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Many perennial plants show mast seeding, characterized by synchronous and highly variable reproduction across years. We propose a general model of masting, integrating proximate factors (environmental variation, weather cues, and resource budgets) with ultimate drivers (predator satiation and pollination efficiency). This general model shows how the relationships between masting and weather shape the diverse responses of species to climate warming, ranging from no change to lower interannual variation or reproductive failure. The role of environmental prediction as a masting driver is being reassessed; future studies need to estimate prediction accuracy and the benefits acquired. Since reproduction is central to plant adaptation to climate change, understanding how masting adapts to shifting environmental conditions is now a central question., Competing Interests: Declaration of interests No competing interests to declare., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

AbstractIn recent years, ecological research has become increasingly synthetic, relying on revolutionary changes in data availability and accessibility. In spite of their strengths, these approaches may cause us to overlook natural history knowledge that is not part of the digitized English-language scientific record. Here, we combine historic and modern documents to quantify species-specific nesting habitat associations of bumblebees ( Bombus spp. Latreille, 1802 Apidae). We compiled nest location data from 316 documents, of which 81 were non-English and 93 were published before 1950. We tested whether nesting traits show phylogenetic signal, examined relationships between habitat associations at different scales, and compared methodologies used to locate nests. We found no clear phylogenetic signals, but we found that nesting habitat associations were somewhat generalizable within subgenera. Landcover associations were related to nesting substrate associations; for example, surface-nesting species also tended to be associated with grasslands. Methodology was associated with nest locations; community scientists were most likely and researchers using nest boxes were least likely to report nests in human-dominated environments. These patterns were not apparent in past syntheses based only on the modern digital record. Our findings highlight the tremendous value of historic accounts for quantifying species' traits and other basic biological knowledge needed to interpret global-scale patterns.

Background: While interactions in nature are inherently local, ecological models often assume homogeneity across space, allowing for generalization across systems and greater mathematical tractability. Density-dependent disease models are a prominent example of models that assume homogeneous interactions, leading to the prediction that disease transmission will scale linearly with population density. In this study, we examined how the scale of larval butterfly movement interacts with the resource landscape to influence the relationship between larval contact and population density in the Baltimore checkerspot (Euphydryas phaeton). Our study was inspired by the recent discovery of a viral pathogen that is transmitted horizontally among Baltimore checkerspot larvae., Methods: We used multi-year larvae location data across six Baltimore checkerspot populations in the eastern U.S. to test whether larval nests are spatially clustered. We then integrated these spatial data with larval movement data in different resource contexts to investigate whether heterogeneity in spatially local interactions alters the assumed linear relationship between larval nest density and contact. We used Correlated Random Walk (CRW) models and field observations of larval movement behavior to construct Probability Distribution Functions (PDFs) of larval dispersal, and calculated the overlap in these PDFs to estimate conspecific contact within each population., Results: We found that all populations exhibited significant spatial clustering in their habitat use. Subsequent larval movement rates were influenced by encounters with host plants and larval age, and under many movement scenarios, the scale of predicted larval movement was not sufficient to allow for the "homogeneous mixing" assumed in density dependent disease models. Therefore, relationships between population density and larval contact were typically non-linear. We also found that observed use of available habitat patches led to significantly greater contact than would occur if habitat use were spatially random., Conclusions: These findings strongly suggest that incorporating larval movement and spatial variation in larval interactions is critical to modeling disease outcomes in E. phaeton. Epidemiological models that assume a linear relationship between population density and larval contact have the potential to underestimate transmission rates, especially in small populations that are already vulnerable to extinction., (© 2024. The Author(s).)

Understanding the circumstances under which insect herbivores will adopt a novel host plant is a longstanding question in basic and applied ecology. While geographic variation in host use can arise through differences in both herbivore preference and plant characteristics, there is a tendency to attribute geographic variation in host use to regional differences in herbivore preference alone. This is especially true for herbivores specialized to one or a few plant species. We compared how geographic variation in herbivore preference and host plant origin shape regional differences in host plant use by the specialized herbivore, Euphydryas phaeton. In parts of its range, E. phaeton uses only a native host, Chelone glabra, while in others, it also uses an introduced host, Plantago lanceolata. We offered female butterflies from each region the non-native host plant sourced from both regions and compared their oviposition behavior. The non-native host was almost universally rejected by butterflies in the region where only the native plant is used. In the region where butterflies use both hosts, females accepted non-native plants from their natal region twice as often as non-native plants from the other region where they are not used. Acceptance differed substantially among individual butterflies within regions but not among plants within regions. Thus, both individual preference and regional differences in both the insect and non-native host contributed to the geographic variation in different ways. These results highlight that, in addition to herbivore preference, regional differences in perceived plant suitability may be an important driver of diet breadth., (© 2023. The Author(s).)

