The bright colors of Alpine leaf beetles (Coleoptera, Chrysomelidae) are thought to act as aposematic signals against predation. Within the European Alps, at least six species display a basal color ...of either blue or green, likely configuring a classic case of müllerian mimicry. In this context, intra-population color polymorphism is paradoxical as the existence of numerous color morphs might hamper the establishment of a search image in visual predators. Assortative mating may be one of the main factors contributing to the maintenance of polymorphic populations. Due to the marked iridescence of these leaf beetles, the perceived color may change as the viewing or illumination angle changes. The present study, conducted over three years, involved intensive sampling of a population of Oreina gloriosa from the Italian Alps and applied colorimetry and a decision tree method to identify the color morphs in an objective manner. The tertiary sex ratio of the population was biased in favor of males, suggesting that viviparous females hide to give birth. Seven color morphs were identified, and their frequencies varied significantly over the course of the study. Three different analyses of mating (JMating, QInfomating, and Montecarlo simulations) recognized a general trend for random mating which coexists with some instances of positive and negative assortative mating. This could help explain the pre-eminence of one morph (which would be favored because of positive selection due to positive assortative mating) in parallel with the persistence of six other morphs (maintained due to negative assortative mating).
Many dung beetle species show male horn polyphenism, the ability of males to develop into distinct phenotypes without intermediate forms as a response to the larval growth environment. While males ...with long (majors) and rudimentary (minor) horn have been widely reported in literature, little is known about the existence of individuals with intermediate horn length. Here we investigate the occurrence of intermediates in natural populations of three dung beetle species (Onthophagus furcatus, Copris lunaris and C. hispanus). We analysed the body size-horn length relationship using linear, exponential, and sigmoidal models with different error structures. We inferred the number of individuals in the minor, intermediate, and major groups by combining changepoint analysis and simulation from fitted allometric models. The sigmoidal equation was a better descriptor of the body size-horn length relationship than linear or exponential equations in all the three studied species. Our results indicated that the number of intermediates equals or exceeds the number of minor and major males. This work provides evidence that, at least in the studied species, males with intermediate horn length exist in natural populations. For similar cases we therefore suggest that continuous phenotypic modulation rather than discrete polyphenism can explain variation in male horn allometry.
The gut microbiota of dung beetles comprises bacteria, archaea, and fungi. Most studies have only considered the bacterial element and focused on differences between species reared in the lab. This ...study considered microorganisms from the gut of Trypocopris pyrenaeus and concentrated on differences among wild individuals of an alpine Italian population. We revealed remarkable interindividual variation in the taxonomic composition of microbiota. Core taxa were few, while individuals harboured many exclusive taxa. Therefore, considering only a few individuals to describe the microbiota of a species is not sufficient. The study also revealed that the nMDS ordination of individuals based on prokaryotes was different from that based on fungi, meaning that both groups should be considered in microbiota analyses and that one microbic group cannot be considered a surrogate of the other. We identified many functions potentially provided by the microbiota and found the taxonomic richness of prokaryotes to be positively correlated with their functional richness. The analysis of the functions potentially provided by these microorganisms confirmed that the gut microbiota, in addition to being essential for the health of their host, may also contribute to the effective functioning of the ecosystems of which dung beetles are part.
In the W Italian Alps, an area characterized by considerable environmental complexity, the widespread Platynus were studied by focusing on their species composition. The ecological niche realized by ...the genus in this area encompasses a preference for humid and cold environments, sometimes associated with altitude. Several specimens from private and public collections were investigated by geometric morphometrics, a powerful technique capable of detecting even minor morphological variation. The quantitative analysis was paired to a qualitative survey of anatomical traits. To classify and discriminate species, external traits (head, pronotum, right elytron) and internal structures (male and female genitalia, mouthparts) were evaluated by direct examination and dissection. The results supported the presence of the five species already known from the study area and also helped to identify four new cryptic taxa to which the specific rank was assigned. They are herein described as Platynus maritimus n. sp., Platynus occitanus n. sp., Platynus simonisi n. sp., and Platynus vignai n. sp.
Although personality studies have primarily focused on vertebrates, the evidence showing invertebrates to be capable of displaying personalities has been steadily growing in recent years. In this ...study, we investigated the behavioural repeatability (repetition of a behaviour over time) and behavioural syndromes (a set of correlated behaviours) in
, which is a dung beetle species showing complex sub-social behaviour. We analysed three behaviours (activity, thanatosis and distress call emission) by measuring seven distinct behavioural traits (i.e., three activity-, one thanatosis- and three distress call-related traits). We found moderate to high levels of individual repeatability in all behavioural traits considered. The duration of thanatosis was inversely correlated with two activity traits, hinting a behavioural syndrome for thanatosis and activity, with bolder individuals exhibiting shorter thanatosis and higher locomotor activity in contrast with fearful individuals, which display longer thanatosis and poor locomotor activity. No relationships were found between the behavioural traits and body size or sex. Results of the principal component analysis (PCA) suggested personality differences among individuals. Dung beetles provide an impressive variety of ecosystem services. Since the provision of these services may depend on the personalities represented in local populations and communities, studies on the ecology of personality in dung beetles should be encouraged in future research.
