Heterogeneity in male horn growth and longevity in a highly sexually dimorphic ungulate

In sexually dimorphic ungulates, sexual selection favoring rapid horn growth in males may be counterbalanced by a decrease in longevity if horns are costly to produce and maintain. Alternatively, if early horn growth varied with individual quality, it may be positively correlated with longevity. We studied Alpine ibex Capra ibex in the Gran Paradiso National Park, Italy, to test these alternatives by comparing early horn growth and longevity of 383 males that died from natural causes. After accounting for age at death, total horn length after age 5 was positively correlated with horn growth from two to four years. Individuals with the fastest horn growth as young adults also had the longest horns later in life. Annual horn growth increments between two and six years of age were independent of longevity for ibex whose age at death ranged from 8 to 16 years. Our results suggest that growing long horns does not constrain longevity. Of the variability in horn length, 22% could be explained by individual heterogeneity, suggesting persistent differences in phenotypic quality among males. Research on unhunted populations of sexually dimorphic ungulates documents how natural mortality varies according to horn or antler size, and can help reduce the impact of sport hunting on natural processes.     

Trade-offs during the development of primary and secondary sexual traits in a horned beetle

Resource allocation trade-offs during development affect the final sizes of adult structures and have the potential to constrain the types and magnitude of evolutionary change that developmental processes can accommodate. Such trade-offs can arise when two or more body parts compete for a limited pool of resources to sustain their growth and differentiation. Recent studies on several holometabolous insects suggest that resource allocation trade-offs may be most pronounced in tissues that grow physically close to each other. Here we examine the nature and magnitude of developmental trade-offs between two very distant body parts: head horns and genitalia of males of the horned scarab beetle Onthophagus taurus. Both structures develop from imaginal disklike tissues that undergo explosive growth during late larval development but differ in exactly when they initiate their growth. We experimentally ablated the precursor cells that normally give rise to male genitalia at several time points during late larval development and examined the degree of horn development in these males compared to that of untreated and sham-operated control males. We found that experimental males developed disproportionately larger horns. Horn overexpression was weakest in response to early ablation and most pronounced in males whose genital disks were ablated just before larvae entered the prepupal stage. Our results suggest that even distant body parts may rely on a common resource pool to sustain their growth and that the relative timing of growth may play an important role in determining whether, and how severely, growing organs will affect each other during development. We use our findings to discuss the physiological causes and evolutionary consequences of resource allocation trade-offs.     

A negative association between horn length and survival in a weakly dimorphic ungulate

While all models of sexual selection assume that the development and expression of enlarged secondary sexual traits are costly, males with larger ornaments or weapons generally show greater survival or longevity. These studies have mostly been performed in species with high sexual size dimorphism, subject to intense sexual selection. Here, we examined the relationships between horn growth and several survival metrics in the weakly dimorphic Pyrenean chamois (Rupicapra pyrenaica). In this unhunted population living at high density, males and females were able to grow long horns without any apparent costs in terms of longevity. However, we found a negative relationship between horn growth and survival during prime age in males. This association reduces the potential evolutionary consequences of trophy hunting in male chamois. We also found that females with long horns tended to have lower survival at old ages. Our results illustrate the contrasting conclusions that may be drawn when different survival metrics are used in analyses. The ability to detect trade‐off between the expression of male secondary sexual traits and survival may depend more on environmental conditions experienced by the population than on the strength of sexual selection.     

Mechanical limits to maximum weapon size in a giant rhinoceros beetle

The horns of giant rhinoceros beetles are a classic example of the elaborate morphologies that can result from sexual selection. Theory predicts that sexual traits will evolve to be increasingly exaggerated until survival costs balance the reproductive benefits of further trait elaboration. In Trypoxylus dichotomus, long horns confer a competitive advantage to males, yet previous studies have found that they do not incur survival costs. It is therefore unlikely that horn size is limited by the theoretical cost–benefit equilibrium. However, males sometimes fight vigorously enough to break their horns, so mechanical limits may set an upper bound on horn size. Here, I tested this mechanical limit hypothesis by measuring safety factors across the full range of horn sizes. Safety factors were calculated as the ratio between the force required to break a horn and the maximum force exerted on a horn during a typical fight. I found that safety factors decrease with increasing horn length, indicating that the risk of breakage is indeed highest for the longest horns. Structural failure of oversized horns may therefore oppose the continued exaggeration of horn length driven by male–male competition and set a mechanical limit on the maximum size of rhinoceros beetle horns.     