Migratory animals follow seasonal cycles comprising linked phases often with different habitat requirements and demographic processes. Conservation of migratory species therefore must consider the full seasonal cycle to identify points limiting population viability. For western monarch butterflies, which have experienced significant declines, early spring is considered a critical period in the annual population cycle. However, records of western monarchs in early spring, when overall abundance is lowest, have historically been extremely limited. We used a community science initiative, the Western Monarch Mystery Challenge, to collect data on monarch distribution throughout the western United States between February 14th and April 22nd over 3 years. Using data from the Western Monarch Mystery Challenge and iNaturalist, we identified potential breeding habitat for western monarchs in early spring that spanned a large geographic area and several ecoregions. We observed monarchs in early spring that likely eclosed in the current year, suggesting that population expansion from overwintering sites reflects both movement and population growth. The number of records of western monarchs from early spring was higher during the Mystery Challenge (33.0/year) than earlier years (5.1/year). This study demonstrates the potential for and limitations of community science to increase our understanding of species at points in the life cycle when they are rare., Competing Interests: The authors declare no competition of interest., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Abstract: Bumblebee ( Bombus spp.) queens overwintered in artificial settings tend to have low survival rates, raising concerns that diapause may be a particularly sensitive life cycle stage for this ecologically and economically valuable group of pollinators. However, it remains unclear whether lab-based estimates of diapause survival are comparable to survival rates of natural populations. In this study, we monitored the survival of Bombus impatiens queens overwintering in the field in Ipswich, MA, and conducted a meta-analysis of studies that estimate queen diapause survival in the lab to compare our field-based estimates of survival to those of lab-based studies. We found that queen B. impatiens had relatively high rates of overwintering survival after about six months (> 60%), especially when compared to estimates of six-month survival from lab studies (< 10%). We also observed a trend that broadly corroborates many lab studies of bumblebees, in that overwinter survival of queens was related to colony origin. In addition to providing the first estimate of diapause survival for bumblebee queens in nature, our study emphasizes the need to verify patterns observed in the lab to field-based studies., Implications for Insect Conservation: Although protecting target species during sensitive life cycle stages is a fundamental goal of conservation ecology, it is first necessary to identify at what stages of the life cycle populations are most vulnerable. Our results suggest that, at least in some study systems, diapause survival of queen bumblebees in the field may be higher than suggested by lab studies., Supplementary Information: The online version contains supplementary material available at 10.1007/s10841-023-00478-8., Competing Interests: Conflict of interestThe authors declare no conflicts of interest., (© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2023, Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.)

Across taxa, the timing of life-history events (phenology) is changing in response to warming temperatures. However, little is known about drivers of variation in phenological trends among species. We analysed 168 years of museum specimen and sighting data to evaluate the patterns of phenological change in 70 species of solitary bees that varied in three ecological traits: diet breadth (generalist or specialist), seasonality (spring, summer or fall) and nesting location (above-ground or below-ground). We estimated changes in onset, median, end and duration of each bee species' annual activity (flight duration) using quantile regression. To determine whether ecological traits could explain phenological trends, we compared average trends across species groups that differed in a single trait. We expected that specialist bees would be constrained by their host plants' phenology and would show weaker phenological change than generalist species. We expected phenological advances in spring and delays in summer and fall. Lastly, we expected stronger shifts in above-ground versus below-ground nesters. Across all species, solitary bees have advanced their phenology by 0.43 days/decade. Since 1970, this advancement has increased fourfold to 1.62 days/decade. Solitary bees have also lengthened their flight period by 0.44 days/decade. Seasonality and nesting location explained variation in trends among species. Spring- and summer-active bees tended to advance their phenology, whereas fall-active bees tended to delay. Above-ground nesting species experienced stronger advances than below-ground nesting bees in spring; however, the opposite was true in summer. Diet breadth was not associated with patterns of phenological change. Our study has two key implications. First, an increasing activity period of bees across the flight season means that bee communities will potentially provide pollination services for a longer period of time during the year. And, since phenological trends in solitary bees can be explained by some ecological traits, our study provides insight into mechanisms underpinning population viability of insect pollinators in a changing world., (© 2022 The Authors. Journal of Animal Ecology © 2022 British Ecological Society.)

