Alternative Male Morphs Research Articles

Weapon allometry and shape variation in the Helm’s stag beetle (Geodorcus helmsi)

Abstract Many animals develop and use exaggerated weapons in contests over access to mating opportunities. Stag beetles (family Lucanidae) show striking sexual dimorphism, where the males have heavily exaggerated mandibles that the females do not possess. In some genera, size and shape can also vary drastically between small and large males resulting in discrete alternative male morphs, described as male dimorphism. The genus Geodorcus is a poorly studied and threatened group of stag beetles that are endemic to New Zealand. Geodorcus helmsi is the most widespread species in this genus and presents an opportunity to test for the presence of both sexual and male dimorphism and investigate the scaling relationships (allometry) of mandibles in this group. We used linear and geometric morphometric analyses on wild populations and museum specimens from Stewart Island, New Zealand, to assess whether G. helmsi exhibits male dimorphism. We found that within a single population, there is no evidence for alternative male morphs, but a gradual shift between smaller and larger individuals. We also found that male mandibles exhibit steep positive allometry in contrast to the negative allometry found in the mandibles of females, as is often found for sexually selected weaponry. Moreover, we found evidence that left and right mandibles of G. helmsi are subtly asymmetric. Our results provide context for the competitive behaviour observed in this species, and give insights into potential trade-offs between smaller and larger individuals of the Helm’s stag beetle.

Equal fitness among alternative mating strategies in a harem polygynous insect.

Alternative mating strategies are widespread among animal taxa, with strategies controlled by a genetic polymorphism (Mendelian strategy) being rarer in nature than condition-dependent developmental strategies. Mendelian strategies are predicted to have equal average fitnesses and the proportion of offspring produced by a strategy should equal the equilibrium proportion of individuals representing the strategy in a population. Developmental strategies are not expected to produce offspring in equilibrium proportions; however, whether the alternative phenotypes should have equal average fitness is debated. The Wellington tree wētā (Hemideina crassidens) (Orthoptera: Anostostomatidae) is a harem polygynous insect in which intense sexual competition has favoured the evolution of three alternative mating strategies that differ in weapon size and the ability to fight for control of harems. Here, we use molecular genotyping to test the hypothesis that the alternative strategies in this species are maintained by having equal relative fitness and that morphs produce offspring in equilibrium proportions. As expected, the average relative fitness of the three strategies did not significantly differ and the proportion of offspring produced by each morph is equal to the frequency of that morph in the population. Our results support the hypothesis that the alternative male morphs in H. crassidens represent Mendelian strategies.

Epigenetic regulation of gonadal and brain aromatase expression in a cichlid fish with environmental sex determination

A fit animal must develop testes or ovaries, with brain and physiology to match. In species with alternative male morphs this coordination of development across tissues operates within sexes as well as between. For Pelvicachromis pulcher, an African cichlid in which early pH exposure influences both sex and alternative male morph, we sequence both copies of aromatase (cyp19a1), a key gene for sex determination. We analyze gene expression and epigenetic state, comparing gonad and brain tissue from females, alternative male morphs, and fry. Relative to brain, we find elevated expression of the A-copy in the ovaries but not testes. Methylation analysis suggests strong epigenetic regulation, with one region specifying sex and another specifying tissue. We find elevated brain expression of the B-copy with no sex or male morph differences. B-copy methylation follows that of the A-copy rather than corresponding to B-copy expression. In 30-day old fry, we see elevated B-copy expression in the head, but we do not see the expected elevated A-copy expression in the trunk that would reflect ovarian development. Interestingly, the A-copy epialleles that distinguish ovaries from testes are among the most explanatory patterns for variation among fry, suggesting epigenetic marking of sex prior to differentiation and thus laying the groundwork for mechanistic studies of epigenetic regulation of sex and morph differentiation.

Alternative male morphs solve sperm performance/longevity trade-off in opposite directions.