Male horn dimorphism is a rather common phenomenon in dung beetles, where some adult individuals have well-developed head horns (i.e., major males), while others exhibit diminished horn length (i.e., ...minor males). We focused on horn dimorphism and associated head and pronotum shape variations in
. We examined the allometric relationship between horn length (i.e., cephalic and pronotal horns) and maximum pronotum width (as index of body size) by fitting linear and sigmoidal models for both sexes. We then asked whether head and pronotum shape variations, quantified using the geometric morphometric approach, contributed to this allometric pattern. We found that female cephalic and pronotal horn growth showed a typical isometric scaling with body size. Horn length in males, however, exhibited sigmoidal allometry, where a certain threshold in body size separated males into two distinct morphs as majors and minors. Interestingly, we highlighted the same allometric patterns (i.e., isometric vs. sigmoidal models) by scaling horn lengths with pronotum shape, making evident that male horn dimorphism is not only a matter of body size. Furthermore, the analysis of shape showed that the three morphs had similar heads, but different pronota, major males showing a more expanded, rounded pronotum than minor males and females. These morphological differences in
can ultimately have important functional consequences in the ecology of this species, which should be explored in future work.
The small dung beetle tribe Eucraniini includes extremely specialized species that have been defined as “lifters” according to their food relocation behaviour. They are characterized by the presence ...of well-developed expansions on the head and pronotum, which can be included in the large and varied group of horns, whose presence is usually related to complex reproductive tactics. In this study, two closely related species, Anomiopsoides cavifrons and A. heteroclyta, were examined employing traditional and geometric morphometrics to test whether the Eucraniini has polymorphic males that might exhibit different reproductive tactics, as in the sister tribe Phanaeini, for which a male trimorphism was demonstrated. If also present in Eucraniini polyphenism could be considered a plesiomorphy common to the two clades. The inter- and intraspecific shape variation and object symmetry of the head and the scaling relationships between body size and traits were evaluated. Marked interspecific and small intraspecific differences in shape variation, high symmetry, and similar isometric growth patterns were shown in both species. The hypothesis of male polymorphism in Anomiopsoides was thus rejected. Instead, the results supported the alternative hypothesis that Eucraniini lacks male polymorphism, perhaps due to functional constraints affecting the shape of the structures involved in their peculiar food relocating behaviour.
The effects of specific feeding positions upon the horse's shape variations of the back and neck postures as well as the variations of the mandibular angle have never been objectively studied. For ...this reason, geometric morphometrics was applied. Six horses, aged 14 ± 8 years (mean ± standard deviation, SD), were video-recorded while using three different feeding positions: on the ground-control position (CP); neck held 15 ± 3° below withers height with low hay net position (LP); neck held 15 ± 3° above withers height with high hay net position (HP). Data were analyzed using principal component analysis (PCA), multivariate analysis of variance (MANOVA), and partial least squares (PLS). A mixed model procedure was applied to evaluate differences in the magnitude of the mandibular angle. Whilst differences between individual horses were confirmed by canonical variate analysis (CVA), PCA analysis showed that a characteristic feeding position could also be identified on a group level. During the HP hay net position, the back and neck postures as well the mandibular angle were different compared to those exhibited by horses feeding from CP. In LP hay net position, the back posture more closely resembled those exhibited while feeding from CP; however, no similarity between LP and CP was found for neck posture and mandibular angle. Since only a few degrees of variation of the feeding position can influence back and neck postures, this aspect should be further investigated. The right compromise between horse welfare, horse safety, and management practices need to be further explored and long-term effects should be investigated.
A necro-coprophagous new genus tha is widespread in the whole Sub-Saharan Africa was identified within the tribe Onthophagini and named
n.gen. The new genus, which is well characterized by an ...exclusive set of characters, comprises, at present, 26 species. Twenty species were formerly included in the genus
and six were identified and here described as new species:
n.sp.,
n.sp.,
n.sp.,
n.sp.,
n.sp., and
n.sp. A phylogenetic analysis that is based on a combined matrix, including discrete and landmark characters, was done. The landmark characters were tested using the geometric morphometrics techniques before their inclusion in the matrix. One single, fully resolved tree was obtained, with
constituting a distinct, monophyletic clade within Onthophagini, which was clearly separated from the other genera examined here. The biogeographical analysis identified the Central Africa as the ancestral area of the new genus and it mainly accounted for dispersal events leading to the present distribution. The generic rank that is assigned to the taxon is supported by the results of the morphological, phylogenetic, and biogeographical analyses, and by the comparison to the outgroups.
Dung beetle functional ecology has traditionally focused on studying the relation between traits and ecosystem functions in multispecies assemblages, often ignoring the contribution of behavioral ...interactions and trait variability within species. Here we focus on the factors that affect dung removal at an intraspecific level in two horned dung beetle species with dimorphic males (
Onthophagus taurus
and
Onthophagus verticicornis
). By setting treatments for each species with single individuals (one female, F; one major male, M; one minor male, m) or with pairs of individuals (MF, mF, MM, mm, FF), we examined the effect on dung removal of morphological traits (head, pronotum, leg, horn), sex, and interactions between individuals. Our results showed that dung removal at an intraspecific level depended more on sex and behavioral interactions than on the underlying morphological traits, whose effects on dung removal were negligible. Single females generally removed more dung than single males, which suggests that females are more effective than males. In both species, pairs with at least one female (MF, mF, FF) showed high dung removal efficiency, but did not perform differently from the sum of single treatments (M + F, m + f, F + F). This suggests an additive effect: males and females (or two females) join their efforts when they are together. The pairs with only males (MM and mm) removed less dung than the sum of the single individuals (M + M and m + m), which indicates a mutual inhibition of males. In both species, male morphs performed similarly as they removed the same amount of dung. Despite our results are limited to two
Onthophagus
species, we suggest that the intraspecific functional ecology of dung beetles might be more influenced by behavioral interactions and sex rather than by morphological traits.
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