as well as aerial flight interception traps and pitfall traps (PT), both of which were baited with different odour lures. In total, 111 males and 25 females were captured, and VES was by far the most efficient method, accounting for 93 % of first captures, and 95 % of all captures. Stag beetles were not significantly attracted to any tested odour, and many PT were damaged by wild boars (Sus scrofa). Flying males were the most frequent adults observed during the season. The use of a net is necessary to capture the stag beetles, in order to evaluate the population parameters and to assess the local status of threat for the species. Capture data revealed that body size (i.e. body weight and elytron length) influences the survival probability of stag beetles, showing a lower survival probability for larger males. Felt-tip pen marking on the ventral sclerites of head and pronotum is a reliable and long lasting method for marking beetles, as proved by the use of an independent marking procedure with a small drill. Assessing the presence and threat status of L. cervus across Europe is urgently needed, and with no efficient odour-based strategy available, collection of adult beetles, dead or alive, by VES is the most reliable way to monitor this emblematic species.     

Size, operational sex ratio, and mate-guarding success of the carrion beetle, Necrophila americana

When male insects guard females until oviposition, the benefitsfrom last-male sperm precedence must outweigh the costs of relinquishingadditional fertilizations. The profitability of guarding isincreased when males guard large, fecund females and when femalesare scarce because fewer fertilizations are sacrificed. However,the male reproductive success is not only determined by theprofitability of guarding but also by his ability to maintainguarding. In this study, we used male carrion beetles (Necrophilaamericana) to examine the effects of sex ratio, male relativesize, and female quality on the ability to guard. First, wepresent a model of mate guarding that explores factors, suchas sperm precedence, sex ratio, male size, and female quality,that influence the profitability of postcopulatory riding. Ourmodel predicts that large N. americana males should preferentiallyguard the largest female only when the sex ratio is male biasedand sperm precedence is above 80%. In contrast, small malesgain little from guarding because they are not likely to maintainit and be the last male to mate. Then, we tested these predictionsby manipulating sex ratio, relative male size, and female quality.All males in equal sex ratio and large males in male-biasedsex ratio guarded females significantly longer than did malesin female-biased sex ratio. In male-biased sex ratio, largemales guarded significantly longer and achieved more takeoversthan small males. Large females were guarded longer. The successof guarding males in this beetle depends on their size relativeto other males and the operational sex ratio.     

When morphological identification meets genetic data: the case of Lucanus cervus and L. tetraodon (Coleoptera,Lucanidae)

The European stag beetle Lucanus cervus, widely distributed across Europe and in some Near East countries, is one of the best known coleopteran species listed in the European Habitats Directive 92/43/EEC and it is considered a flagship species for conservation of saproxylic fauna. Lucanus tetraodon is a closely related species whose geographical distribution is still poorly known and debated. The two species have a sympatric occurrence in central Italy, and in some localities of these areas, many individuals show a mosaic of morphological traits that makes species assignment nearly impossible. We used both mitochondrial and nuclear markers to analyse these specimens. The mitochondrial results evidenced that the two species represent well‐defined genetic entities with mitochondrial DNA introgression. This pattern could be the result of either hybridization or of a convergence of morphological characters under local selective pressures in areas of sympatric occurrence. The nuclear marker was polymorphic across the two species and therefore did not reveal hybridization, even if many are the supports to this phenomenon. The most plausible explanation for this genetic pattern is a very recent divergence of the two species which share a common origin and thus a common wg genotype.     

Body size influences mating success of the Eucalyptus longhorned borer (Coleoptera: Cerambycidae)

Both sexes of adultPhoracantha semipunctata F. (Coleoptera: Cerambycidae) congregate on stressedEucalyptus that are the larval hosts. In a field study, 721 adultP. semipunctata captured on host trees varied considerably in body size with the largest individuals being about twice the length of the smallest. Females that were paired with a mate were similar in size to solitary females, suggesting that the probability of a female being mated was not affected by her size. However, large males had greater success than smaller males in obtaining mates. MaleP. semipunctata rely on antennal contact to locate and identify females on the larval host. Therefore, the rate at which males search for mates is a function of the area swept by their antennae per unit time. Because of their greater antennal spread, large males were able to search for females at double the rate of the smallest males. Large males also dominated in aggressive contests for females. The superior abilities of large maleP. semipunctata in both locating and defending mates account for the influence of body size on mating success.     