Males pursuing alternative reproductive tactics have been predicted to face a trade-off between maximizing either swimming performance or endurance of their sperm. However, empirical evidence for this trade-off is equivocal, which may be due to simplistic assumptions. In the shell-brooding cichlid fish Lamprologus callipterus, two Mendelian male morphs compete for fertilization by divergent means: Bourgeois nest males ejaculate sperm, on average, about six times farther from the unfertilized ova than do parasitic dwarf males. This asymmetry is opposite to the usual situation, in which bourgeois males typically benefit from superior fertilization opportunities, suggesting that nest males' sperm should persist longer than dwarf male sperm. The assumed trade-off between sperm swimming performance and longevity predicts that, in turn, sperm of dwarf males should outperform that of nest males in swimming efficiency. Measurement of sperm performance and endurance reveals that dwarf male spermatozoa swim straighter initially than those of nest males, but their motility declines earlier and their velocity slows down more abruptly. Nest male sperm survives longer, which relates to a larger sperm head plus midpiece, implying more mitochondria. Thus, the trade-off between sperm performance and endurance is optimized in opposite directions by alternative male morphs. We argue that the relative success of alternative sperm performance strategies can be influenced strongly by environmental factors such as the time window between gamete release and fertilization, and the position of gamete release. This is an important yet little understood aspect of gametic adaptations to sperm competition.

Doublesex alters aggressiveness as a function of social context and sex in the polyphenic beetle Onthophagus taurus

Despite sharing nearly the same genome, individuals within the same species can vary drastically in both morphology and behaviour as a function of developmental stage, sex or developmental plasticity. Thus, regulatory processes must exist that enable the stage-, sex- or environment-specific expression of traits and their integration during ontogeny, yet exactly how trait complexes are co-regulated and integrated is poorly understood. In this study, we explore the developmental genetic basis of the regulation and integration of environment-dependent sexual dimorphism in behaviour and morphology in the horn-polyphenic dung beetle Onthophagus taurus through the experimental manipulation of the transcription factor doublesex (dsx). The gene dsx plays a profound role in the developmental regulation of morphological differences between sexes as well as alternative male morphs by inhibiting horn formation in females but enabling nutrition-responsive horn growth in males. Specifically, we investigated whether experimental downregulation of dsx expression affects male and female aggressive and courtship behaviours in two social contexts: interactions between individuals of the same sex and interactions between males and females. We find that dsx downregulation significantly alters aggressiveness in both males and females, yet does so differently for both sexes as a function of social context: dsxRNAi males exhibited elevated aggression towards males but showed reduced aggression towards females, whereas dsxRNAi females became more aggressive towards males, while their aggressiveness towards other females was unaffected. Moreover, we document unexpectedly high levels of female aggression independent of dsx treatment in both wild-type and control-injected individuals. Lastly, we found no effects of dsxRNAi on courtship and mating behaviours. We discuss the role of dsx in the regulation of sex-specific and plastic behaviours, the unexpectedly high levels of aggression of hornless dsxRNAi males in relation to the well-established description of the hornless sneaker phenotype and the potential ecological function of high female aggression.

Healthier or bigger? Trade‐off mediating male dimorphism in the black scavenger fly Sepsis thoracica (Diptera: Sepsidae)

1. Life history trade‐offs emerge when limited resources are allocated to multiple functions of an organism. Under highly competitive conditions trade‐offs can result in alternative phenotypes that differ morphologically and physiologically. Such is the case in insect species that grow under high densities, where competition for resources but also the risk of disease contagion is high, prompting important adjustments in immune response and melanic cuticular pigmentation, with consequent sacrifices in other fitness‐related traits.2. In the present study, the potential trade‐offs between total‐ and active phenoloxidase (PO), body size and body pigmentation in Sepsis thoracica black scavenger flies that show alternative male morphs differing in cuticular pigmentation, and body size were evaluated.3. As expected, small/dark (obsidian) males showed higher total‐PO activity than larger/orange (amber) males. A negative relationship was found between total‐PO activity and body size in females and obsidian but not amber males, suggesting that growth and immunity are more costly for the former. In contrast, density did not affect PO activity, as predicted by the density‐dependent prophylaxis hypothesis, which had not been tested in dipterans before. However, rearing density did affect the body size negatively in females and amber but not obsidian males, showing that male morph is largely determined by condition‐dependent plasticity rather than genes.4. This study provides good evidence that trade‐offs between different life‐history traits can result in alternative resource allocation strategies, even within one species. These strategies can produce strikingly different alternative phenotypes, evincing that there is not only one optimal solution to address fitness optimisation.

