Life histories are not just fast or slow.

Life history strategies, which combine schedules of survival, development, and reproduction, shape how natural selection acts on species' heritable traits and organismal fitness. Comparative analyses have historically ranked life histories along a fast-slow continuum, describing a negative association between time allocation to reproduction and development versus survival. However, higher-quality, more representative data and analyses have revealed that life history variation cannot be fully accounted for by this single continuum. Moreover, studies often do not test predictions from existing theories and instead operate as exploratory exercises. To move forward, we offer three recommendations for future investigations: standardizing life history traits, overcoming taxonomic siloes, and using theory to move from describing to understanding life history variation across the Tree of Life.

High-resolution data are necessary to understand the effects of climate on plant population dynamics of a forest herb.

Climate is assumed to strongly influence species distribution and abundance. Although the performance of many organisms is influenced by the climate in their immediate proximity, the climate data used to model their distributions often have a coarse spatial resolution. This is problematic because the local climate experienced by individuals might deviate substantially from the regional average. This problem is likely to be particularly important for sessile organisms like plants and in environments where small-scale variation in climate is large. To quantify the effect of local temperature on vital rates and population growth rates, we used temperature values measured at the local scale (in situ logger measures) and integral projection models with demographic data from 37 populations of the forest herb Lathyrus vernus across a wide latitudinal gradient in Sweden. To assess how the spatial resolution of temperature data influences assessments of climate effects, we compared effects from models using local data with models using regionally aggregated temperature data at several spatial resolutions (≥1 km). Using local temperature data, we found that spring frost reduced the asymptotic population growth rate in the first of two annual transitions and influenced survival in both transitions. Only one of the four regional estimates showed a similar negative effect of spring frost on population growth rate. Our results for a perennial forest herb show that analyses using regionally aggregated data often fail to identify the effects of climate on population dynamics. This emphasizes the importance of using organism-relevant estimates of climate when examining effects on individual performance and population dynamics, as well as when modeling species distributions. For sessile organisms that experience the environment over small spatial scales, this will require climate data at high spatial resolutions.

Anyone Can Get Old-All You Have to Do Is Live Long Enough: Understanding Mortality and Life Expectancy in European Hedgehogs (Erinaceus europaeus).

The European hedgehog is in decline, triggering a need to monitor population dynamics to optimise conservation initiatives directed at this species. By counting periosteal growth lines, we determined the age of 388 dead European hedgehogs collected through citizen science in Denmark. The overall mean age was 1.8 years (1.6 years for females and 2.1 years for males), ranging between 0 and 16 years. We constructed life tables showing life expectancies at 2.1 years for females and 2.6 years for males. We discovered that male hedgehogs were more likely to have died in traffic than females, but traffic-related deaths peaked in July for both sexes. A sex difference was detected for non-traffic deaths, as most males died in July, and most females died in September. We created empirical survivorship curves and hazard curves showing that the risk of death for male hedgehogs remains approximately constant with age. In contrast, the risk of death for females increases with age. Most of the collected road-killed individuals died in rural habitats. The degree of inbreeding did not influence longevity. These new insights are important for preparing conservation strategies for the European hedgehog.

Climate causes shifts in grey seal phenology by modifying age structure.

There are numerous examples of phenological shifts that are recognized both as indicators of climate change and drivers of ecosystem change. A pressing challenge is to understand the causal mechanisms by which climate affects phenology. We combined annual population census data and individual longitudinal data (1992-2018) on grey seals, Halicheorus grypus, to quantify the relationship between pupping season phenology and sea surface temperature. A temperature increase of 2°C was associated with a pupping season advance of approximately seven days at the population level. However, we found that maternal age, rather than sea temperature, accounted for changes in pupping date by individuals. Warmer years were associated with an older average age of mothers, allowing us to explain phenological observations in terms of a changing population age structure. Finally, we developed a matrix population model to test whether our observations were consistent with changes to the stable age distribution. This could not fully account for observed phenological shift, strongly suggesting transient modification of population age structure, for example owing to immigration. We demonstrate a novel mechanism for phenological shifts under climate change in long-lived, age- or stage-structured species with broad implications for dynamics and resilience, as well as population management.

The effect of nest temperature on growth and survival in juvenile Great Tits Parus major.

