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1.
Ecol Evol ; 14(5): e11356, 2024 May.
Artículo en Inglés | MEDLINE | ID: mdl-38694748

RESUMEN

The house sparrow (Passer domesticus) is a small passerine known to be highly sedentary. Throughout a 30-year capture-mark-recapture study, we have obtained occasional reports of recoveries far outside our main metapopulation study system, documenting unusually long dispersal distances. Our records constitute the highest occurrence of long-distance dispersal events recorded for this species in Scandinavia. Such long-distance dispersals radically change the predicted distribution of dispersal distances and connectedness for our study metapopulation. Moreover, it reveals a much greater potential for colonization than formerly recorded for the house sparrow, which is an invasive species across four continents. These rare and occasional long-distance dispersal events are challenging to document but may have important implications for the genetic composition of small and isolated populations and for our understanding of dispersal ecology and evolution.

2.
Evol Appl ; 17(4): e13684, 2024 Apr.
Artículo en Inglés | MEDLINE | ID: mdl-38617828

RESUMEN

Harvesting and culling are methods used to monitor and manage wildlife diseases. An important consequence of these practices is a change in the genetic dynamics of affected populations that may threaten their long-term viability. The effective population size (N e) is a fundamental parameter for describing such changes as it determines the amount of genetic drift in a population. Here, we estimate N e of a harvested wild reindeer population in Norway. Then we use simulations to investigate the genetic consequences of management efforts for handling a recent spread of chronic wasting disease, including increased adult male harvest and population decimation. The N e/N ratio in this population was found to be 0.124 at the end of the study period, compared to 0.239 in the preceding 14 years period. The difference was caused by increased harvest rates with a high proportion of adult males (older than 2.5 years) being shot (15.2% in 2005-2018 and 44.8% in 2021). Increased harvest rates decreased N e in the simulations, but less sex biased harvest strategies had a lower negative impact. For harvest strategies that yield stable population dynamics, shifting the harvest from calves to adult males and females increased N e. Population decimation always resulted in decreased genetic variation in the population, with higher loss of heterozygosity and rare alleles with more severe decimation or longer periods of low population size. A very high proportion of males in the harvest had the most severe consequences for the loss of genetic variation. This study clearly shows how the effects of harvest strategies and changes in population size interact to determine the genetic drift of a managed population. The long-term genetic viability of wildlife populations subject to a disease will also depend on population impacts of the disease and how these interact with management actions.

3.
Nature ; 626(8000): 725-726, 2024 Feb.
Artículo en Inglés | MEDLINE | ID: mdl-38321158
4.
J Anim Ecol ; 93(1): 8-20, 2024 01.
Artículo en Inglés | MEDLINE | ID: mdl-37740526

RESUMEN

We propose that the ecological resilience of communities to permanent changes of the environment can be based on how variation in the overall abundance of individuals affects the number of species. Community sensitivity is defined as the ratio between the rate of change in the log expected number of species and the rate of change in the log expected number of individuals in the community. High community sensitivity means that small changes in the total abundance strongly impact the number of species. Community resistance is the proportional reduction in expected number of individuals that the community can sustain before expecting to lose one species. A small value of community resistance means that the community can only endure a small reduction in abundance before it is expected to lose one species. Based on long-term studies of four bird communities in European deciduous forests at different latitudes large differences were found in the resilience to environmental perturbations. Estimating the variance components of the species abundance distribution revealed how different processes contributed to the community sensitivity and resistance. Species heterogeneity in the population dynamics was the largest component, but its proportion varied among communities. Species-specific response to environmental fluctuations was the second major component of the variation in abundance. Estimates of community sensitivity and resistance based on data only from a single year were in general larger than those based on estimates from longer time series. Thus, our approach can provide rapid and conservative assessment of the resilience of communities to environmental changes also including only short-term data. This study shows that a general ecological mechanism, caused by increased strength of density dependence due to reduction in resource availability, can provide an intuitive measure of community resilience to environmental variation. Our analyses also illustrate the importance of including specific assumptions about how different processes affect community dynamics. For example, if stochastic fluctuations in the environment affect all species in a similar way, the sensitivity and resistance of the community to environmental changes will be different from communities in which all species show independent responses.


