Genetic Effects of Multiple Generations of Supportive Breeding

Abstract: The practice of supporting weak wild populations by capturing a fraction of the wild individuals, bringing them into captivity for reproduction, and releasing their offspring into the natural habitat to mix with wild ones is called supportive breeding and has been widely applied in the fields of conservation biology and fish and wildlife management. This procedure is intended to increase population size without introducing exogenous genes into the managed population. Previous work examining the genetic effects of a single generation of supportive breeding has shown that although a successful program increases the census population size, it may reduce the genetically effective population size and thereby induce excessive inbreeding and loss of genetic variation. We expand and generalize previous analyses of supportive breeding and consider the effects of multiple generations of supportive breeding on rates of inbreeding and genetic drift. We derived recurrence equations for the inbreeding coefficient and coancestry, and thereby equations for inbreeding and variance effective sizes, under three models for selecting captive breeders: at random, preferentially among those born in captivity, and preferentially among those born in the wild. Numerical examples indicate that supportive breeding, when carried out successfully over multiple generations, may increase not only the census but also the effective size of the supported population as a whole. If supportive breeding does not result in a substantial and continuous increase of the census size of the breeding population, however, it might be genetically harmful because of elevated rates of inbreeding and genetic drift.     

Role of Familiarity and Preference in Reproductive Success in Ex Situ Breeding Programs

Abstract: Success of captive‐breeding programs centers on consistent reproduction among captive animals. However, many individuals do not reproduce even when they are apparently healthy and presented with mates. Mate choice can affect multiple parameters of reproductive success, including mating success, offspring production, offspring survival, and offspring fecundity. We investigated the role of familiarity and preference on reproductive success of female Columbia Basin pygmy rabbits (Brachylagus idahoensis) as measured by litter production, litter size, average number of young that emerged from the burrow, and average number of young that survived to 1 year. We conducted these studies on pygmy rabbits at the Oregon Zoo (Portland, Oregon, U.S.A.) and Washington State University (Pullman, Washington, U.S.A.) from February to June 2006, 2007, and 2008. Before mating, we housed each female adjacent to 2 males (neighbors). Female preference for each potential mate was determined on the basis of behavioral interactions observed and measured between the rabbits. We compared reproductive success between females mated with neighbor and non‐neighbor males and between females mated with preferred and nonpreferred males. Our findings suggest that mating with a neighbor compared with a non‐neighbor and mating with a preferred neighbor compared with a nonpreferred neighbor increased reproductive success in female pygmy rabbits. Litter production, average number of young that emerged, and average number of young that survived to 1 year were higher in rabbits that were neighbors before mating than in animals who were not neighbors. Pairing rabbits with a preferred partner increased the probability of producing a litter and was significantly associated with increased litter size. In captive breeding programs, mates are traditionally selected on the basis of genetic parameters to minimize loss of genetic diversity and inbreeding coefficients. Our results suggest that integrating genetic information with social dynamics and behavioral measures of preference may increase the reproductive output of the pygmy rabbit captive‐breeding program. Our findings are consistent with the idea that allowing mate choice and familiarity increase the reproductive success of captive‐breeding programs for endangered species.     

Supportive Breeding and Variance Effective Population Size

The practice of supporting weak, wild populations through release of individuals bred in captivity is becoming an increasingly important conservation measure. A frequently recommended form of such breeding-release activity refers to supportive breeding: a fraction of the target population is brought into captivity for reproduction, and the resulting progeny are released to mix with the wild segment of the population. We derived an expression for the variance effective size of a population managed through supportive breeding and discuss its relationship to previously published equations that are based on the assumption of random mating. We show that the effect of supportive breeding may be quite different on the inbreeding and the variance effective sizes. Whereas supportive breeding always results in a reduction of the inbreeding effective number, the variance effective number may either decrease, increase, or remain unchanged. We discuss these observations in relation to conservation management and suggest some general guidelines for supportive breeding situations. Our recommendations include making a distinction between inbreeding and variance effective numbers; taking particular care when dealing with organisms with high reproductive potential; assuring that the amount of drift be no larger than it would be without supportive breeding; and focusing primarily on the variance effective size of a population-that is, on the effective number directly related to the rate of loss of gene diversity.     

