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Predicting the evolutionary effects of hunting requires an understanding of genetics: Evolutionary Responses to Hunting

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... Kardos et al. (2018) helped clarify details of the mechanisms behind selective harvest and evolutionary effect. This dialogue is important but does not alter the primary premise of Heffelfinger (2018), which was to call attention to the multitude of previously neglected genetic and nongenetic factors that make hunter selection inefficient and uncommon in wild ungulates, and to warn against generalizing the results from specific studies. ...
... This dialogue is important but does not alter the primary premise of Heffelfinger (2018), which was to call attention to the multitude of previously neglected genetic and nongenetic factors that make hunter selection inefficient and uncommon in wild ungulates, and to warn against generalizing the results from specific studies. Kardos et al. (2018) misrepresent my paper by repeatedly creating a false argument that the factors I discuss do not preclude rapid evolution in response to selective hunting. These factors were offered as obstacles, not absolute obstructions, to harvest-based selection causing evolutionary change in the size of horns or antlers. ...
... Heffelfinger (2018) provides a detailed discussion of the inappropriateness of applying results from Ram Mountain widely to other wild sheep (and other ungulate) populations under different management paradigms and selection intensities. Heffelfinger (2018) provided a poor example of linked loci confounding an evolutionary response to selection, but Kardos et al. (2018) acknowledged this concept is correct and well supported by theoretical and empirical genetics research. The point remains valid even if the empirical example provided was not. ...
... For instance, Lindsey (2008) estimated that trophy hunting in sub-Saharan Africa generated profits of approximately $201 million/year (US$). Although intense and deregulated trophy hunting can trigger phenotypic and evolutionary trait changes in wild bovids (Coltman et al. 2003, Pigeon et al. 2016, it remains largely unexplored how individual performance, population structure and composition, and a multitude of environmental factors, including climate and habitat changes, weaken, counterbalance, or strengthen these effects (Heffelfinger 2018, Kardos et al. 2018. Understanding the effect of biotic and abiotic variables on phenotypic-trait changes is essential to manage wild bovid populations more effectively, guaranteeing the long-term sustainability of hunting systems. ...
... First, our data regarding the number of harvested ibex is limited in time and the effects of selective harvesting on phenotypic traits can take several decades to be recorded (Coulson et al. 2018). Second, our data merely represents a proxy of hunting pressure and no genetic information (e.g., selection differential, heritability, pedigree) is available for our population (Kardos et al. 2018). Even though we did not estimate the potential phenotypic variance related to genetic differences, there are some aspects that can weaken the phenotypic and evolutionary responses to trophy hunting. ...
Article
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. The encroachment of coniferous forests and the consequent loss of natural pastures had a negative effect on the annual horn growth of male ibex inhabiting a Mediterranean ecosystem. The selective harvesting of small‐horned males had a positive effect on annual horn growth. The expansion of grass‐rich open areas should be prioritized to reduce the negative effects of habitat deterioration on the phenotypic traits of male ibex.
... Wildlife management agencies also regulate hunting to enhance hunter opportunity and experience on public lands by manipulating season lengths, opening dates, and hunter density to delay animal movement to private lands (Conner et al. 2001, Vieira et al. 2003. Hunting can directly alter movements and distribution (Proffitt et al. 2010, Cleveland et al. 2012, Neumann and Ericsson 2018, social organization (Singer and Zeigenfuss 2002), and genetics (Kardos et al. 2018) of ungulate populations, and indirectly affect disease spread (Davidson et al. 2012, Apollonio et al. 2017) and population performance (Davidson et al. 2012, Festa-Bianchet et al. 2017. ...