Floral constancy of foraging bees influences plant reproduction. Constancy as observed in nature arises from at least four distinct mechanisms frequently confounded in the literature: context-independent preferences for particular plant species, preferential visitation to the same species as the previous plant visited (simple constancy), the spatial arrangement of plants, and the relative abundances of co-flowering species. To disentangle these mechanisms, we followed individual bee flight paths within patches where all flowering plants were mapped, and we used step selection models to estimate how each mechanism influences the probability of selecting any particular plant given the available plants in a multi-species community. We found that simple constancy was positive: bees preferred to visit the same species sequentially. In addition, bees preferred to travel short distances and maintain their direction of travel between plants. After accounting for distance, we found no significant effect of site-level plant relative abundances on bee foraging choices. To explore the importance of the spatial arrangement of plants for bee foraging choices, we compared our full model containing all parameters to one with spatial arrangement removed. Due to bees' tendency to select nearby plants, combined with strong intraspecific plant clumping, spatial arrangement was responsible for about 50% of the total observed constancy. Our results suggest that floral constancy may be overestimated in studies that do not account for the spatial arrangement of plants, especially in systems with intraspecific plant clumping. Plant spatial patterns at within-site scales are important for pollinator foraging behavior and pollination success., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

In the western United States, the population of migratory monarch butterflies is on the brink of collapse, having dropped from several million butterflies in the 1980s to ~2000 butterflies in the winter of 2020-2021. At the same time, a resident (non-migratory) monarch butterfly population in urban gardens has been growing in abundance. The new resident population is not sufficient to make up for the loss of the migratory population; there are still orders of magnitude fewer butterflies now than in the recent past. The resident population also probably lacks the demographic capacity to expand its range inland during summer months. Nonetheless, the resident population may have the capacity to persist. This sudden change emphasises the extent to which environmental change can have unexpected consequences, and how quickly these changes can happen. We hope it will provoke discussion about how we define resilience and viability in changing environments., (© 2021 John Wiley & Sons Ltd.)

Life history theories analyze and predict variation in vital rates, such as survival and reproduction, based on age. The age-from-stage method to derive age-specific vital rates from stage data was developed because age-specific data are rarely obtained for plants. Age-specific vital rates derived by this method might underestimate effects of age on vital rates, because the models assume that vital rates do not vary within stage classes. Consequently, population models and life history summaries relying on these vital rates could be biased against detecting senescence. Here, we perform a comparative study of methods to estimate age-specific vital rates using monitoring data with known age and stage. We derived age-, stage-, and age-and-stage-specific vital rates with demographic data from a long-lived perennial, Silene spaldingii. Then, we derived three age-specific population matrix models (age, age-from-stage, and age-and-stage). For each model, we derived life history summaries commonly used in ecology: population growth rate, net reproductive value, relative reproductive values, stable age distribution, generation time, and sensitivity and elasticity of population growth rate. Many vital rates depended on both age and stage in S. spaldingii. However, this species does not senesce; in fact, the number of flowers increased with age. As expected, the age-from-stage method was not able to accurately recreate the age dependence in some life history summaries, such as relative reproductive value. The age-from-stage model suggested faster reproductive dynamics in S. spaldingii than the models based on known age, i.e., plants started to reproduce earlier, and fertility remained constant thereafter, which may lead to biased predictions about evolutionary consequences of age-dependent life history traits. However, population growth rate, generation time, and net reproductive rate did not differ significantly among the models. Our study demonstrated that some metrics are robust to imprecision in model structure, while others are more sensitive. In spite of these biases, this case study provides another example of the diversity of aging patterns in plants. Age can be essential information when studying senescence in plants, but demographic metrics that were not about age per se were similar across model structures., (© 2021 by the Ecological Society of America.)

Ecologists often collect data with the aim of determining which of many variables are associated with a particular cause or consequence. Unsupervised analyses (e.g. principal components analysis, PCA) summarize variation in the data, without regard to the response. Supervised analyses (e.g., partial least squares, PLS) evaluate the variables to find the combination that best explain a causal relationship. These approaches are not interchangeable, especially when the variables most responsible for a causal relationship are not the greatest source of overall variation in the data-a situation that ecologists are likely to encounter. To illustrate the differences between unsupervised and supervised techniques, we analyze a published dataset using both PCA and PLS and compare the questions and answers associated with each method. We also use simulated datasets representing situations that further illustrate differences between unsupervised and supervised analyses. For simulated data with many correlated variables that were unrelated to the response, PLS was better than PCA at identifying which variables were associated with the response. There are many applications for both unsupervised and supervised approaches in ecology. However, PCA is currently overused, at least in part because supervised approaches, such as PLS, are less familiar.

