Nonlinear life table response experiment analysis: Decomposing nonlinear and nonadditive population growth responses to changes in environmental drivers.

Life table response experiments (LTREs) decompose differences in population growth rate between environments into separate contributions from each underlying demographic rate. However, most LTRE analyses make the unrealistic assumption that the relationships between demographic rates and environmental drivers are linear and independent, which may result in diminished accuracy when these assumptions are violated. We extend regression LTREs to incorporate nonlinear (second-order) terms and compare the accuracy of both approaches for three previously published demographic datasets. We show that the second-order approach equals or outperforms the linear approach for all three case studies, even when all of the underlying vital rate functions are linear. Nonlinear vital rate responses to driver changes contributed most to population growth rate responses, but life history changes also made substantial contributions. Our results suggest that moving from linear to second-order LTRE analyses could improve our understanding of population responses to changing environments.

Inclusive Fitness May Explain Some but Not All Benefits Derived from Helping Behavior in a Cooperatively Breeding Bird.

AbstractIn cooperative breeding systems, inclusive fitness theory predicts that nonbreeding helpers more closely related to the breeders should be more willing to provide costly alloparental care and thus have more impact on breeder fitness. In the red-cockaded woodpecker (Dryobates borealis), most helpers are the breeders' earlier offspring, but helpers do vary within groups in both relatedness to the breeders (some even being unrelated) and sex, and it can be difficult to parse their separate impacts on breeder fitness. Moreover, most support for inclusive fitness theory has been positive associations between relatedness and behavior rather than actual fitness consequences. We used functional linear models to evaluate the per capita effects of helpers of different relatedness on eight breeder fitness components measured for up to 41 years at three sites. In support of inclusive fitness theory, helpers more related to the breeding pair made greater contributions to six fitness components. However, male helpers made equal contributions to increasing prefledging survival regardless of relatedness. These findings suggest that both inclusive fitness benefits and other direct benefits may underlie helping behaviors in the red-cockaded woodpecker. Our results also demonstrate the application of an underused statistical approach to disentangle a complex ecological phenomenon.

Conflicting constraints on male mating success shape reward size in pollen-rewarding plants.

PREMISE: Pollen-rewarding plants face two conflicting constraints: They must prevent consumptive emasculation while remaining attractive to pollen-collecting visitors. Small pollen packages (the quantity of pollen available in a single visit) may discourage visitors from grooming (reducing consumptive loss) but may also decrease a plant's attractiveness to pollen-collecting visitors. What package size best balances these two constraints? METHODS: We modeled the joint effects of pollinators' grooming behaviors and package size preferences on the optimal package size (i.e., the size that maximizes pollen donation). We then used this model to examine Darwin's conjecture that selection should favor increased pollen production in pollen-rewarding plants. RESULTS: When package size preferences are weak, minimizing package size reduces grooming losses and should be favored (as in previous theoretical studies). Stronger preferences select for larger packages despite the associated increase to grooming loss because loss associated with nonremoval of smaller packages is even greater. Total pollen donation increases with production (as Darwin suggested). However, if floral visitation declines or packages size preference increases with overall pollen availability, the fraction of pollen donated may decline as per-plant pollen production increases. Hence, increasing production may result in diminishing returns. CONCLUSIONS: Pollen-rewarding plants can balance conflicting constraints on pollen donation by producing intermediate-sized pollen packages. Strictly pollen-rewarding plants may have responded to past selection to produce more pollen in total, but diminishing returns may limit the strength of that selection.

Biotic and anthropogenic forces rival climatic/abiotic factors in determining global plant population growth and fitness.

