Abstract. Both ornaments and weapons of sexual selection frequently exhibit prolific interspecific diversity of form. Yet, most studies of this diversity have focused on ornaments involved with female mate choice, rather than on the weapons of male competition. With few exceptions, the mechanisms of divergence in weapon morphology remain largely unexplored. Here, we characterize the evolutionary radiation of one type of weapon: beetle horns. We use partial sequences from four nuclear and three mitochondrial genes to develop a phylogenetic hypothesis for a worldwide sample of 48 species from the dung beetle genus Onthophagus (Coleoptera: Scarabaeidae). We then use these data to test for multiple evolutionary origins of horns and to characterize the evolutionary radiation of horns. Although our limited sampling of one of the world’s most species-rich genera almost certainly underestimates the number of evolutionary events, our phylogeny reveals prolific evolutionary lability of these exaggerated sexually selected weapons (more than 25 separate gains and losses of five different horn types). We discuss these results in the context of the natural history of these beetles and explore ways that sexual selection and ecology may have interacted to generate this extraordinary diversity of weapon morphology.

Abstract: In a variety of organisms morphological variation is discrete rather than continuous. Discrete variation within a sex has attracted particular interest as it is thought to reflect the existence of alternative adaptations to a heterogeneous selection environment. The beetle Onthophagus taurus shows a dimorphism for male horns: males that exceed a critical body size develop a pair of long, curved horns on their heads, while smaller males remain hornless. In this study we report on the alternative reproductive tactics used by males with these two morphologies, and present experimental and behavioural data suggesting that these alternative tactics selectively favour discretely different male phenotypes. Horned males aggressively defended tunnel entrances containing breeding females. Fights involved the use of horns, and males with longer horns were more likely to win fights. In contrast, hornless males employed nonaggressive sneaking behaviours when faced with competitively superior males. Sneaking behaviours appeared to require high degrees of manoeuvrability inside tunnels to access and mate with females despite the presence of a guarding male. Comparisons of running performances of males with identical body sizes but different horn lengths suggest that the possession of horns reduces male agility inside tunnels. Thus, horn possession confers a clear advantage to males using fighting behaviours to access females, whereas hornlessness may be favoured in males that rely primarily on sneaking behaviours. Combined, the two alternative reproductive tactics used by male O. taurus appear to favour opposite horn phenotypes, which may explain the paucity of intermediate morphologies in natural populations of O. taurus.

Abstract: A survey of 117 species of arthropods and 17 species of vertebrates showed a strong trend for male genitalia to have relatively low static allometric values. This trend contrasts with the allometry of other structures under sexual selection, which usually show
steep allometric slopes. The trend to low allometric genital values is less consistent in mammals than in arthropods. Data not in accord with the previous the “one-size-fits-all” explanation for low allometric slopes in genitalia, which was based on sexual selection by female choice, suggest a more general version that includes both natural selection and sexual selection, and involves both mechanical fit and stimulation. Less-complete data on the female genitalia of arthropods suggest a trend to similar low allometric slopes, and may also be explained by mechanical fit and stimulatory one-size-fits-all arguments.

Abstract: Horned beetles and beetle horns are emerging as amodel system suited to address fundamental questions in evolutionary developmental biology. Here we briefly review the biology of horned beetles and highlight the unusual opportunities they provide for evo-devo research. We then summarize recent advances in the development of new approaches and techniques that are now available to scientists interested in working with these organisms. We end by discussing ways to implement and combine these new approaches to explore new frontiers in evo-devo research previously unavailable to reseachers working outside traditional model organisms.

Abstract: Large males of Golofa porteri possess several striking secondary sexual characteristics: the head has a long, curving horn with serrations along its edges; the prothorax has an even longer, thinner, nearly vertical horn which is smooth and covered on its anterior surface with a thick mat of golden hairs; and the front legs, especially the robust tibiae and tarsi, are monstrously elongate, with the last tarsomeres sporting thick growths of golden hair on their ventral surfaces. Howden and Campbell (1974) observed two struggles between males in nature at apparent feeding sites on long thin stalks of a bamboo-like plant. Wille (1943) also noted G. aegeon (especially males) on sugar cane stalks. This study is a follow-up of these observations to determine the functions of the males’ bizarre secondary sexual structures.

