Phenotypic plasticity allows organisms to cope with rapid environmental change. Yet exactly when during ontogeny plastic responses are elicited, whether plastic responses produced in one generation influence phenotypic variation and fitness in subsequent generations, and the role of plasticity in shaping population divergences, remains overall poorly understood. Here, we use the dung beetle Onthophagus taurus to assess plastic responses to temperature at several life stages bridging three generations and compare these responses across three recently diverged populations. We find that beetles reared at hotter temperatures grow less than those reared at mild temperatures, and that this attenuated growth has transgenerational consequences by reducing offspring size and survival in subsequent generations. However, we also find evidence that plasticity may mitigate these consequences in two ways: 1) mothers modify the temperature of their offspring's developmental environment via behavioral plasticity and 2) in one population, offspring exhibit accelerated growth when exposed to hot temperatures during very early development (‘developmental programming’). Lastly, our study reveals that offspring responses to temperature diverged among populations in fewer than 100 generations, possibly in response to range‐specific changes in climatic or social conditions.

Many dung beetle communities are characterized by species that share very similar morphological, ecological, and behavioral traits and requirements yet appear to be stably maintained. Here, we document that the morphologically nearly indistinguishable, sympatric, and syntopic tunneling sister species Onthophagus taurus and Onthophagus illyricus may be avoiding competitive exclusion by nesting at remarkably different soil depths. Intriguingly, we also find rapid divergence in preferred nesting depth across native and recently established O. taurus populations. Furthermore, geometric morphometric analyses reveal that both inter- and intraspecific divergences in nesting depth are paralleled by similar changes in the shape of the primary digging appendages, the fore tibiae. Collectively, our results identify preferred nesting depth and tibial shape as surprisingly evolutionarily labile and with the potential to ease interspecific competition and/or to facilitate adaptation to local climatic conditions.

Genitalia are among the fastest evolving morphological traits in arthropods. Among the many hypotheses aimed at explaining this observation, some explicitly or implicitly predict concomitant male and female changes of genital traits that interact during copulation (i.e., lock and key, sexual conflict, cryptic female choice and pleiotropy). Testing these hypotheses requires insights into whether male and female copulatory structures that physically interact during mating also affect each other's evolution and patterns of diversification. Here we compare and contrast size and shape evolution of male and female structures that are known to interact tightly during copulation using two model systems: (a) the sister species O. taurus (1 native, 3 recently established populations) and O. illyricus, and (b) the species-complex O. fracticornis-similis-opacicollis. Partial Least Squares analyses indicated very little to no correlation between size and shape of copulatory structures, both in males and females. Accordingly, comparing shape and size diversification patterns of genitalia within each sex showed that the two components diversify readily - though largely independently of each other - within and between species. Similarly, comparing patterns of divergence across sexes showed that relative sizes of male and female copulatory organs diversify largely independent of each other. However, performing this analysis for genital shape revealed a signature of parallel divergence. Our results therefore suggest that male and female copulatory structures that are linked mechanically during copulation may diverge in concert with respect to their shapes. Furthermore, our results suggest that genital divergence in general, and co-divergence of male and female genital shape in particular, can evolve over an extraordinarily short time frame. Results are discussed in the framework of the hypotheses that assume or predict concomitant evolutionary changes in male and female copulatory organs.

Horn polyphenism and related head shape variation in a single-horned dung beetle: Onthophagus (Palaeonthophagus) fracticornis (Coleoptera: Scarabaeidae) Abstract Horns of Onthophagus beetles are typical examples of phenotypically plastic traits: they are expressed as a function of environmental (nutritional) stimuli, and their reaction norm (i.e. the full set of horn lengths expressed as a response to different degrees of nutritional states) can be either linear or threshold-dependent. Horned males of Onthophagus (Palaeonthophagus) fracticornis (Preyssler, 1790) bear a single triangular cephalic protrusion of vertex carina, which has received phylogenetic support as the most primitive horn shape in the genus. Inter-and intra-sexual patterns of horn expression were studied in O. fracticornis by means of static allometries while associated variations in head shape were assessed using geometric morphometric techniques. The relation between log-transformed measurements of body size and vertex carina supported an isometric scaling in females. On the contrary, a sigmoidal model described better the horn length-body size allometry in males, with a switch point between alternative morphs at a pronotum width of 3.88 mm. Sigmoidal static allometries of horns in Onthophagus populations arise from a threshold-dependent developmental process of horn growth. This process underlies the expression of both plesiomorphic and apomorphic horn shapes in the genus. Given that the single-horn model has been identified as primitive, we propose that such a developmental process giving rise to it may be evolutionarily ancient as well. Horn expression was accompanied by a deformation of the head which makes minor and major morphs appear even more different. Therefore, in this species both horn and head shape expression contribute to male dimorphism.