Multiple, simultaneous environmental changes, in climatic/abiotic factors, interacting species, and direct human influences, are impacting natural populations and thus biodiversity, ecosystem services, and evolutionary trajectories. Determining whether the magnitudes of the population impacts of abiotic, biotic, and anthropogenic drivers differ, accounting for their direct effects and effects mediated through other drivers, would allow us to better predict population fates and design mitigation strategies. We compiled 644 paired values of the population growth rate (λ) from high and low levels of an identified driver from demographic studies of terrestrial plants. Among abiotic drivers, natural disturbance (not climate), and among biotic drivers, interactions with neighboring plants had the strongest effects on λ However, when drivers were combined into the 3 main types, their average effects on λ did not differ. For the subset of studies that measured both the average and variability of the driver, λ was marginally more sensitive to 1 SD of change in abiotic drivers relative to biotic drivers, but sensitivity to biotic drivers was still substantial. Similar impact magnitudes for abiotic/biotic/anthropogenic drivers hold for plants of different growth forms, for different latitudinal zones, and for biomes characterized by harsher or milder abiotic conditions, suggesting that all 3 drivers have equivalent impacts across a variety of contexts. Thus, the best available information about the integrated effects of drivers on all demographic rates provides no justification for ignoring drivers of any of these 3 types when projecting ecological and evolutionary responses of populations and of biodiversity to environmental changes.

Latitudinal gradients in population growth do not reflect demographic responses to climate.

Spatial gradients in population growth, such as across latitudinal or elevational gradients, are often assumed to primarily be driven by variation in climate, and are frequently used to infer species' responses to climate change. Here, we use a novel demographic, mixed-model approach to dissect the contributions of climate variables vs. other latitudinal or local site effects on spatiotemporal variation in population performance in three perennial bunchgrasses. For all three species, we find that performance of local populations decreases with warmer and drier conditions, despite latitudinal trends of decreasing population growth toward the cooler and wetter northern portion of each species' range. Thus, latitudinal gradients in performance are not predictive of either local or species-wide responses to climate. This pattern could be common, as many environmental drivers, such as habitat quality or species' interactions, are likely to vary with latitude or elevation, and thus influence or oppose climate responses.

Aridity weakens population-level effects of multiple species interactions on Hibiscus meyeri.

Predicting how species' abundances and ranges will shift in response to climate change requires a mechanistic understanding of how multiple factors interact to limit population growth. Both abiotic stress and species interactions can limit populations and potentially set range boundaries, but we have a poor understanding of when and where each is most critical. A commonly cited hypothesis, first proposed by Darwin, posits that abiotic factors (e.g., temperature, precipitation) are stronger determinants of range boundaries in apparently abiotically stressful areas ("stress" indicates abiotic factors that reduce population growth), including desert, polar, or high-elevation environments, whereas species interactions (e.g., herbivory, competition) play a stronger role in apparently less stressful environments. We tested a core tenet of this hypothesis-that population growth rate is more strongly affected by species interactions in less stressful areas-using experimental manipulations of species interactions affecting a common herbaceous plant, Hibiscus meyeri (Malvaceae), across an aridity gradient in a semiarid African savanna. Population growth was more strongly affected by four distinct species interactions (competition with herbaceous and shrubby neighbors, herbivory, and pollination) in less stressful mesic areas than in more stressful arid sites. However, contrary to common assumptions, this effect did not arise because of greater density or diversity of interacting species in less stressful areas, but rather because aridity reduced sensitivity of population growth to these interactions. Our work supports classic predictions about the relative strength of factors regulating population growth across stress gradients, but suggests that this pattern results from a previously unappreciated mechanism that may apply to many species worldwide.

Incorporating local adaptation into forecasts of species' distribution and abundance under climate change.

Populations of many species are genetically adapted to local historical climate conditions. Yet most forecasts of species' distributions under climate change have ignored local adaptation (LA), which may paint a false picture of how species will respond across their geographic ranges. We review recent studies that have incorporated intraspecific variation, a potential proxy for LA, into distribution forecasts, assess their strengths and weaknesses, and make recommendations for how to improve forecasts in the face of LA. The three methods used so far (species distribution models, response functions, and mechanistic models) reflect a trade-off between data availability and the ability to rigorously demonstrate LA to climate. We identify key considerations for incorporating LA into distribution forecasts that are currently missing from many published studies, including testing the spatial scale and pattern of LA, the confounding effects of LA to nonclimatic or biotic drivers, and the need to incorporate empirically based dispersal or gene flow processes. We suggest approaches to better evaluate these aspects of LA and their effects on species-level forecasts. In particular, we highlight demographic and dynamic evolutionary models as promising approaches to better integrate LA into forecasts, and emphasize the importance of independent model validation. Finally, we urge closer examination of how LA will alter the responses of central vs. marginal populations to allow stronger generalizations about changes in distribution and abundance in the face of LA.

