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.     

Compensatory Growth Limits Opportunities for Artificial Selection in Alpine Chamois

ABSTRACT In ungulates, big males with large weapons typically outcompete other males over access to estrous females. In many species, rapid early growth leads to large adult mass and weapon size. We compared males in one hunted and one protected population of Alpine chamois (Rupicapra rupicapra) to examine the relationship between horn length and body mass. We assessed whether early development and hunter selectivity affected age-specific patterns of body and horn size and whether sport hunting could be an artificial selection pressure favoring smaller horns. Adult horn length was mostly independent of body mass. For adult males, the coefficient of variation of horn length (0.06) was <50% of that for body mass (0.16), suggesting that horn length presents a lower potential for selection and may be less important for male mating success than is body mass. Surprisingly, early development did not affect adult mass because of apparent compensatory growth. We found few differences in body and horn size between hunted and protected populations, suggesting the absence of strong effects of hunting on male phenotype. If horn length has a limited role in male reproductive success, hunter selectivity for males with longer horns is unlikely to lead to an artificial selective pressure on horn size. These results imply that the potential evolutionary effects of selective hunting depend on how the characteristics selected by hunters affect individual reproductive success.     

Environmental Influences on Dall's Sheep Survival

Understanding how species respond to environmental conditions can assist with conservation strategies and harvest management, especially in arctic and boreal regions that are experiencing rapid climate change. Although climatic influences on species distributions have been studied, broad-scale effects of climate on survival are less well known. We examined the interactive effects of meteorological and remotely sensed environmental variables on survival of Dall's sheep (Ovis dalli dalli) lambs and adults by synthesizing radio-telemetry data across their range. We used data from 9 studies of adult sheep and 2 studies of lambs that were conducted between 1997 and 2012 at sites spanning the species' range in Alaska, USA, and northwestern Canada. We obtained environmental variables throughout the range of Dall's sheep, including the normalized difference vegetation index (NDVI) from optical remote sensing, freeze-thaw frequency (FTF) from passive microwave remote sensing, and gridded climate variables such as snow water equivalent, temperature, and precipitation. We used Cox proportional hazard regression to investigate the effects of environmental variables recorded during summer, winter, and the previous winter on annual survival rates of Dall's sheep lambs and adults. Summer NDVI was the most influential environmental factor affecting lamb survival, with improved lamb survival occurring in years with a high maximum NDVI. Also, lamb predation by coyotes (Canis latrans) and golden eagles (Aquila chrysaetos) decreased substantially with increasing NDVI. The previous winter FTF had the strongest effect on adult survival, with decreased survival occurring after winters with high FTF. In addition, these remotely sensed environmental factors interacted with meteorological factors to affect survival, such that effects of winter temperature depended on summer NDVI and winter FTF. Warm winters increased lamb survival only when preceded by summers with high NDVI, and warm winters increased adult survival only when winter FTF was low. Thus, potential benefits of climate warming may be counteracted if wintertime freeze-thaw events markedly increase. Correlations among environmental variables across sites were low, and regional climate cycles such as the Pacific Decadal Oscillation (PDO) had weak effects, indicating substantial local variability in climatic conditions experienced by Dall's sheep across their range. These findings can help managers anticipate how Dall's sheep populations will respond to changes in local environmental conditions. Our results also highlight the utility of multiple remotely sensed environmental conditions for ungulate management, especially passive microwave products that provide valuable information on winter icing events. © 2020 The Wildlife Society.     

Long-term data reveal effects of climate,road access,and latitude on mountain goat horn size

Potential negative artificial selection on horn size is a concern for many harvested ungulates. The mountain goat (Oreamnos americanus) has distinct black horns, but targeting animals based on horn size in the field can be challenging. We analyzed over 23,000 horn records that included base circumference and total length, from which we also derived horn volume, from mountain goats harvested in Alaska, British Columbia, and the Northwest Territories from 1980 to 2016. We tested 3 potential drivers of horn size variation: geographical location, environmental conditions, and artificial selection. We found no support for a latitudinal effect with surprisingly little variation across the sampling distribution. The Pacific Decadal Oscillation had the largest effect outside age in the model, suggesting a role of climate in shaping variation. Mountain goats harvested closer to roads had larger horns, indicating that ease of access might allow hunters to be more selective, though the effect size was small. Our findings reinforce the value of accurate and complete record keeping on horn size, age, and sex of harvested animals, and highlight the importance of explicitly considering climate and accessibility when devising management strategies for the mountain goat.     