Evidence for ontogenetically and morphologically distinct alternative reproductive tactics in the invasive Round Goby Neogobius melanostomus.

Alternative reproductive tactics are characterized by the occurrence of discrete alternative morphs that differ in behavioural, morphological and physiological traits within the same sex. Although much effort has been made to describe the behaviour, morphology and physiology of such alternative morphs, less effort has been invested investigating how much overlap there is in the characteristics of such morphs in natural populations. We studied random population samples of the invasive Round Goby Neogobius melanostomus from five different localities in the river Rhine system in the Netherlands. We found two morphologically and physiologically distinct male morphs which likely represent alternative reproductive tactics. Almost all mature males under 9.35 cm total length had a gonadosomatic index > 3%, suggestive of a sneaker tactic, while nearly all males above 9.35 cm has a gonadosomatic index of < 3%, suggestive of a parental tactic. Cheek size and eye diameter alone were sufficient to distinguish the two morphs. Gonads had a different relationship with size in the two morphs, indicating separate growth trajectories. The gonad mass of sneaker morphs would be ca. 7.5 times as high as the gonad mass of parental morphs of the same total length after extrapolation. Few (9%) intermediates were found, suggesting that the expression of alternative reproductive tactics is determined before the first breeding season. This contrasts with studies on other goby species, which show evidence of plastic tactics that can be affected by social circumstances. We conclude that it is possible to distinguish two alternative male morphs in the Dutch Round Goby population using morphological measurements alone. Although behavioural observations are needed to provide conclusive evidence, the difference in GSI between these morphs indicates that these morphs reflect alternative reproductive tactics.

Transcriptomics of Intralocus Sexual Conflict: Gene Expression Patterns in Females Change in Response to Selection on a Male Secondary Sexual Trait in the Bulb Mite.

Intralocus sexual conflict (IASC) prevents males and females from reaching their disparate phenotypic optima and is widespread, but little is known about its genetic underpinnings. In Rhizoglyphus robini, a mite species with alternative male morphs, elevated sexual dimorphism of the armored fighter males (compared to more feminized scramblers males) was previously reported to be associated with increased IASC. Because IASC persists if gene expression patterns are correlated between sexes, we compared gene expression patterns of males and females from the replicate lines selected for increased proportion of fighter or scrambler males (F- and S-lines, respectively). Specifically, we tested the prediction that selection for fighter morph caused correlated changes in gene expression patterns in females. We identified 532 differentially expressed genes (FDR < 0.05) between the F-line and S-line males. Consistent with the prediction, expression levels of these genes also differed between females from respective lines. Thus, significant proportion of genes differentially expressed between sexually selected male phenotypes showed correlated expression levels in females, likely contributing to elevated IASC in F-lines reported in a previous study.

A supergene determines highly divergent male reproductive morphs in the ruff.

Three strikingly different alternative male mating morphs (aggressive “Independents”, semi-cooperative “Satellites” and female mimic “Faeders”) coexist as a balanced polymorphism in the ruff, Philomachus pugnax, a lek-breeding wading bird1,2,3. Major differences in body size, ornamentation, and aggressive and mating behaviour are inherited as an autosomal polymorphism4,5. We show that development into Satellites and Faeders is determined by a supergene6,7,8 consisting of divergent alternative, dominant, non-recombining haplotypes of an inversion on chromosome 11, which contains 125 predicted genes. Independents are homozygous for the ancestral sequence. One breakpoint of the inversion disrupts the essential Centromere protein N (CENP-N) gene, and pedigree analysis confirms lethality of inversion homozygotes. We describe novel behavioural, testes size, and steroid metabolic differences among morphs, and identify polymorphic genes within the inversion that are likely to contribute to the differences among morphs in reproductive traits.