Geographic location, local environment, and individual size mediate the effects of climate warming and neighbors on a benefactor plant.

Predictions of plant responses to global warming frequently ignore biotic interactions and intraspecific variation across geographical ranges. Benefactor species play an important role in plant communities by protecting other taxa from harsh environments, but the combined effects of warming and beneficiary species on their performance have been largely unexamined. We analyzed the joint effects of elevated temperature and neighbor removal on the benefactor plant Silene acaulis, in factorial experiments near its low- and high-latitude range limits in Europe. We recorded growth, probability of reproduction and fruit set during 3 years. The effects of enhanced temperature were positive near the northern limit and negative in the south for some performance measures. This pattern was stronger in the presence of neighbors, possibly due to differential thermal tolerances between S. acaulis and beneficiary species in each location. Neighbors generally had a negative or null impact on S. acaulis, in agreement with previous reviews of overall effects of plant-plant interactions on benefactors. However, small S. acaulis individuals in the north showed higher growth when surrounded by neighbors. Finally, the local habitat within each location influenced some effects of experimental treatments. Overall, we show that plant responses to rising temperatures may strongly depend on their position within the geographic range, and on species interactions. Our results also highlight the need to consider features of the interacting taxa, such as whether they are benefactor species, as well as local-scale environmental variation, to predict the joint effects of global warming and biotic interactions on species and communities.

Both life-history plasticity and local adaptation will shape range-wide responses to climate warming in the tundra plant Silene acaulis.

Many predictions of how climate change will impact biodiversity have focused on range shifts using species-wide climate tolerances, an approach that ignores the demographic mechanisms that enable species to attain broad geographic distributions. But these mechanisms matter, as responses to climate change could fundamentally differ depending on the contributions of life-history plasticity vs. local adaptation to species-wide climate tolerances. In particular, if local adaptation to climate is strong, populations across a species' range-not only those at the trailing range edge-could decline sharply with global climate change. Indeed, faster rates of climate change in many high latitude regions could combine with local adaptation to generate sharper declines well away from trailing edges. Combining 15 years of demographic data from field populations across North America with growth chamber warming experiments, we show that growth and survival in a widespread tundra plant show compensatory responses to warming throughout the species' latitudinal range, buffering overall performance across a range of temperatures. However, populations also differ in their temperature responses, consistent with adaptation to local climate, especially growing season temperature. In particular, warming begins to negatively impact plant growth at cooler temperatures for plants from colder, northern populations than for those from warmer, southern populations, both in the field and in growth chambers. Furthermore, the individuals and maternal families with the fastest growth also have the lowest water use efficiency at all temperatures, suggesting that a trade-off between growth and water use efficiency could further constrain responses to forecasted warming and drying. Taken together, these results suggest that populations throughout species' ranges could be at risk of decline with continued climate change, and that the focus on trailing edge populations risks overlooking the largest potential impacts of climate change on species' abundance and distribution.

Mechanism matters: the cause of fluctuations in boom-bust populations governs optimal habitat restoration strategy.

Many populations exhibit boom-bust dynamics in which abundance fluctuates dramatically over time. Past research has focused on identifying whether the cause of fluctuations is primarily exogenous, e.g., environmental stochasticity coupled with weak density dependence, or endogenous, e.g., over-compensatory density dependence. Far fewer studies have addressed whether the mechanism responsible for boom-bust dynamics matters with respect to at-risk species management. Here, we ask whether the best strategy for restoring habitat across a landscape differs under exogenously vs. endogenously driven boom-bust dynamics. We used spatially explicit individual-based models to assess how butterfly populations governed by the two mechanisms would respond to habitat restoration strategies that varied in the level of resource patchiness, from a single large patch to multiple patches spaced at different distances. Our models showed that the restoration strategy that minimized extinction risk and boom-bust dynamics would be markedly different depending on the governing mechanism. Exogenously governed populations fared best in a single large habitat patch, whereas for endogenously driven populations, boom-bust dynamics were dampened and extinction risk declined when the total restored area was split into multiple patches with low to moderate inter-patch spacing. Adding environmental stochasticity to the endogenous model did not alter this result. Habitat fragmentation lowered extinction risk in the endogenously driven populations by reducing their growth rate, precluding both "boom" phases and, more importantly, "bust" phases. Our findings suggest that (1) successful restoration will depend on understanding the causes of fluctuations in at-risk populations, (2) the level and pattern of spatiotemporal environmental heterogeneity will also affect the ideal management approach, and (3) counterintuitively, for at-risk species with endogenously governed boom-bust dynamics, lowering the intrinsic population growth rate may decrease extinction risk.