For birds, maintaining an optimal nest temperature is critical for early-life growth and development. Temperatures deviating from this optimum can affect nestling growth and fledging success with potential consequences on survival and lifetime reproductive success. It is therefore particularly important to understand these effects in relation to projected temperature changes associated with climate change.Targets set by the 2015 Paris Agreement aim to limit temperature increases to 2°C, and, with this in mind, we carried out an experiment in 2017 and 2018 where we applied a treatment that increased Great Tit Parus major nest temperature by approximately this magnitude (achieving an increase of 1.6°C, relative to the control) during the period from hatching to fledging to estimate how small temperature differences might affect nestling body size and weight at fledging and fledging success.We recorded hatching and fledging success and measured skeletal size (tarsus length) and body mass at days 5, 7, 10, and 15 posthatch in nestlings from two groups of nest boxes: control and heated (+1.6°C).Our results show that nestlings in heated nest boxes were 1.6% smaller in skeletal size at fledging than those in the cooler control nests, indicating lower growth rates in heated boxes, and that their weight was, in addition, 3.3% lower.These results suggest that even fairly small changes in temperature can influence phenotype and postfledging survival in cavity-nesting birds. This has the potential to affect the population dynamics of these birds in the face of ongoing climatic change, as individuals of reduced size in colder winters may suffer from decreased fitness.

The myriad of complex demographic responses of terrestrial mammals to climate change and gaps of knowledge: A global analysis.

Approximately 25% of mammals are currently threatened with extinction, a risk that is amplified under climate change. Species persistence under climate change is determined by the combined effects of climatic factors on multiple demographic rates (survival, development and reproduction), and hence, population dynamics. Thus, to quantify which species and regions on Earth are most vulnerable to climate-driven extinction, a global understanding of how different demographic rates respond to climate is urgently needed. Here, we perform a systematic review of literature on demographic responses to climate, focusing on terrestrial mammals, for which extensive demographic data are available. To assess the full spectrum of responses, we synthesize information from studies that quantitatively link climate to multiple demographic rates. We find only 106 such studies, corresponding to 87 mammal species. These 87 species constitute <1% of all terrestrial mammals. Our synthesis reveals a strong mismatch between the locations of demographic studies and the regions and taxa currently recognized as most vulnerable to climate change. Surprisingly, for most mammals and regions sensitive to climate change, holistic demographic responses to climate remain unknown. At the same time, we reveal that filling this knowledge gap is critical as the effects of climate change will operate via complex demographic mechanisms: a vast majority of mammal populations display projected increases in some demographic rates but declines in others, often depending on the specific environmental context, complicating simple projections of population fates. Assessments of population viability under climate change are in critical need to gather data that account for multiple demographic responses, and coordinated actions to assess demography holistically should be prioritized for mammals and other taxa.

Herbaceous perennial plants with short generation time have stronger responses to climate anomalies than those with longer generation time.

There is an urgent need to synthesize the state of our knowledge on plant responses to climate. The availability of open-access data provide opportunities to examine quantitative generalizations regarding which biomes and species are most responsive to climate drivers. Here, we synthesize time series of structured population models from 162 populations of 62 plants, mostly herbaceous species from temperate biomes, to link plant population growth rates (λ) to precipitation and temperature drivers. We expect: (1) more pronounced demographic responses to precipitation than temperature, especially in arid biomes; and (2) a higher climate sensitivity in short-lived rather than long-lived species. We find that precipitation anomalies have a nearly three-fold larger effect on λ than temperature. Species with shorter generation time have much stronger absolute responses to climate anomalies. We conclude that key species-level traits can predict plant population responses to climate, and discuss the relevance of this generalization for conservation planning.

Bridging gaps in demographic analysis with phylogenetic imputation.

Phylogenetically informed imputation methods have rarely been applied to estimate missing values in demographic data but may be a powerful tool for reconstructing vital rates of survival, maturation, and fecundity for species of conservation concern. Imputed vital rates could be used to parameterize demographic models to explore how populations respond when vital rates are perturbed. We used standardized vital rate estimates for 50 bird species to assess the use of phylogenetic imputation to fill gaps in demographic data. We calculated imputation accuracy for vital rates of focal species excluded from the data set either singly or in combination and with and without phylogeny, body mass, and life-history trait data. We used imputed vital rates to calculate demographic metrics, including generation time, to validate the use of imputation in demographic analyses. Covariance among vital rates and other trait data provided a strong basis to guide imputation of missing vital rates in birds, even in the absence of phylogenetic information. Mean NRMSE for null and phylogenetic models differed by <0.01 except when no vital rates were available or for vital rates with high phylogenetic signal (Pagel's λ > 0.8). In these cases, including body mass and life-history trait data compensated for lack of phylogenetic information: mean normalized root mean square error (NRMSE) for null and phylogenetic models differed by <0.01 for adult survival and <0.04 for maturation rate. Estimates of demographic metrics were sensitive to the accuracy of imputed vital rates. For example, mean error in generation time doubled in response to inaccurate estimates of maturation time. Accurate demographic data and metrics, such as generation time, are needed to inform conservation planning processes, for example through International Union for Conservation of Nature Red List assessments and population viability analysis. Imputed vital rates could be useful in this context but, as for any estimated model parameters, awareness of the sensitivities of demographic model outputs to the imputed vital rates is essential.