Androgens regulate sex differences in signaling but are not associated with male variation in morphology in the weakly electric fish Parapteronotus hasemani

Sexually dimorphic signaling is widespread among animals and can act as an honest indicator of mate quality. Additionally, differences in signaling and morphology within a sex can be associated with different strategies for acquiring mates. Weakly electric fish communicate via self-generated electrical fields that transmit information about sex, reproductive state, and social status. The weakly electric knifefish Parapteronotus hasemani exhibits sexual dimorphism in body size as well as substantial within-male variation in body size and jaw length. We asked whether P. hasemani exhibits hormonally mediated sexual dimorphism in electrocommunication behavior. We also asked whether males with short versus long jaws differed significantly from each other in morphology, behavior, hormone levels, or reproductive maturity. Males produced longer chirps than females, but other signal parameters (electric organ discharge frequency; chirp rate and frequency modulation) were sexually monomorphic. Pharmacologically blocking androgen receptors in males reduced chirp duration, suggesting that this sexually dimorphic trait is regulated at least in part by the activational effects of androgens. Males sorted into two distinct morphological categories but did not differ in circulating 11-ketotestosterone or testosterone. Short-jawed males and long-jawed males also did not differ in any aspects of signaling. Thus, chirping and high levels of 11-ketotestosterone were reliably associated with reproductively active males but do not necessarily indicate male type or quality. This contrasts with other alternative male morph systems in which males that differ in morphology also differ in androgen profiles and signaling behavior.

Evidence for alternative male morphs in a Tanganyikan cichlid fish

AbstractTeleost fish show tremendous variation in breeding systems. In particular, indeterminate growth and external fertilization create great disparities in reproductive success among males, which drive the evolution of male reproductive polymorphisms. Here, we present evidence for alternative male reproductive tactics in Neolamprologus modestus, a substrate‐spawning African cichlid fish. We observed two types of males in our study site: (1) males that guarded large territories comprised of smaller subterritories of several females and (2) males that did not hold territories and were vigorously attacked by the guarding males upon intrusion into the guarding male's territory. We hypothesized that these intruder males constitute an alternative male phenotype. To test this hypothesis, we collected both territorial males and these intruder males to determine if there were differences in overall body size, gonad size, age or hormone profiles between the two male phenotypes that would be consistent with alternative reproductive tactics in this species. We also collected guarded offspring from territorial male and female pairs to determine if there was any extra‐pair paternity in N. modestus that could indicate the possibility of alternative reproductive tactics. We found that intruder males were significantly smaller in body size and had significantly larger testes in absolute and relative mass compared with territorial paired males. While we assigned no paternity to any collected intruder males, we found that extra‐pair paternity occurred in eight of the 12 collected broods and accounted for c. 27% of all offspring across all broods. Finally, the two sets of males did not differ significantly in age or in circulating androgen levels, suggesting that individuals may not change morphs within their lifetime, but instead may adopt distinct life history strategies. Collectively, we provide strong evidence that intruder N. modestus males represent an alternative small‐bodied male morph that may practice alternative reproductive tactics.