Optimal control of Atlantic population Canada geese

Management of Canada geese (Branta canadensis) can be a balance between providing sustained harvest opportunity while not allowing populations to become overabundant and cause damage. In this paper, we focus on the Atlantic population of Canada geese and use stochastic dynamic programming to determine the optimal harvest strategy over a range of plausible models for population dynamics. There is evidence to suggest that the population exhibits significant age structure, and it is possible to reconstruct age structure from surveys. Consequently the harvest strategy is a function of the age composition, as well as the abundance, of the population. The objective is to maximize harvest while maintaining the number of breeding adults in the population between specified upper and lower limits. In addition, the total harvest capacity is limited and there is uncertainty about the strength of density-dependence. We find that under a density-independent model, harvest is maximized by maintaining the breeding population at the highest acceptable abundance. However if harvest capacity is limited, then the optimal long-term breeding population size is lower than the highest acceptable level, to reduce the risk of the population growing to an unacceptably large size. Under the proposed density-dependent model, harvest is maximized by maintaining the breeding population at an intermediate level between the bounds on acceptable population size; limits to harvest capacity have little effect on the optimal long-term population size. It is clear that the strength of density-dependence and constraints on harvest significantly affect the optimal harvest strategy for this population. Model discrimination might be achieved in the long term, while continuing to meet management goals, by adopting an adaptive management strategy.     

Long-term demographic and genetic effects of releasing captive-born individuals into the wild

Because of continued habitat destruction and species extirpations, the need to use captive breeding for conservation purposes has been increasing steadily. However, the long-term demographic and genetic effects associated with releasing captive-born individuals with varied life histories into the wild remain largely unknown. To address this question, we developed forward-time, agent-based models for 4 species with long-running captive-breeding and release programs: coho salmon (Oncorhynchus kisutch), golden lion tamarin (Leontopithecus rosalia), western toad (Anaxyrus boreas), and Whooping Crane (Grus americana). We measured the effects of supplementation by comparing population size and neutral genetic diversity in supplemented populations to the same characteristics in unaltered populations 100 years after supplementation ended. Releasing even slightly less fit captive-born individuals to supplement wild populations typically resulted in reductions in population sizes and genetic diversity over the long term when the fitness reductions were heritable (i.e., due to genetic adaptation to captivity) and populations continued to be regulated by density-dependent mechanisms over time. Negative effects for species with longer life spans and lower rates of population replacement were smaller than for species with shorter life spans and higher rates of population replacement. Programs that released captive-born individuals over fewer years or that avoided breeding individuals with captive ancestry had smaller reductions in population size and genetic diversity over the long term. Relying on selection in the wild to remove individuals with reduced fitness mitigated some negative demographic effects, but at a substantial cost to neutral genetic diversity. Our results suggest that conservation-focused captive-breeding programs should take measures to prevent even small amounts of genetic adaptation to captivity, quantitatively determine the minimum number of captive-born individuals to release each year, and fully account for the interactions among genetic adaptation to captivity, population regulation, and life-history variation.     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Population Viability Analysis for a Small Population of Red-Cockaded Woodpeckers and an Evaluation of Enhancement Strategies