Cerrando Brechas en los Análisis Demográficos con Imputación Filogenética Resumen Los métodos de imputación guiados filogenéticamente se han aplicado con poca frecuencia para estimar los valores faltantes en los datos demográficos, aunque pueden ser una herramienta poderosa para la reconstrucción de tasas vitales de supervivencia, maduración y fecundidad de especies de importancia para la conservación. Las tasas vitales imputadas podrían usarse para generar parámetros en los modelos demográficos para explorar cómo responden las poblaciones cuando se perturban las tasas vitales. Utilizamos estimaciones de tasas vitales estandarizadas para 50 especies de aves para analizar el uso de la imputación filogenética para llenar los vacíos en los datos demográficos. Calculamos la certeza de imputación para las tasas vitales de las especies focales excluidas del conjunto de datos por sí solas o en combinación y con y sin datos de filogenia, masa corporal y características de historia de vida. Usamos las tasas vitales imputadas para calcular las medidas demográficas, incluyendo el tiempo de generación, y así validar el uso de la imputación en los análisis demográficos. La covarianza entre las tasas vitales y otros datos de características proporcionó una base sólida para orientar la imputación de tasas vitales faltantes en las aves, incluso la ausencia de información filogenética. El NRMSE medio para los modelos nulo y filogenético difirió por <0.01 salvo cuando no hubo tasas vitales disponibles o para tasas vitales con una señal filogenética alta (λ de Pagel > 0.8). En estos casos, la inclusión de la masa corporal y las características de historia de vida compensó la falta de información filogenética: el error cuadrático medio de la raíz normalizada media (NRMSE) para los modelos nulo y filogenéticos difirió por <0.01 para la supervivencia adulta y <0.04 para la tasa de maduración. Las estimaciones de las medidas demográficas fueron sensibles a la certeza de las tasas vitales imputadas. Por ejemplo, el error medio en el tiempo generacional se duplicó en respuesta a las estimaciones imprecisas del tiempo de maduración. Las medidas y datos demográficos certeros, como el tiempo generacional, son necesarios para orientar los procesos de planeación de la conservación; por ejemplo, a través de las valoraciones de la Lista Roja de la Unión Internacional para la Conservación de la Naturaleza y los análisis de viabilidad poblacional. Las tasas vitales imputadas podrían ser útiles en este contexto, pero como para cualquier tipo de parámetro de modelo estimado, el conocimiento de las sensibilidades del rendimiento del modelo demográfico es esencial para las tasas vitales imputadas.

Census data aggregation decisions can affect population-level inference in heterogeneous populations.

Conservation and population management decisions often rely on population models parameterized using census data. However, the sampling regime, precision, sample size, and methods used to collect census data are usually heterogeneous in time and space. Decisions about how to derive population-wide estimates from this patchwork of data are complicated and may bias estimated population dynamics, with important implications for subsequent management decisions.Here, we explore the impact of site selection and data aggregation decisions on pup survival estimates, and downstream estimates derived from parameterized matrix population models (MPMs), using a long-term dataset on grey seal (Halichoerus grypus) pup survival from southwestern Wales. The spatiotemporal and methodological heterogeneity of the data are fairly typical for ecological census data and it is, therefore, a good model to address this topic.Data were collected from 46 sampling locations (sites) over 25 years, and we explore the impact of data handling decisions by varying how years and sampling locations are combined to parameterize pup survival in population-level MPMs. We focus on pup survival because abundant high-quality data are available on this developmental stage.We found that survival probability was highly variable with most variation being at the site level, and poorly correlated among sampling sites. This variation could generate marked differences in predicted population dynamics depending on sampling strategy. The sample size required for a confident survival estimate also varied markedly geographically.We conclude that for populations with highly variable vital rates among sub-populations, site selection and data aggregation methods are important. In particular, including peripheral or less frequently used areas can introduce substantial variation into population estimates. This is likely to be context-dependent, but these choices, including the use of appropriate weights when summarizing across sampling areas, should be explored to ensure that management actions are successful.