Asunto(s)
Bosques , Modelos Biológicos , Humanos , Animales , Dinámica Poblacional , Factores de Tiempo
6.
J Anim Ecol ; 92(11): 2214-2227, 2023 11.
Artículo en Inglés | MEDLINE | ID: mdl-37750026

RESUMEN

Disentangling empirically the many processes affecting spatial population synchrony is a challenge in population ecology. Two processes that could have major effects on the spatial synchrony of wild population dynamics are density dependence and variation in environmental conditions like temperature. Understanding these effects is crucial for predicting the effects of climate change on local and regional population dynamics. We quantified the direct contribution of local temperature and density dependence to spatial synchrony in the population dynamics of nine fish species inhabiting the Barents Sea. First, we estimated the degree to which the annual spatial autocorrelations in density are influenced by temperature. Second, we estimated and mapped the local effects of temperature and strength of density dependence on annual changes in density. Finally, we measured the relative effects of temperature and density dependence on the spatial synchrony in changes in density. Temperature influenced the annual spatial autocorrelation in density more in species with greater affinities to the benthos and to warmer waters. Temperature correlated positively with changes in density in the eastern Barents Sea for most species. Temperature had a weak synchronizing effect on density dynamics, while increasing strength of density dependence consistently desynchronised the dynamics. Quantifying the relative effects of different processes affecting population synchrony is important to better predict how population dynamics might change when environmental conditions change. Here, high degrees of spatial synchrony in the population dynamics remained unexplained by local temperature and density dependence, confirming the presence of additional synchronizing drivers, such as trophic interactions or harvesting.


Asunto(s)
Ecología , Ecosistema , Animales , Temperatura , Dinámica Poblacional
7.
Sci Rep ; 13(1): 15181, 2023 09 13.
Artículo en Inglés | MEDLINE | ID: mdl-37704641

RESUMEN

Demographic consequences of rapid environmental change and extreme climatic events (ECEs) can cascade across trophic levels with evolutionary implications that have rarely been explored. Here, we show how an ECE in high Arctic Svalbard triggered a trophic chain reaction, directly or indirectly affecting the demography of both overwintering and migratory vertebrates, ultimately inducing a shift in density-dependent phenotypic selection in migratory geese. A record-breaking rain-on-snow event and ice-locked pastures led to reindeer mass starvation and a population crash, followed by a period of low mortality and population recovery. This caused lagged, long-lasting reductions in reindeer carrion numbers and resultant low abundances of Arctic foxes, a scavenger on reindeer and predator of migratory birds. The associated decrease in Arctic fox predation of goose offspring allowed for a rapid increase in barnacle goose densities. As expected according to r- and K-selection theory, the goose body condition (affecting reproduction and post-fledging survival) maximising Malthusian fitness increased with this shift in population density. Thus, the winter ECE acting on reindeer and their scavenger, the Arctic fox, indirectly selected for higher body condition in migratory geese. This high Arctic study provides rare empirical evidence of links between ECEs, community dynamics and evolution, with implications for our understanding of indirect eco-evolutionary impacts of global change.


Asunto(s)
Zorros , Reno , Animales , Patos , Gansos , Carne
8.
Proc Biol Sci ; 290(2002): 20230511, 2023 07 12.
Artículo en Inglés | MEDLINE | ID: mdl-37403509

RESUMEN

The slow-fast continuum is a commonly used framework to describe variation in life-history strategies across species. Individual life histories have also been assumed to follow a similar pattern, especially in the pace-of-life syndrome literature. However, whether a slow-fast continuum commonly explains life-history variation among individuals within a population remains unclear. Here, we formally tested for the presence of a slow-fast continuum of life histories both within populations and across species using detailed long-term individual-based demographic data for 17 bird and mammal species with markedly different life histories. We estimated adult lifespan, age at first reproduction, annual breeding frequency, and annual fecundity, and identified the main axes of life-history variation using principal component analyses. Across species, we retrieved the slow-fast continuum as the main axis of life-history variation. However, within populations, the patterns of individual life-history variation did not align with a slow-fast continuum in any species. Thus, a continuum ranking individuals from slow to fast living is unlikely to shape individual differences in life histories within populations. Rather, individual life-history variation is likely idiosyncratic across species, potentially because of processes such as stochasticity, density dependence, and individual differences in resource acquisition that affect species differently and generate non-generalizable patterns across species.