We performed a series of population and pedigree analyses to examine the viability of a small Red-cockaded Woodpecker ( Picoides borealis ) population located at the Savannah River Site, in Barnwell and Aiken counties of South Carolina. The population's existence and future survival are precarious. As few as four individuals, including just one breeding pair, comprised this population in 1985. Now, primarily because of experimental transformation of birds from other areas, the population has increased to 25. As of 1990, genealogy pedigree analysis showed that the respective contribution of 14 founders to the extant population has not been equal. Founder gender equivalents are low (5.4) but could reach 9.2 if poorly-represented founders were to produce offspring. The fraction of founder gene diversity retained in the current population is 0.91. Successful recovery strategies would ensure 95% probability of population survival while maintaining 90% heterozygosity for 200 years. Viability analyses indicated that, depending on relative effects of inbreeding depression and stochastic environmental events, the Savannah River Site population has a 68–100% chance of extinction during this period. Annual translocation into the population of at least three females and two males for a 10-year period will achieve a 96% probability of survival for 200 years. Even with translocation of numerous males and females per year (up to 50 of each), the 90% heterozygosity goal may not be achieved. We discuss recommendations for choosing individuals for translocation logistical constraints on achieving recovery objectives, and limitations of our modeling approach.     

Intra-group relatedness affects parental and helper investment rules in offspring care

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.     

Mitochondrial DNA Variability in the Endangered Razorback Sucker (Xyrauchen texanus): Analysis of Hatchery Stocks and Implications for Captive Propagation

The razorback sucker ( Xyrauchen texanus ) is a large, long-lived catostomid fish endemic to the Colorado River drainage of western North America, endangered because of recruitment failure. Efforts to preserve the species have emphasized artificial propagation and reintroduction. Given the importance of maintaining genetic diversity in such a program, we examined mitochondrial DNA diversity in a source population (Lake Mohave, Arizona-Nevada) and three hatchery-produced year classes (1987, 1989, 1990). The source contained considerable variation, indicated by high haplotype diversity ( ĥ = 0.97) and a large number of unique haplotypes (17 in 25 individuals). Diversity also was high in the 1987 ( ĥ = 0.89, 6 haplotypes in 10 individuals) and 1989 hatchery-produced year classes ( ĥ = 0.91, 7 in 11), but significantly lower in the 1990 year class ( ĥ = 0.71, 4 in 10). Low diversity in the last class was likely because of differences among females in fecundity, viability of progeny, or both. Because natural populations have collapsed throughout the species' range, we must identify methods that preserve the most diversity. We examined three potential alternatives: standard hatchery propagation, natural spawning in predator-free environments, and protective custody of larvae collected from the lake with reintroduction after growth to a size likely to survive. The last is the preferred alternative and should be pursued as the most cost-effective option for preserving genetic diversity in the razorback sucker.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

On the meta-analysis of response ratios for studies with correlated and multi-group designs

A common effect size metric used to quantify the outcome of experiments for ecological meta-analysis is the response ratio (RR): the log proportional change in the means of a treatment and control group. Estimates of the variance of RR are also important for meta-analysis because they serve as weights when effect sizes are averaged and compared. The variance of an effect size is typically a function of sampling error; however, it can also be influenced by study design. Here, I derive new variances and covariances for RR for several often-encountered experimental designs: when the treatment and control means are correlated; when multiple treatments have a common control; when means are based on repeated measures; and when the study has a correlated factorial design, or is multivariate. These developments are useful for improving the quality of data extracted from studies for meta-analysis and help address some of the common challenges meta-analysts face when quantifying a diversity of experimental designs with the response ratio.     

Population dynamics of the endangered fan mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis> in a marine lake: a metapopulation matrix modeling approach