Does climate variability influence the demography of wild primates? Evidence from long-term life-history data in seven species.

Earth's rapidly changing climate creates a growing need to understand how demographic processes in natural populations are affected by climate variability, particularly among organisms threatened by extinction. Long-term, large-scale, and cross-taxon studies of vital rate variation in relation to climate variability can be particularly valuable because they can reveal environmental drivers that affect multiple species over extensive regions. Few such data exist for animals with slow life histories, particularly in the tropics, where climate variation over large-scale space is asynchronous. As our closest relatives, nonhuman primates are especially valuable as a resource to understand the roles of climate variability and climate change in human evolutionary history. Here, we provide the first comprehensive investigation of vital rate variation in relation to climate variability among wild primates. We ask whether primates are sensitive to global changes that are universal (e.g., higher temperature, large-scale climate oscillations) or whether they are more sensitive to global change effects that are local (e.g., more rain in some places), which would complicate predictions of how primates in general will respond to climate change. To address these questions, we use a database of long-term life-history data for natural populations of seven primate species that have been studied for 29-52 years to investigate associations between vital rate variation, local climate variability, and global climate oscillations. Associations between vital rates and climate variability varied among species and depended on the time windows considered, highlighting the importance of temporal scale in detection of such effects. We found strong climate signals in the fertility rates of three species. However, survival, which has a greater impact on population growth, was little affected by climate variability. Thus, we found evidence for demographic buffering of life histories, but also evidence of mechanisms by which climate change could affect the fates of wild primates.

Demographic compensation and tipping points in climate-induced range shifts.

To persist, species are expected to shift their geographical ranges polewards or to higher elevations as the Earth's climate warms. However, although many species' ranges have shifted in historical times, many others have not, or have shifted only at the high-latitude or high-elevation limits, leading to range expansions rather than contractions. Given these idiosyncratic responses to climate warming, and their varied implications for species' vulnerability to climate change, a critical task is to understand why some species have not shifted their ranges, particularly at the equatorial or low-elevation limits, and whether such resilience will last as warming continues. Here we show that compensatory changes in demographic rates are buffering southern populations of two North American tundra plants against the negative effects of a warming climate, slowing their northward range shifts, but that this buffering is unlikely to continue indefinitely. Southern populations of both species showed lower survival and recruitment but higher growth of individual plants, possibly owing to longer, warmer growing seasons. Because of these and other compensatory changes, the population growth rates of southern populations are not at present lower than those of northern ones. However, continued warming may yet prove detrimental, as most demographic rates that improved in moderately warmer years declined in the warmest years, with the potential to drive future population declines. Our results emphasize the need for long-term, range-wide measurement of all population processes to detect demographic compensation and to identify nonlinear responses that may lead to sudden range shifts as climatic tipping points are exceeded.

Reproductive aging patterns in primates reveal that humans are distinct.

Women rarely give birth after ∼45 y of age, and they experience the cessation of reproductive cycles, menopause, at ∼50 y of age after a fertility decline lasting almost two decades. Such reproductive senescence in mid-lifespan is an evolutionary puzzle of enduring interest because it should be inherently disadvantageous. Furthermore, comparative data on reproductive senescence from other primates, or indeed other mammals, remains relatively rare. Here we carried out a unique detailed comparative study of reproductive senescence in seven species of nonhuman primates in natural populations, using long-term, individual-based data, and compared them to a population of humans experiencing natural fertility and mortality. In four of seven primate species we found that reproductive senescence occurred before death only in a small minority of individuals. In three primate species we found evidence of reproductive senescence that accelerated throughout adulthood; however, its initial rate was much lower than mortality, so that relatively few individuals experienced reproductive senescence before death. In contrast, the human population showed the predicted and well-known pattern in which reproductive senescence occurred before death for many women and its rate accelerated throughout adulthood. These results provide strong support for the hypothesis that reproductive senescence in midlife, although apparent in natural-fertility, natural-mortality populations of humans, is generally absent in other primates living in such populations.