Age-dependent genetic effects on a secondary sexual trait in male Alpine ibex, Capra ibex

Secondary sexual traits, such as horns in ungulates, may be good indicators of genetic quality because they are costly to develop. Genetic effects on such traits may be revealed by examining correlations between multilocus heterozygosity (MLH) and trait value. Correlations between MLH and fitness traits, termed heterozygosity-fitness correlations (HFC), may reflect inbreeding depression or associative overdominance of neutral microsatellite loci with loci directly affecting fitness traits. We investigated HFCs for horn growth, body mass and faecal counts of nematode eggs in wild Alpine ibex (Capra ibex). We also tested if individual inbreeding coefficients (f') estimated from microsatellite data were more strongly correlated with fitness traits than MLH. MLH was more strongly associated with trait variation than f'. We found HFC for horn growth but not for body mass or faecal counts of nematode eggs. The effect of MLH on horn growth was age-specific. The slope of the correlation between MLH and yearly horn growth changed from negative to positive as males aged, in accordance with the mutation accumulation theory of the evolution of senescence. Our results suggest that the horns of ibex males are an honest signal of genetic quality.     

Climate-dependent allocation of resources to secondary sexual traits in red deer

Fitness in highly polygynous male ungulates is related both to body size, weight and antler size. Males must therefore allocate resources both to growth of the body and growth of the antlers, which may lead to tradeoffs whenever resource levels are in limited supply. Several studies have reported how (absolute) growth of antlers and horns are related to environmental conditions, but few have looked for the relative allocation patterns (i.e. relative to body size and weight). We analyzed how the influence of variation in climate (the North Atlantic oscillations, NAO) and population density affected the allocation of resources to antlers, based on data from 2720 red deer stags two years or older harvested during 1965–2002 along the west coast of Norway. Number of antler tines increased up to six years of age, remained stable until 12 years of age, and then decreased significantly (>12 years, n=45). The NAO was positively related to number of antler tines in prime aged males (6–12 years, n=629), also after controlling for both body size and weight. Our study thus suggests that deer have a higher allocation of resources to antlers relative to body weight during favorable environmental conditions.     

Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic

1. Age-specific survival of 215 males and 117 females of the highly sexually dimorphic Alpine ibex Capra ibex (L.) was assessed from a 21-year capture-mark-recapture (CMR) programme (1983-2004). The study covered two contrasted periods of population performance (high performance from 1983 to 1997 vs. low performance from 1998 onwards). 2. Based on current life-history theories for sexually dimorphic species, we expected that survival should decrease with age in both sexes, female survival should be buffered against environmental variations, male survival should decrease during the low performance period, and adult survival should be lower in males than females during the low performance period. 3. Survival of both sexes was strongly affected by age, with the four age classes (yearling, prime-aged adults of 2-8 years of age, old adults of 8-13 years of age, and senescent adults from 13 years of age onwards) generally reported for large herbivores. 4. Survival of females at all ages, and of yearling and prime-aged males, was buffered against environmental variations and was the same during periods of high and low population performance. The survival of old males decreased in years of low population performance. 5. All marked yearlings (32 females, 56 males) survived to age 2. Survival of prime-aged females (0.996 +/- 0.011) was higher than for other large herbivores, but similarly to other large herbivore species, it declined slowly and regularly with increasing age afterwards. Male survival was 5-15% higher each year than that of males of other large herbivores. Males enjoyed very high survival when prime-aged (0.981 +/- 0.009) and as old adults (high-performance period: 0.965 +/- 0.028, low-performance period: 0.847 +/- 0.032). 6. The very high survival of males, coupled with their prolonged mass gain, suggests a highly conservative reproductive tactic. Male ibex differ from similar-sized herbivores by showing a nearly indeterminate growth in horn size and body mass. By surviving to an advanced age, males may enjoy high reproductive success because of their large size.     