A metapopulation, time-invariant, stage-classified matrix model was developed to assess the dynamics of an important Pinna nobilis population in the marine Lake Vouliagmeni (Korinthiakos Gulf, Greece). The main aim of the study was to provide insight on the life cycle of the fan mussel and reveal influential factors for its population dynamics, with a special focus on the effect of poaching. The size of the fan mussel shell was selected as a state variable, and the model consisted of five size classes. The lake was divided in two regions, a shallow region of high (illegal) fishing mortality and high recruitment (region 1) and a deeper region of low mortality and low recruitment (region 2). The estimation of the transition matrix (stage-specific growth and mortality probabilities) was based on a tagging survey between 2005 and 2006, while independent annual surveys for abundance estimation using distance sampling techniques were utilized for the estimation of recruitment and stage-specific fertilities. The population was found to be increasing with an intrinsic rate of increase r = 0.038; however, r was not statistically different from zero. The life expectancy and expected lifetime offspring production of individuals in region 1 was markedly lower than that of individuals in region 2. Due to poaching, the life expectancy of a yearling fan mussel was less than 2.5 years in region 1, while it was almost 12 years in region 2. The highest expected annual natural mortality of fan mussels occurred on their first year of life after settlement (~43%) and greatly declined at greater sizes. Perturbation analysis revealed that the population growth rate was most sensitive to the vital rates of the larger size classes in region 2 and to fertilities corresponding to offspring that settled in the same region. The spatial distribution and abundance of the species was greatly dependent on the extent of poaching, which caused a size segregation of individuals, with small and young individuals being abundant in region 1, and larger and older individuals being restricted in region 2. If poaching ceased, the fan mussel population would be increasing with a significantly higher intrinsic rate of increase (r = 0.186), while if region 2 was also illegally exploited at the same intensity as region 1, the fan mussel population would be decreasing with r = −0.364 and would eventually collapse. The existence of refuge areas, where fan mussels may grow and reproduce, providing adjacent areas with offspring, seems crucial for the viability of local populations. Transplantation of fan mussels from high mortality areas to low mortality refuges might prove to be an effective measure to protect local populations of the species.     

Genealogical Analysis of a Closed Herd of Black Hairless Iberian Pigs

Abstract: Livestock breeds are recognized as important components of world biodiversity. The Iberian pig is a swine breed well adapted to the Mediterranean forest ecosystem and provides cured products of high quality. The ancient population of Iberian pigs (  Sus scrofa meridionalis ) was differentiated in several local types, the black hairless pigs representing the fattest genetic type. The conservation program of the Guadyerbas strain has maintained this germplasm isolated since 1945 as a closed population in an experimental herd. The complete pedigree, with 1000 breeding animals descending from 24 founders, has been used to measure along the successive cohorts of breeding animals the effective number of founders, effective number of nonfounders, founder genome equivalents, and expected number of founders' surviving alleles. For the last cohort, the values were 10.34, 1.42, 1.25, and 4.06, respectively. The evolution of inbreeding and coancestry and its components attributable to each founder were also studied. The rate of increase in inbreeding and coancestry was 2.21% per cohort, or 0.906% per year. Finally, the effect of family structure and mating tactics on the evolution of coancestry was also analyzed. The greatest rates of coancestry per cohort were attributable to unbalanced family sizes, and the use of minimum coancestry matings effectively delayed the increase in inbreeding.     

A role for selective contraception of individuals in conservation

Contraception has an established role in managing overabundant populations and preventing undesirable breeding in zoos. We propose that it can also be used strategically and selectively in conservation to increase the genetic and behavioral quality of the animals. In captive breeding programs, it is becoming increasingly important to maximize the retention of genetic diversity by managing the reproductive contribution of each individual and preventing genetically suboptimal breeding through the use of selective contraception. Reproductive suppression of selected individuals in conservation programs has further benefits of allowing animals to be housed as a group in extensive enclosures without interfering with breeding recommendations, which reduces adaptation to captivity and facilitates the expression of wild behaviors and social structures. Before selective contraception can be incorporated into a breeding program, the most suitable method of fertility control must be selected, and this can be influenced by factors such as species life history, age, ease of treatment, potential for reversibility, and desired management outcome for the individual or population. Contraception should then be implemented in the population following a step‐by‐step process. In this way, it can provide crucial, flexible control over breeding to promote the physical and genetic health and sustainability of a conservation dependent species held in captivity. For Tasmanian devils (Sarcophilus harrisii), black‐flanked rock wallabies (Petrogale lateralis), and burrowing bettongs (Bettongia lesueur), contraception can benefit their conservation by maximizing genetic diversity and behavioral integrity in the captive breeding program, or, in the case of the wallabies and bettongs, by reducing populations to a sustainable size when they become locally overabundant. In these examples, contraceptive duration relative to reproductive life, reversibility, and predictability of the contraceptive agent being used are important to ensure the potential for individuals to reproduce following cessation of contraception, as exemplified by the wallabies when their population crashed and needed females to resume breeding.     