Fecundity is a fundamental determinant of fitness, yet the proximate developmental and physiological mechanisms that enable its often rapid evolution in natural populations are poorly understood. Here, we investigated two populations of the dung beetle Onthophagus taurus that were established in exotic ranges in the early 1970s. These populations are subject to drastically different levels of resource competition in the field, and have diverged dramatically in female fecundity. Specifically, Western Australian O. taurus experience high levels of resource competition, and exhibit greatly elevated reproductive output compared to beetles from the Eastern US, where resource competition is minimal and female fecundity is low. We compared patterns of ovarian maturation, relative investment into and timing of egg production, and potential trade-offs between ovarian investment and the duration of larval development and adult body size between populations representative of both exotic ranges. We found that the rapid divergence in fecundity between exotic populations is associated with striking differences in several aspects of ovarian development: (1) Western Australian females exhibit accelerated ovarian development, (2) produce more eggs, (3) bigger eggs, and (4) start laying eggs earlier compared to their Eastern US counterparts. At the same time, divergence in ovarian maturation patterns occurred alongside changes in (5) larval developmental time, and (6) adult body size, and (7) mass. Western Australian females take longer to complete larval development and, surprisingly, emerge into smaller yet heavier adults than size-matched Eastern US females. We discuss our results in the context of the evolutionary developmental biology of fecundity in exotic populations.

Abstract. In insects, allometries of exaggerated traits such as horns or mandibles are often considered species specific and constant during a season. However, given that constraints imposed by the advancing season affect the developmental processes of organisms, these allometries may not be fixed, and the switch point between morphs may vary between populations and within populations during a season. The hypothesis of such a seasonal variation in exaggerated traits was tested using the dimorphic males of the beetle Lucanus cervus. The remains of specimens killed by predators were collected along forest tracks from mid May to late August 2008 in a protected lowland forest in northern Italy. The largest beetles were collected in mid May and average size thereafter decreased. Males collected early in the season mostly had large mandibles (i.e. they belonged to the major morph). In contrast, late in the season the probability of finding males with large mandibles was very low. The threshold body size determining morph expression also shifted during the season. Early in the season, the threshold pronotum width for a 50% chance of developing into the major morph was 1.74 cm, whereas later in the season it was 1.90 cm. This shift in the threshold body size was interpreted as the effect of phenotypic plasticity in a population exposed to constraints imposed by the advancing season.

Developmental processes transduce diverse influences during phenotype formation, thereby biasing and structuring amount and type of phenotypic

1. Microbial symbionts play a crucial role in the development, health, and homeostasis of their hosts. However, the eco-evolutionary conditions shaping these relationships and the evolutionary scale at which host-microbiome interactions may diverge warrant further investigation, especially in non-model systems. This study examines the impact of reciprocal gut microbiome transplants between two ecologically very similar, sympatric, and syntopic dung beetle sister species.2. Onthophagus vacca and Onthophagus medius were specifically used to compare the growth, development, and fitness outcomes of individuals that were either (i) reared in the presence of a microbiome provided by a mother of the same species ("self-inoculated"), (ii) forced to develop with a microbiome derived from a heterospecific mother ("cross-inoculated"), or (iii) reared without a maternally transmitted microbiome.3. This study found that individuals reared in the absence of a maternally derived gut microbiome incur detrimental changes in survival, as well as in several metrics signalling normative development. Furthermore, such negative effects are only partly rescued through inoculation with a heterologous microbiome.4. Collectively, this study's results suggest that inoculation with a species-specific, maternally transmitted microbiome is critical for normative development, that the significance of maternally derived microbiota for host survival differs across species, and that the phenotypic outcomes resulting from host-microbiome interactions may diverge even between closely related, ecologically similar host species.