Clinal variation in morphology and assortative mating in the soldier beetle, Chauliognathus pennsylvanicus (Goleoptera: Cantharidae)

In north Georgia populations of the soldier beetle, Chauliognathus pennsylvanicus , the length of the elytral spot varies clinally. At the southern end of a 200 km cline the distribution of spot length is unimodal with longer spot lengths predominating while at the northern end of the cline the distribution is bimodal but with shorter spot lengths being more frequent. North of the cline only short elytral spot lengths are observed, while the converse is true south of the cline. The strength of assortative mating on the basis of elytral spot length increases from south to north along the cline, resulting in complete pre-mating isolation between short and long spot length morphs at the north end of the cline. Laboratory mate choice tests indicate that assortative mating in the field is not the result of differential timing of activity or microhabitat choice but rather that it represents a real behavioural preference. Individuals from monomorphic populations on either side of the cline do not mate assortatively in the laboratory, indicating that reproductive isolation has evolved on the cline.     

Diversification in subtropical mountains: phylogeography, Pleistocene demographic expansion, and evolution of polyphenic mandibles in Taiwanese stag beetle, Lucanus formosanus

Pleistocene glacial oscillations have had profound impacts on the historical population dynamics of extant species. However, the genetic consequences of past climatic changes depend largely on the latitude and topography of the regions in question. This study investigates the effect of Pleistocene glacial periods and the Central Mountain Range on the phylogeography, historical demography, and phenotypic differentiation of a montane forest-dwelling stag beetle, Lucanus formosanus (Coleoptera: Lucanidae), which exhibits extensive mandible variations across mountain ranges in subtropical Taiwan. Analyses of mitochondrial (cox1) and nuclear (wg) loci reveal that L. formosanus originated nearly 1.6 million years ago (Mya) in the early Pleistocene period and consisted of geographically overlapping Alishan and Widespread clades. A drastic population expansion starting approximately 0.2 Mya in the Widespread clade likely resulted from altitudinal range shift of the temperate forests, which was closely tied to the arrival of the Riss glacial period in the late Middle Pleistocene. A ring-like pattern of historical gene flow among neighboring populations in the vicinity of the Central Mountain Range indicates that the mountains constitute a strong vicariant barrier to the east-west gene flow of L. formosanus populations. A geographic cline of decreasing mandible size from central to north and south, and onto southeast of Taiwan is inconsistent with the low overall phylogeographic structures. The degree of mandible variation does not correlate with the expected pattern of neutral evolution, indicating that the evolutionary diversification of this morphological weapon is most likely subject to sexual or natural selection. We hypothesize that the adaptive evolution of mandibles in L. formosanus is shaped largely by the habitat heterogeneity.     

Variation in male mating success in the paternal nest brooder, Rhinogobius goby, along a river course

Rhinogobius sp. CB (cross band type) is an amphidromous freshwater goby which receives paternal nest care under stones on river beds. This goby is known to migrate to the upper reaches of rivers as it grows. In the present study, the relationships among male mating success of Rhinogobius sp. CB, male body size, the size frequency distribution of river bed stones and the presence of the sympatric goby Tridentiger brevispinis (which uses nest sites similar to those of Rhinogobius sp. CB) were investigated along the course of the Aizu River, Wakayama Prefecture, Japan. In the lower reach, where T. brevispinis was present and large stones were scarce, the sizes of the nest stones and the egg clusters of male Rhinogobius sp. CB were smaller than those of T. brevispinis. In the middle and the upper reaches, where T. brevispinis was absent and large stones were abundant, males of Rhinogobius sp. CB used larger nest stones than in the lower reach. In both gobies, there were positive correlations among male body size, nest size and egg cluster size in all reaches. Nest-choice experiments carried out in the laboratory, with or without a rival, showed that males of both gobies preferentially used large nest sites and that large males had an advantage in male-male competition for large nest sites. These results indicate that large male body size can increase male mating success and that male mating success increases in the upper reach in this goby. The migration pattern of Rhinogobius sp. CB was also discussed in relation to these findings.     

Flight dimorphism is related to survival,reproduction and mating success in the leaf beetle Oreina cacaliae

1. Alternative life histories may be maintained in populations due to variation in the costs and benefits of the underlying strategies. In this study, potential costs of dispersal by flight were investigated as an alternative life‐history strategy in the mountain‐living chrysomelid beetle Oreina cacaliae. 2. In this species, previous mark–recapture studies showed a dispersal dimorphism in both males and females. While a fraction of the population engages in flight in autumn and spring (in the following referred to as ‘flyers’), the other part does not fly (non‐flyers). Flyers emerge earlier than non‐flyers and feed on a spring host plant before the emergence of the main host plant. 3. In this study, the overwintering and dispersal locations were recorded over 7 years in the field, flyers from the spring host plant were collected, and morphology and lifetime reproductive output and survival of collected flyers and non‐flyers were compared. 4. A potential trade‐off between flight and life‐history traits was observed: flyers were smaller in size, lighter in body mass, had a lower lifetime fecundity and a higher mortality. 5. Mating experiments of field‐caught beetles in the laboratory showed that larger beetles had a higher (multiple) mating success, but there was no evidence for size‐assortative mating. It is hypothesized that one reason for small beetles to disperse by flight might be to escape competition for mates with larger non‐flyers. 6. The overwhelming quantity of beetles found on the spring host every year reveals that the flying strategy is successful, despite the costs and risks.     