Asunto(s)
Rasgos de la Historia de Vida , Reproducción , Humanos , Animales , Mamíferos , Aves
9.
Sci Rep ; 13(1): 4272, 2023 03 15.
Artículo en Inglés | MEDLINE | ID: mdl-36922555

RESUMEN

Telomeres, the nucleotide sequences that protect the ends of eukaryotic chromosomes, shorten with each cell division and telomere loss may be influenced by environmental factors. Telomere length (TL) decreases with age in several species, but little is known about the sources of genetic and environmental variation in the change in TL (∆TL) in wild animals. In this study, we tracked changes in TL throughout the natural lifespan (from a few months to almost 9 years) of free-living house sparrows (Passer domesticus) in two different island populations. TL was measured in nestlings and subsequently up to four times during their lifetime. TL generally decreased with age (senescence), but we also observed instances of telomere lengthening within individuals. We found some evidence for selective disappearance of individuals with shorter telomeres through life. Early-life TL positively predicted later-life TL, but the within-individual repeatability in TL was low (9.2%). Using genetic pedigrees, we found a moderate heritability of ∆TL (h2 = 0.21), which was higher than the heritabilities of early-life TL (h2 = 0.14) and later-life TL measurements (h2 = 0.15). Cohort effects explained considerable proportions of variation in early-life TL (60%), later-life TL (53%), and ∆TL (37%), which suggests persistent impacts of the early-life environment on lifelong telomere dynamics. Individual changes in TL were independent of early-life TL. Finally, there was weak evidence for population differences in ∆TL that may be linked to ecological differences in habitat types. Combined, our results show that individual telomere biology is highly dynamic and influenced by both genetic and environmental variation in natural conditions.


Asunto(s)
Animales Salvajes , Longevidad , Animales , Animales Salvajes/genética , Longevidad/genética , Aves/genética , Homeostasis del Telómero , Acortamiento del Telómero/genética , Telómero/genética
10.
J Evol Biol ; 36(4): 650-662, 2023 04.
Artículo en Inglés | MEDLINE | ID: mdl-36811205

RESUMEN

An organism's energy budget is strongly related to resource consumption, performance, and fitness. Hence, understanding the evolution of key energetic traits, such as basal metabolic rate (BMR), in natural populations is central for understanding life-history evolution and ecological processes. Here we used quantitative genetic analyses to study evolutionary potential of BMR in two insular populations of the house sparrow (Passer domesticus). We obtained measurements of BMR and body mass (Mb ) from 911 house sparrows on the islands of Leka and Vega along the coast of Norway. These two populations were the source populations for translocations to create an additional third, admixed 'common garden' population in 2012. With the use of a novel genetic group animal model concomitant with a genetically determined pedigree, we differentiate genetic and environmental sources of variation, thereby providing insight into the effects of spatial population structure on evolutionary potential. We found that the evolutionary potential of BMR was similar in the two source populations, whereas the Vega population had a somewhat higher evolutionary potential of Mb than the Leka population. BMR was genetically correlated with Mb in both populations, and the conditional evolutionary potential of BMR (independent of body mass) was 41% (Leka) and 53% (Vega) lower than unconditional estimates. Overall, our results show that there is potential for BMR to evolve independently of Mb , but that selection on BMR and/or Mb may have different evolutionary consequences in different populations of the same species.


Asunto(s)
Animales Salvajes , Metabolismo Basal , Animales , Aves , Fenotipo
11.
Ecology ; 104(2): e3908, 2023 02.
Artículo en Inglés | MEDLINE | ID: mdl-36314902

RESUMEN

Identifying the environmental drivers of variation in fitness-related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness-related trait values (i.e., correlated temporal trait fluctuations across populations) is poorly understood. Using data from long-term monitored populations of blue tits (Cyanistes caeruleus, n = 31), great tits (Parus major, n = 35), and pied flycatchers (Ficedula hypoleuca, n = 20) across Europe, we assessed the influence of two local climatic variables (mean temperature and mean precipitation in February-May) on spatial synchrony in three fitness-related traits: laying date, clutch size, and fledgling number. We found a high degree of spatial synchrony in laying date but a lower degree in clutch size and fledgling number for each species. Temperature strongly influenced spatial synchrony in laying date for resident blue tits and great tits but not for migratory pied flycatchers. This is a relevant finding in the context of environmental impacts on populations because spatial synchrony in fitness-related trait values among populations may influence fluctuations in vital rates or population abundances. If environmentally induced spatial synchrony in fitness-related traits increases the spatial synchrony in vital rates or population abundances, this will ultimately increase the risk of extinction for populations and species. Assessing how environmental conditions influence spatiotemporal variation in trait values improves our mechanistic understanding of environmental impacts on populations.