Environmental and evolutionary effects on horn growth of male bighorn sheep

The development of male secondary sexual characters such as antlers or horns has substantial biological and socio‐economic importance because in many species these traits affect male fitness positively through sexual selection and negatively through trophy hunting. Both environmental conditions and selective hunting can affect horn growth but their relative importance remains unexplored. We first examined how a large‐scale climate index, the Pacific Decadal Oscillation (PDO), local weather and population density influenced both absolute and relative annual horn growth from birth to three years of male bighorn sheep Ovis canadensis over 42 years. We then examined the relative influence of environmental conditions and evolution mainly driven by trophy hunting on male horn length at three years of age. Horn growth was positively influenced by low population density and warm spring temperature, suggesting that ongoing climate change should lead to larger horns. Seasonal values of PDO were highly correlated. Horn growth increased with PDO in spring or summer at low density, but was weak at high density regardless of PDO. The interaction between population density and PDO in spring or summer accounted for a similar proportion of the observed annual variation in horn growth (32% or 37%) as did the additive effects of spring temperature and density (34%). When environmental conditions deteriorated, males allocated relatively more resources to summer mass gain than to horn growth, suggesting a conservative strategy favoring maintenance of condition over allocation to secondary sexual characters. Population density explained 27% of the variation in horn length, while evolutionary effects explained 9% of the variance. Thus, our study underlines the importance of both evolution and phenotypic plasticity on the development of a secondary sexual trait.     

Harvest regulations and artificial selection on horn size in male bighorn sheep

Wild sheep in North America are highly prized by hunters and most harvest regulations restrict legal harvest to males with a specified minimum horn curl. Because reproductive success is skewed toward larger males that are socially dominant, these regulations may select against high-quality, fast-growing males. To evaluate potential selective effects of alternative management strategies, we analyzed horn increment measures of males harvested over 28 yr (1975–2003) in 2 bighorn sheep (Ovis canadensis) ecotypes in British Columbia, Canada. Using mixed-effect models we examined variation in hunter selection for horn size, early horn growth, and male age under different harvest regulations (Full Curl, Three Quarter Curl, Any Ram). Under all regulations, males with the greatest early horn growth were harvested at the youngest ages, before the age at which large horns influence reproductive success. Early growth decreased with harvest age and until ≥7 yr of age it was greatest in males harvested under Full Curl regulation. Permit type (General vs. Limited Entry Hunt) and hunter origin (British Columbia Resident vs. Non-Resident) had little effect on horn size of harvested males. Full Curl regulations increased the average age of harvested males by <1 yr relative to Three-Quarter Curl regulations. Age-specific horn measures in the California ecotype harvested under Three-Quarter Curl regulations declined over time but we observed no temporal declines in the Rocky Mountain ecotype, primarily harvested under Full Curl regulations. Management strategies that protect some males with greater early horn growth or provide harvest refuges to maintain genetic diversity are likely to reduce potential for negative effects of artificial selection. © 2010 The Wildlife Society     

When does selective hunting select,how can we tell,and what should we do about it?

Potential evolutionary consequences of selective hunting of mammals are controversial because of limited evidence and important socio‐economic impacts. Several ecological and management variables facilitate evolutionary responses to selection for horn, tusk or antler size, including strong selective hunting pressure; harvest of males with large horns, tusks or antlers before they can breed; unavailable or ineffective sources of unselected immigrants; and age‐dependent relationships between horn, tusk or antler size and male mating success. Plastic responses of male horns, tusks and antlers to environment are probably more common than evolutionary changes. Evidence for evolutionary effects of selective hunting is strong for large mammals where biological characteristics and hunting regulations combine to favour them.     

Parallel Lasers for Remote Measurements of Morphological Traits

Abstract: Animal ecology research could benefit from the measurement of individual morphological traits. In bovids, male horn size often correlates with annual reproductive success, is sensitive to resource abundance, and could be a predictor of survival. However, live captures are costly, involve some risk of injury or substantial disturbance to the animals, and are impossible in many situations. To remotely measure horn growth of free-ranging Alpine ibex (Capra ibex), I designed an aluminum frame that holds parallel laser pointers and a digital camera. I took digital pictures of ibex horns and calculated horn growth based on the fixed distance between the 2 laser points. This simple and accurate technique could benefit many ecological studies that require linear measurements, such as shoulder height, body length, leg length, or fin length. It could also help measure body features (e.g., fur or skin patterns, scars), increasing the reliability of individual photographic identification.     

