Population dynamics and comparative demographics in sympatric populations of the round-leaved orchids Platanthera macrophylla and P. orbiculata

Orchids (Orchidaceae) are a family of flowering plants with a high proportion of threatened taxa making them an important focus of plant conservation. Orchid conservation efforts are most effective when informed by reliable demographic research. We utilized transition matrix models to examine the population dynamics and demography within sympatric populations of a species pair of terrestrial round-leaved orchids, Platanthera macrophylla and P. orbiculata. The models were parameterized from a large data set spanning 9 years from field observations of over 1,000 orchids. Life table response experiments (LTRE) were used to identify which life history transitions, and which vital rates within those transitions, most contributed to observed differences between the two species and most contributed to interannual variation within each species. Results from mean transition matrices projected finite rates of population growth that were not significantly different between the two species, with P. macrophylla near the replacement rate and P. orbiculata below it. LTRE revealed that the difference in population growth rates between the two species was mostly due to differences in fecundity (flowering adult to protocorm transition) driven by differences in fruit set and seed germination into protocorm, which were much greater for P. macrophylla. The two primary contributors to interannual variation in population growth rates for both orchids were adult survival and fruit set, respectively. These findings indicate that any environmental disturbances harming adult survival or fecundity will have a disproportionately negative effect on the orchid populations.     

Density regulation in the Mediterranean leaf-toed gecko <Emphasis Type="Italic">Euleptes europaea</Emphasis>

Isolated populations are particularly prone to extinction, and understanding their temporal dynamics is relevant for conservation and management. In this study, the abundance of a population of the nocturnal leaf-toed gecko Euleptes europaea was estimated by mark-recapture over a 12-year period in northwest Italy. Simulation tests showed the presence of density-dependence, and autoregressive analyses indicated that direct density dependence was responsible for a large part of the variation in population growth rates. Density-dependent recruitment was suggested as the main demographic mechanism controlling population dynamics, which was also affected by solar radiation measured during the active gecko season. These results may contribute to implement conservation strategies in other small and isolated leaf-toed gecko populations.     

Site‐to‐site variation in the demography of a fire‐affected perennial. Acacia suaveolens,at Ku‐ring‐gai Chase National Park,New South Wales,Australia

Abstract Acacia suaveolens (Sm.) Willd is a perennial shrub that forms even‐aged stands, recruited from a soil seed‐bank following fire. It has previously been subject to demographic studies, which used a space‐for‐time substitution to investigate temporal patterns following fire. In the present study the potential for spatial variation across sites was investigated by sampling at several similarly aged populations in Ku‐ring‐gai Chase National Park, northern Sydney, Australia. Significant variation in mean size and fecundity of A. suaveolens individuals was observed among sites, over a 2‐4.6‐fold range in plant size, and a sevenfold range in mean fecundity. The observed variation at 3 years after fire encapsulated most of the variation previously observed among sites 0‐17 years since fire, emphasizing the importance of spatial variation in this species. For each site a two‐stage (seed, plant) matrix model was constructed, and projected from 3 to 25 years following fire. Population growth was measured as number of seeds per 3‐year‐old plant, and found to vary 1.4‐fold across models for different sites. This site‐to‐site variation, as well as that in size, fecundity and survival, was statistically significant. Variation in projected seeds per plant could mostly be attributed to differences in fecundity rather than plant survival. Sensitivity analyses emphasized the biological significance of the variation in fecundity. Whereas previous studies have focused on temporal variation, this work demonstrates the importance of extending our understanding of a species to include the spatial component of population dynamics.     