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.     

Using distance sampling and hierarchical models to improve estimates of Dall's sheep abundance

Management of large mammal populations has often been based on aerial minimum count surveys that are uncorrected for incomplete detection and lack estimates of precision. These limitations can be particularly problematic for Dall's sheep (Ovis dalli dalli) due to the high cost of surveys and variation in detection probability across time and space. The limitations of these methods have been recognized for some time, but previously proposed alternatives for sheep surveys proved to be too costly and logistically unfeasible in most circumstances (Udevitz et al. 2006). We assessed the potential for a combination of distance sampling surveys and a hierarchical modeling approach to provide a more efficient means for estimating Dall's sheep abundance by conducting aerial contour transect surveys over all sheep habitat in Gates of the Arctic National Park and Preserve (GAAR), Alaska in 2009 and 2010. We estimated the population of Dall's sheep was 8,412 (95% CI: 6,517–11,090) and 10,072 (95% CI 8,081–12,520) in 2009 and 2010, respectively. Abundance within the Itkillik Preserve area within GAAR was 1,898 (95% CI: 1,421–2,578) and 1,854 (95% CI: 1,342–2,488) in 2009 and 2010, respectively. Estimates of lamb abundance in 2010 were more than double those of 2009 after correcting for detection bias related to group size, suggesting that the apparent estimate of lambs in the population may be biased in some years depending on the degree of aggregation. Overall, the contour transect surveys were feasible logistically, cost 70–80% less than minimum count surveys, and produced precise estimates of abundance, indicating that the application of these methods could be used effectively to increase the statistical rigor and spatial extent of Dall's sheep abundance surveys throughout Alaska. These methods could be used to improve the assessment of long-term trends in populations and productivity and provide valuable information for harvest management at both local and landscape scales at reduced costs in comparison to traditional minimum count surveys. © 2011 The Wildlife Society.     

Reproductive,behavioral, and endocrine characteristics of the Dall's sheep (Ovis dalli dalli)

Failed interspecific embryo transfer between Dall's sheep (Ovis dalli dalli) and domestic ewes (Ovis aries), and a paucity of physiological data available for the Dall's sheep, provided incentive for developing a reproductive database in Dall's sheep. Urine samples were collected on a regular basis from a captive herd of Dall's sheep during the breeding to lambing interval over a 3 year period. To provide comparative endocrine data during pregnancy, samples also were collected during a single gestation period from five Suffolk ewes. All urine samples were analyzed for pregnanediol-3-glucuronide (PdG) and estrone conjugates (EC). Behavioral observations from Dall's sheep were made each year during the rut and lambing periods to calculate estrous cycle and gestation lengths. Placentas from Dall's sheep (n = 20) and Suffolk ewes (n = 10) were collected, the total number of cotyledons counted, and the length, width, area, and weight of eight cotyledons/horn from each placenta calculated. Endocrine and behavioral results indicated that the Dall's sheep is seasonally polyestrous and monovulatory, with a mean estrous cycle length of 18.2 ± 0.3 days, a mean luteal phase of 11.8 ± 0.5 days, and an average gestation length of 171.6 ± 0.9 days. While hormonal patterns for pregnancy generally were similar between the two species, there was a pronounced difference in the magnitude of hormone concentrations, particularly PdG, and an absence of a marked preparturition EC rise in domestic sheep. Cotyledon numbers, weight, and area were smaller (P < 0.05) in the Dall's sheep, compared to Suffolk ewes. These data suggest that unsuccessful interspecies embryo transfer attempts may have been due to failed fetal/maternal communications resulting from hormonal and/or placental differences. © 1996 Wiley-Liss, Inc.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.     

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.     