SELECTION FOR ALTERNATIVE MALE REPRODUCTIVE TACTICS ALTERS INTRALOCUS SEXUAL CONFLICT

Intralocus sexual conflict (IASC) arises when fitness optima for a shared trait differ between the sexes; such conflict may help maintain genetic variation within populations. Sex-limited expression of sexually antagonistic traits may help resolve the conflict, but the extent of this resolution remains a subject of debate. In species with alternative male reproductive tactics, unresolved conflict should manifest more in a more sexually dimorphic male phenotype. We tested this prediction in the bulb mite (Rhizoglyphus robini), a species in which aggressive fighters coexist with benign scramblers. To do this, we established replicated lines in which we increased the proportion of each of the alternative male morphs using artificial selection. After approximately 40 generations, the proportion of fighters and scramblers stabilized at >0.9 in fighter- and scrambler-selected lines, respectively. We then measured several female fitness components. As predicted by IASC theory, female fecundity and longevity were lower in lines selected for fighters and higher in lines selected for scramblers. This finding indicates that sexually selected phenotypes are associated with an ontogenetic conflict that is not easily resolved. Furthermore, we suggest that IASC may be an important mechanism contributing to the maintenance of genetic variation in the expression of alternative reproductive tactics.

Trade-off between resource allocation and acquisition in anadromous adult male Atlantic salmon (Salmo salar L.)

Trade-off between resource allocation and acquisition in anadromous adult male Atlantic salmon (Salmo salar L.)

Genetic mapping of the female mimic morph locus in the ruff

BackgroundRuffs (Aves: Philomachus pugnax) possess a genetic polymorphism for male mating behaviour resulting in three permanent alternative male reproductive morphs: (i) territorial ‘Independents’, (ii) non-territorial ‘Satellites’, and (iii) female-mimicking ‘Faeders’. Development into independent or satellite morphs has previously been shown to be due to a single-locus, two-allele autosomal Mendelian mode of inheritance at the Satellite locus. Here, we use linkage analysis to map the chromosomal location of the Faeder locus, which controls development into the Faeder morph, and draw further conclusions about candidate genes, assuming shared synteny with other birds.ResultsSegregation data on the Faeder locus were obtained from captive-bred pedigrees comprising 64 multi-generation families (N = 381). There was no evidence that the Faeder locus was linked to the Satellite locus, but it was linked with microsatellite marker Ppu020. Comparative mapping of ruff microsatellite markers against the chicken (Gallus gallus) and zebra finch (Taeniopygia guttata) genomes places the Ppu020 and Faeder loci on a region of chromosome 11 that includes the Melanocortin-1 receptor (MC1R) gene, which regulates colour polymorphisms in numerous birds and other vertebrates. Melanin-based colouration varies with life-history strategies in ruffs and other species, thus the MC1R gene is a strong candidate to play a role in alternative male morph determination.ConclusionTwo unlinked loci appear to control behavioural development in ruffs. The Faeder locus is linked to Ppu020, which, assuming synteny, is located on avian chromosome 11. MC1R is a candidate gene involved in alternative male morph determination in ruffs.

Male dimorphism of a neotropical arachnid: harem size, sneaker opportunities, and gonadal investment

Serracutisoma proximum is a harvestman with alternative male morphs. Large males use sexually dimorphic second legs in fights for the possession of territories on the vegetation, where females oviposit. Small males have short second legs and do not fight but rather sneak into the territories and copulate with egg-guarding females. We investigated the presence of male dimorphism across 10 populations of S. proximum, compared gonadal investment between male morphs, and assessed if the distribution of the sneakers is influenced by harem size. In all populations, there was male dimorphism, indicated by the bimodal distribution of the leg II length/body length. Gonadal investment did not differ between morphs and was not affected by male size, second leg length, and morph relative frequency in the populations. We found 361 territories, 90.0% containing 1 male, 9.7% containing 2 males (dyads), and 0.3% containing 3 males. The probability of encountering dyads increased with the number of females present in the territories. Moreover, the proportion of sneakers in territories containing dyads was higher than would be expected by chance. One possible reason for the ubiquity of alternative morphs in S. proximum could be the high mating opportunities experienced by sneakers in spatially structured populations with a resource defense polygyny system. Additionally, the high frequency of successful invasions by sneakers and hence the high sperm competition risk for both morphs may explain the similarity in gonadal investment between male morphs. Key words: alternative mating tactics, resource defense polygyny, sperm competition, testis mass. [Behav Ecol]