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Sport hunting of ungulates is a predominant recreational pursuit and the primary tool for managing their populations in North America and beyond, given its influence on ungulate distributions, social organization, and population performance. Similarly, land management, such as motorized vehicle access, influences ungulate distributions during and outside hunting seasons. Although research on ungulate responses to hunting and land use is widespread, knowledge gaps persist about space use of hunters and what landscape features discriminate among hunt types and between successful and unsuccessful hunters. We used telemetry location data from hunters (n = 341) to estimate space use from 2008–2013 during 3 types of controlled, 5‐day hunts for antlered mule deer (Odocoileus hemionus) and elk (Cervus canadensis) in northeastern Oregon, USA: archery elk, rifle deer, and rifle elk. To evaluate space use, we developed utilization distributions for each hunter, created core areas (50% contours) for groups of hunters, and derived several metrics of space‐use overlap between successful and unsuccessful hunters. We also modeled predictors of space use using resource utilization functions with beta regression and stepwise model building. Hunter space use was compressed, with even the largest core area (unsuccessful rifle elk hunters) encompassing <16% (1,178 ha) of the area. We found strong similarities in space use of rifle hunters compared to archers, and core areas of successful hunters were markedly smaller than those of unsuccessful hunters (e.g., = 104 ha vs. 681 ha, respectively, for archers). Percentage cover and distance from open roads were the most consistent covariates in the 6 final models (successful vs. unsuccessful for each of 3 hunts) but with different signs. For example, predicted use of archery and rifle elk hunters increased with cover but decreased for rifle deer hunters. Although the same covariates were in the final models for unsuccessful and successful rifle elk hunters, their negligible spatial overlap suggested they sought those features in different locales, a pattern also documented for rifle deer hunters. Our models performed well (Spearman's rank correlation coefficients = 0.99 for 5 of 6 models), reflecting their utility for managing hunters and landscapes. Our results suggest that strategic management of open roads and forest cover can benefit managers seeking to balance hunter opportunity and satisfaction with harvest objectives, especially for species of special concern such as mule deer, and that differences in space use among hunter groups should be accounted for in hunting season designs. © 2021 The Wildlife Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA. We evaluated space use of successful and unsuccessful elk and mule deer hunters, contrasting archers with rifle hunters, and found minimal spatial overlap between groups but similarities in environmental features influencing space use. Integrated management of forest cover in tandem with roads and trails can aid in providing hunter opportunity while maintaining herd numbers and composition at desired levels.
... Selection would be stronger if it affected both sexes but in most species only males are hunted selectively, reducing the strength of selection by half (Kardos et al. 2018). For populations with highly polygynous mating systems, removal of a few very dominant males could have major effects on the distribution of reproductive success. ...
Article
Intense selective harvest of large mammals who carry the largest weapons may lead to an evolutionary shrinkage of those weapons. Currently, evidence suggesting evolutionary effects of harvest is limited to a few species of Bovidae and only 1 study has obtained data indicating a genetic effect. To have an evolutionary impact, harvest must be intense, persistent over time, similar over a large area without an effective source of unselected immigrants, and remove large individuals before they have a chance to breed. Many current harvest schemes do not fulfill all of these requirements, and they are unlikely to cause evolution. Before changes in weapon size over time are attributed to evolution, potential environmental sources of change, mainly density and climate, must be considered. We suggest that the role of weapon size in determining reproductive success, especially in interaction with male age, will determine whether or not intensive selective harvests may have evolutionary consequences. Age at harvest is a very important variable to consider. Changes in age structure over time may reveal underlying changes in harvest pressure or selectivity. A lack of data hampers our ability to assess the potential evolutionary effects of selective hunting. We provide a list of research hypotheses required to advance our ability to assess the evolutionary sustainability of current management practices.
... canadensis) at Ram Mountain, Alberta, Canada are typical and should shape harvest management for mountain sheep more broadly (Heffelfinger 2018b). We find that evidence for evolutionary response to hunter selection at Ram Mountain is overshadowed by environmental influences (Postma 2006, Pigeon et al. 2016, and there is little support for meaningful selection in most hunter-harvest schemes (Heffelfinger 2018a), a point reinforced by a reply to Kardos et al. (2018) by Heffelfinger (2018b). Boyce and Krausman (2018) did not dismiss the possibility that hunting might cause selection for smaller horn size in mountain sheep and it would be a surprise if this did not happen over long periods of intense selection. ...
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We respond to Festa‐Bianchet (2019) and caution against using interpretations from the unique Ram Mountain history to justify management of mountain sheep throughout their range. Because harvest management at Ram Mountain is atypical, it is not useful in informing the management of most mountain sheep herds.
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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. Management of bighorn sheep for horn size and for population maintenance has focused on factors including nutrition, environmental conditions, and genetic diversity. We demonstrate that disease plays an important role in horn size: pneumonia disease outbreak events significantly reduced horn growth in male bighorn sheep, and continued horn stunting occurred when chronically infected individuals remained present in the population.
