High-quality Individuals Research Articles

Breeding experience, but not mate retention, determines the breeding performance in a passerine bird

Individuals that retain their former partners often perform better than conspecifics that switch partners. This may be due to high-quality individuals being more faithful to their partners and more productive. Investigations of the fitness benefits of mate retention that also control for potential confounding effects of individuals are scarce. We studied the influence of mate retention and breeding experience on breeding performance of the hair-crested drongo (Dicrurus hottentottus) by carrying out both cross-sectional and longitudinal analyses. Pairs with longer pair-bond duration did not fledge more young or fledglings of better body condition nor did they produce more or better fledglings than newly formed pairs consisting of at least 1 experienced breeder, that is, individuals that had bred before. Individuals produced fewer fledglings when they were paired with an inexperienced breeder, especially when females were paired with inexperienced males. Although clutch size was not affected by mate retention or breeding experience, pairs consisting of inexperienced breeder(s) had a relatively higher predation rate of eggs and/or nestlings because they may be less effective in nest defense. The onset of breeding was advanced in the year following mate retention, but not in the second year thereafter, when pairs still remained together. Furthermore, only the breeding experience of the male determined the onset of breeding: Pairs consisting of inexperienced males bred later in the season. Our results suggest that breeding experience, and particularly the breeding experience of the male, but not mate retention, is important in determining the breeding performance of hair-crested drongos.

Plasma carotenoid concentrations of incubating American kestrels (Falco sparverius) show annual, seasonal, and individual variation and explain reproductive outcome.

In wild birds, the proximate and ultimate factors that affect circulating carotenoid concentrations remain poorly understood. We studied variation in plasma carotenoid concentrations across several scales: annual, seasonal, pair, territory and individual, and evaluated whether carotenoid levels explained reproductive outcome of wild American kestrels (Falco sparverius). We sampled plasma carotenoid concentrations of 99 female and 80 male incubating kestrels from April-June in 2008-2012. Plasma carotenoid concentrations were explained by an interaction between year and sex, date, and random effects for pair and individual identity. In general, plasma carotenoid concentrations of males were significantly higher than females, but this depended on year. Within a breeding season, earlier nesting kestrels had higher carotenoid concentrations than later nesting kestrels, a pattern that is coincident with seasonal trends in local fitness. Pair and individual identity explained variation in carotenoid concentrations suggesting that carotenoid concentrations of mated birds were correlated, and some individuals consistently maintained higher carotenoid levels than others. Male carotenoid concentrations were positively associated with number of young fledged per pair. These results are consistent with the hypothesis that higher quality individuals have higher carotenoid levels compared to lower quality individuals, despite annual variations in carotenoid availability.

Song repertoire size, not territory location, predicts reproductive success and territory tenure in a migratory songbird

In territorial animals occupying environments that vary in quality over the landscape, high-quality individuals are predicted to monopolize high-quality territories. Thus, in many cases it may be difficult to disentangle the relative effects of individual quality from those of territory quality on long-term fitness. We used a 9-year field data set from a migratory population of Eastern Song Sparrows (Melospiza melodia melodia (A. Wilson, 1810)) to evaluate the relative contributions of male song quality (as measured by song repertoire size) and territory location to fitness components including annual reproductive success, overwinter return rates, and between-year territory tenure. Song repertoire size did not predict territory location, allowing us to evaluate territory location and song quality separately. Song repertoire size, but not territory location, predicted annual reproductive success. Moreover, males with larger repertoires moved smaller distances between subsequent breeding seasons, suggesting more successful territory tenure. There was no effect of either repertoire size or territory location on overwinter return. We conclude that intrinsic male phenotype, indicated by song repertoire size, is an important predictor of male fitness, independent of breeding-territory location in this migratory population, and that the value of specific territories may depend largely on previous experience.

Quantifying the influence of measured and unmeasured individual differences on demography.