Abstract: To investigate the causes of striking sexual dimorphism in the harlequin beetle (Acrocinus longimanus), we carried out a study of the behavior and morphology of two widely separated populations (in French Guiana and Panama). Males of this species possess greatly elongated forelegs which exhibit strong positive dometry with body size (elytra length). Males use their forelegs in fights with other males and in guarding females and oviposition sites. Field data on fighting and mating success suggest that sexual selection favors large size in males. However, the direct targets of selection were masked by high correlations between traits of the elytra, forelegs, and antennae. Sexual selection appears to be driven by intense male competition to monopolize suitable sites for egg deposition. Female harlequin beetles are highly selective in choosing only recently dead or dying trees for oviposition (Moraceae and Apocynaceae). Despite marked geographic variation in coloration, host trees, and dimate, the two populations did not differ in mean size and extent of sexual dimorphism. We suggest that sexual selection in this species is sufficiently intense to override any effects of differing ecological factors.

Abstract: We discuss a framework for studying the evolution of morphology in insects, based on the concepts of ‘‘phenotypic plasticity’’ and ‘‘reaction norms.’’ We illustrate this approach with the evolution of some of the most extreme morphologies in insects: exaggerated, sexually selected male ornaments and weapons, and elaborate social insect soldier castes. Most of these traits scale with body size, and these scaling relationships are often nonlinear. We argue that scaling relationships are best viewed as reaction norms, and that the evolution of exaggerated morphological traits results from genetic changes in the slope and/or shape of these scaling relationships. After reviewing literature on sexually selected and caste-specific structures, we suggest two possible routes to the evolution of exaggerated trait dimensions: (a) the evolution of steeper scaling relationship slopes and (b) the evolution of sigmoid or discontinuous scaling relationship shapes. We discuss evolutionary implications of these two routes to exaggeration and suggest why so many of the most exaggerated insect structures scale nonlinearly with body size. Finally, we review literature on insect development to provide a comprehensive picture of how scaling relationships arise and to suggest how they may be modified through evolution.

Abstract: Sexual selection has traditionally been divided into competition over mates and mate choice. Currently, models of sexual selection predict that sexual traits are expressed in proportion to the condition of their bearer. In horned beetles, male contest
competition is well established, but studies on female preferences are scarce. Here I present data on male mating success and condition dependence of courtship rate in three species of horn-dimorphic dung beetles, Onthophagus taurus, Onthophagus binodis, and Onthophagus australis. I found that in the absence of male contest competition, mating success of O. taurus and O. australis was unrelated to their horn length and body size, whereas in O. binodis horn size had a negative effect but body size had a positive effect on male mating success. Overall, in O. binodis major morph males had greater mating success than minor morph males. In all three species male mating success was affected by courtship rate, and the courtship rate was condition dependent such that when males were manipulated to be in poor condition they had lower courtship rates than males that were manipulated to be in good condition. My findings provide new insight into the mating systems of horned dung beetles and support an important assumption in indicator models of sexual selection. Key words: condition dependence, horn dimorphism, horned dung beetles, logistic regression, mating success, Onthophagus, selection gradient, sexual selection. 

Abstract. Although it has been the subject of verbal theory since Darwin, the evolution of morphological trait allometries remains poorly understood, especially in the context of sexual selection. Here we present an allocation trade-off model that predicts the optimal pattern of allometry under different selective regimes. We derive a general solution that has a simple and intuitive interpretation and use it to investigate several examples of fitness functions. Verbal arguments have suggested cost or benefit scenarios under which sexual selection on signal or weapon traits may favor larger individuals with disproportionately larger traits (i.e., positive allometry). However, our results suggest that this is necessarily true only under a precisely specified set of conditions: positive allometry will evolve when the marginal fitness gains from an increase in relative trait size are greater for large individuals than for small ones. Thus, the optimal allometric pattern depends on the precise nature of net selection, and simple examples readily yield isometry, positive or negative allometry, or polymorphisms corresponding to sigmoidal scaling. The variety of allometric patterns predicted by our model is consistent with the diversity of patterns observed in empirical studies on the allometries of sexually selected traits. More generally, our findings highlight the difficulty of inferring complex underlying processes from simple emergent patterns.