Small-scale demographic variability of the biocolor damselfish, <Emphasis Type="Italic">Stegastes partitus</Emphasis>, in the Florida Keys USA

The demographic responses of reef fish to their environment can be complex and in many cases, quite strong. Growth, mortality, longevity, and even reproductive effort have been demonstrated to vary for the same species of reef fish over scales of 100s to 1,000s of kilometers due to physiological and ecological interactions. Though few studies have explicitly documented it, this sort of habitat-mediated demography can also exist at very local scales. Here we present the results of a 2-year study of the bicolor damselfish, Stegastes partitus, in the Florida Keys, USA. We measured density and distribution, calculated key demographic rates (growth, survival, and fecundity), and characterized the environment (resident fish assemblage, substrate type and complexity, and food availability) of populations living in two adjacent but different habitats, the continuous fore reef and patchy back reef. Fish on the fore reef had an elevated growth rate and asymptotic size, increased mortality, and higher fecundity than fish on the back reef. We identified four potential causative mechanisms for these differences: food availability; competition; intraspecific density-dependent effects; and predation risk. Our data did not support an effect of either food availability or intraspecific density-dependence, but rather suggested that demographic responses are affected by both competition and predation risk.     

Prediction of cutting effects on a population of Chaerophyllum aureum– a demographic approach

Abstract. A demographic study of the undesirable perennial weed Chaerophyllum aureum in extensified French Pyrenean hay meadows was carried out. For two successive years the effect of date and intensity of cutting on the population demography of this colonizer. Population density is sensitive to cutting practices essentially through fecundity, seedling survival rate and seed dispersal between fields. Cutting reduces fecundity by preventing seed formation (early cutting) or by exportation of some mature seeds with hay (late cutting). Nevertheless, the number of seeds transported between field populations by hay harvest, organic manuring and associated cutting practices compensates for the lack of seed production in early‐cut populations, maintaining them at relative high densities. For each of three cutting regimes, the number of immigrant and emigrant seeds has been indirectly estimated from a prediction of population density at equilibrium from demographic parameters measured in field populations and compared with observed population densities.     

Plant performance in central and northern peripheral populations of the widespread Plantago coronopus

Peripheral populations have long been predicted to show lower vital rates, higher demographic fluctuations, and lower densities than central populations. However, recent research has questioned the existence of clear patterns across species’ ranges. To test these hypotheses, we monitored five central and six northern peripheral populations of the widespread herb Plantago coronopus along the European Atlantic coast during 5 yr. We estimated population density, and calculated mean values and temporal variability of four vital rates (survival, individual growth, fecundity and recruitment) in hundreds of plants in permanent plots. Central populations showed higher fecundity, whereas peripheral populations had higher recruitment per reproductive plant, indicating a higher overall reproductive success in the periphery. Central populations showed a marginally significant tendency for higher growth, and there were no differences between range positions in survival. Fecundity and growth were affected by intraspecific competition, and recruitment was affected by precipitation, highlighting the importance of local environmental conditions for population performance. Central and peripheral populations showed no significant differences in temporal variability of vital rates. Finally, density was significantly higher in peripheral than in central populations, in discrepancy with the abundant‐centre model. Density was correlated to seedling recruitment, which would counterbalance in peripheral populations the lower fecundity and the tendency for lower growth of established plants. Such compensations among vital rates might be particularly common in widespread plants, and advise against simplistic assumptions of population performance across ranges. The whole species’ life cycle should be considered, since different arrangements of vital rates are expected to maximize fitness in local environments. Our results show also the importance of discerning between geographical periphery and ecological marginality. In a context of climate‐induced range shifts, these considerations are crucial for the reliability of niche‐models and the management of plant peripheral populations.     

Variation in northern bobwhite demography along two temporal scales

Quantification and understanding of demographic variation across intra- and inter-annual temporal scales can benefit from the development of theoretical models of evolution and applied conservation of species. We used long-term survey data for northern bobwhites (Colinus virginianus) collected at the northern and southern extent of its geographic range to develop matrix population models which would allow investigation of intra- and inter-annual patterns in bobwhite population dynamics. We first evaluated intra-annual patterns in the importance of a seasonal demographic rate to asymptotic population growth rate with prospective perturbation analysis (elasticity analysis). We then conducted retrospective analysis (life table response experiments) of inter-annual patterns in the contribution of observed changes in demography to the observed change in population growth rate. Survival in the earliest age class during the nonbreeding season had the greatest potential influence in both the northern and southern populations. Examination of inter-annual variation in demography indicated that variation in nonbreeding season survival in the earliest age class contributed the most to observed changes in population growth rate in the northern population. In contrast, changes in fertility in the earliest age class in the southern population had the greatest influence on changes in population growth rate. Prospective elasticity analyses highlight the similarities in bobwhite demography throughout different parts of its geographic range, while retrospective life table response experiments revealed important patterns in the temporal differences of bobwhite life history at the northern and southern extent of its geographic range.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