1. Demographic rates can vary not only with measured individual characters like age, sex and mass but also with unmeasured individual variables like behaviour, genes and health. 2. Predictions from population models that include measured individual characteristics often differ from models that exclude them. Similarly, unmeasured individual differences have the potential to impact predictions from population models. However, unmeasured individual differences are rarely included in population models. 3. We construct stage- and age-structured models (where stage is mass) of a roe deer population, which are parameterized from statistical functions that either include, or ignore, unmeasured individual differences. 4. We found that mass and age structures substantially impacted model parameters describing population dynamics, as did temporal environmental variation, while unmeasured individual differences impacted parameters describing population dynamics to a much smaller extent once individual heterogeneity related to mass and age has been included in the model. We discuss how our assumptions (unmeasured individual differences only in mean trait values) could have influenced our findings and under what circumstances unmeasured individual differences could have had a larger impact on population dynamics. 5. There are two reasons explaining the relative small influence of unmeasured individual differences on population dynamics in roe deer. First, individual body mass and age both capture a large amount of individual differences in roe deer. Second, in large populations of long-lived animals, the average quality of individuals (independent of age and mass) within the population is unlikely to show substantial variation over time, unless rapid evolution is occurring. So even though a population consisting of high-quality individuals would have much higher population growth rate than a population consisting of low-quality individuals, the probability of observing a population consisting only of high-quality individuals is small.

High Standards Enhance Inequality in Idealized Labor Markets

We built a simple model of an idealized labor market, in which there is no objective difference in average quality between groups and hiring decisions are not biased in favor of any particular group. Our results show that inequality in employment emerges necessarily also in such idealized situations due to the limited supply of high quality individuals and asymmetric information. Inequalities are exacerbated when employers have high standards and keep only the best workers in house. We found that ambitious workers get higher quality jobs even if ambition does not correlate or even negatively correlates with internal quality. Our findings help to corroborate empirical findings on higher employment discrepancies in high rather than low status jobs.

Breeding performance and survival in the peregrine falcon Falco peregrinus support an age‐related competence improvement hypothesis mediated via an age threshold

We studied the effects of age on breeding performance and survival probability in a peregrine falcon population, using data from a long term monitoring programme (carried out over 16 yr), in which we were able to identify individual birds. We compared the breeding performance and survival of yearling breeders, first‐time adult breeders and adult breeders. We found significant differences in breeding performance but not in survival. Yearling breeders had lower breeding success than older individuals but the breeding performance of inexperienced adults did not differ from that of experienced adults. We did not find changes in terminal breeding success since peregrines in their last year of life sustained the performance levels shown in previous years although with increasing variability. We found no evidence that attempting to breed affected survival probability in any age group. We argue that differences in breeding performance are related to age, not to breeding experience, and that there is an age threshold, coincident with the development of adult plumage, after which breeding performance is not affected either by age or experience. Peregrines that start breeding as yearlings are likely to have greater lifetime reproductive success than birds entering the breeding pool as adults. Consequently, such birds may represent a set of high quality individuals. Our results support the age‐related competence improvement hypothesis as being the relevant explanation for the increase in breeding performance with age.

Senescence or selective disappearance? Age trajectories of body mass in wild and captive populations of a small-bodied primate

Classic theories of ageing consider extrinsic mortality (EM) a major factor in shaping longevity and ageing, yet most studies of functional ageing focus on species with low EM. This bias may cause overestimation of the influence of senescent declines in performance over condition-dependent mortality on demographic processes across taxa. To simultaneously investigate the roles of functional senescence (FS) and intrinsic, extrinsic and condition-dependent mortality in a species with a high predation risk in nature, we compared age trajectories of body mass (BM) in wild and captive grey mouse lemurs (Microcebus murinus) using longitudinal data (853 individuals followed through adulthood). We found evidence of non-random mortality in both settings. In captivity, the oldest animals showed senescence in their ability to regain lost BM, whereas no evidence of FS was found in the wild. Overall, captive animals lived longer, but a reversed sex bias in lifespan was observed between wild and captive populations. We suggest that even moderately condition-dependent EM may lead to negligible FS in the wild. While high EM may act to reduce the average lifespan, this evolutionary process may be counteracted by the increased fitness of the long-lived, high-quality individuals.

Dead before detection: addressing the effects of left truncation on survival estimation and ecological inference for neonates