Effective Population Size in Winter-Run Chinook Salmon

Winter-run chinook salmon from the Sacramento River, California, is federally listed as endangered. Since 1989 there has been aprogram to augment the natural population by capturing adults, artificially spawning them, raising tine young and releasing the smolt. Here we estimate the effective population size of these captive-raised fish, the natural run, and the combination of both groups over the three-year period from 1991 to 1993. We find that the most appropriate estimate of the effective population size of the captive-raised progeny is a variance estimate of effective population size standardized so that the number of released smolts returning to spawn was the same as the number of spawners used to produce the smolts originally. We have generated 10,000 random samples to simulate returns from these released progeny. The estimates of the effective population sizes in 1991, 1992, and 1993 were only 7.02, 19.07, and 7.74, respectively. We then determined limits on the effective population size of the natural run based on 0.1 and 0.333 of the run-size estimates. Using estimates of the captive proportion of the run, the minimum estimates of the effective population size of the overall run for the three years were 21.9, 127.3, and 39.0, and the maximum estimates were 61.6, 401.0, and 108.7. It does not appear that the hatchery program has reduced the overall effective population size. The run sizes in each year are extremely low, however, and it is possible that fish will be lost from this run in one of the years in the immediate future, making reestablishment of a healthy run even more difficult.     

Preserving Population Allele Frequencies in Ex Situ Conservation Programs

Abstract: Optimization of contributions of parents to progeny by minimizing the average coancestry of the progeny is an effective strategy for maintaining genetic diversity in ex situ conservation programs, but its application on the basis of molecular markers has the negative collateral effect of homogenizing the allelic frequencies at each locus. Because one of the objectives of a conservation program is to preserve the genetic composition of the original endangered population, we devised a method in which markers are used to maintain the allele frequency distribution at each locus as closely as possible to that of the native population. Contributions of parents were obtained so as to minimize changes in allele frequency for a set of molecular markers in a population of reduced size. We used computer simulations, under a range of scenarios, to assess the effectiveness of the method in comparison with methods in which contributions of minimum coancestry are sought, either making use of molecular markers or genealogical information. Our simulations indicated that the proposed method effectively maintained the original distribution of allele frequencies, particularly under strong linkage, and maintained acceptable levels of genetic diversity in the population. Nevertheless, contributions of minimum coancestry determined from pedigree information but ignoring the genealogy previous to the conservation program, was the most effective method for maintaining allelic frequencies in realistic situations.     

Modeling Problems in Conservation Genetics Using Captive Drosophila Populations: Consequences of Equalization of Family Sizes

Equalization of family sizes is recommended for use in captive breeding programs, as it is predicted to double effective population sizes, reduce inbreeding, and slow the loss of genetic variation. The effects of maintaining small captive populations with equalization of family sizes versus random choice of parents on levels of inbreeding genetic variation, reproductive fitness, and effective population sizes ( N _e) were evaluated in 10 lines of each treatment maintained with four pairs of parents per generation. The mean inbreeding coefficient ( F ) increased at a significantly slower rate with equalization than with random choice (means of 0.35 and 0.44 at generation 10). Average heterozygosities at generation 10, based on six polymorphic enzyme loci, were significantly higher with equalization (0.149) than with random choice (0.085), compared to the generation 0 level of 0.188. The competitive index measure of reproductive fitness at generation 11 was more than twice as high with equalization as with random choice, both being much lower than in the outbred base population. There was considerable variation among replicate lines within treatments in all the above measures and considerable overlap between lines from the two treatments. Estimates of N _e for equalization were greater than those for random choice, whether estimated from changes in average heterozygosities or from changes in F. Equalization of family sizes can be unequivocally recommended for use in the genetic management of captive populations.     