Fitness trade‐offs and the maintenance of alternative male morphs in the bulb mite (Rhizoglyphus robini)

Alternative reproductive phenotypes (ARPs) occur across a wide range of taxa. Most ARPs are conditionally expressed in response to a cue, for example body size, that reliably correlates with the status of the environment: individuals below the (body size) threshold then develop into one morph, and individuals above the threshold develop into the alternative morph. The environmental threshold model provides a theoretical framework to understand the evolution and maintenance of such ARPs, yet no study has examined the underlying fitness functions that are necessary to realize this. Here, we empirically examined fitness functions for the two male morphs of the bulb mite (Rhizoglyphus robini). Fitness functions were derived in relation to male size for solitary males and in relation to female size under competition. In both cases, the fitness functions of the two morphs intersected, and the resulting fitness trade-offs may play a role in the maintenance of this male dimorphism. We furthermore found that competition was strongest between males of the same morph, suggesting that fitness trade-off in relation to male size may persist under competition. Our results are a first step towards unravelling fitness functions of ARPs that are environmentally cued threshold traits, which is essential for understanding their maintenance and in explaining the response to selection against alternative morphs.

Reproductive investment of giants and dwarfs: specialized tactics in a cichlid fish with alternative male morphs

Summary 1. Adopting alternative reproductive tactics may require divergent solutions to reproductive competition among individuals of a population. Often investment in reproduction differs substantially between individuals pursuing bourgeois and parasitic tactics, which may result in different trade‐offs and limitations. 2. Here we identify divergent behavioural, morphological and physiological traits of bourgeois and parasitic male morphs in Lamprologus callipterus, a Lake Tanganyika cichlid with an extreme size dimorphism among males. We focus on limiting factors and compare these between large, nest‐building males and dwarf males parasitizing their reproductive effort. 3. Only nest males invest in courtship, and they exhibit much more aggression than dwarf males. In contrast, dwarf males spend 20% of their time feeding, whereas nest males hardly ever feed. 4. Nest males accumulate reserves before breeding and use these up before taking a reproductive break, thereby performing a ‘capital breeder’ strategy. In contrast, dwarf males use assimilated energy immediately for reproduction, thus acting as ‘income breeders’. This is a requirement of their spawning tactic, which only works out with a small and slim body. 5. A field experiment showed that nest males lose weight by their restricted feeding opportunities while holding a nest, which would allow them to hold a territory for 103 days on average. Due to their reproductive investment, however, they held territories only for a mean period of 33 days, which reveals the relative importance of opportunity costs and reproductive expenditure. 6. Nest males are also limited by the requirement to fertilize each egg of a clutch with a separate ejaculate. Their ejaculation rate and the number of sperm released both decline sharply after 5 h, whereas undisturbed spawning lasts 2–4 h longer than that. 7. There is a strong allometric relationship between body mass and gonad weight, with smaller males of both tactics investing disproportionately more in testes than large males. The major limitation of dwarf males is apparently access to spawning females, which is prevented by the monopolization of nest owners and becomes more difficult with increasing size of dwarf males. 8. Our results show that different males in a population may act as capital or income breeders depending on tactic and may face very different limitations, which is a direct result of highly divergent spawning tactics and resulting body sizes. 9. We argue that capital and income breeding are useful concepts to understand divergent life history decisions associated with alternative reproductive tactics, i.e. behavioural polymorphisms within a species and within one sex. It might turn out that in general, bourgeois tactics rather adopt a capital breeding strategy whereas parasitic tactics are inclined to perform as income breeders, due to the diverging constraints faced by these types of reproduction, although we discuss possible exceptions.