SummaryNeonate survival is a key life history trait, yet remains challenging to measure in wild populations because neonates can be difficult to capture at birth. Estimates of survival from neonates that are opportunistically captured might be inaccurate because some individuals die before sampling, resulting in data that are left truncated. The resulting overestimation of survival rates can further affect ecological inference through biased estimates of covariate effects in survival models, yet is not addressed in most studies of animal survival. Here, we quantify the effects of left truncation on survival estimates and subsequent ecological inference.Vaginal implant transmitters (VITs) enable capture of ungulates at birth, yielding data without left truncation. The effects of left truncation on survival estimation were quantified using age‐dependent survival models for VIT and opportunistically captured neonatal deer. Differences in daily survival rates (DSRs) and cumulative survival probability were calculated for the first 70 days of life. In addition, left truncation was simulated by removing fawns that died during the first 1 or 2 days of life from the VIT‐caught sample, isolating the effect of left truncation.Cumulative probability of survival during the first 70 days of life was overestimated by 7–23% for fawns caught opportunistically compared with those caught by VIT, depending on model design. Differences in DSRs were large at age 1 day, but had converged by age 30 days. Simulated left truncation resulted in overestimates of survival of up to 31%. Model selection and covariate coefficients were strongly affected by left truncation, producing spurious ecological inference, including changes to sign and/or magnitude of inferred effects of all covariates.We recommend (i) every effort be made to capture neonates; (ii) consistent capture methods, using at least in part non‐truncating techniques, be implemented across years and study areas; and (iii) exclusion of left‐truncated data from survival estimates until DSRs converge with those calculated from non‐truncated data. This work emphasizes the importance of accounting for left truncation in survival estimation for any species with strong age‐dependent survival in order to prevent biased conclusions produced by sampling method rather than true ecological effects.

Factors influencing double brooding in Eurasian Hoopoes Upupa epops

Double brooding may be a good strategy for short‐lived species to maximize annual and lifetime reproductive success (ARS and LRS, respectively). Nevertheless, there is typically individual variation in the probability of producing a second clutch. Here we evaluate factors that influence the decision to double brood in the Eurasian Hoopoe Upupa epops. Analyses of an 11‐year dataset showed that 36% of the females and 21% of the males produced a second clutch after successfully raising a first clutch. Double‐brooded females had higher ARS (9.1 ± 1.9 fledglings; mean ± se) and LRS (0.93 ± 0.08 recruits) than single‐brooded females (ARS: 4.5 ± 2.1 fledglings; LRS: 0.36 ± 0.03 recruits). This suggests that double brooding is adaptive in Hoopoes, and raises the question of why most individuals only produce one clutch per season. The probability of double brooding varied only slightly between years, suggesting that it is influenced by individual characteristics rather than by external, population‐level environmental factors. In both sexes, the probability of double brooding increased with earlier timing of the first clutch, and the timing of reproduction was the most important factor influencing reproductive success. The latter is likely to be mediated by changes in resources during the season. The probability of double brooding also increased slightly with female age, due to differences in intrinsic quality among females rather than to a gain in experience. In contrast to many other studies, the probability of double brooding increased with an increasing number of fledglings from the first clutch, suggesting that it is a strategy of individuals of high quality. Taken together, we show that the individual quality of the breeder and the timing of their first clutches are key factors influencing the decision to double brood, and thereby that they are important determinants of reproductive performance in Eurasian Hoopoes.

Global Warming May Depress Avian Population Fecundity by Selecting Against Early-Breeding, High-Quality Individuals in Northern Populations of Single-Brooded, Long-Lived Species

Global climate is changing at an unprecedented rate. Adjustments to breeding phenology represent responses to current climate change, and some climatic effects have negatively affected population reproductive performances. Here we simulated the possibility that climate warming-induced changes in the timing of egg-laying may modify the phenotype composition (i.e. proportion of high- vs. low-quality phenotypes) of avian populations of single-brooded, long-lived species in northern countries (where asymmetric changes of weather conditions are more pronounced), therefore affecting the internal structure and long-term stability of populations. In northern countries, prelaying temperatures have risen and laying and hatching are expected to occur earlier. However, because post-hatching temperatures have remained stable, early chicks hatch under conditions of low temperature and great precipitation, and may face increased mortality. Because early breeders are generally high-quality individuals, their contribution to the future recruitment of the breeding population will decrease, engendering a doubly negative effect: (1) the number of offspring in a population will be lower than in the past because of higher mortality in the largest broods; and (2) the population will increasingly be composed of the offspring of low-quality individuals, which will consequently decrease fitness of the entire population.

The spectacular human nose: an amplifier of individual quality?