Variation in life history and demography of the American black bear

Variation in life history and demography across a species' range informs researchers about regional adaptations and affects whether managers can borrow information from other populations in decision-making. The American black bear (Ursus americanus) is a long-lived game species whose continued persistence depends on management of harvest and removal of habituated bears that come into conflict with humans. Understanding the demography of black bears guides efforts at management and conservation, yet detailed knowledge of many populations is typically lacking. I performed a hierarchical Bayesian meta-analysis of black bear demographic studies across the geographic range of the species to explore how vital rates vary across the range, what information they give us about population growth, and whether managers can justify borrowing information from other studies to inform management decisions. Cub, yearling, and adult survival and fecundity varied between eastern and western North America, whereas subadult survival did not show geographic structuring. Adult survival and fecundity appeared to trade off, with higher survival in the western portions of bears' range and higher fecundity in the east. Although adult survival had the highest elasticity, differences in reproduction drove differences in population growth rate. Mean population growth rate was higher in the east (0.99; 95% credible interval [CrI]: 0.96, 1.03) than the west (0.97; 95%CrI: 0.93, 1.01). Despite declining trends in the west, 34% of the distribution of population growth rate was >1, compared to 55% in the east. Further work needs to be done to address the cause of the apparent trade-off between adult survival and fecundity and explore how the estimated growth rates are likely to affect population status of black bears. Because population growth rates are close to 1 and small deviations could impact whether a population is considered increasing or decreasing, managers need to employ caution in borrowing vital rates from other populations. © 2011 The Wildlife Society.     

Seed production and pre-dispersal reproductive losses in the narrow endemic <Emphasis Type="Italic">Euphorbia pedroi</Emphasis> (Euphorbiaceae)

Euphorbia pedroi is a narrow endemic species with three known populations located in coastal areas of western Portugal. This study focused on the reproductive biology of this species from flowering to dispersal, aiming to identify the factors causing decrease in seed production potential and to assess the spatio-temporal patterns of seed production at the individual and population levels. The abortion of reproductive structures, particularly seeds, represented a major fraction of losses in the potential seed production of E. pedroi. Moth larvae destroyed a variable proportion of cyathia in a large number of plants from the two populations regardless of their degree of isolation. Furthermore, generalist and specialist pre-dispersal seed predators were responsible for temporally variable seed losses unrelated with variables indicative of plant size and fecundity, and showing no consistency at the individual level. Specialist seed-wasps inflicted the highest losses to E. pedroi and their impact was intimately associated with the magnitude of yearly variation in seed production. This finding highlights the role of the inter-annual variation in seed production as a key feature in this plant-seed predator system. The effect of the two groups of seed predators on the reproductive output of E. pedroi was additive and those insects do not seem to exert an important selective pressure on the traits studied. The proportion of intact seeds produced by E. pedroi differed between locations, but not between individuals within each population, highlighting the major contribution of larger plants to the seed pool.     

Population size,center–periphery,and seed dispersers’ effects on the genetic diversity and population structure of the Mediterranean relict shrub Cneorum tricoccon