Article
Estimating heritability (h2) is required to predict the response to selection and is useful in species that are managed or farmed using trait information. Estimating h2 in free-ranging populations is challenging due to the need for pedigrees; genomic-relatedness matrices (GRMs) circumvent this need and can be implemented in nearly any system where phenotypic and genome-wide single nucleotide polymorphism (SNP) data are available. We estimated the heritability of five body and three antler traits in a free-ranging population of white-tailed deer (Odocoileus virginianus) on Anticosti Island, Quebec, Canada. We generated classic and robust GRMs from >10,000 SNPs: hind foot length, dressed body mass and peroneus muscle mass had high h2 values of 0.62, 0.44 and 0.55, respectively. Heritability in male-only antler features ranged from 0.07 to 0.33 and had high standard errors. We explored the influence of filtering by minor allele frequency and data completion on h2: GRMs derived from fewer SNPs had reduced h2 estimates and the relatedness coefficients significantly deviated from those generated with more SNPs. As a corollary, we discussed limitations to the application of GRMs in the wild, notably how skewed GRMs, specifically many unrelated individuals, can increase variance around h2 estimates. This is the first study to estimate h2 on a free-ranging population of white-tailed deer and should be informative for breeding designs and management as these traits should respond to selection.
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Changing environmental conditions cause changes in the distributions of phenotypic traits in natural populations. However, determining the mechanisms responsible for these changes—and, in particular, the relative contributions of phenotypic plasticity versus evolutionary responses—is difficult. To our knowledge, no study has yet reported evidence that evolutionary change underlies the most widely reported phenotypic response to climate change: the advancement of breeding times. In a wild population of red deer, average parturition date has advanced by nearly 2 weeks in 4 decades. Here, we quantify the contribution of plastic, demographic, and genetic components to this change. In particular, we quantify the role of direct phenotypic plasticity in response to increasing temperatures and the role of changes in the population structure. Importantly, we show that adaptive evolution likely played a role in the shift towards earlier parturition dates. The observed rate of evolution was consistent with a response to selection and was less likely to be due to genetic drift. Our study provides a rare example of observed rates of genetic change being consistent with theoretical predictions, although the consistency would not have been detected with a solely phenotypic analysis. It also provides, to our knowledge, the first evidence of both evolution and phenotypic plasticity contributing to advances in phenology in a changing climate.
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Differentially harvesting individual animals with specific traits has led some to argue that such selection can cause evolutionary change that may be detrimental to the species, especially if those traits are related positively to individual fitness. Most hunters are not selective in the type of animal they take, satisfied instead to harvest any legal animal. In a few exceptions, however, regulations may limit hunters to harvest animals of a minimum size or age regardless of their personal choice. Using information from a broad range of aquatic and terrestrial systems exposed to a myriad of potential and operational selective pressures, several authors have made expansive generalizations about selective harvest and its applicability to ungulates. Harvest-based selection can potentially be intensive enough to be relevant in an evolutionary sense, but phenotypic changes consistent with hunter selection are otherwise confounded with multiple environmental influences. Factors such as age, genetic contribution of females, nutrition, maternal effects, epigenetics, patterns of mating success, gene linkage, gene flow, refugia, date of birth, and other factors affecting selection interact with harvest to impede unidirectional evolution of a trait. The intensity of selection determines potential for evolutionary change in a meaningful temporal framework. Indeed, only under severe intensity, and strict selection on a trait, could human harvest prompt evolutionary changes in that trait. Broad generalizations across populations or ecological systems can yield erroneous extrapolations and inappropriate assumptions. Removal of males expressing a variety of horn or antler sizes, including some very large males, does not inevitably represent directional artificial selection unless the selective pressures are intensive enough to cause a unidirectional shift in allele frequencies that may act on some relevant life-history trait or process. Here I review the topic of harvest-based selection in male ungulates and discuss the inefficiency of trophy hunting in changing genetic expression of phenotype. © 2017 The Wildlife Society.
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Trophy hunting, the selective removal of animals for human recreation, can contribute to conservation when appropriately managed. Yet, little is known about how harvest rates or different definitions of trophy affect age structure and trophy size in harvested animals and in survivors because no controlled studies exist. To investigate the impacts of different management regimes, we developed an individual-based model for bighorn sheep (Ovis canadensis), based on empirical data on survival from a protected population and data on horn growth from 2 populations that differed in their growth rates. One population showed slow horn growth and the other population fast horn growth. We subjected these model populations to varying harvest rates and 2 different hunting regulations: 4/5 curl and full-curl definitions of a trophy male. We found that the effect of hunting regulations depends on horn growth rate. In populations with fast horn growth, the effects of trophy hunting on male age structure and horn size were greater and the effect of a change in the definition of legal male smaller than in populations with slow growth rates. High harvest rates led to a younger age structure and smaller horn size. Both effects were weakened by a more restrictive definition of trophy male. As harvest rates increased past 40% of legal males, the number of males harvested increased only marginally because an increasing proportion of the harvested males included those that had just become legal. Although our simulation focused on bighorn sheep, the link between horn growth rate and harvest effects may be applicable for any size-selective harvest regime.