Asunto(s)
Passeriformes , Pájaros Cantores , Animales , Temperatura , Estaciones del Año , Reproducción
12.
Ecol Evol ; 12(8): e9144, 2022 Aug.
Artículo en Inglés | MEDLINE | ID: mdl-35923948

RESUMEN

Environmental conditions during early-life development can have lasting effects shaping individual heterogeneity in fitness and fitness-related traits. The length of telomeres, the DNA sequences protecting chromosome ends, may be affected by early-life conditions, and telomere length (TL) has been associated with individual performance within some wild animal populations. Thus, knowledge of the mechanisms that generate variation in TL, and the relationship between TL and fitness, is important in understanding the role of telomeres in ecology and life-history evolution. Here, we investigate how environmental conditions and morphological traits are associated with early-life blood TL and if TL predicts natal dispersal probability or components of fitness in 2746 wild house sparrow (Passer domesticus) nestlings from two populations sampled across 20 years (1994-2013). We retrieved weather data and we monitored population fluctuations, individual survival, and reproductive output using field observations and genetic pedigrees. We found a negative effect of population density on TL, but only in one of the populations. There was a curvilinear association between TL and the maximum daily North Atlantic Oscillation index during incubation, suggesting that there are optimal weather conditions that result in the longest TL. Dispersers tended to have shorter telomeres than non-dispersers. TL did not predict survival, but we found a tendency for individuals with short telomeres to have higher annual reproductive success. Our study showed how early-life TL is shaped by effects of growth, weather conditions, and population density, supporting that environmental stressors negatively affect TL in wild populations. In addition, shorter telomeres may be associated with a faster pace-of-life, as individuals with higher dispersal rates and annual reproduction tended to have shorter early-life TL.

13.
Proc Biol Sci ; 289(1976): 20220296, 2022 06 08.
Artículo en Inglés | MEDLINE | ID: mdl-35642371

RESUMEN

The capacity of natural selection to generate adaptive changes is (according to the fundamental theorem of natural selection) proportional to the additive genetic variance in fitness. In spite of its importance for development of new adaptations to a changing environment, processes affecting the magnitude of the genetic variance in fitness-related traits are poorly understood. Here, we show that the red-white colour polymorphism in female barn owls is subject to density-dependent selection at the phenotypic and genotypic level. The diallelic melanocortin-1 receptor gene explained a large amount of the phenotypic variance in reddish coloration in the females ([Formula: see text]). Red individuals (RR genotype) were selected for at low densities, while white individuals (WW genotype) were favoured at high densities and were less sensitive to changes in density. We show that this density-dependent selection favours white individuals and predicts fixation of the white allele in this population at longer time scales without immigration or other selective forces. Still, fluctuating population density will cause selection to fluctuate and periodically favour red individuals. These results suggest how balancing selection caused by fluctuations in population density can be a general mechanism affecting the level of additive genetic variance in natural populations.


Asunto(s)
Estrigiformes , Animales , Color , Femenino , Genotipo , Polimorfismo Genético , Selección Genética , Estrigiformes/genética
14.
Ecol Lett ; 25(7): 1640-1654, 2022 Jul.
Artículo en Inglés | MEDLINE | ID: mdl-35610546

RESUMEN

Temporal correlations among demographic parameters can strongly influence population dynamics. Our empirical knowledge, however, is very limited regarding the direction and the magnitude of these correlations and how they vary among demographic parameters and species' life histories. Here, we use long-term demographic data from 15 bird and mammal species with contrasting pace of life to quantify correlation patterns among five key demographic parameters: juvenile and adult survival, reproductive probability, reproductive success and productivity. Correlations among demographic parameters were ubiquitous, more frequently positive than negative, but strongly differed across species. Correlations did not markedly change along the slow-fast continuum of life histories, suggesting that they were more strongly driven by ecological than evolutionary factors. As positive temporal demographic correlations decrease the mean of the long-run population growth rate, the common practice of ignoring temporal correlations in population models could lead to the underestimation of extinction risks in most species.