The effect of population size on population genetic diversity and structure has rarely been studied jointly with other factors such as the position of a population within the species’ distribution range or the presence of mutualistic partners influencing dispersal. Understanding these determining factors for genetic variation is critical for conservation of relict plants that are generally suffering from genetic deterioration. Working with 16 populations of the vulnerable relict shrub Cneorum tricoccon throughout the majority of its western Mediterranean distribution range, and using nine polymorphic microsatellite markers, we examined the effects of periphery (peripheral vs. central), population size (large vs. small), and seed disperser (introduced carnivores vs. endemic lizards) on the genetic diversity and population structure of the species. Contrasting genetic variation (H_E: 0.04–0.476) was found across populations. Peripheral populations showed lower genetic diversity, but this was dependent on population size. Large peripheral populations showed high levels of genetic diversity, whereas small central populations were less diverse. Significant isolation by distance was detected, indicating that the effect of long‐distance gene flow is limited relative to that of genetic drift, probably due to high selfing rates (F_IS = 0.155–0.887), restricted pollen flow, and ineffective seed dispersal. Bayesian clustering also supported the strong population differentiation and highly fragmented structure. Contrary to expectations, the type of disperser showed no significant effect on either population genetic diversity or structure. Our results challenge the idea of an effect of periphery per se that can be mainly explained by population size, drawing attention to the need of integrative approaches considering different determinants of genetic variation. Furthermore, the very low genetic diversity observed in several small populations and the strong among‐population differentiation highlight the conservation value of large populations throughout the species’ range, particularly in light of climate change and direct human threats.     

Forest Fragmentation Alters the Population Dynamics of a Late‐successional Tropical Tree

Tropical late‐successional tree species are at high risk of local extinction due to habitat loss and fragmentation. Population‐growth rates in fragmented populations are predicted to decline as a result of reduced fecundity, survival and growth. We examined the demographic effects of habitat fragmentation by comparing the population dynamics of the late‐successional tree Poulsenia armata (Moraceae) in southern Mexico between a continuous forest and several forest fragments using integral projection models (IPMs) during 2010–2012. Forest fragmentation did not lead to differences in population density and even resulted in a higher population‐growth rate (λ) in fragments compared to continuous forests. Habitat fragmentation had drastic effects on the dynamics of P. armata, causing the population structure to shift toward smaller sizes. Fragmented populations experienced a significant decrease in juvenile survival and growth compared to unaltered populations. Adult survival and growth made the greatest relative contributions to λ in both habitat types during 2011–2012. However, the relative importance of juvenile survival and growth to λ was highest in the fragmented forest in 2010–2011. Our Life Table Response Experiment analysis revealed that positive contributions of adult fecundity explained most of the variation of λ between both habitats and annual periods. Finally, P. armata has a relatively slow speed of recovery after disturbances, compromising persistence of fragmented populations. Developing a mechanistic understanding of how forest fragmentation affects plant population dynamics, as done here, will prove essential for the preservation of natural areas.     

The role of demography,intra‐species variation,and species distribution models in species' projections under climate change

Organisms are projected to shift their distribution ranges under climate change. The typical way to assess range shifts is by species distribution models (SDMs), which predict species’ responses to climate based solely on projected climatic suitability. However, life history traits can impact species’ responses to shifting habitat suitability. Additionally, it remains unclear if differences in vital rates across populations within a species can offset or exacerbate the effects of predicted changes in climatic suitability on population viability. In order to obtain a fuller understanding of the response of one species to projected climatic changes, we coupled demographic processes with predicted changes in suitable habitat for the monocarpic thistle Carlina vulgaris across northern Europe. We first developed a life history model with species‐specific average fecundity and survival rates and linked it to a SDM that predicted changes in habitat suitability through time with changes in climatic variables. We then varied the demographic parameters based upon observed vital rates of local populations from a translocation experiment. Despite the fact that the SDM alone predicted C. vulgaris to be a climate ‘winner’ overall, coupling the model with changes in demography and small‐scale habitat suitability resulted in a matrix of stable, declining, and increasing patches. For populations predicted to experience declines or increases in abundance due to changes in habitat suitability, altered fecundity and survival rates can reverse projected population trends.     