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The potential for selective harvests to induce rapid evolutionary change is an important question for conservation and evolutionary biology, with numerous biological, social and economic implications. We analyze 39 years of phenotypic data on horn size in bighorn sheep (Ovis canadensis) subject to intense trophy hunting for 23 years, after which harvests nearly ceased. Our analyses revealed a significant decline in genetic value for horn length of rams, consistent with an evolutionary response to artificial selection on this trait. The probability that the observed change in male horn length was due solely to drift is 9.9%. Female horn length and male horn base, traits genetically correlated to the trait under selection, showed weak declining trends. There was no temporal trend in genetic value for female horn base circumference, a trait not directly targeted by selective hunting and not genetically correlated with male horn length. The decline in genetic value for male horn length stopped, but was not reversed, when hunting pressure was drastically reduced. Our analysis provides support for the contention that selective hunting led to a reduction in horn length through evolutionary change. It also confirms that after artificial selection stops, recovery through natural selection is slow. This article is protected by copyright. All rights reserved.
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Long-term data (1974–2011) from harvested bighorn rams (Ovis canadensis) in Alberta, Canada, suggested a reduction in horn size and in the proportion of trophy rams in the provincial population over time. Age at harvest increased over time, suggesting slower horn growth. Rams that experienced favorable environmental conditions early in life had rapid horn growth and were harvested at a younger age than rams with slower horn growth. Guided nonresident hunters did not harvest larger rams than residents, suggesting that few large rams were available. Resident hunter success declined in recent years. Despite an apparently stable population, successive cohorts produced a decreasing harvest of trophy rams. We suggest that unrestricted harvest based on a threshold horn size led to a decline in the availability of trophy rams. That decline is partly an inevitable consequence of selective hunting that removes larger rams. Although our analysis does not establish that evolution of smaller horns caused the observed decline in both horn size and harvest of trophy rams, we suggest that intensive trophy hunting may have artificially selected for a decrease in horn growth rate. © 2013 The Wildlife Society.
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Sexual selection, through intra-male competition or female choice, is assumed to be a source of strong and sustained directional selection in the wild. In the presence of such strong directional selection, alleles enhancing a particular trait are predicted to become fixed within a population, leading to a decrease in the underlying genetic variation. However, there is often considerable genetic variation underlying sexually selected traits in wild populations, and consequently, this phenomenon has become a long-discussed issue in the field of evolutionary biology. In wild Soay sheep, large horns confer an advantage in strong intra-sexual competition, yet males show an inherited polymorphism for horn type and have substantial genetic variation in their horn size. Here we show that most genetic variation in this trait is maintained by a trade-off between natural and sexual selection at a single gene, relaxin-like receptor 2 (RXFP2). We found that an allele conferring larger horns, Ho(+), is associated with higher reproductive success, whereas a smaller horn allele, Ho(P), confers increased survival, resulting in a net effect of overdominance (that is, heterozygote advantage) for fitness at RXFP2. The nature of this trade-off is simple relative to commonly proposed explanations for the maintenance of sexually selected traits, such as genic capture ('good genes') and sexually antagonistic selection. Our results demonstrate that by identifying the genetic architecture of trait variation, we can determine the principal mechanisms maintaining genetic variation in traits under strong selection and explain apparently counter-evolutionary observations.
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Life-history trade-offs are well known in female mammals, but have seldom been quantified for males in polygynous species. I compared age-specific mass, weapon size, survival, and reproductive success of males in eight species of ungulates, and found weak interspecific correlations among life-history traits. Young males tended to have higher reproductive success in rapidly-growing than in slow-growing species, and in species where horns or antlers reached near-asymptotic size over the first few years of life. There was no clear interspecific trade-off between early reproduction and early survival. Reproductive senescence was evident in most species. Generation length, calculated as the mean age of fathers, was negatively correlated with the reproductive success of young males and positively with life expectancy of 3-year-olds, but not with early mortality. The main determinant of male reproductive success in polygynous ungulates is the ability to prevail against competing males. Consequently, the number and age structure of competitors should strongly affect an individual’s ability to reproduce, making classic trade-offs among life-history traits very context-dependent. Most fitness costs of reproduction in male ungulates likely arise from energy expenditure and injuries sustained while attempting to mate. Individual costs may be weakly correlated with fitness returns.