Genetic structure of the European hedgehog (Erinaceus europaeus) in Denmark.

OBJECTIVES: Low genetic diversity can lead to reduced average fitness in a population or even extinction. Preserving genetic connectivity across fragmented landscapes is therefore vital to counteract the negative consequences of genetic drift and inbreeding. This study aimed to assess the genetic composition and consequently the conservation status of a nationwide sample of European hedgehogs (Erinaceus europaeus) in Denmark. METHODS: We applied an adaptation of the genotyping by sequencing (GBS) technique to 178 individuals from six geographically distinct populations. We used a Bayesian clustering method to subdivide individuals into genetically distinct populations. We estimated individual observed (iHO), observed (HO), and unbiased expected (uHE) heterozygosity, inbreeding coefficient (FIS), percentage of polymorphic loci (P%) and tested for deviations from Hardy-Weinberg equilibrium (HWE). We used linear models to test for potential anthropogenic effects on the genetic variability of hedgehogs with iHO, uHE, P% and FIS as response variables, and assessed the demographic history of the population. RESULTS: The Danish hedgehog population is composed of three genetic clusters. We found a mean P% of 54.44-94.71, a mean uHE of 0.126-0.318 and a mean HO of 0.124-0.293 in the six populations. The FIS was found to be significantly positive for three of the six populations. We detected a large heterogeneity of iHO values within populations, which can be due to inbreeding and/or fragmentation. FIS values decreased with increasing farmland density, but there was no significant association with human population or road density. CONCLUSIONS: We found a low level of genetic variability and evidence for genetic substructure and low effective population size, which are all consequences of habitat fragmentation. We failed to detect signs of a recent population bottleneck or population increase or decline. However, because the test only identifies recent changes in population size, we cannot reject the possibility of a longer-term decline in the Danish hedgehog population.

The ecology of suburban juvenile European hedgehogs (Erinaceus europaeus) in Denmark.

European hedgehog (Erinaceus europaeus) populations are widespread across diverse habitats but are declining in Western Europe. Drastic declines have been described in the UK, with the most severe declines occurring in rural areas. Hedgehogs are widely distributed in Denmark, but their status remains unknown.Fieldwork on hedgehogs has tended to focus on rural areas, leaving their ecology in suburban habitats largely unexplored, with clear implications for conservation initiatives. Here, we study the ecology of 35 juvenile hedgehogs using radio tracking during their first year of life in the suburbs of western Copenhagen.We use radio-tracking data to estimate (a) home range sizes in autumn and spring/summer, (b) survival during their first year of life, (c) the body mass changes before, during, and after hibernation, and (d) the hibernation behavior of the juvenile hedgehogs.We show that males and females have small home ranges compared with previous studies. The 95% MCP home range sizes in autumn were 1.33 ha (95% CI = 0.88-2.00) for males and 1.40 ha (95% CI = 0.84-2.32) for females; for spring/summer they were 6.54 ha (95% CI = 3.76-11.38) for males and 1.51 ha (95% CI = 0.63-3.63) for females. The juvenile survival probabilities during the study period from September 2014 to July 2015 were .56 for females and .79 for males. All healthy individuals gained body mass during the autumn and survived hibernation with little body mass loss thus demonstrating that the juveniles in the study were capable of gaining sufficient weight in the wild to survive their first hibernation.The climate is changing, but there is a lack of knowledge on how this affects mammal ecology. The exceptionally mild autumn of 2014 caused the juvenile hedgehogs to delay hibernation for up to a month compared with previous studies in Denmark.

European hedgehogs (Erinaceus europaeus) as a natural reservoir of methicillin-resistant Staphylococcus aureus carrying mecC in Denmark.