Male reproductive pattern in a polygynous ungulate with a slow life-history: the role of age, social status and alternative mating tactics

According to life-history theory age-dependent investments into reproduction are thought to co-vary with survival and growth of animals. In polygynous species, in which size is an important determinant of reproductive success, male reproduction via alternative mating tactics at young age are consequently expected to be the less frequent in species with higher survival. We tested this hypothesis in male Alpine ibex (Capra ibex), a highly sexually dimorphic mountain ungulate whose males have been reported to exhibit extremely high adult survival rates. Using data from two offspring cohorts in a population in the Swiss Alps, the effects of age, dominance and mating tactic on the likelihood of paternity were inferred within a Bayesian framework. In accordance with our hypothesis, reproductive success in male Alpine ibex was heavily biased towards older, dominant males that monopolized access to receptive females by adopting the ‘tending’ tactic, while success among young, subordinate males via the sneaking tactic ‘coursing’ was in general low and rare. In addition, we detected a high reproductive skew in male Alpine ibex, suggesting a large opportunity for selection. Compared with other ungulates with higher mortality rates, reproduction among young male Alpine ibex was much lower and more sporadic. Consistent with that, further examinations on the species level indicated that in polygynous ungulates the significance of early reproduction appears to decrease with increasing survival. Overall, this study supports the theory that survival prospects of males modulate the investments into reproduction via alternative mating tactics early in life. In the case of male Alpine ibex, the results indicate that their life-history strategy targets for long life, slow and prolonged growth and late reproduction.     

Is there a trade‐off between horn growth and survival in adult female chamois?

Life‐history theory predicts trade‐offs in energy allocation between different life‐history traits when resources are limited, i.e. certain traits should be negatively correlated. However, individuals differ in their ability to acquire resources, which can lead to positive correlations between traits at the population level. Here, we investigated the consequences of the allocation in horn growth and body mass on survival in a bovid (Rupicapra rupicapra) with capture‐mark re‐sighting data on 161 females. In female ungulates, body mass often covaries positively with demographic performance and the few studies on horn size suggest that this trait could be a signal of individual quality. Thus, we expected to measure positive correlations between the allocation in these traits and female survival. However, body mass was not correlated to female survival and there was only a negative, though marginal, effect of horn growth. Hence, it seems that the allocation in growth is not an indicator of female quality. Future studies could investigate the importance of growth on female reproduction to evaluate its effect on lifetime reproductive success. Moreover, it is important to confirm in other populations our result that suggests a cost of the allocation in horn growth to better understand the presence of horns in female bovids. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 516–521.     

Habitat and Harvesting Practices Influence Horn Growth of Male Ibex

Size-selective harvesting of wild ungulates can trigger a range of ecological and evolutionary consequences. It remains unclear how environmental conditions, including changes in habitat, climate, and local weather conditions, dilute or strengthen the effects of trophy hunting. We analyzed horn length measurements of 2,815 male ibex (Capra pyrenaica) that were harvested from 1995 to 2017 in Els Ports de Tortosa i Beseit National Hunting Reserve in northeastern Spain. We used linear mixed models to determine the magnitude of inter-individual horn growth variability and partial least square path models to evaluate long-term effects of environmental change, population size, and hunting strategy on horn growth. Age-specific horn length significantly decreased over the study period, and nearly a quarter (23%) of its annual variation was attributed to individual heterogeneity among males. The encroachment of pine (Pinus spp.) forests had a negative effect on annual horn growth, possibly through nutritional impoverishment. The harvesting of trophy and selective individuals (e.g., small-horned males) from the entire population increased horn growth, probably because it reduced the competition for resources and prevented breeding of these smaller males. Local weather conditions and population size did not influence horn growth. Our study demonstrates how habitat changes are altering the horn growth of male ibex. We suggest that habitat interventions, such the thinning of pine forests, can contribute to securing the sustainability of trophy hunting. Even in situations where size-selective harvesting is not causing a detectable phenotypic response, management actions leading to the expansion of preferred land cover types, such as grass-rich open areas, can have a positive effect on ungulate fitness. Forest encroachment on open meadows and heterogeneous grasslands is pervasive throughout Mediterranean ecosystems. Therefore, our management recommendations can be extended to the landscape level, which will have the potential to mitigate the side effects of habitat deterioration on the phenotypic traits of wild ibex. © 2020 The Wildlife Society.     

Disease and secondary sexual traits: effects of pneumonia on horn size of bighorn sheep

Secondary sexual traits (e.g., horns and antlers) have ecological and evolutionary importance and are of management interest for game species. Yet, how these traits respond to emerging threats like infectious disease remains underexplored. Infectious pneumonia threatens bighorn sheep (Ovis canadensis) populations across North America and we hypothesized it may also reduce horn growth in male sheep. We assess the effect of pneumonia on horn size in male bighorn sheep using 12 herd datasets from across the western United States that had horn growth and disease data. Disease resulted in 12–35% reduction in increment (yearly) length and 3–13% reduction in total horn length in exposed individuals. The disease effect was prolonged when pathogens continued to circulate in sheep populations. Further, disease likely delays the age at which horns reach ¾-curl and prevents achievement of full-curl. This is further evidenced with 6 of the 12 herds experiencing an increase in average age at harvest following die-off events.     

