Morph‐dependent form of asymmetry in mandibles of the stag beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Abstract 1. The form of asymmetry in bilateral organs usually follows the same pattern within single populations. However, some exceptions may occur when a population consists of different phenotypes that are from different ontogenic backgrounds and under different selective pressures. We investigated the asymmetric patterns of mandibles of larvae, females, and males in the stag beetle Prosopocoilus inclinatus. 2. Larval mandibles exhibited directional asymmetry both in length and cross direction, whereas female mandibles showed directional asymmetry in cross direction. These asymmetric structures might be more effective in cutting wood fibres. 3. For the relation of male mandible length to body size, a model with a switch point showed a better fit to the data than a convex curve model. This shows that the males are dimorphic with two distinct morphs. 4. The form of asymmetry in male mandible length differed between the morphs. The smaller males exhibited left‐biased directional asymmetry in common with larvae, whereas the larger males exhibited fluctuating asymmetry. 5. This is a novel finding of a morph‐dependent asymmetry. The morph‐dependent asymmetry in males may be as a result of different selection on each morph or a developmental constraint from larval mandibles to adult ones.     

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     

Distinctive developmental variability of genital parts in the sexually dimorphic beetle, Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Recent comparative studies have revealed that the rapid diversity of genitalia is closely related to sexual selection and that genital development interacts with the development of different body parts. Hypotheses about developmental stability due to selection to genital parts were tested by estimating allometric relations in a sexually dimorphic stag beetle Prosopocoilus inclinatus . All genital parts of males scaled to body size with a slope of less than 1 and all but the median lobe (male intromittent organ) showed smaller variability than other body parts. This supported the 'one-size-fits-all' hypothesis, which suggests broad copulation opportunity by males of any size with females within a population. Nevertheless, we found large variation among different genital parts in coefficients of variation and in values of the switch point where the allometric relations varied significantly. These results strongly support the view that developmental trajectories of genital traits are not necessarily integrated. Among the genitalic traits, male intromittent organ and female genitalia exhibited large variability, suggesting a high responsiveness to the selective regimes and physical interaction during copulation. This may account for rapid diversification of genital morphology, even in closely-related populations in beetle species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 573–581.     

Effect of morph types, body size and prior residence on food-site holding by males of the male-dimorphic stag beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Mate-securing tactics of small males in male-polymorphic species exhibiting male–male combat is an important issue in behavioral ecology. While most studies have focused on the outcomes of such combat encounters, the holding of a mating resource like a feeding site has a greater impact for obtaining reproductive success. We examined the effects of the prior residence at a feeding site on resource acquisition in the male-dimorphic stag beetle, Prosopocoilus inclinatus. More than 70 % of encounters did not result in combat. While larger males tended to occupy a food site after a combat, smaller males with prior residence tended to occupy food sites when no fighting occurred. Morph types or body size have no effect on the occurrence of combat, meaning that small males do not hesitate to fight with large males. These findings show that, under experimental conditions, the prior residence has a positive effect to hold food site in P. inclinatus.     

Taxonomic revision of Prosopocoilus gracilis(Saunders,1854) and its allied species from China(Coleoptera:Lucanidae)

Prosopocoilns gracilis(Saunders,1854) and its seven allied species from China are revised.Lectotype,paralectotypes of P.gracilis(Saunders,1854) and P.denticulatus(Boileau,1901) are designated.The doubtful species,P.piceipennis(Westwood,1855) is evidently confirmed as a valid species with verification of its type depository.Two highly similar species,P.crenulidens(Fairmaire,1895) and P.denticulatus(Boileau,1901) are compared in detail for correcting longstanding wrong diagnoses of them.P.similis Schenk,2009 is illustrated with male genitalia firstly.Two recently-published species,P.andreasi Schenk,2009 and P.katsurai Fujita,2010 are treated as new junior synonyms.All the five valid species in this paper are redescribed,with illustrations of adult(including type)habitus,male and female genitalia if they are available.A key to large males is provided for identifying them briefly.     