Trophic dimorphism in alternative male reproductive morphs of the acarid mite Sancassania berlesei

Presence of alternative male morphologies within a species has traditionally been related to different reproductive behaviors of morphs, and sexually selected weapons, that is, structures often present in a major male morph of a species, have been associated exclusively with competition between males over females. In this study, I show that alternative male phenotypes of the acarid mite Sancassania berlesei, one of the model male dimorphic species, differ in their foraging behaviors and diets. When food is deficient, aggressive fighter males use their weapons--thickened and sharply clawed third pair of legs, to kill conspecific juveniles and females, as well as individuals from at least one other species, which are subsequently consumed. Unarmed and benign scrambler males have no means of feeding in this way. The ecological importance and evolutionary consequences of this trophic dimorphism are unknown, but it might play an important role in fitness trade-offs between alternative male morphologies and, therefore, in evolution of conditional strategies. It might also increase the species' potential for rapid diversification and speciation. Copyright 2010, Oxford University Press.

Pupal remodeling and the evolution and development of alternative male morphologies in horned beetles

BackgroundHow novel morphological traits originate and diversify represents a major frontier in evolutionary biology. Horned beetles are emerging as an increasingly popular model system to explore the genetic, developmental, and ecological mechanisms, as well as the interplay between them, in the genesis of novelty and diversity. The horns of beetles originate during a rapid growth phase during the prepupal stage of larval development. Differential growth during this period is either implicitly or explicitly assumed to be the sole mechanism underlying differences in horn expression within and between species. Here I focus on male horn dimorphisms, a phenomenon at the center of many studies in behavioral ecology and evolutionary development, and quantify the relative contributions of a previously ignored developmental process, pupal remodeling, to the expression of male dimorphism in three horned beetle species.ResultsPrepupal growth is not the only determinant of differences in male horn expression. Instead, following their initial prepupal growth phase, beetles may be extensively remodeled during the subsequent pupal stage in a sex and size-dependent manner. Specifically, male dimorphism in the three Onthophagus species studied here was shaped not at all, partly or entirely by such pupal remodeling rather than differential growth, suggesting that pupal remodeling is phylogenetically widespread, evolutionarily labile, and developmentally flexible.ConclusionThis study is the first to document that male dimorphism in horned beetles is the product of two developmentaly dissociated processes: prepupal growth and pupal remodeling. More generally, adult morphology alone appears to provide few clues, if any, as to the relative contributions of both processes to the expression of alternative male morphs, underscoring the importance of developmental studies in efforts aimed at understanding the evolution of adult diversity patterns.

Horn possession reduces maneuverability in the horn-polyphenic beetle, Onthophagus nigriventris

Alternative male morphologies are common in a wide range of organisms and particularly extreme in horned beetles. Here, large males (majors) commonly develop extravagant weaponry such as horns or enlarged mandibles, whereas small males (minors) develop only rudimentary traits. In some taxa, including the genus Onthophagus, the transition from minors to majors occurs over a very small range of body sizes causing intermediate morphologies to be rare or absent from natural populations. Several studies have shown that majors use horns as weapons during male combat over females and that the possession of horns increases male fighting success, and presumably fitness. However, the advantages of a hornless morphology, if any, have remained elusive. Here the alternative male morphs are examined in the horn-polyphenic beetle Onthophagus nigriventris. In particular, the hypothesis was tested that lack of horns in minors increases their maneuverability inside tunnel systems in which these males sneak matings from major males. Using a simple behavioral assay the effects of horn possession on maneuverability were quantified inside an artificial tunnel. Minors were found to be significantly more mobile compared to majors. No such differences were found in mobility between similarly small and large females, which always lack horns. This suggests that mobility differences observed among male morphs are due to the presence or absence of horns rather than differences in body size. This notion was further supported in a second experiment in which surgical removal of horns significantly improved maneuverability, while subsequent re-attachment of horns reversed this effect. These results suggest that lack of horns increases male maneuverability inside tunnels and may thus be advantageous in the context of the particular social niche inhabited by minor males. The results are discussed in the context of the evolutionary ecology of horn-polyphenic beetles.