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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.
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Phenotype-based selective harvests, including trophy hunting, can have important implications for sustainable wildlife management if they target heritable traits. Here we show that in an evolutionary response to sport hunting of bighorn trophy rams (Ovis canadensis) body weight and horn size have declined significantly over time. We used quantitative genetic analyses, based on a partly genetically reconstructed pedigree from a 30-year study of a wild population in which trophy hunting targeted rams with rapidly growing horns, to explore the evolutionary response to hunter selection on ram weight and horn size. Both traits were highly heritable, and trophy-harvested rams were of significantly higher genetic 'breeding value' for weight and horn size than rams that were not harvested. Rams of high breeding value were also shot at an early age, and thus did not achieve high reproductive success. Declines in mean breeding values for weight and horn size therefore occurred in response to unrestricted trophy hunting, resulting in the production of smaller-horned, lighter rams, and fewer trophies.
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In recent years, scientists have realized that evolution can occur on timescales much shorter than the “long lapse of ages” emphasized by Darwin—in fact, evolutionary change is occurring all around us all the time. This book provides an authoritative and accessible introduction to eco-evolutionary dynamics, a cutting-edge new field that seeks to unify evolution and ecology into a common conceptual framework focusing on rapid and dynamic environmental and evolutionary change. The book covers key aspects of evolution, ecology, and their interactions. Topics range from natural selection, adaptive divergence, ecological speciation, and gene flow to population and community dynamics, ecosystem function, plasticity, and genomics. The book evaluates conceptual and methodological approaches, and draws on empirical data from natural populations—including those in human-disturbed environments—to tackle a number of classic and emerging research questions. It also discusses exciting new directions for future research at the intersection of ecology and evolution. The book reveals how evolution and ecology interact strongly on short timescales to shape the world we see around us.
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(1) The paper presents a theory of selection limits in artificial selection. It is, however, developed primarily in terms of single genes. (2) For a single gene with selective advantage s, the chance of fixation (the expected gene frequency at the limit) is a function only of Ns, where N is the effective population size. In artificial selection based on individual measurements, where the selection differential is {imath} standard deviations, the expected limit of individual selection in any population is a function only of N{imath}. (3) For low values of N{imath}, the total advance by selection is, for additive genes, 2N times the gain in the first generation but may be much greater than this for recessives, particularly if their initial frequency is low. (4) The half-life of any selection process will, for additive genes, not be greater than 1\cdot 4 N generations but may for rare recessives equal 2N. (5) The effect of an initial period of selection or inbreeding or of both together on the limits in further selection is discussed. It appears that the effects of restrictions in population size on the selection limit may be a useful diagnostic tool in the laboratory. (6) The treatment can be extended to deal with the limits of further selection after the crossing of replicate lines from the same population when the initial response has ceased. (7) In a selection programme of individual selection of equal intensity in both sexes, the furthest limit should be attained when half the population is selected from each generation. (8) The treatment can also be extended to include selection based on progeny or family records. Consideration of the optimum structure, as far as the limit is concerned, shows that the use of the information on relatives is always a sacrifice on the eventual limit for the sake of immediate gain in the early generations. The loss may, however, be small in large populations.