Asunto(s)
Crecimiento Demográfico , Reproducción , Animales , Evolución Biológica , Aves , Mamíferos , Dinámica Poblacional
15.
Ecology ; 103(8): e3731, 2022 08.
Artículo en Inglés | MEDLINE | ID: mdl-35416286

RESUMEN

Our knowledge of the factors affecting species abundances is mainly based on time-series analyses of a few well-studied species at single or few localities, but we know little about whether results from such analyses can be extrapolated to the community level. We apply a joint species distribution model to long-term time-series data on British bird communities to examine the relative contribution of intra- and interspecific density dependence at different spatial scales, as well as the influence of environmental stochasticity, to spatiotemporal interspecific variation in abundance. Intraspecific density dependence has the major structuring effect on these bird communities. In addition, environmental fluctuations affect spatiotemporal differences in abundance. In contrast, species interactions had a minor impact on variation in abundance. Thus, important drivers of single-species dynamics are also strongly affecting dynamics of communities in time and space.


Asunto(s)
Aves , Ecosistema , Animales , Dinámica Poblacional
16.
Ecol Lett ; 25(4): 863-875, 2022 Apr.
Artículo en Inglés | MEDLINE | ID: mdl-35103374

RESUMEN

Harvesting can magnify the destabilising effects of environmental perturbations on population dynamics and, thereby, increase extinction risk. However, population-dynamic theory predicts that impacts of harvesting depend on the type and strength of density-dependent regulation. Here, we used logistic population growth models and an empirical reindeer case study to show that low to moderate harvesting can actually buffer populations against environmental perturbations. This occurs because of density-dependent environmental stochasticity, where negative environmental impacts on vital rates are amplified at high population density due to intra-specific resource competition. Simulations from our population models show that even low levels of harvesting may prevent overabundance, thereby dampening population fluctuations and reducing the risk of population collapse and quasi-extinction following environmental perturbations. Thus, depending on the species' life history and the strength of density-dependent environmental drivers, low to moderate harvesting can improve population resistance to increased climate variability and extreme weather expected under global warming.


Asunto(s)
Dinámica Poblacional , Modelos Logísticos , Densidad de Población
17.
Mol Ecol ; 31(23): 6224-6238, 2022 12.
Artículo en Inglés | MEDLINE | ID: mdl-34997994

RESUMEN

Telomere dynamics could underlie life-history trade-offs among growth, size and longevity, but our ability to quantify such processes in natural, unmanipulated populations is limited. We investigated how 4 years of artificial selection for either larger or smaller tarsus length, a proxy for body size, affected early-life telomere length (TL) and several components of fitness in two insular populations of wild house sparrows over a study period of 11 years. The artificial selection was expected to shift the populations away from their optimal body size and increase the phenotypic variance in body size. Artificial selection for larger individuals caused TL to decrease, but there was little evidence that TL increased when selecting for smaller individuals. There was a negative correlation between nestling TL and tarsus length under both selection regimes. Males had longer telomeres than females and there was a negative effect of harsh weather on TL. We then investigated whether changes in TL might underpin fitness effects due to the deviation from the optimal body size. Mortality analyses indicated disruptive selection on TL because both short and long early-life telomeres tended to be associated with the lowest mortality rates. In addition, there was a tendency for a negative association between TL and annual reproductive success, but only in the population where body size was increased experimentally. Our results suggest that natural selection for optimal body size in the wild may be associated with changes in TL during growth, which is known to be linked to longevity in some bird species.


Asunto(s)
Longevidad , Passeriformes , Humanos , Masculino , Femenino , Animales , Longevidad/genética , Selección Genética , Telómero , Passeriformes/genética , Acortamiento del Telómero/genética
18.
Mol Ecol ; 31(23): 6360-6381, 2022 12.
Artículo en Inglés | MEDLINE | ID: mdl-34825754