Abstract: One of the most pervasive ideas in the sexual selection literature is the belief that sexually selected traits almost universally exhibit positive static allometries (i.e., within a sample of conspecific adults, larger individuals have disproportionally larger traits). In this review, I show that this idea is contradicted by empirical evidence and theory. Although positive allometry is a typical attribute of some sexual traits in certain groups, the preponderance of positively allometric sexual traits in the empirical literature apparently results from a sampling bias reflecting a fascination with unusually exaggerated (bizarre) traits. I review empirical examples from a broad range of taxa illustrating the diversity of allometric patterns exhibited by signal, weapon, clasping and genital traits, as well as nonsexual traits. This evidence suggests that positive allometry may be the exception rather than the rule in sexual traits, that directional sexual selection does not necessarily lead to the evolution of positive allometry and, conversely, that positive allometry is not necessarily a consequence of sexual selection, and that many sexual traits exhibit sex differences in allometric intercept rather than slope. Such diversity in the allometries of secondary sexual traits is to be expected, given that optimal allometry should reflect resource allocation trade-offs, and patterns of sexual and viability selection on both trait size and body size. An unbiased empirical assessment of the relation between sexual selection and allometry is an essential step towards an understanding of this diversity.

Abstract. Detailed contest behaviour of the Japanese horned beetle, Trypoxylus dichotomus septentrionalis, was examined in the field. Male-male interactions have three sequences, and in these sequences four behavioural stages were recognized. After two males encountered (Stage 1), they always faced and shoved each other with their horns (Stage 2: ‘Shoving’). Then, if the horn length or body size difference between the contestantswas large, one male began to retreat and was chased by the other male, and the interactions terminated (Stage 4A: ‘Chasing’). If the difference was small, the interactions proceeded to the escalated fighting stage (Stage 3: ‘Pry’), in which two males put their horns under their opponents and push and try to flip them up each other. The interactions,which proceeded to Stage 3, have two ways of termination. If the body size differencewas large, one male was flipped up by the other male, and the interactions was terminated quickly (Stage 4B). If the difference was small, the interactions was not terminated so quickly and continued until one male began to retreat, proceeding to Stage 4A. It is suggested that males with shorter horns relative to the opponents avoid the escalated fighting stage, ‘Pry’, after perceiving the horn length difference during ‘Shoving’, which would be an appraising behaviour. Thus, ‘Shoving’ is the most important stage among all the interaction processes in that the highest proportion of judgement is made here. The great allometric variation of horn length would presently functionmore greatly for enhancing the efficiency of mutual appraisal than that in actual fighting.

Abstract. I conducted a detailed morphological analysis of the Japanese horned beetle Trypoxylus dichotomus septentrionalis to clarify the allometric relationship between horn length and body size and examined its mating success and reproductive behaviour in the field. The relationship between horn and body size was not discontinuous at the switch point body size, but the slope of the linear relationship changed at the switch point. Shape of the allometric relationship was initially steep and became flatten around the switch point in both linear and log scales; that is, minor males showed a positive relationship and major males showed a negative one. Major males gained more mating success than minor males. Within major males, individuals with larger horn or body size had higher mating success than individuals with smaller ones. Within minor males there were no differences in horn and body size between mated and unmated individuals. Although sneak-like behaviours were exhibited by both morphs, it is likely that these behaviours rarely lead to direct benefit. These results suggest that dimorphic allometry of T. dichotomus is consistent with the hypothesis of a continuous reaction norm that meets a ceiling, which restrains further allometric growth.