Predicting changes in the distribution and abundance of species under environmental change.

Environmental changes are expected to alter both the distribution and the abundance of organisms. A disproportionate amount of past work has focused on distribution only, either documenting historical range shifts or predicting future occurrence patterns. However, simultaneous predictions of abundance and distribution across landscapes would be far more useful. To critically assess which approaches represent advances towards the goal of joint predictions of abundance and distribution, we review recent work on changing distributions and on effects of environmental drivers on single populations. Several methods have been used to predict changing distributions. Some of these can be easily modified to also predict abundance, but others cannot. In parallel, demographers have developed a much better understanding of how changing abiotic and biotic drivers will influence growth rate and abundance in single populations. However, this demographic work has rarely taken a landscape perspective and has largely ignored the effects of intraspecific density. We advocate a synthetic approach in which population models accounting for both density dependence and effects of environmental drivers are used to make integrated predictions of equilibrium abundance and distribution across entire landscapes. Such predictions would constitute an important step forward in assessing the ecological consequences of environmental changes.

Demographic compensation among populations: what is it, how does it arise and what are its implications?

Most species are exposed to significant environmental gradients across their ranges, but vital rates (survival, growth, reproduction and recruitment) need not respond in the same direction to those gradients. Opposing vital rate trends across environments, a phenomenon that has been loosely called 'demographic compensation', may allow species to occupy larger geographical ranges and alter their responses to climate change. Yet the term has never been precisely defined, nor has its existence or strength been assessed for multiple species. Here, we provide a rigorous definition, and use it to develop a strong test for demographic compensation. By applying the test to data from 26 published, multi-population demographic studies of plants, we show that demographic compensation commonly occurs. We also investigate the mechanisms by which this phenomenon arises by assessing which demographic processes and life stages are most often involved. In addition, we quantify the effect of demographic compensation on variation in population growth rates across environmental gradients, a potentially important determinant of the size of a species' geographical range. Finally, we discuss the implications of demographic compensation for the responses of single populations and species' ranges to temporal environmental variation and to ongoing environmental trends, e.g. due to climate change.

The demographic consequences of mutualism: ants increase host-plant fruit production but not population growth.

The impact of mutualists on a partner's demography depends on how they affect the partner's multiple vital rates and how those vital rates, in turn, affect population growth. However, mutualism studies rarely measure effects on multiple vital rates or integrate them to assess the ultimate impact on population growth. We used vital rate data, population models and simulations of long-term population dynamics to quantify the demographic impact of a guild of ant species on the plant Ferocactus wislizeni. The ants feed at the plant's extrafloral nectaries and attack herbivores attempting to consume reproductive organs. Ant-guarded plants produced significantly more fruit, but ants had no significant effect on individual growth or survival. After integrating ant effects across these vital rates, we found that projected population growth was not significantly different between unguarded and ant-guarded plants because population growth was only weakly influenced by differences in fruit production (though strongly influenced by differences in individual growth and survival). However, simulations showed that ants could positively affect long-term plant population dynamics through services provided during rare but important events (herbivore outbreaks that reduce survival or years of high seedling recruitment associated with abundant precipitation). Thus, in this seemingly clear example of mutualism, the interaction may actually yield no clear benefit to plant population growth, or if it does, may only do so through the actions of the ants during rare events. These insights demonstrate the value of taking a demographic approach to studying the consequences of mutualism.

Defining and evaluating the umbrella species concept for conserving and restoring landscape connectivity.

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species-a bird (the umbrella), a butterfly, and a frog-inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements.

Variation in stochastic demography between and within central and peripheral regions in a widespread short-lived herb.