RESUMEN

Early-life telomere length (TL) is associated with fitness in a range of organisms. Little is known about the genetic basis of variation in TL in wild animal populations, but to understand the evolutionary and ecological significance of TL it is important to quantify the relative importance of genetic and environmental variation in TL. In this study, we measured TL in 2746 house sparrow nestlings sampled across 20 years and used an animal model to show that there is a small heritable component of early-life TL (h2  = 0.04). Variation in TL among individuals was mainly driven by environmental (annual) variance, but also brood and parental effects. Parent-offspring regressions showed a large maternal inheritance component in TL ( h maternal 2  = 0.44), but no paternal inheritance. We did not find evidence for a negative genetic correlation underlying the observed negative phenotypic correlation between TL and structural body size. Thus, TL may evolve independently of body size and the negative phenotypic correlation is likely to be caused by nongenetic environmental effects. We further used genome-wide association analysis to identify genomic regions associated with TL variation. We identified several putative genes underlying TL variation; these have been inferred to be involved in oxidative stress, cellular growth, skeletal development, cell differentiation and tumorigenesis in other species. Together, our results show that TL has a low heritability and is a polygenic trait strongly affected by environmental conditions in a free-living bird.


Asunto(s)
Estudio de Asociación del Genoma Completo , Passeriformes , Animales , Longevidad/genética , Telómero/genética , Passeriformes/genética
19.
Glob Chang Biol ; 28(7): 2236-2258, 2022 Apr.
Artículo en Inglés | MEDLINE | ID: mdl-34931401

RESUMEN

Climate impacts are not always easily discerned in wild populations as detecting climate change signals in populations is challenged by stochastic noise associated with natural climate variability, variability in biotic and abiotic processes, and observation error in demographic rates. Detection of the impact of climate change on populations requires making a formal distinction between signals in the population associated with long-term climate trends from those generated by stochastic noise. The time of emergence (ToE) identifies when the signal of anthropogenic climate change can be quantitatively distinguished from natural climate variability. This concept has been applied extensively in the climate sciences, but has not been explored in the context of population dynamics. Here, we outline an approach to detecting climate-driven signals in populations based on an assessment of when climate change drives population dynamics beyond the envelope characteristic of stochastic variations in an unperturbed state. Specifically, we present a theoretical assessment of the time of emergence of climate-driven signals in population dynamics ( ToE pop ). We identify the dependence of ToE pop on the magnitude of both trends and variability in climate and also explore the effect of intrinsic demographic controls on ToE pop . We demonstrate that different life histories (fast species vs. slow species), demographic processes (survival, reproduction), and the relationships between climate and demographic rates yield population dynamics that filter climate trends and variability differently. We illustrate empirically how to detect the point in time when anthropogenic signals in populations emerge from stochastic noise for a species threatened by climate change: the emperor penguin. Finally, we propose six testable hypotheses and a road map for future research.


Asunto(s)
Cambio Climático , Spheniscidae , Animales , Dinámica Poblacional , Reproducción
20.
Ecol Evol ; 11(21): 14789-14804, 2021 Nov.
Artículo en Inglés | MEDLINE | ID: mdl-34765141

RESUMEN

Despite their importance in shaping life history tactics and population dynamics, individual growth trajectories have only been rarely explored in the wild because their analysis requires multiple measurements of individuals throughout their lifetime and some knowledge of age, a key timer of body growth. The availability of long-term longitudinal studies of two wild boar populations subjected to contrasting environments (rich vs. poor) provided an opportunity to analyze individual growth trajectories. We quantified wild boar growth trajectories at both the population and the individual levels using standard growth models (i.e., Gompertz, logistic, and monomolecular models) that encompass the expected range of growth shapes in determinate growers. Wild boar is a rather altricial species, with a polygynous mating system and is strongly sexually dimorphic in size. According to current theories of life history evolution, we thus expect wild boar to display a sex-specific Gompertz type growth trajectory and lower sexual size dimorphism in the poorer environment. While wild boar displayed the expected Gompertz type trajectory in the rich site at the population level, we found some evidence for potential differences in growth shapes between populations and individuals. Asymptotic body mass, growth rate and timing of maximum growth rate differed as well, which indicates a high flexibility of growth in wild boar. We also found a cohort effect on asymptotic body mass, which suggests that environmental conditions early in life shape body mass at adulthood in this species. Our findings demonstrate that body growth trajectories in wild boar are highly diverse in relation to differences of environmental context, sex and year of birth. Whether the intermediate ranking of wild boar along the precocial-altricial continuum of development at birth may explain the ability of this species to exhibit this high diversity of growth patterns remains to be investigated.

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