Drivers of megaherbivore demographic fluctuations: inference from elephants

Environmentally induced variation in survival and fecundity generates demographic fluctuations that affect population growth rate. However, a general pattern of the comparative influence of variation in fecundity and juvenile survival on elephant population dynamics has not been investigated at a broad scale. We evaluated the relative importance of conception, gestation, first year survival and subsequent survivorship for controlling demographic variation by exploring the relationship between past environmental conditions determined by integrated normalized difference vegetation index (INDVI) and the shape of age distributions at 17 sites across Africa. We showed that, generally, INDVI during gestation best explained anomalies in age structure. However, in areas with low mean annual rainfall, INDVI during the first year of life was critical. The results challenge Eberhardt's paradigm for population analysis that suggests that populations respond to limited resource availability through a sequential decrease in juvenile survival, reproductive rate and adult survival. Contrastingly, elephants appear to respond first through a reduction in reproductive rate. We conclude that this discrepancy is likely due to the evolutionary significance of extremely large body size – an adaptation that increases survival rate but decreases reproductive potential. Other megaherbivores may respond similarly to resource limitation due to similarities in population dynamics. Knowing how vital rates vary with changing environmental conditions will permit better forecasts of the trajectories of megaherbivore populations.     

Local population dynamics and the impact of scale and isolation: a study on different little owl populations

The understanding of how variation of demographic rates translates into variation of population growth is a central aim in population ecology. Besides stochastic and deterministic factors, the spatial extent and the isolation of a local population may have an impact on the contribution of the different demographic components. Using long-term demographic data we performed retrospective population analyses of four little owl ( Athene noctua ) populations with differential spatial extent and degree of isolation to assess the contribution of demographic rates to the variation of the growth rate (λ) of each local population and to the difference of λ among populations. In all populations variation of fecundity contributed least to variation of λ, and variation of adult survival contributed most to variation of λ in three of four populations. Between population comparisons revealed that differences mainly stem from differences of immigration and juvenile local survival. The relative importance of immigration to λ tended to decrease with increasing spatial extent and isolation of the local populations. None of the four local populations was self-sustainable. Because the local populations export and import individuals, they can be considered as open recruitment systems in which part of the recruited breeding birds are not produced locally. The spatial extent and the degree of isolation of a local population have an impact on local population dynamics; hence these factors need to be considered in studies about local population dynamics and for deriving conservation measures.     

Spatiotemporal variation in predispersal seed predation intensity

The effect of predispersal seed predation by Bruchus atomarius (Bruchidae, Coleoptera) on individual performance and population dynamics of the perennial forest herb, Lathyrus vernus (Leguminosae), was investigated in 11 permanent plots over 4 years. Seed predation and parameters describing intra-specific neighbour distance, plant size, inflorescence size, flowering phenology and current and previous herbivore damage were measured on all plants. In addition, demographic information from all plots was analysed using transition matrix population models in order to estimate the influence of seed predation on population growth rates. Predispersal seed predation rates differed significantly among years. Plot averages ranged from 0 to 83.7%. However, most of the variation occurred among individuals. Within individuals there was no consistency in predation rates among years. Exposure to herbivory, plant size and flowering phenology did not affect predation rates but individuals with larger inflorescences suffered from significantly higher predation. Seed predation in L. vernus was not influenced by neighbour distances of individual plants but it was positively correlated with the average density of seeds within plots, suggesting that seed predation is density dependent at the patch level. The reduction in population growth rate due to seed predation ranged from 0 to 7.6%. The sensitivity of population growth rate to reductions in seed production varied considerably among years and plots. This variation was mainly due to differences in the reproductive value of seeds and seedlings. The intensity of seed predation over the range found was not correlated with changes in population growth rate. The results of this study suggest that the influence of external factors, like seed predation, on population growth rate largely depends on the demographic transition rates in the investigated population.     

The demographic drivers of local population dynamics in two rare migratory birds

The exchange of individuals among populations can have strong effects on the dynamics and persistence of a given population. Yet, estimation of immigration rates remains one of the greatest challenges for animal demographers. Little empirical knowledge exists about the effects of immigration on population dynamics. New integrated population models fitted using Bayesian methods enable simultaneous estimation of fecundity, survival and immigration, as well as the growth rate of a population of interest. We applied this novel analytical framework to the demography of two populations of long-distance migratory birds, hoopoe Upupa epops and wryneck Jynx torquilla, in a study area in south-western Switzerland. During 2002–2010, the hoopoe population increased annually by 11%, while the wryneck population remained fairly stable. Apparent juvenile and adult survival probability was nearly identical in both species, but fecundity and immigration were slightly higher in the hoopoe. Hoopoe population growth rate was strongly correlated with juvenile survival, fecundity and immigration, while that of wrynecks strongly correlated only with immigration. This indicates that demographic components impacting the arrival of new individuals into the populations were more important for their dynamics than demographic components affecting the loss of individuals. The finding that immigration plays a crucial role in the population growth rates of these two rare species emphasizes the need for a broad rather than local perspective for population studies, and the development of wide-scale conservation actions.     