Relative allocation to horn and body growth in bighorn rams varies with resource availability

Males may allocate a greater proportion of metabolic resourcesto maintenance than to the development of secondary sexual characterswhen food is scarce, to avoid compromising their probabilityof survival. We assessed the effects of resource availabilityon body mass and horn growth of bighorn rams (Ovis canadensis)at Ram Mountain, Alberta, Canada over 30 years. The number ofadult ewes in the population tripled during our study, and theaverage mass of yearling females decreased by 13%. We used theaverage mass of yearling females as an index of resource availability.Yearling female mass was negatively correlated with the bodymass of rams of all ages, but it affected horn growth only duringthe first three years of life. Yearly horn growth was affectedby a complex interaction of age, body mass, and resource availability.Among rams aged 2–4 years, the heaviest individuals hadsimilar horn growth at high and at low resource availability,but as ram mass decreased, horn growth for a given body massbecame progressively smaller with decreasing resource availability.For rams aged 5–9 years, horn growth was weakly but positivelycorrelated with body mass, and rams grew slightly more hornfor a given body mass as resource availability decreased. Whenfood is limited, young rams may direct more resources to bodygrowth than to horn growth, possibly trading long-term reproductivesuccess for short-term survival. Although horn growth of olderrams appeared to be greater at low than at high resource availability,we found no correlation between early and late growth in hornlength for the same ram, suggesting that compensatory horn growthdoes not occur in our study population. Young rams with longerhorns were more likely to be shot by sport hunters than thosewith shorter horns. Trophy hunting could select against ramswith fast-growing horns.     

Resource allocation during ontogeny is influenced by genetic,developmental and ecological factors in the horned beetle,Onthophagus taurus

Resource allocation trade-offs arise when developing organs are in competition for a limited pool of resources to sustain growth and differentiation. Such competition may constrain the maximal size to which structures can grow and may force a situation in which the evolutionary elaboration of one structure may only be possible at the expense of another. However, recent studies have called into question both the consistency and evolutionary importance of resource allocation trade-offs. This study focuses on a well-described trade-off between the horns and eyes of Onthophagus beetles and assesses the degree to which it is influenced by genetic, developmental and ecological conditions. Contrary to expectations, we observed that trade-off signatures (i) were mostly absent within natural populations, (ii) mostly failed to match naturally evolved divergences in horn investment among populations, (iii) were subject to differential changes in F₁ populations derived from divergent field populations and (iv) remained largely unaffected by developmental genetic manipulations of horn investment. Collectively, our results demonstrate that populations subject to different ecological conditions exhibit different patterns of, and differential plasticity in, resource allocation. Further, variation in ecological conditions, rather than canalized developmental mechanisms, may determine whether and to what degree morphological structures engage in resource allocation trade-offs.     

Live fast, die young: trade-offs between fitness components and sexually antagonistic selection on weaponry in Soay sheep

Males are predicted to compete for reproductive opportunities, with sexual selection driving the evolution of large body size and weaponry through the advantage they confer for access to females. Few studies have explored potential trade-offs of investment in secondary sexual traits between different components of fitness or tested for sexually antagonistic selection pressures. These factors may provide explanations for observed polymorphisms in both form and quality of secondary sexual traits. We report here an analysis of selection on horn phenotype in a feral population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, Scotland. Soay sheep display a phenotypic polymorphism for horn type with males growing either normal or reduced (scurred) horns, and females growing either normal, scurred, or no (polled) horns; further variation in size exists within horn morphs. We show that horn phenotype and the size of the trait displayed is subject to different selection pressures in males and females, generating sexually antagonistic selection. Furthermore, there was evidence of a trade-off between breeding success and longevity in normal-horned males, with both the normal horn type and larger horn size being associated with greater annual breeding success but reduced longevity. Therefore, selection through lifetime breeding success was not found to act upon horn phenotype in males. In females, a negative association of annual breeding success within the normal-horned phenotype did not result in a significant difference in lifetime fitness when compared to scurred individuals, as no significant difference in longevity was found. However, increased horn size within this group was negatively associated with breeding success and longevity. Females without horns (polled) suffered reduced longevity and thus reduced lifetime breeding success relative the other horn morphs. Our results therefore suggest that trade-offs between different components of fitness and antagonistic selection between the sexes may maintain genetic variation for secondary sexual traits within a population.     