Size and mating success in males of the western harvester ant,Pogonomyrmex occidentalis (Hymenoptera: Formicidae)

Mating success in males of the lek mating ant species,Pogonomyrmex occidentalis, increases with increased body size. We estimated the magnitude of the selection coefficients on components of size by collecting males in copula and comparing their morphology to that of males that were collected at the lek but that were not mating. Four characters, body mass, head width, wing length, and leg length, were measured for a sample of 225 mating and 324 nonmating males and 225 females. Significant direct selection favors increased wing length and leg length. Multiple regression of transformed variables (principal components) indicated that the increased mating success of larger males is a function of all four characters. We found no evidence of positive assortative mating on the basis of any individual character or on the multivariate general size variable (the first principal component).     

Size variation and reproductive success in the mosquito Aedes cantans

Abstract. A link between density-dependent larval competition and adult size of the mosquito Aedes cantans was demonstrated in northern England. Ponds containing high larval densities produced smaller larvae which, in turn, resulted in smaller adults at emergence.
In both 1989 and 1990, parous mosquitoes caught at human bait were larger than nulliparous ones, suggesting that larger mosquitoes are more successful at host location and egg-laying and also that they may be longer lived.
Larger mosquitoes produced larger egg clutches than smaller females: however, there was no difference in the size of eggs laid by large and small females.     

Morphological variation associated with dispersal capacity in a tree‐killing bark beetle Dendroctonus ponderosae Hopkins

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Microsatellite loci for the small hive beetle, Aethina tumida, a nest parasite of honey bees

Aethina tumida, a beetle parasite of honey bee colonies, has recently and dramatically expanded its range and now parasitizes honey bees on three continents. Polymorphic microsatellite loci for this beetle species will help map this continuing range expansion, and will also prove useful for exploring the mating system and local gene flow patterns for this important parasite. We describe 15 loci that are polymorphic in both the native and introduced ranges of this species, showing from two to 22 alleles.     

Relationship between size and shape in the sexually dimorphic beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Male characters that are used for male−male combat are often developed and exaggerated, whereas female equivalent characters are vestigial or vanished. In order to assess whether the characters common to both sexes share the same phenotypic variability due to common genetic architecture, we compared males and females of the stag beetle Prosopocoilus inclinatus using recently developed geometric morphometric methods. Elliptic Fourier analysis was used to compare shape variation between male characters (including exaggerated mandibles) and developmentally homologous female characters. A significant positive correlation was found between the size or between the weight of different body parts in both sexes, but a conspicuous difference was detected in the frequency distribution of the weight of all the body parts. Elliptic Fourier analysis demonstrated that there was marked discontinuous variation in mandibles in males, whereas such a discontinuity was not clear in females. The shape of a character in males exhibited some similarity with that of other characters, but this was not found in females. In a character, growth trajectory of shape was significantly affected by both size and weight in males, whereas size and shape tended to vary independently in female characters. These results support the hypothesis that a large sexual dimorphism in variation in shape is due to alleles accumulating in tight linkage with a sex-determining gene.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 219–233     

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Abstract:  Stag beetles are xylophagous insects that feed mainly on dead wood. They play an important role in the decomposition of dead wood in forest ecosystems. Most dead wood contains 1% nitrogen at most. It is suspected that stag beetles can utilize atmospheric nitrogen. We show that the larvae of Dorcus ( Macrodorcus ) rectus exposed to nitrogen reduce acetylene to ethylene in a time-dependent fashion. No reaction was detected with the dead wood or autoclaved larvae, suggesting that living larvae use the reaction for fixing nitrogen. Acetylene reduction to ethylene by larvae increased with incubation time. This effect was not seen using decayed wood only, autoclaved wood only or autoclaved larvae. Acetylene reduction by the larva proceeded at 1.25 ± 0.37 nmol acetylene/h/g (fresh wt), corresponding to the fixation of 0.25  μ g nitrogen per day per larva.     

Body size and mating success inDrosophila willistoni are uncorrelated under laboratory conditions

Mating activity and wing length were investigated in the F₁ progeny ofDrosophila willistoni females collected in the field to examine any possible relationship between body size and mating success. The flies were observed in a mating chamber under laboratory conditions. No significant differences in wing length were observed between copulating and noncopulating flies, and there was no significant correlation between wing length and copulation latency for both males and females. These results therefore suggest that the commonly accepted view that large body size is positively correlated with mating success inDrosophila does not always hold true. The results support the view that the extent of environmentally induced variation in body size may be an important factor in determining whether an association between body size and mating success is observed inDrosophila species.     

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