Amplifiers are signals that improve the perception of underlying differences in quality. They are cost free and advantageous to high quality individuals, but disadvantageous to low quality individuals, as poor quality is easier perceived because of the amplifier. For an amplifier to evolve, the average fitness benefit to the high quality individuals should be higher than the average cost for the low quality individuals. The human nose is, compared to the nose of most other primates, extraordinary large, fragile and easily broken—especially in male–male interactions. May it have evolved as an amplifier among high quality individuals, allowing easy assessment of individual quality and influencing the perception of attractiveness? We tested the latter by manipulating the position of the nose tip or, as a control, the mouth in facial pictures and had the pictures rated for attractiveness. Our results show that facial attractiveness failed to be influenced by mouth manipulations. Yet, facial attractiveness increased when the nose tip was artificially centered according to other facial features. Conversely, attractiveness decreased when the nose tip was displaced away from its central position. Our results suggest that our evaluation of attractiveness is clearly sensitive to the centering of the nose tip, possibly because it affects our perception of the face’s symmetry and/or averageness. However, whether such centering is related to individual quality remains unclear.

Intra-sexual competition alters the relationship between testosterone and ornament expression in a wild territorial bird

In a reliable signalling system, individual quality is expected to mediate the costs associated with ornamental displays, with relatively lower costs being paid by individuals of higher quality. These relative costs should depend not only on individual quality, but also on levels of intra-sexual competition. We explored the current and delayed effects that testosterone implants have on bird ornamentation in populations with contrasted population densities, as a proxy for intra-sexual competition. In a replicated experiment, we manipulated testosterone in 196 yearling male red grouse Lagopus lagopus scoticus in autumn in populations of high and low levels of intra-sexual competition. Males were assigned to one of three exogenous testosterone (T) treatments: empty implants (T0), small T implants (T1) or larger T implants (T2). We monitored subsequent changes in testosterone levels, ornament size and carotenoid-based colouration, carotenoid levels and body condition from autumn to spring. Testosterone implants increased testosterone levels, comb redness and comb size, and decreased body condition but these effects depended on levels of intra-sexual competition. Specifically, T2-implanted birds increased testosterone levels and comb size more, and reduced body condition more, in populations where intra-sexual competition was low. In the following spring, testosterone levels of T2-treated birds kept increasing in populations where intra-sexual competition was high but not in populations where intra-sexual competition was low. Our results highlight that levels of intra-sexual competition alter the relationship between testosterone levels and ornament expression, influencing their condition-dependence; they also indicate that the outcome of standard hormone manipulation conducted in free-living animals vary depending on the population context.

OPTIMAL INVESTMENT IN SOCIAL SIGNALS

This study is an attempt to determine how much individuals should invest in social communication, depending on the type of relationships they may form. Two simple models of social relationships are considered. In both models, individuals emit costly signals to advertise their "quality" as potential friends. Relationships are asymmetrical or symmetrical. In the asymmetrical condition (first model), we observe that low-quality individuals are discouraged from signaling. In the symmetrical condition (second model), all individuals invest in communication. In both models, high-quality individuals (elite) do not compete and signal uniformly. The level of this uniform signal and the size of the "elite" turn out to be controlled by the accuracy of signals. The two models may be relevant to several aspects of animal and human social communication.

Evolution of within-colony distribution patterns of birds in response to habitat structure.

It has long been suggested that habitat structure affects how colonial birds are distributed within their nesting aggregations, but this hypothesis has never been formally tested. The aim of this study was to test for a correlated evolution between habitat heterogeneity and within-colony distributions of Ciconiiformes by using Pagel’s general method of comparative analysis for discrete variables. The analysis indicated that central-periphery gradients of distribution (high-quality individuals occupying central nesting locations) prevail in species breeding in homogeneous habitats. These were mainly ground-nesting larids and spheniscids, where clear central-periphery patterns were recorded in ca. 80 % of the taxa. Since homogeneous habitats provide little variation in the physical quality of nest sites, central nesting locations should be largely preferred because they give better protection against predators by means of more efficient predator detection and deterrence. By contrast, central-periphery gradients tended to be disrupted in heterogeneous habitats, where 75 % of colonial Ciconiiform species showed uniform patterns of distribution. Under this model of distribution, edge nest sites of high physical quality confer higher fitness benefits in comparison to low-quality central sites, and thus, high-quality pairs are likely to choose nest sites irrespectively of their within-colony location. Breeding in homogeneous habitats and uniform distribution patterns were identified as probable ancestral states in Ciconiiformes, but there was a significant transition rate from uniform to central-periphery distributions in species occupying homogeneous habitats.