Effects of harvest,culture, and climate on trends in size of horn-like structures in trophy ungulates

Hunting remains the cornerstone of the North American model of wildlife conservation and management. Nevertheless, research has indicated the potential for hunting to adversely influence size of horn-like structures of some ungulates. In polygynous ungulates, mating success of males is strongly correlated with body size and size of horn-like structures; consequently, sexual selection has favored the development of large horns and antlers. Horn-like structures are biologically important and are of great cultural interest, both of which highlight the need to identify long-term trends in size of those structures, and understand the underlying mechanisms responsible for such trends. We evaluated trends in horn and antler size of trophy males (individuals exhibiting exceptionally large horns or antlers) recorded from 1900 to 2008 in Records of North American Big Game, which comprised >22,000 records among 25 trophy categories encompassing the geographic extent of species occupying North America. The long-term and broad-scale nature of those data neutralized localized effects of climate and population dynamics, making it possible to detect meaningful changes in size of horn-like structures among trophy males over the past century; however, ages of individual specimens were not available, which prevented us from evaluating age-class specific changes in size. Therefore, we used a weight-of-evidence approach based on differences among trophy categories in life-history characteristics, geographic distribution, morphological attributes, and harvest regimes to discriminate among competing hypotheses for explaining long-term trends in horn and antler size of trophy ungulates, and provide directions for future research. These hypotheses were young male age structure caused by intensive harvest of males (H1), genetic change as a result of selective male harvest (H2), a sociological effect (H3), effects of climate (H4), and habitat alteration (H5). Although the number of entries per decade has increased for most trophy categories, trends in size of horn-like structures were negative and significant for 11 of 17 antlered categories and 3 of 8 horned categories. Mean predicted declines during 1950–2008 were 1.87% and 0.68% for categories of trophy antlers and horns, respectively. Our results were not consistent with a sociological effect (H3), nutritional limitation imposed by climate (H4), or habitat alteration (H5) as potential explanations for long-term trends in size of trophies. In contrast, our results were consistent with a harvest-based explanation. Two of the 3 species that experienced the most conservative harvest regimes in North America (i.e., bighorn sheep [Ovis canadensis] and bison [Bison bison]) did not exhibit a significant, long-term trend in horn size. In addition, horn size of pronghorn (Antilocapra americana), which are capable of attaining peak horn size by 2–3 years of age, increased significantly over the past century. Both of those results provide support for the intensive-harvest hypothesis, which predicts that harvest of males has gradually shifted age structure towards younger, and thus smaller, males. The absence of a significant trend for mountain goats (Oreamnos americanus), which are difficult to accurately judge size of horns in the field, provided some support for the selective-harvest hypothesis. One other prediction that followed from the selective-harvest hypothesis was not supported; horned game were not more susceptible to reductions in size. A harvest-induced reduction in age structure can increase the number of males that are harvested prior to attaining peak horn or antler size, whereas genetic change imposed by selective harvest may be less likely to occur in free-ranging populations when other factors, such as age and nutrition, can override genetic potential for size. Long-term trends in the size of trophy horn-like structures provide the incentive to evaluate the appropriateness of the current harvest paradigm, wherein harvest is focused largely on males; although the lack of information on age of specimens prevented us from rigorously differentiating among causal mechanisms. Disentangling potential mechanisms underpinning long-term trends in horn and antler size is a daunting task, but one that is worthy of additional research focused on elucidating the relative influence of nutrition and effects (both demographic and genetic) of harvest. © 2013 The Wildlife Society.     

Diet alters male horn allometry in the beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae)

Darwin considered the horns of male beetles to be among the most striking examples of sexual selection. As with antlers in deer or elk, beetle horns scale positively with male body size, with the result that large males have disproportionately longer horns than small males. It is generally assumed that such scaling relationships (''static allometries'') are insensitive to short-term changes in the environment, and for this reason they are regularly used as diagnostic attributes of populations or species. Here I report breeding experiments on horned beetles that demonstrate that the scaling relationship between male horn length and body size changes when larval nutrition changes. Males reared on a low-quality diet had longer horn lengths at any given body size than sibling males reared on a high-quality diet. Such ''allometry plasticity'' may explain seasonal changes observed in this same scaling relationship in a natural population. These experiments demonstrate that scaling relationships of sexually selected traits can respond facultatively to variation in the environment, thereby revealing a new mechanism by which males regulate the production of exaggerated secondary sexual traits.