Data from selective harvests underestimate temporal trends in quantitative traits

Human harvests can select against phenotypes favoured by natural selection, and natural resource managers should evaluate possible artificial selection on wild populations. Because the required genetic data are extremely difficult to gather, however, managers typically rely on harvested animals to document temporal trends. It is usually unknown whether these data are unbiased. We explore our ability to detect a decline in horn size of bighorn sheep (Ovis canadensis) by comparing harvested males with all males in a population where evolutionary changes owing to trophy hunting were previously reported. Hunting records underestimated the temporal decline, partly because of an increasing proportion of rams that could not be harvested because their horns were smaller than the threshold set by hunting regulations. If harvests are selective, temporal trends measured from harvest records will underestimate the magnitude of changes in wild populations.     

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     

Does selection on horn length of males and females differ in protected and hunted populations of a weakly dimorphic ungulate?

Weaponry in ungulates may be costly to grow and maintain, and different selective pressures in males and females may lead to sex‐biased natural survival. Sexual differences in the relationship between weapon growth and survival may increase under anthropogenic selection through culling, for example because of trophy hunting. Selection on weaponry growth under different scenarios has been largely investigated in males of highly dimorphic ungulates, for which survival costs (either natural or hunting related) are thought to be greatest. Little is known, however, about the survival costs of weaponry in males and females of weakly dimorphic species. We collected information on horn length and age at death/shooting of 407 chamois Rupicapra rupicapra in a protected population and in two hunted populations with different hunting regimes, to explore sexual differences in the selection on early horn growth under contrasting selective pressures. We also investigated the variation of horn growth and body mass in yearling males (n = 688) and females (n = 539) culled in one of the hunted populations over 14 years. The relationship between horn growth and survival showed remarkable sexual differences under different evolutionary scenarios. Within the protected population, under natural selection, we found no significant trade‐off in either males or females. Under anthropogenic pressure, selection on early horn growth of culled individuals showed diametrically opposed sex‐biased patterns, depending on the culling regime and hunters’ preferences. Despite the selective bias between males and females in one of the hunted populations, we did not detect significant sex‐specific differences in the long‐term pattern of early growth. The relationship between early horn growth and natural survival in either sex might suggest stabilizing selection on horn size in chamois. Selection through culling can be strongly sex‐biased also in weakly dimorphic species, depending on hunters’ preferences and hunting regulations, and long‐term data are needed to reveal potential undesirable evolutionary consequences.     

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.     

Can phenotypic rescue from harvest refuges buffer wild sheep from selective hunting?

Human harvests can unwittingly drive evolution on morphology and life history, and these selective effects may be detrimental to the management of natural resources. Although theory suggests that harvest refuges, as sources of unselected animals, could buffer the effects of human exploitation on wild populations, few studies have assessed their efficiency. We analyzed records from >7000 trophy bighorn rams (Ovis canadensis) harvested in Alberta, Canada, between 1974 and 2011 to investigate if the movement of rams from refuges toward harvested areas reduced the effects of selective harvesting on horn size through phenotypic rescue. Rams taken near refuges had horns on average about 3% longer than rams shot far from refuges and were slightly older, suggesting migration from refuges into hunted areas. Rams from areas adjacent to and far from harvest refuges, however, showed similar declines in horn length and increases in age at harvest over time, indicating a decreasing rate of horn growth. Our study suggests that the influx of rams from refuges is not sufficient to mitigate the selective effects of sheep trophy harvest. Instead, we suggest that selective hunting of highly mobile animals may affect the genetic structure of populations that spend part of the year inside protected areas.     

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.     

DENSITY-DEPENDENT SEXUAL SELECTION IN THE FUNGUS BEETLE,BOLITOTHERUS CORNUTUS

The hypothesis that population density can affect sexual selection on male horn size was tested in a three-year study of a fungus beetle, Bolitotherus cornutus. Males of this species have horns that vary greatly in length. These horns are used in fights over females; longer-horned males win the majority of fights, regardless of population density. However, density does affect the relationship between horn length and access to females. In six populations of naturally and experimentally varying densities, longer-horned males gained a greater advantage in access to females in low-density populations than at high density. This increase in access to females causes an increase in the number of females inseminated by longer-horned males; thus, sexual selection for longer horns is stronger at lower densities.     

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.     