Analyzing intraspecific variation in population dynamics in relation to environmental factors is crucial to understand the current and future distributions of plant species. Across ranges, peripheral populations are often expected to show lower and more temporally variable vital rates than central populations, although it remains unclear how much any differences in vital rates actually contribute to differences in population growth rates. Moreover, few demographic studies accounting for environmental stochasticity have been carried out both at continental and regional scales. In this study we calculated stochastic growth rates in five central and six northern peripheral populations of the widespread shortlived herb Plantago coronopus along the Atlantic Coast in Europe. To evaluate at two spatial scales how mean values and variability of vital rates (i.e., fecundity, recruitment, survival, growth, and shrinkage) contributed to the differences in stochastic growth rates, we performed Stochastic Life Table Response Experiment (SLTRE) analyses between and within central and peripheral regions. Additionally, we searched for correlations between vital rate contributions and local environmental conditions. Lower mean values and greater variability for some vital rates in peripheral than in central populations had an overall negative but nonsignificant effect on the stochastic growth rates in the periphery. Different life cycle components accounted for differences in population growth depending on spatial scale, although recruitment was the vital rate with the highest influence both between and within regions. Interestingly, the same pattern of differentiation among populations was found within central and peripheral areas: in both regions, one group of populations displayed positive contributions of growth and shrinkage and negative contributions of recruitment and survival; the opposite pattern was found in the remaining populations. These differences in vital rate contributions among populations within regions were correlated with precipitation regime, whereas at the continental scale, differences in contribution patterns were related to temperature. Altogether, our results show how populations of P. coronopus exhibit life cycle differences that may enable the species to persist in locations with widely varying environmental conditions. This demographic flexibility may help to explain the success of widespread plants across large and heterogeneous ranges.

Ability of matrix models to explain the past and predict the future of plant populations.

Uncertainty associated with ecological forecasts has long been recognized, but forecast accuracy is rarely quantified. We evaluated how well data on 82 populations of 20 species of plants spanning 3 continents explained and predicted plant population dynamics. We parameterized stage-based matrix models with demographic data from individually marked plants and determined how well these models forecast population sizes observed at least 5 years into the future. Simple demographic models forecasted population dynamics poorly; only 40% of observed population sizes fell within our forecasts' 95% confidence limits. However, these models explained population dynamics during the years in which data were collected; observed changes in population size during the data-collection period were strongly positively correlated with population growth rate. Thus, these models are at least a sound way to quantify population status. Poor forecasts were not associated with the number of individual plants or years of data. We tested whether vital rates were density dependent and found both positive and negative density dependence. However, density dependence was not associated with forecast error. Forecast error was significantly associated with environmental differences between the data collection and forecast periods. To forecast population fates, more detailed models, such as those that project how environments are likely to change and how these changes will affect population dynamics, may be needed. Such detailed models are not always feasible. Thus, it may be wiser to make risk-averse decisions than to expect precise forecasts from models.

Seed type and origin-dependent seedling emergence patterns in Danthonia californica, a species commonly used in grassland restoration.

Danthonia californica Bolander (Poaceae)is a native perennial bunchgrass commonly used in the restoration of prairie ecosystems in the western United States. Plants of this species simultaneously produce both chasmogamous (potentially outcrossed) and cleistogamous (obligately self-fertilized) seeds. Restoration practitioners almost exclusively use chasmogamous seeds for outplanting, which are predicted to perform better in novel environments due to their greater genetic diversity. Meanwhile, cleistogamous seeds may exhibit greater local adaptation to the conditions in which the maternal plant exists. We performed a common garden experiment at two sites in the Willamette Valley, Oregon, to assess the influence of seed type and source population (eight populations from a latitudinal gradient) on seedling emergence and found no evidence of local adaptation for either seed type. Cleistogamous seeds outperformed chasmogamous seeds, regardless of whether seeds were sourced directly from the common gardens (local seeds) or other populations (nonlocal seeds). Furthermore, average seed weight had a strong positive effect on seedling emergence, despite the fact that chasmogamous seeds had significantly greater mass than cleistogamous seeds. At one common garden, we observed that seeds of both types sourced from north of our planting site performed significantly better than local or southern-sourced seeds. We also found a significant seed type and distance-dependent interaction, with cleistogamous seedling emergence peaking approximately 125 km from the garden. These results suggest that cleistogamous seeds should be considered for greater use in D. californica restoration.