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.     

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.     

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.     

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.     

Source populations in carnivore management: cougar demography and emigration in a lightly hunted population

Wildlife agencies typically attempt to manage carnivore numbers in localized game management units through hunting, and do not always consider the potential influences of immigration and emigration on the outcome of those hunting practices. However, such a closed population structure may not be an appropriate model for management of carnivore populations where immigration and emigration are important population parameters. The closed population hypothesis predicts that high hunting mortality will reduce numbers and densities of carnivores and that low hunting mortality will increase numbers and densities. By contrast, the open population hypothesis predicts that high hunting mortality may not reduce carnivore densities because of compensatory immigration, and low hunting mortality may not result in more carnivores because of compensatory emigration. Previous research supported the open population hypothesis with high immigration rates in a heavily hunted (hunting mortality rate=0.24) cougar population in northern Washington. We test the open population hypothesis and high emigration rates in a lightly hunted (hunting mortality rate=0.11) cougar population in central Washington by monitoring demography from 2002 to 2007. We used a dual sex survival/fecundity Leslie matrix to estimate closed population growth and annual census counts to estimate open population growth. The observed open population growth rate of 0.98 was lower than the closed survival/fecundity growth rates of 1.13 (deterministic) and 1.10 (stochastic), and suggests a 12–15% annual emigration rate. Our data support the open population hypothesis for lightly hunted populations of carnivores. Low hunting mortality did not result in increased numbers and densities of cougars, as commonly believed because of compensatory emigration.     

Dall's sheep horn growth and harvest management in the Mackenzie Mountains,Northwest Territories,Canada

Across most of their native North American range, the horns of mountain sheep (Ovis spp.) males are getting smaller, a pattern attributed to selective hunting pressure. We measured the horns of 755 Dall's sheep males (Ovis dalli dalli) in the southern Mackenzie Mountains, Northwest Territories, between 2002 and 2017. For each male, we measured the circumference and length of each annulus for the right horn and calculated horn volume for each year. We examined changes in horn size in 4 different outfitter areas, using age at harvest as a covariate. Hunting pressure across years in the study area was consistently low, and this population did not experience the decline in horn size observed in several other mountain sheep populations in Canada. Over the 16-year period, the average horn volume of harvested males was stable and even increased in 1 outfitter area. Local management of Dall's sheep delivered independently by the guide outfitters in the Mackenzie Mountains appears to contribute to maintaining a population of males that has not been adversely affected by strong selective hunting pressure. The resilience of this management strategy may be challenged by environmental changes associated with rapid warming in northern mountain environments.     

Horn growth patterns in Alpine chamois

The analysis of horn growth may provide important information about the allocation of metabolic resources to secondary sexual traits. Depending on the selective advantages offered by horn size during intra- and inter-specific interactions, ungulates may show different investment in horn development, and growth variations within species may be influenced by several parameters, such as sex, age, or resource availability. We investigated the horn growth patterns in two hunted populations of Alpine chamois (Rupicapra r. rupicapra) in the Central Italian Alps. We tested the role of individual heterogeneity on the growth pattern and explored the variation in annulus length as a function of different factors (sex, age, hunting location, cohort). We then investigated the mechanisms underlying horn growth trajectories to test for the occurrence of compensatory or recovery growth and their potential differences between sexes and populations. Annulus length varied as a function of sex, age of individuals and, marginally, hunting location; no effect of cohort or individual heterogeneity was detected. Male and female chamois showed compensatory horn growth within the first 5½ years of life, though the partial convergence of horn trajectories in chamois suggests that this mechanisms would best be described as ‘recovery growth’. Compensation rates were greater in males than in females, while only compensatory growth rates up to 2½ years of age were different in the two populations. Besides confirming the sex- and age-dependent pattern of horn development, our study suggests that the mechanism of recovery growth supports the hypothesis of horn size as a weakly selected sexual trait in male and female chamois. Furthermore, the greater compensation rates in horn growth shown by male chamois possibly suggest selective effects of hunting on age at first reproduction, while different compensation rates between populations may suggest the occurrence of some plasticity in resource allocation to sexual traits in relation to different environments.     