Genome-wide association mapping identifies the genetic basis of discrete and quantitative variation in sexual weaponry in a wild sheep population

Understanding the genetic architecture of phenotypic variation in natural populations is a fundamental goal of evolutionary genetics. Wild Soay sheep (Ovis aries) have an inherited polymorphism for horn morphology in both sexes, controlled by a single autosomal locus, Horns. The majority of males have large normal horns, but a small number have vestigial, deformed horns, known as scurs; females have either normal horns, scurs or no horns (polled). Given that scurred males and polled females have reduced fitness within each sex, it is counterintuitive that the polymorphism persists within the population. Therefore, identifying the genetic basis of horn type will provide a vital foundation for understanding why the different morphs are maintained in the face of natural selection. We conducted a genome-wide association study using ～36000 single nucleotide polymorphisms (SNPs) and determined the main candidate for Horns as RXFP2, an autosomal gene with a known involvement in determining primary sex characters in humans and mice. Evidence from additional SNPs in and around RXFP2 supports a new model of horn-type inheritance in Soay sheep, and for the first time, sheep with the same horn phenotype but different underlying genotypes can be identified. In addition, RXFP2 was shown to be an additive quantitative trait locus (QTL) for horn size in normal-horned males, accounting for up to 76% of additive genetic variation in this trait. This finding contrasts markedly from genome-wide association studies of quantitative traits in humans and some model species, where it is often observed that mapped loci only explain a modest proportion of the overall genetic variation.     

Horn growth rate and longevity: implications for natural and artificial selection in thinhorn sheep (Ovis dalli)

We used horn measurements from natural and hunted mortalities of male thinhorn sheep Ovis dalli from Yukon Territory, Canada, to examine the relationship between rapid growth early in life and longevity. We found that rapid growth was associated with reduced longevity for sheep aged 5 years and older for both the hunted and natural mortality data sets. The negative relationship between growth rate and longevity in hunted sheep can at least partially be explained by morphologically biased hunting regulations. The same trend was evident from natural mortalities from populations that were not hunted or underwent very limited hunting, suggesting a naturally imposed mortality cost directly or indirectly associated with rapid growth. Age and growth rate were both positively associated with horn size at death for both data sets, however of the two growth rate appeared to be a better predictor. Large horn size can be achieved both by individuals that grow horns rapidly and by those that have greater longevity, and the trade-off between growth rate and longevity could limit horn size evolution in this species. The similarity in the relationship between growth rate and longevity for hunted and natural mortalities suggests that horn growth rate should not respond to artificial selection. Our study highlights the need for the existence and study of protected populations to properly assess the impacts of selective harvesting.     

Linear and threshold-dependent expression of secondary sexual traits in the same individual: insights from a horned beetle (Coleoptera: Scarabaeidae)

Elaborate horns or horn‐like structures in male scarab beetles commonly scale with body size either (a) in a linear fashion with horn size increasing relatively faster than body size or (b) in a threshold‐dependent, sigmoid fashion; that is, males smaller than a certain critical body size develop no or only rudimentary horns, whereas males larger than the threshold size express fully developed horns. The development of linear vs. sigmoid scaling relationships is thought to require fundamentally different regulatory mechanisms. Here we show that such disparate regulatory mechanisms may co‐occur in the same individual. Large males of the south‐east Asian Onthophagus (Proagoderus) watanabei (Ochi & Kon) (Scarabaeidae, Onthophagini) develop a pair of long, curved head horns as well as a single thoracic horn. We show that unlike paired head horns in a large number of Onthophagus species, in O. watanabei the relationship between head horns and body size is best explained by a linear model. Large males develop disproportionately longer horns than small males, but the difference in relative horn sizes across the range of body sizes is small compared to other Onthophagus species. However, the scaling relationship between the thoracic horn and body size is best explained by a strongly sigmoid model. Only males above a certain body size threshold express a thoracic horn and males smaller than this threshold express no horn at all. We found a significant positive correlation between head and thoracic horn length residuals, contrary to what would be expected if a resource allocation tradeoff during larval development would influence the length of both horn types. Our results suggest that the scaling relationship between body size and horn length, and the developmental regulation underlying these scaling relationships, may be quite different for different horns, even though these horns may develop in the same individual. We discuss our results in the context of the developmental biology of secondary sexual traits in beetles. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 473–480.