Abstract. Male competition for mates can occur through contests or a scramble to locate females. We examined the significance of contests for mates in the leaf beetle Chrysomela aeneicollis, which experiences a short breeding season. During peak mating season, 18–52% of beetles are found in male-female pairs, and nearly half of these are copulating. Sex ratios do not differ from parity, females are larger than males, and positive size-assortative mating occurs. Males fight (2–4% of beetles) over access to females, and disruption of mating usually follows these contests. In the laboratory, we compared mating and fighting frequencies for males found in mating pairs (field-paired) and single males placed into an arena with a field-paired female. Mating pairs were switched in half of arenas (new male-female pairs) and maintained in the other half. For 2 days, each male was free to move about and fight; thereafter males were tethered to prevent contests. Mating frequencies were significantly greater for field-paired than single males in both situations. Male size was not related to mating frequency; however, large females received more matings than small ones. These data suggest that males fight for high quality females, but otherwise search for as many matings as possible.

Abstract: Weapons used in male fighting can be costly to males and are often reported to trade off with other characters such as wings or spermatogenic investment. This study investigated whether increased investment into weapons can generate evolutionary changes in mating strategy for armed males. Male flour beetles, Gnatocerus cornutus, have enlarged mandibles that are used in male–male competition. We subjected these weapons to 12 generations of bidirectional selection and found trade-offs between weapons and two other male characters: wing and testis size. In addition, probably as a consequence of the observed changes in investment, dispersal ability and ejaculatory volume differ significantly between the lines. This indicates that the exaggeration of a weapon can be associated with dispersal and ejaculatory strategies. Thus, altered investment into weapons can lead to correlated changes in life-history traits.

Abstract: Male horns in several groups of beetles represent a special class of secondary sexual characters in which condition-dependent, alternate developmental programs produce not only a bimodal horn-size distribution, but also discrete male mating behaviours correlated with these alternate phenotypes. While these intrasexual dimorphisms have recently received theoretical and experimental attention concerning how they are produced and modified, less has been focussed on the macroevo-lutionary behaviour of the ontogenetic mechanism that produces them. The developmental program that produces alternate male morphologies is manifested by a non-linear horn-size allometry that has been noted to vary within and among various taxa according to its shape and position. The purpose of the present study is to produce a preliminary measure of the macroevolutionary behaviour of these allometric characters as a function of defined phylogenetic scale among the rhinoceros beetles of the widespread genus Xylotrupes. A phylogenetic analysis performed herein suggests that Xylotrupes is monophyletic and is composed of six lineages, which are treated as discrete species. The taxon Xylotrupes gideon of previous literature is shown to constitute five species. Explicit rationale, including morphological diagnoses and evidence of reproductive isolation, supports a new, readily testable taxonomic scheme that recognises the following species: Xylotrupes florensis in the Lesser Sunda and Tanimbar Islands, Indonesia; X. meridionalis in Sri Lanka and India; X. ulysses in Sulawesi, Moluccas, Australia, Papua New Guinea and Melanesia; X. pubescens in the Philippines, Sumatra and Sulawesi; X. mniszechi in south-central and south-east Asia and China; and X. gideon in west Malaysia, Borneo and the Indonesian archipelago from Sumatra through the Lesser Sunda Islands. Subspecies are recognised in some of these taxa and are based upon geographic and phylogenetic partitioning. As in other groups of beetles, the sigmoidal allometric relationship of horn size to body size produces bimodal horn-size distributions in the males of all the species of Xylotrupes in which adequate samples were obtained. The present data show that there is more variation in allometric shape but less variation in allometric position in Xylotrupes than in dung beetles of the genus Onthophagus. Moreover, the phylogenetic patterns of variation in horn allometry among the taxa of Xylotrupes indicate that as much variation in allometric position and shape occurs among the subspecies of a single species, X. ulysses, as occurs among the remainder of the species in this genus. Evidence is provided that allometric position in Xylotrupes is responsive to interspecific competition inasmuch as character displacement of body size relative to horn size occurs in newly discerned sympatric populations of X. gideon and X. pubescens zideki. Further, major evolutionary modifications in allometric shape in two subspecies of X. ulysses have apparently occurred independently and involve fundamentally different adaptive mechanisms. These results suggest that modifications in the developmental program that controls male horn dimorphism are a principal feature of diversification in the beetles of the genus Xylotrupes.