From recruitment to senescence: food shapes the age‐dependent pattern of breeding performance in a long‐lived bird

We used a long-term data set (26 years) from Audouin's Gull (Larus audouinii), a long-lived seabird, to address the relationship between the age-dependent pattern of reproductive performance and environmental conditions during breeding. Although theoretical models predict that the youngest and oldest breeders (due to inexperience and senescence, respectively) will perform less well than intermediate age classes, few empirical data exist regarding how this expected pattern varies with food availability. To assess the influence of age and food availability (corrected by population size of the main consumers to take into account density dependence) on a number of breeding parameters (laying dates, egg volume, clutch size, and hatching success), we modeled mean and variances of these parameters by incorporating heterogeneity into generalized linear models. All parameters varied with age and to different degrees, depending on food availability. As expected, performance improved with increased food supply, and the observed age pattern was quadratic, with poorer breeding performances occurring in extreme ages. For most parameters (except for laying dates, for which age and food did not interact), the pattern changed with food somewhat unexpectedly; the differences in performance between age classes were higher (i.e., the quadratic pattern was more noticeable) when food was more readily available than when food availability was lower. We suggest that, under poor environmental conditions, only high-quality individuals of the younger and older birds bred and that the differences in breeding performance between age classes were smaller. Although variances for egg volume were constant, variances for laying dates were highest for the youngest breeders and tended to decrease with age, either due to the selection of higher-quality individuals or to a greater frequency of birds skipping breeding with age, especially when food was in low supply. Our results show that mean and variances of breeding parameters changed with age, but that this pattern was different for each parameter and also varied according to food availability. It is likely that, other than food, certain additional factors (e.g., sex, cohort effects, density dependence) also influence changes in breeding performance with age, and this may preclude the finding of a common pattern among traits and among studies on different taxa.

Age-dependent relationships between multiple sexual pigments and condition in males and females

Author(s): Grunst, AS; Rotenberry, JT; Grunst, ML | Abstract: The reliability of sexual signaling may change across age classes due to shifts in resource allocation patterns. Two contrasting hypotheses exist regarding how the condition dependence of ornaments may shift with age, and both have received empirical support. On one hand, ornaments may more reliably reflect condition and quality in older individuals, because younger individuals of high quality invest in survival over signaling effort. On the other hand, the condition dependence of ornaments may decline with age, if older individuals in poor condition terminally invest in ornaments, or if resource constraints decline with age. Further, the expression and condition dependence of different ornaments may shift with age in unique ways, such that multifaceted sexual displays maintain reliable signaling across age classes. In yellow warblers (Setophaga petechia) of both sexes, we assessed how relationships between carotenoid-and phaeomelanin-based sexual pigmentation, prenesting body reserves, and condition at molt (reflected by growth bars and feather quality) vary across age classes. Melanin coverage correlated with condition at molt across age classes in males and showed high repeatability in both sexes. In contrast, carotenoid saturation increased longitudinally with age in males and correlated with condition at molt in different age classes in the 2 sexes. Specifically, carotenoid saturation correlated positively with condition at molt in younger, but not older males, whereas in females, the situation was reversed, with a positive correlation present only in older females. Results suggest that age-dependent signaling may promote maintenance of multifaceted sexual displays and that agedependent signaling dynamics depend on sex. © The Author 2013. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. All rights reserved.

Follicular dynamics in Plains zebra (Equus burchelli): preliminary results

The current knowledge on reproductive biology of zebras is very vague and based primarily on observations of free roaming animals in nature or captivity, or obtained from genital tracts collected after culling. Information regarding zebra estrous cycles and ovarian follicular dynamics is lacking [Nunez et al., In: Equine Reproduction, 2nd Edition, 2011, v. 2, 2851-65]. A preliminary report on slaughtered mountain zebras (Equus zebra hartmannae) showed the presence of 20-40-mm follicles during the proestrus phase and corpora lutea in pregnant females [Lilian et al., J Reprod Fertil 1988;84:505-11]. Ovum pick-up from follicles 25 mm performed in captive reared or slaughtered animals produced high quality oocytes from Plains zebras (Equus burchelli) and Mountain zebras [Meintjes et al., J Zoo Wildl Med 1997;28:251-59]. Embryo transfer from zebras to mares has produced viable, healthy foals as a method of genetic preservation and production of high quality individuals [Bennett et al., Equine Vet J 1985;17:78-9; Summers et al., J Reprod Fertil 1987;80:1320]. These assisted reproductive techniques have been carried out utilizing just the beginning and end of estrus dates for calculating the day of embryo flushing [Allen et al., J Heredity 1997;88:384-92]. The objective of this study was to characterize follicular dynamics during spontaneous (Experiment 1) and induced (Experiment 2) estrous cycles