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(i) A computer simulation study has been made of selection on two linked loci in small populations, where both loci were assumed to have additive effects on the character under selection with no interaction between loci. If N is the effective population size, i the intensity of selection in standard units, α and β measure the effects of the two loci on the character under selection as a proportion of the pheno-typic standard deviation and c is the crossover distance between them, it was shown that the selection process can be completely specified by Ni α, Ni βand Nc and the initial gene frequencies and linkage disequilibrium coefficient. It is then easily possible to generalize from computer runs at only one population size. All computer runs assumed an initial population at linkage equilibrium between the two loci. Analysis of the results was greatly simplified by considering the influence of segregation at the second locus on the chance of fixation at the first (defined as the proportion of replicate lines in which the favoured allele was eventually fixed). (ii) The effects of linkage are sufficiently described by Nc. The relationship between chance of fixation at the limit and linkage distance (expressed as 2Nc /( 2Nc + 1)) was linear in the majority of computer runs. (iii) When gene frequency changes under independent segregation were small, linkage had no effect on the advance under selection. In general, segregation at the second locus had no detectable influence on the chance of fixation at the first if the gene effects at the second were less than one-half those at the first. With larger gene effects at the second locus, the chance of fixation passed through a minimum and then rose again. For two loci to have a mutual influence on one another, their effects on the character under selection should not differ by a factor of more than two. (iv) Under conditions of suitable relative gene effects, the influence of segregation at the second locus was very dependent on the initial frequency of the desirable allele. The chance of fixation at the first, plotted against initial frequency of the desirable allele at the second, passed through a minimum when the chance of fixation at the second locus was about 0·8. (v) A transformation was found which made the influence of segregation at the second locus on the chance of fixation at the first almost independent of initial gene frequency at the first and of gene effects at the first locus when these are small. (vi) In the population of gametes at final fixation, linkage was not at equilibrium and there was an excess of repulsion gametes. (vii) The results were extended to a consideration of the effect of linkage on the limits under artificial selection. Linkage proved only to be of importance when the two loci had roughly equal effects on the character under selection. The maximum effect on the advance under selection occurred when the chance of fixation at both of the loci was between 0·7 and 0·8. When the advance under selection is most sensitive to changes in recombination value, a doubling of the latter in no case increased the advance under selection by more than about 6%. The proportion selected to give maximum advance under individual selection (0·5 under independent segregation) was increased, but only very slightly, when linkage is important. (viii) These phenomena could be satisfactorily accounted for in terms of the time scale of the selection process and the effective size of the population within which changes of gene frequency at the locus with smaller effect must take place.
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The ability to predict individual breeding values in natural populations with known pedigrees has provided a powerful tool to separate phenotypic values into their genetic and environmental components in a nonexperimental setting. This has allowed sophisticated analyses of selection, as well as powerful tests of evolutionary change and differentiation. To date, there has, however, been no evaluation of the reliability or potential limitations of the approach. In this article, I address these gaps. In particular, I emphasize the differences between true and predicted breeding values (PBVs), which as yet have largely been ignored. These differences do, however, have important implications for the interpretation of, firstly, the relationship between PBVs and fitness, and secondly, patterns in PBVs over time. I subsequently present guidelines I believe to be essential in the formulation of the questions addressed in studies using PBVs, and I discuss possibilities for future research.
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Exploitation of fish populations can induce evolutionary responses in life histories. For example, fisheries targeting large individuals are expected to select for early maturation at smaller sizes, leading to reduced fecundity and thus also reduced fisheries yield. These predicted phenotypic shifts have been observed in several fish stocks, but disentangling the environmental and genetic causes behind them has proved difficult. Here, we review recent studies investigating phenotypic shifts in exploited populations and strategies for minimizing fisheries-induced evolution. Responses to selective harvesting will depend on species-specific life-history traits, and on community-level and environmental processes. Therefore, the detection of fisheries-induced evolution and successful fish stock management requires routine population monitoring, and a good understanding of genetics, relevant ecological processes and changing environmental conditions.
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Harvesting of wildlife populations by humans is usually targeted by sex, age or phenotypic criteria, and is therefore selective. Selective harvesting has the potential to elicit a genetic response from the target populations in several ways. First, selective harvesting may affect population demographic structure (age structure, sex ratio), which in turn may have consequences for effective population size and hence genetic diversity. Second, wildlife-harvesting regimes that use selective criteria based on phenotypic characteristics (e.g. minimum body size, horn length or antler size) have the potential to impose artificial selection on harvested populations. If there is heritable genetic variation for the target characteristic and harvesting occurs before the age of maturity, then an evolutionary response over time may ensue. Molecular ecological techniques offer ways to predict and detect genetic change in harvested populations, and therefore have great utility for effective wildlife management. Molecular markers can be used to assess the genetic structure of wildlife populations, and thereby assist in the prediction of genetic impacts by delineating evolutionarily meaningful management units. Genetic markers can be used for monitoring genetic diversity and changes in effective population size and breeding systems. Tracking evolutionary change at the phenotypic level in the wild through quantitative genetic analysis can be made possible by genetically determined pedigrees. Finally, advances in genome sequencing and bioinformatics offer the opportunity to study the molecular basis of phenotypic variation through trait mapping and candidate gene approaches. With this understanding, it could be possible to monitor the selective impacts of harvesting at a molecular level in the future. Effective wildlife management practice needs to consider more than the direct impact of harvesting on population dynamics. Programs that utilize molecular genetic tools will be better positioned to assess the long-term evolutionary impact of artificial selection on the evolutionary trajectory and viability of harvested populations.
  • Hendry