OBJECTIVES: A recent study from Sweden showed that European hedgehogs may constitute a reservoir for methicillin-resistant Staphylococcus aureus (MRSA), but this host-parasite relationship remains to be investigated in other countries. In this study, we therefore sought to: 1) determine the dissemination of MRSA in European hedgehogs throughout Denmark; 2) investigate determinants of MRSA carriage in hedgehogs; 3) determine the potential for zoonotic transmission of MRSA from hedgehogs to humans; and 4) characterise the detected MRSA on both a phenotypic and molecular level. METHODS: Nasal swabs were taken from 188 dead hedgehogs collected by volunteers throughout Denmark to determine the occurrence of MRSA. Additionally, 16 hedgehog rehabilitators were tested for potential zoonotic transmission of MRSA from hedgehogs to humans. The swabs were incubated in tryptic soy broth supplemented with 6.5% NaCl, followed by spread of 10 µl on Brilliance MRSA 2 agar. One presumptive MRSA colony from each plate was subcultured on 5% blood agar. All S. aureus subcultures were verified by a PCR assay detecting mecA, mecC, lukF-PV, scn, and spa, followed by spa typing. RESULTS: A total of 114 (61%) hedgehogs carried mecC-MRSA, whereas none carried mecA-MRSA. The detected mecC-MRSA belonged to two genetic lineages CC130 (spa-types: t528, t843, t1048, t3256, t3570, t6220, t17133) and CC1943 (spa-types: t978, t2345, t3391, t8835, t16868), 52% of which were spa-type t843 (CC130).The detection rate of mecC-MRSA in the hedgehogs was similar regardless of cause of death, sex, region and habitat type. None of the hedgehog rehabilitators carried MRSA. CONCLUSIONS: This nationwide study confirms a high occurrence of mecC-MRSA in hedgehogs, which could serve as a natural reservoir for this specific type of MRSA. Furthermore, our study did not find signs of zoonotic transmission of mecC-MRSA to hedgehog rehabilitators.

Animal life history is shaped by the pace of life and the distribution of age-specific mortality and reproduction.

Animals exhibit an extraordinary diversity of life history strategies. These realized combinations of survival, development and reproduction are predicted to be constrained by physiological limitations and by trade-offs in resource allocation. However, our understanding of these patterns is restricted to a few taxonomic groups. Using demographic data from 121 species, ranging from humans to sponges, we test whether such trade-offs universally shape animal life history strategies. We show that, after accounting for body mass and phylogenetic relatedness, 71% of the variation in animal life history strategies can be explained by life history traits associated with the fast-slow continuum (pace of life) and with a second axis defined by the distribution of age-specific mortality hazards and the spread of reproduction. While we found that life history strategies are associated with metabolic rate and ecological modes of life, surprisingly similar life history strategies can be found across the phylogenetic and physiological diversity of animals.

Data gaps and opportunities for comparative and conservation biology.

Biodiversity loss is a major challenge. Over the past century, the average rate of vertebrate extinction has been about 100-fold higher than the estimated background rate and population declines continue to increase globally. Birth and death rates determine the pace of population increase or decline, thus driving the expansion or extinction of a species. Design of species conservation policies hence depends on demographic data (e.g., for extinction risk assessments or estimation of harvesting quotas). However, an overview of the accessible data, even for better known taxa, is lacking. Here, we present the Demographic Species Knowledge Index, which classifies the available information for 32,144 (97%) of extant described mammals, birds, reptiles, and amphibians. We show that only 1.3% of the tetrapod species have comprehensive information on birth and death rates. We found no demographic measures, not even crude ones such as maximum life span or typical litter/clutch size, for 65% of threatened tetrapods. More field studies are needed; however, some progress can be made by digitalizing existing knowledge, by imputing data from related species with similar life histories, and by using information from captive populations. We show that data from zoos and aquariums in the Species360 network can significantly improve knowledge for an almost eightfold gain. Assessing the landscape of limited demographic knowledge is essential to prioritize ways to fill data gaps. Such information is urgently needed to implement management strategies to conserve at-risk taxa and to discover new unifying concepts and evolutionary relationships across thousands of tetrapod species.

The diversity of population responses to environmental change.

The current extinction and climate change crises pressure us to predict population dynamics with ever-greater accuracy. Although predictions rest on the well-advanced theory of age-structured populations, two key issues remain poorly explored. Specifically, how the age-dependency in demographic rates and the year-to-year interactions between survival and fecundity affect stochastic population growth rates. We use inference, simulations and mathematical derivations to explore how environmental perturbations determine population growth rates for populations with different age-specific demographic rates and when ages are reduced to stages. We find that stage- vs. age-based models can produce markedly divergent stochastic population growth rates. The differences are most pronounced when there are survival-fecundity-trade-offs, which reduce the variance in the population growth rate. Finally, the expected value and variance of the stochastic growth rates of populations with different age-specific demographic rates can diverge to the extent that, while some populations may thrive, others will inevitably go extinct.

Towards global data products of Essential Biodiversity Variables on species traits.