Variation in litter size: a test of hypotheses in Richardson's ground squirrels

We studied litter size variation in a population of Richardson's ground squirrels (Spermophilus richardsonii) in Alberta, Canada, from 1987 to 2004. Litter size at first emergence of juveniles from the natal burrow ranged from 1 to 14; the most common litter sizes, collectively accounting for 41.0% of 999 litters, were 6 and 7. The number of offspring surviving to adulthood (attained on emergence from hibernation as yearlings) increased with increasing litter size, a result that was not predicted by Lack's "optimal litter size" hypothesis, Mountford's "cliff-edge" effect, or the "bad-years" effect. Contrary to the negative effects predicted by the "cost of reproduction" hypothesis, litter size had no significant influence on survival of mothers to the subsequent year or on the size of the subsequent litter. Rather, our results best fit the predictions of the "individual optimization" hypothesis, which suggests that litter size is determined by the body condition and environmental circumstances of each mother. Supporting this hypothesis, survival of individual offspring was not significantly associated with litter size. Additionally, year-to-year changes in maternal body mass at mating were positively associated with concurrent changes in litter size (r = 0.56), suggesting that litter size depends on the body condition of the mother. Because the mean number of recruits to adulthood increased as litter size increased (r2 = 0.96) and litter size increased with maternal condition, offspring productivity was greater for mothers in better body condition.     

Cooperatively breeding carrion crows adjust offspring sex ratio according to group composition

In cooperatively breeding species, parents should bias offspring sex ratio towards the philopatric sex to obtain new helpers (helper repayment hypothesis). However, philopatric offspring might increase within-group competition for resources (local resource competition hypothesis), diluting the benefits of helper acquisition. Furthermore, benefits of offspring sex bias on parents’ fitness may depend on different costs of production and/or different breeding opportunities of sons and daughters. Because of these counteracting factors, strategies of offspring sex allocation in cooperative species are often difficult to investigate. In carrion crows Corvus corone corone in northern Spain, sons are more philopatric and more helpful at the nest than daughters, which disperse earlier and have higher chances to find a breeding vacancy. Consistent with the helper repayment hypothesis, we found that crows fledged more sons in groups short of subordinate males than in groups with sufficient helper contingent, where daughters were preferred. Crow females also proved able to bias primary sex ratio, allocating offspring sex along the hatching sequence in a way that provided the highest fledging probability to sons in the first breeding attempt and to daughters in the following ones. The higher cost of producing male offspring may explain this pattern, with breeding females shifting to the cheapest sex (female) as a response to the costs generated by previous reproductive attempts. Our results suggest complex adjustments of offspring sex ratio that allowed crows to maximize the value of daughters and sons.     

Effectiveness of Predator Removal for Enhancing Bird Populations

Abstract: Predation pressure on vulnerable bird species has made predator control an important issue for international nature conservation. Predator removal by culling or translocation is controversial, expensive, and time‐consuming, and results are often temporary. Thus, it is important to assess its effectiveness from all available evidence. We used explicit systematic review methodology to determine the impact of predator removal on four measurable responses in birds: breeding performance (hatching success and fledging success) and population size (breeding and postbreeding). We used meta‐analysis to summarize results from 83 predator removal studies from six continents. We also investigated whether characteristics of the prey, predator species, location, and study methodology explained heterogeneity in effect sizes. Removing predators increased hatching success, fledging success, and breeding populations. Removing all predator species achieved a significantly larger increase in breeding population than removing only a subset. Postbreeding population size was not improved on islands, or overall, but did increase on mainlands. Heterogeneity in effect sizes for the four population parameters was not explained by whether predators were native or introduced; prey were declining, migratory, or game species; or by the study methodology. Effect sizes for fledging success were smaller for ground‐nesting birds than those that nest elsewhere, but the difference was not significant. We conclude that current evidence indicates that predator removal is an effective strategy for the conservation of vulnerable bird populations. Nevertheless, the ethical and practical problems associated with predator removal may lead managers to favor alternative, nonlethal solutions. Research is needed to provide and synthesize data to determine whether these are effective management practices for future policies on bird conservation.