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.     

Active female courtship behavior and male nutritional contribution to female fecundity in Bruchidius dorsalis (Fahraeus) (Coleoptera: Bruchidae)

Mating behavior and the male's contribution to female fecundity were studied in the bean weevil Bruchidius dorsalis (Fahraeus) (Coleoptera: Bruchidae) in comparison with two other species, Callosobruchus chinensis (which infests stored beans) and Kytorhinus sharpianus (which feeds on wild legumes). Only females of B. dorsalis showed multiple mating and characteristic precopulatory behavior that appeared to solicit the male's nutritious secretion. In contrast, all females of the other two species did not copulate multiply and did not show such precopulatory behavior. In B. dorsalis, the decrement of male body weight just after copulation indicated that seminal fluid weighing as much as approximately 7% of the male's body weight was transferred to the female. Fecundity was more than eight times higher in females that had copulated ten times than in females that had copulated only once, indicating that males paid most of the nutritional cost of egg production. These facts suggest that the sex role is reversed in B. dorsalis. Received: May 22, 1998 / Accepted: July 19, 1999     

Nitrogen fixation in the stag beetle, Dorcus (Macrodorcus) rectus (Motschulsky) (Col., Lucanidae)

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.     

Abiotic variables dictate the best monitoring times for the endangered Table Mountain stag beetle (Colophon westwoodi Gray 1832, Coleoptera: Lucanidae)

There is often a lack of basic ecological data needed to implement effective conservation management programmes for endangered arthropods. This is particularly true for the highly localized and rare Colophon, a genus of beetles of which all members are narrow range endemics of conservation concern. The genus is confined to the highest mountain peaks in the Cape Floristic Region of South Africa and each of the 17 known species is endemic to a particular mountain or range. We investigated the influence of selected abiotic variables on adult Colophon westwoodi activity in Table Mountain National Park, South Africa, which will aid the development of an effective monitoring programme. Weekly surveys conducted on Table Mountain showed that adult numbers peaked during early summer. Adults were active at dusk on clear days, although earlier when misty. Unexpectedly, relative humidity and air temperature had no significant effect on Colophon abundance, while illuminance was the most important predictor. Contrary to general consensus, C. westwoodi activity is not strictly crepuscular but also appears nocturnal. Thus, Colophon monitoring programmes need to be conducted in early summer after sunset and monitoring can continue later than previously assumed. Little is still known about C. westwoodi population size, area of occupancy, general behaviour and the impact tourists have on it, or indeed why the genus is restricted to mountain tops. Therefore this research is important for the design of monitoring protocols for this flagship species, while at the same time directing future research.     

Phylogenetic position of a threatened stag beetle, Lucanus datunensis (Coleoptera: Lucanidae) in Taiwan and implications for conservation

Among nine endemic Lucanus beetles in Taiwan, L. datunensis is the island’s smallest and most threatened species. It currently exists as only one population located in tall grasslands of Mt. Datun in the Yangmingshan National Park. Given the isolated population, unique subtropical grassland, and the threats resulting from human activities, L. datunensis raises immediate conservation concern for its long-term survival. Phylogenies reconstructed from combined mitochondrial cytochrome oxidase subunit 1 (1310 bps) and nuclear wingless (436 bps) genes were resolved and placed L. datunensis as a phylogenetically distinct species sister to L. fortunei from China. All 13 examined individuals of L. datunensis shared just one mitochondrial haplotype suggesting extremely low mitochondrial DNA diversity and a small effective population size. L. datunensis and morphologically closest L. miwai were distantly related and appear to have evolved in parallel the life history traits of a small body size and diurnal mate-searching behavior. We hypothesize that these habitat-associated characters are convergent adaptations that have evolved in response to shifts from forests to grasslands.     