Effects of hunting on cougar spatial organization

The effects of increased mortality on the spatial dynamics of solitary carnivores are not well understood. We examined the spatial ecology of two cougar populations that differed in hunting intensity to test whether increased mortality affected home range size and overlap. The stability hypothesis predicts that home range size and overlap will be similar for both sexes among the two areas. The instability hypothesis predicts that home range size and overlap will be greater in the heavily hunted population, although may differ for males versus females due to behavior strategies. We marked 22 adult resident cougars in the lightly hunted population and 20 in the heavily hunted population with GPS collars from 2002 to 2008. Cougar densities and predation rates were similar among areas, suggesting no difference in per capita resources. We compared home range size, two‐dimensional home range overlap, and three‐dimensional utilization distribution overlap index (UDOI) among annual home ranges for male and female cougars. Male cougars in the heavily hunted area had larger sized home ranges and greater two‐dimensional and three‐dimensional UDOI overlap than those in the lightly hunted area. Females showed no difference in size and overlap of home range areas between study populations – further suggesting that differences in prey quantity and distribution between study areas did not explain differences in male spatial organization. We reject the spatial stability hypothesis and provide evidence to support the spatial instability hypothesis. Increased hunting and ensuing increased male home range size and overlap may result in negative demographic effects for cougars and potential unintended consequences for managers.     

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.     

Does hunting affect the demography and genetic structure of the greywing francolin Francolinus africanus?

Hunted and unhunted populations of greywing francolin Francolinus africanus have been studied in the eastern Cape Province of South Africa in order to understand the effects of hunting on the demography and genetic structure of these populations. Greywing population density cycled annually for both hunted and unhunted populations. However, there was an apparent pulse of immigration of sub-dominant birds, and earlier reproduction, in the hunted populations immediately after the winter hunting season. Average levels of allozyme heterozygosity (H) for hunted and unhunted populations were both 0.076, and although the proportion of polymorphic loci per sample and the mean number of alleles per locus for each sample were lower for the hunted populations than for the unhunted populations, these differences were not significant. However, the hunted populations displayed higher levels of outbreeding (lower F _IS and F _IT values) than those for unhunted populations. Therefore, it is concluded that although greywing francolin populations contain relatively high levels of genetic heterogeneity, it is probably the increased levels of local immigration following hunting which reduces the effects of any reduction in genetic variation due to a decrease in local population size from hunting.     

Optimal growth strategies under divergent predation pressure

The conditions leading to gigantism in nine‐spined sticklebacks Pungitius pungitius were analysed by modelling fish growth with the von Bertalanffy model searching for the optimal strategy when the model's growth constant and asymptotic fish size parameters are negatively related to each other. Predator‐related mortality was modelled through the increased risk of death during active foraging. The model was parameterized with empirical growth data of fish from four different populations and analysed for optimal growth strategy at different mortality levels. The growth constant and asymptotic fish size were negatively related in most populations. Optimal fish size, fitness and life span decreased with predator‐induced mortality. At low mortality, the fitness of pond populations was higher than that of sea populations. The differences disappeared at intermediate mortalities, and sea populations had slightly higher fitness at extremely high mortalities. In the scenario where all populations mature at the same age, the pond populations perform better at low mortalities and the sea populations at high mortalities. It is concluded that a trade‐off between growth constant and asymptotic fish size, together with different mortality rates, can explain a significant proportion of body size differentiation between populations. In the present case, it is a sufficient explanation of gigantism in pond P. pungitius.     