Linking intraspecific variation in territory size, cone supply, and survival of North American red squirrels

Abstract In territorial species, competitive asymmetries can result in the uneven distribution of food resources as high-quality individuals force young or subordinates to occupy smaller or lower-quality sites, or both. However, spatiotemporal variation in the production of resources also can influence an individual's ability to monopolize resources and, consequently, affect survival. We examined how spatial and temporal variation in food supply affects the distribution of resources among territorial food-hoarding red squirrels (Tamiasciurus hudsonicus) by mapping cone production over 62.5 ha of white spruce (Picea glauca) forest and measuring the interrelationships between territory size, cone supply, and survival during 4 years of low cone abundance in Yukon, Canada. Territory size and cone production within a territory varied 10- and 520-fold, respectively, with juvenile squirrels occupying smaller territories with fewer cones. Because of small-scale heterogeneity in the distribution of cones, territor...

Apparent survival of Snowy Plovers (Charadrius nivosus) varies with reproductive effort and year and between sexes

Abstract. Life history theory predicts a tradeoff between reproductive effort and survival, which suggests that some management practices aimed at increasing productivity may compromise population growth. We analyzed a 10-year data set of 225 individually marked Snowy Plovers (Charadrius nivosus), a threatened shorebird, to determine whether individual reproductive effort was correlated with low apparent survival. Most adults resided in the population an average of 2 years (range: 1–10 years), during which females laid 3–60 eggs, and both males and females invested considerable time in incubation and brooding. Apparent survival varied annually and was higher for males than for females. Contrary to theory, we found no evidence that increased reproductive effort, either current or cumulative, compromised survival. Instead, apparent survival was correlated positively with incubation time, which may be related to either high-quality individuals having high reproductive rates and high survival or permanent emi...

Individual heterogeneity in black brant survival and recruitment with implications for harvest dynamics

We examined individual heterogeneity in survival and recruitment of female Pacific black brant (Branta bernicla nigricans) using frailty models adapted to a capture–mark–recapture context. Our main objectives were (1) to quantify levels of heterogeneity and examine factors affecting heterogeneity, and (2) model the effects of individual heterogeneity on harvest dynamics through matrix models. We used 24 years of data on brant marked and recaptured at the Tutakoke River colony, AK. Multievent models were fit as hidden Markov chain using program E-SURGE with an adequate overdispersion coefficient. Annual survival of individuals marked as goslings was heterogeneous among individuals and year specific with about 0.23 difference in survival between “high” (0.73)- and “low” (0.50)-quality individuals at average survival probability. Adult survival (0.85 ± 0.004) was homogeneous and higher than survival of both groups of juveniles. The annual recruitment probability was heterogeneous for brant >1-year-old; 0.56 (±0.21) and 0.31 (±0.03) for high- and low-quality individuals, respectively. Assuming equal clutch sizes for high- and low-quality individuals and that 80% of offspring were in the same quality class as the breeding female resulted in reproductive values about twice as high for high-quality individuals than low-quality individual for a given class of individuals producing differential contributions to population growth among groups. Differences in reproductive values greatly increased when we assumed high-quality individuals had larger clutch sizes. When we assumed that 50% of offspring were in the same quality class as their mothers and clutches were equal, differences in reproductive values between quality classes were greatly reduced or eliminated (breeders [BRs]). We considered several harvest scenarios using the assumption that 80% of offspring were in the same quality class as their mothers. The amount of compensation for harvest mortality declined as the proportion of high-quality individuals in the harvest increased, as differences in clutch sizes between groups decreased and as the proportion of BRs in the harvest increased. Synthesis and applications. Harvest at the same proportional level of the overall population can result in variable responses in population growth rate when heterogeneity is present in a population. λ was <1.0 under every scenario when harvest rates were >10%, and heterogeneity caused as much as +2% difference in growth rates at the highest levels of proportional harvest for low-quality individuals and the greatest differences in qualities between classes of individuals, a critical difference for a population with λ near 1.0 such as the brant. We observed less response in overall survival in the presence of heterogeneity because we did not observe heterogeneity in the annual survival of BRs. This analysis provides a comprehensive view of overall compensation at the population level and also constitutes the first example of a survival-recruitment model with heterogeneity. Individual heterogeneity should be more explicitly considered in harvest management of vertebrates.