Spermatophore expulsion in the carrion beetle Silpha perforata (Coleoptera: Silphidae)

Silphinae (Coleoptera: Silphidae) is an abundant decomposer that plays important roles in the ecosystem. However, there is little information about the life history of this taxon. We found sperm displacement behavior in carrion beetle Silpha perforata. Copulating males bit the female's antenna strongly and inserted the penis into the partner's genital organ more than once. We found a white substance on the tip of penis during copulation. We examined whether this white substance is a previous male's spermatophore, which was removed from the mating partner. When females were dissected just after mating, the same substance that often presents on the penis of mating males was found in the bursa copulatrix of females, although the bursa copulatrix of virgin females was empty. Male behavior during copulation with females of different mating history was also observed to confirm that the removal of spermatophores was observed only in copulation with females that have the spermatophores of previous males. Consequently, we estimated that S. perforata males removed spermatophores of previous males from mating partners. In addition, we dissected the males frozen during copulation, and inspected the penis morphology. This observation revealed that the apical part of the penis was usually hidden in the basal part of penis, but expanded and appeared during insertion. This apical part had many spines, which play an important role in sperm displacement and sexual conflict in some species. These results indicate that there is the sperm competition in S. perforata. This is the first report on sperm competition in Silphinae.     

Bark-carving behavior of the Japanese horned beetle Trypoxylus dichotomus septentrionalis (Coleoptera: Scarabaeidae)

The Japanese horned beetle, Trypoxylus dichotomus septentrionalis, usually aggregates at the wounds caused by boring insects on Quercus oak trees to feed on exuding sap. Here, a unique behavior of the beetle, which enables it to independently obtain sap without any activities of other insects, is reported. Beetles carved intact bark of Fraxinus griffithii, where no sap was exuding, using clypeus. As a result of carving behavior, bark was stripped, and sap exuded from these scars temporarily, which beetles immediately began to lick. A digital video image relating to the article is available at ().     

贵州省黔东南州锹甲科(鞘翅目:金龟总科)昆虫调查初报

对贵州省黔东南州的锹甲科Lucanidae(鞘翅目:金龟总科)昆虫进行调查研究,查视锹甲科昆虫标本100余号,鉴定得到锹甲8属19种(亚种),其中包括15个贵州新纪录种(亚种).所有查视标本保存在凯里学院昆虫标本室.     

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.     

The importance of streamside sandbars to ground beetle (Coleoptera, Carabidae) communities in a deciduous forest

We used pitfall traps to sample ground beetles on sandbars along a small woodland stream and in the adjacent floodplain forest (Oglethorpe Co., GA, USA). We captured a total of 1,477 ground beetles representing 41 species. Twenty-two species were exclusive to sandbars, while eight were found only in the forested habitat. Ground beetles were captured in significantly greater numbers from sandbars, especially Brachinus janthinipennis and Omophron americanum. The B. janthinipennis record represents a new state record for the species. This study demonstrates that many unique species can be found in specialized microhabitats and emphasizes the need for biodiversity assessment surveys to include a wide range of these microhabitats within a survey area. In addition, it appears that many generalist forest species might use sandbars seasonally to exploit available resources. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Electraesalopsis beuteli gen.& sp.nov., the first lucanid beetle from the Cretaceous Burmese amber (Coleoptera: Scarabaeoidea)

The first stag beetle found in Cretaceous Myanmar amber,Electraesalopsis beuteli Bai,Zhang & Qiu gen.& sp.nov.,is described and illustrated on the basis of one well-preserved specimen.This specimen provides more detailed morphological characters compared with compression fossils,e.g.the highly setose antennae and the punctured elytra.However,the systematic position of the new genus is still questionable and we provisionally place it as Lucanidae incertae sedis.In addition,the discovery ofElectraesalopsis Bai,Zhang & Qiu gen.nov.also indicates that there is very high biodiversity in Burmese amber.The new discoveries in future studies on Burmese amber will provide more information and improve our knowledge of biogeography and early evolution of the Lucanidae.     