Quantifying anthropogenic wolf mortality in relation to hunting regulations and landscape attributes across North America

Understanding the types and magnitude of human‐caused mortality is essential for maintaining viable large carnivore populations. We used a database of cause‐specific mortality to examine how hunting regulations and landscape configurations influenced human‐caused mortality of North American gray wolves (Canis lupus). Our dataset included 21 studies that monitored the fates of 3564 wolves and reported 1442 mortalities. Human‐caused mortality accounted for 61% of mortality overall, with 23% due to illegal harvest, 16% due to legal harvest, and 12% the result of management removal. The overall proportion of anthropogenic wolf mortality was lowest in areas with an open hunting season compared to areas with a closed hunting season or mixed hunting regulations, suggesting that harvest mortality was neither fully additive nor compensatory. Proportion of mortality from management removal was reduced in areas with an open hunting season, suggesting that legal harvest may reduce human‐wolf conflicts or alternatively that areas with legal harvest have less potential for management removals (e.g., less livestock depredation). Proportion of natural habitat was negatively correlated with the proportion of anthropogenic and illegal harvest mortality. Additionally, the proportion of mortality due to illegal harvest increased with greater natural habitat fragmentation. The observed association between large patches of natural habitat and reductions in several sources of anthropogenic wolf mortality reiterate the importance of habitat preservation to maintain wolf populations. Furthermore, effective management of wolf populations via implementation of harvest may reduce conflict with humans. Effective wolf conservation will depend on holistic strategies that integrate ecological and socioeconomic factors to facilitate their long‐term coexistence with humans.     

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.     

Culling Versus Density Effects in Management of a Deer Population

Abstract: Wildlife managers often manipulate hunting regulations to control deer populations. However, few empirical studies have examined the level of hunting effort (hunter-days) required to limit population growth and demographic effects through harvesting of females. Moreover, the relative importance of density effects on population growth has not been quantified. We reconstructed a sika deer [Cervus nippon] population over a period of 12 years (1990–2001) using age- and sex-specific harvest data. Using cohort analysis, we analyzed population dynamics, focusing on 1) the relationship between hunting effort and hunting-induced mortality rate, 2) relative contributions of hunting mortality and recruitment of yearlings to annual changes in population growth rate, and 3) annual variation in recruitment rate. Population size increased until 1998 and declined thereafter. The population growth rate changed more in response to annual changes in recruitment rate than hunting mortality rate. Temporal variation in recruitment rate was not controlled by birth rate alone; direct density dependence, intensities of hunting mortality for fawns, and for females (≥2 yr of age), which accounted for the fawn survival rate, were required as factors to explain temporal variation. Density effects on the recruitment rate were not strong enough to regulate the population within the study period; high hunting mortality, with intensive female harvesting, was necessary to prevent population growth. Hunting effort was a good predictor of the hunting mortality rate, and female harvest had a negative effect on the recruitment rate through fawn survival. We suggest that >3,500 hunter-days and prioritization of female harvesting are required to prevent increases in this deer population.     

On being the right size: increased body size is associated with reduced telomere length under natural conditions

Evolution of body size is likely to involve trade-offs between body size, growth rate and longevity. Within species, larger body size is associated with faster growth and ageing, and reduced longevity, but the cellular processes driving these relationships are poorly understood. One mechanism that might play a key role in determining optimal body size is the relationship between body size and telomere dynamics. However, we know little about how telomere length is affected when selection for larger size is imposed in natural populations. We report here on the relationship between structural body size and telomere length in wild house sparrows at the beginning and end of a selection regime for larger parent size that was imposed for 4 years in an isolated population of house sparrows. A negative relationship between fledgling size and telomere length was present at the start of the selection; this was extended when fledgling size increased under the selection regime, demonstrating a persistent covariance between structural size and telomere length. Changes in telomere dynamics, either as a correlated trait or a consequence of larger size, could reduce potential longevity and the consequent trade-offs could thereby play an important role in the evolution of optimal body size.     