Essential Biodiversity Variables (EBVs) allow observation and reporting of global biodiversity change, but a detailed framework for the empirical derivation of specific EBVs has yet to be developed. Here, we re-examine and refine the previous candidate set of species traits EBVs and show how traits related to phenology, morphology, reproduction, physiology and movement can contribute to EBV operationalization. The selected EBVs express intra-specific trait variation and allow monitoring of how organisms respond to global change. We evaluate the societal relevance of species traits EBVs for policy targets and demonstrate how open, interoperable and machine-readable trait data enable the building of EBV data products. We outline collection methods, meta(data) standardization, reproducible workflows, semantic tools and licence requirements for producing species traits EBVs. An operationalization is critical for assessing progress towards biodiversity conservation and sustainable development goals and has wide implications for data-intensive science in ecology, biogeography, conservation and Earth observation.

Senescence is not inevitable.

Senescence, the physiological deterioration resulting in an increase in mortality and decline in fertility with age, is widespread in the animal kingdom and has often been regarded as an inescapable feature of all organisms. This essay briefly describes the history of the evolutionary theoretical ideas on senescence. The canonical evolutionary theories suggest that increasing mortality and decreasing fertility should be ubiquitous. However, increasing empirical data demonstrates that senescence may not be as universal a feature of life as once thought and that a diversity of demographic trajectories exists. These empirical observations support theoretical work indicating that a wide range of mortality and fertility trajectories is indeed possible, including senescence, negligible senescence and even negative senescence (improvement). Although many mysteries remain in the field of biogerontology, it is clear that senescence is not inevitable.

Individual heterogeneity determines sex differences in mortality in a monogamous bird with reversed sexual dimorphism.

Sex differences in mortality are pervasive in vertebrates, and usually result in shorter life spans in the larger sex, although the underlying mechanisms are still unclear. On the other hand, differences in frailty among individuals (i.e. individual heterogeneity), can play a major role in shaping demographic trajectories in wild populations. The link between these two processes has seldom been explored. We used Bayesian survival trajectory analysis to study age-specific mortality trajectories in the Eurasian sparrowhawk (Accipiter nisus), a monogamous raptor with reversed sexual size dimorphism. We tested the effect of individual heterogeneity on age-specific mortality, and the extent by which this heterogeneity was determined by average reproductive output and wing length as measures of an individual's frailty. We found that sex differences in age-specific mortality were primarily driven by the differences in individual heterogeneity between the two sexes. Females were more heterogeneous than males in their level of frailty. Thus, a larger number of females with low frailty are able to survive to older ages than males, with life expectancy for the least frail adult females reaching up to 4·23 years, while for the least frail adult males it was of 2·68 years. We found that 50% of this heterogeneity was determined by average reproductive output and wing length in both sexes. For both, individuals with high average reproductive output had also higher chances to survive. However, the effect of wing length was different between the two sexes. While larger females had higher survival, larger males had lower chances to survive. Our results contribute a novel perspective to the ongoing debate about the mechanisms that drive sex differences in vital rates in vertebrates. Although we found that variables that relate to the cost of reproduction and sexual dimorphism are at least partially involved in determining these sex differences, it is through their effect on the level of frailty that they affect age patterns of mortality. Therefore, our results raise the possibility that observed differences in age-specific demographic rates may in fact be driven by differences in individual heterogeneity.

Actuarial senescence in a long-lived orchid challenges our current understanding of ageing.

The dominant evolutionary theory of actuarial senescence-an increase in death rate with advancing age-is based on the concept of a germ cell line that is separated from the somatic cells early in life. However, such a separation is not clear in all organisms. This has been suggested to explain the paucity of evidence for actuarial senescence in plants. We used a 32 year study of Dactylorhiza lapponica that replaces its organs each growing season, to test whether individuals of this tuberous orchid senesce. We performed a Bayesian survival trajectory analysis accounting for reproductive investment, for individuals under two types of land use, in two climatic regions. The mortality trajectory was best approximated by a Weibull model, showing clear actuarial senescence. Rates of senescence in this model declined with advancing age, but were slightly higher in mown plots and in the more benign climatic region. At older ages, senescence was evident only when accounting for a positive effect of reproductive investment on mortality. Our results demonstrate actuarial senescence as well as a survival-reproduction trade-off in plants, and indicate that environmental context may influence senescence rates. This knowledge is crucial for understanding the evolution of demographic senescence and for models of plant population dynamics.