Fitness components and the determinants of fecundity in populations of a native perennial grass (Tridens flavus)

Understanding intraspecific variation in traits that determine fitness is foundational to a trait-based approach to plant ecology. This study examined fitness components during 3 years of reproduction in a polycarpic perennial bunchgrass (Tridens flavus) native to eastern North America that could prove useful in revegetating disturbed habitats. Plants were cultured from seeds of five populations in central New Jersey, USA, and planted in July 2015 into two undisturbed gardens 30 m apart that differed in availability of sunlight and soil moisture. Following flowering in 2016, 2017 and 2018, the number of panicles, seed set, seed number (fecundity) and seed mass were recorded. Final dry aboveground mass was determined. Seed set was high (>70%) in all populations and gardens. Panicle production varied with population and was strongly correlated with fecundity, but populations were not differentiated for other fitness components. Panicle and seed number were greatest in the drier garden with greater daily light availability. Mass per seed was reduced as more seeds were produced in the second and third year but showed low variation compared to fecundity. Vegetative mass was the most important variable determining fecundity. Close proximity of sampled sites and an outcrossed, wind-pollinated mating strategy may have precluded detection of differentiation among T. flavus populations in the common gardens. High seed set, prodigious seed production on multiple panicles and high seed germinability and overwinter survival account for the occurrence of large populations of this native grass along roadsides and within successional fields and young woodlands throughout the region.     

Comparative demography of two giant caulescent rosettes (Espeletia timotensis and E. spicata) from the high tropical Andes

Using field data from previous studies we built matrix models for two populations of giant rosettes, Espeletia timotensis Cuatrec. and E. spicata Sch. Bip. Wedd., from the Andes Cordillera in Mérida, Venezuela. We analysed the models and calculated population growth rate (λ), sensitivities, elasticities and the sensitivity of the elasticities to changes in the vital rates. The analysis showed that the two species behave alike in general demographic terms. In both models, population growth rate is positive and sensitivities of λ to changes in vital rates decrease markedly in this order: plant establishment, progression of juvenile–adult, germination and survival. The relative contributions of vital rates to λ (elasticities) are very similar to those of other woody plant species: a higher contribution of survival and a very low contribution of fecundity. Transition from seedling to juvenile is most important and the younger established stages (juveniles and young adults) play a predominant demographic role in both populations. Seed banks and older adults are playing a relatively minor role in the dynamics of both populations. However, they may be important in relation to unpredictable, favourable or detrimental events. Perturbation analysis of elasticities showed that increasing the rate of plant establishment will decrease the relative importance of stasis. We conclude that both species are demographically very close, and similar to other long‐lived woody plant species. However, the two species differ in the role of the seed bank, which seems more important in the demography of E. spicata than in E. timotensis.     

IS SURVIVORSHIP A BETTER FITNESS SURROGATE THAN FECUNDITY?

Although fitness depends on both survivorship and fecundity, we tend to assume fecundity relates to fitness more directly than survivorship. In fact, several recent ecological studies suggest fitness depends more heavily on annual survivorship than annual fecundity for most taxa with lifespans longer than one year. These studies review elasticities of transition matrices for a broad range of taxa. Elasticities covary monotonically with selection gradients for demographic rates and are identical to selection gradients for traits rescaled to have mean values of zero and variance of one. For all taxa except semelparous perennial plants, adult survivorship has consistently higher elasticity than other suites of demographic rates. Fecundity only rarely has the highest elasticity. Thus, differences in yearly survival affect fitness disproportionately more than differences in yearly fecundity, even in many exponentially growing populations. This pattern reinforces the importance of interpreting the contribution of vital rates to fitness in the context of life history and population dynamics.