Evaluating how hunters see and react to telemetry collars on white-tailed deer

Fates of individuals outfitted with radiotransmitters commonly are used for estimating survival rates in populations of large animals that are hunted. Despite precautions, this practice may be subject to complex biases associated with hunter reaction to presence of radiotransmitters. To assess this potential bias we conducted an experiment using artificial deer (i.e., decoys) to measure hunters' abilities to see deer and determine if deer seen were wearing radiocollars. We used logistic regression to quantify probabilities that seeing deer and subsequently seeing radiocollars might be influenced by distance, percent visual obstruction, body orientation, hunter experience, and antler characteristics of deer. Additionally, we evaluated how experience and antler characteristics of deer might influence a hunter's decision to harvest a radiocollared deer. We found that 25.8% of the potentially observable collared deer (n = 663) were subsequently observed by hunters. Odds of observing deer and radiocollars increased 95% and 230%, respectively, for each additional log(yr) of hunting experience. Willingness to harvest radiocollared deer increased 89% for each additional log(yr) of hunting experience and 144% for large-antlered deer relative to antlerless deer. When hunting is an important source of mortality, analysts need to understand how potential biases associated with observing deer are associated with hunters' reactions to and subsequent decisions to harvest radiocollared animals. Our study suggested that presence of radiocollars may influence a deer's potential risk of being harvested and in turn bias telemetry-based estimates of survival, given that hunting mortality is the largest component of total mortality in hunted deer populations. Collar-based telemetry is used nearly universally by wildlife managers and researchers throughout North America and elsewhere to estimate and monitor the survival of big game populations that are managed through hunting. Our findings demonstrate that these estimates are likely subject to complex and systematic biases that managers should consider when evaluating future population-level effects of managed hunting. © 2011 The Wildlife Society     

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.     

Prehistoric bird extinctions and human hunting

Holocene fossils document the extinction of hundreds of bird species on Pacific islands during prehistoric human occupation. Human hunting is implicated in these extinctions, but the impact of hunting is difficult to disentangle from the effects of other changes induced by humans, including habitat destruction and the introduction of other mammalian predators. Here, we use data from bones collected at a natural sand dune site and associated archaeological middens in New Zealand to show that, having controlled for differences in body mass and family membership (and hence for variation in life-history traits related to population growth rate), birds that were more intensively hunted by prehistoric humans had a higher probability of extinction. This result cannot be attributed to preservation biases and provides clear evidence that selective hunting contributed significantly to prehistoric bird extinctions at this site.     

Hunting and the fate of French breeding waterbirds

Capsule French breeding populations of hunted waterbirds have more negative long-term population trends than closely related non-hunted species.

Aims To assess the relative fate of breeding populations of hunted and non-hunted waterbird species in France.

Methods We compared long-term population growth rates of hunted versus non-hunted waterbirds using two large-scale and long-term national surveys.

Results Waterbirds globally displayed long-term increases in population sizes most likely driven by their positive response to the implementation of numerous protected areas over remnant wetlands across the country since the early 1970s. In contrast, hunted species revealed more negative trends compared to non-hunted species.

Conclusion A causal relationship between hunting and population dynamics is not demonstrated here, but the results are consistent with species' breeding populations being negatively affected, on average, if they are hunted.     

Effects of hunting on guianan forest game birds

To assess the effect of on-going and of previous or near by hunting pressure on game birds in a neotropical rain forest (French Guiana), I compared species abundances between six hunted and disturbed areas, seven non-hunted and pristine areas and eight intermediate areas, undisturbed and not currently hunted, but formerly hunted or close to current hunting areas. I recorded all birds detected within 100 m-wide strip transects, walked at random through every forest type all day long. The frequency of records per 10 h and flock sizes on each transect were averaged over all surveys in each study site (N = 3025 h). Censused groups included terrestrial (tinamous, woodquails, curassow and trumpeter) and arboreal species (guans, macaws, parrots, toucans), but not waterbirds. The abundance of all game birds was higher in non-hunted than in hunted areas, though not always significantly for secretive understorey species (tinamous, guans, wood-quail) or canopy frugivores (parrots, macaws, toucans). The Black Curassow and Gray-winged Trumpeter exhibited the highest and most consistent increases (7–10-fold) and more so between intermediate and non-hunted sites than between hunted and intermediate sites. The mean flock size also increased along the hunting gradient, especially in flocking species (macaws, parrots, trumpeter). Low reproductive rates and/or seasonal movements may explain that hunting pressure could still be felt after hunting has ceased or when it persists only away from a given area. Such a persistent effect would affect some populations in small protected areas.