Captive breeding reveals larval performance and adult body size differences between two geographical populations of the stag beetle Aegus chelifer chelifer (Coleoptera: Lucanidae)

Stag beetles usually have great intraspecific variation in their body sizes, which can be affected by both environmental and genetic factors. However, direct studies on wild-caught specimens may be insufficient to clarify such variation due to the confounding effects of ecological variance in natural habitats. To evaluate this, the stag beetle Aegus chelifer chelifer MacLeay, 1819 was collected from within two localities (Bangkok metropolitan area and Chanthaburi province) in Thailand and then reared under the same condition to investigate the differences in morphological characteristics between the wild-caught and captive-bred beetles and between the two geographical populations. Narrow-sense heritabilities (h²) of the observed traits in adults were not significant. Variation in the body size of captive-bred specimens was less than in the wild-caught specimens and the overlap of the body size variation between the two populations was lower in the captive-bred beetles. The Chanthaburi population had a significantly larger body size than the Bangkok population. Allometric slopes and intercepts were also significantly different between the two geographic populations. Captive-bred larvae showed similar relative growth rates, but male larvae from the Chanthaburi population had a longer feeding period, and so a larger adult body size, than those from the Bangkok population. The differences between the two populations could be explained by adaptation through larval performances and body size in order to respond to their habitats.     

A review of the primary types of the Hawaiian stag beetle genus Apterocyclus Waterhouse (Coleoptera,Lucanidae, Lucaninae), with the description of a new species

The species of the Hawaiian stag beetle genus Apterocyclus Waterhouse (Coleoptera: Lucanidae) are reviewed following an examination of all primary types. Although the continued existence of the species is unknown and some possibly are extinct there are five recently extant species, including one species that is described here as new. The holotypes for all available names are pictured, and synonymies discussed and updated. Lectotypes are designated for Apterocyclus honoluluensis Waterhouse and A. munroi Sharp. A key to species and a revised catalog for the genus are provided.     

Female dimorphism in Japanese diving beetle Dytiscus marginalis czerskii (Coleoptera: Dytiscidae) evidenced by mitochondrial gene sequence analysis

Three species of the Japanese diving beetle Dytiscus have been identified: D. dauricus Gebler, 1832; D. marginalis czerskii Zeitzev, 1953; and D. sharpi. At present, the latter consists of the subspecies D. sharpi sharpi Wehncke, 1875 and D. sharpi validus Régimbart, 1899 based on the comparative data of mitochondrial DNA cytochrome‐c oxidase I (COI) sequences. Many Dytiscus species have smooth and grooved elytra, which are female dimorphic traits. For many years it has been thought that Japanese D. marginalis czerskii has a single morph, that is, only grooved females, although there were some collecting reports of smooth females occurring at the foot of Mt. Chokaisan in Yamagata and Akita Prefectures. However, the population of smooth females (smooth population) has not yet been identified by DNA markers. To understand the species status of the smooth population, we sequenced 769 bp of COI of a male derived from a smooth mother insect and compared it with the sequence from a known grooved female. The sequences of 769 bp of the COI gene in the smooth population were identical to that in the grooved female, indicating that Japanese D. marginalis czerskii has female dimorphic traits.     

Variation in the number of sperm transferred during mating among males of the Colorado potato beetle (Coleoptera: Chrysomelidae)

We suggest an explanation for the bimodal distribution in the number of sperm transferred into the female during mating by males of the Colorado potato beetle (Leptinotarsa decemlineata). While in most matings females receive 20,000 sperm, in others they receive less than 1000. A variety of potential proximate causes for this bimodal distribution including experimental procedures and several physiological and behavioral causes were examined: male body size, female body size, male mating history, socio-sexual environments, duration and number of mounts, and duration and number of copulations. Surprisingly, none of these explained the observed distribution. However, we did find that the total time spent in copulation was significantly less variable among males that transferred more than 20,000 sperm than among males that passed less than 1000 sperm. In addition, males transferring large amounts of sperm resulted in proportionally more sperm inside the female’s spermatheca within a short period of time than males passing less than 1000 sperm. The lack of pattern in male sperm ejaculate suggests definite differences in sperm delivery tactics and may be related to the female condition rather than the male.