Behavior and organization of social groups is thought to be vital to the functioning of societies, yet the contributions of various roles within social groups toward population growth and dynamics have been difficult to quantify. A common approach to quantifying these role-based contributions is evaluating the number of individuals conducting certain roles, which ignores how behavior might scale up to effects at the population-level. Manipulative experiments are another common approach to determine population-level effects, but they often ignore potential feedbacks associated with these various roles.Here, we evaluate the effects of worker size distribution in bumblebee colonies on worker production in 24 observational colonies across three environments, using functional linear models. Functional linear models are an underused correlative technique that has been used to assess lag effects of environmental drivers on plant performance. We demonstrate potential applications of this technique for exploring high-dimensional ecological systems, such as the contributions of individuals with different traits to colony dynamics.We found that more larger workers had mostly positive effects and more smaller workers had negative effects on worker production. Most of these effects were only detected under low or fluctuating resource environments suggesting that the advantage of colonies with larger-bodied workers becomes more apparent under stressful conditions.We also demonstrate the wider ecological application of functional linear models. We highlight the advantages and limitations when considering these models, and how they are a valuable complement to many of these performance-based and manipulative experiments., Competing Interests: We have no conflicts of interest to declare., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Phenological shifts are well-documented in the ecological literature. However, their significance for changes in demography and abundance is less clear. We used 27 years of citizen science monitoring to quantify trends in phenology and relative abundance across 89 butterfly species. We calculated shifts in phenology using quantile regression and shifts in relative abundance using list length analysis and counts from field trips. Elongated activity periods within a year were the strongest predictor of increases in relative abundance. These changes may be driven in part by changes in voltinism, as this association was stronger in multivoltine species. Some species appear to be adding a late-season generation, whereas other species appear to be adding a spring generation, revealing a possible shift from vagrant to resident. Our results emphasise the importance of evaluating phenological changes throughout species' flight period and understanding the consequences for such climate-related changes on viability or population dynamics., (© 2020 John Wiley & Sons Ltd.)

Background and Aims: In a range of plant species, the distribution of individual mean fecundity is skewed and dominated by a few highly fecund individuals. Larger plants produce greater seed crops, but the exact nature of the relationship between size and reproductive patterns is poorly understood. This is especially clear in plants that reproduce by exhibiting synchronized quasi-periodic variation in fruit production, a process called masting., Methods: We investigated covariation of plant size and fecundity with individual-plant-level masting patterns and seed predation in 12 mast-seeding species: Pinus pinea, Astragalus scaphoides, Sorbus aucuparia, Quercus ilex, Q. humilis, Q. rubra, Q. alba, Q. montana, Chionochloa pallens, C. macra, Celmisia lyallii and Phormium tenax., Key Results: Fecundity was non-linearly related to masting patterns. Small and unproductive plants frequently failed to produce any seeds, which elevated their annual variation and decreased synchrony. Above a low fecundity threshold, plants had similar variability and synchrony, regardless of their size and productivity., Conclusions: Our study shows that within-species variation in masting patterns is correlated with variation in fecundity, which in turn is related to plant size. Low synchrony of low-fertility plants shows that the failure years were idiosyncratic to each small plant, which in turn implies that the small plants fail to reproduce because of plant-specific factors (e.g. internal resource limits). Thus, the behaviour of these sub-producers is apparently the result of trade-offs in resource allocation and environmental limits with which the small plants cannot cope. Plant size and especially fecundity and propensity for mast failure years play a major role in determining the variability and synchrony of reproduction in plants., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

A rapidly changing climate has the potential to interfere with the timing of environmental cues that ectothermic organisms rely on to initiate and regulate life history events. Short-lived ectotherms that exhibit plasticity in their life history could increase the number of generations per year under warming climate. If many individuals successfully complete an additional generation, the population experiences an additional opportunity to grow, and a warming climate could lead to a demographic bonanza. However, these plastic responses could become maladaptive in temperate regions, where a warmer climate could trigger a developmental pathway that cannot be completed within the growing season, referred to as a developmental trap. Here we incorporated detailed demography into commonly used photothermal models to evaluate these demographic consequences of phenological shifts due to a warming climate on the formerly widespread, multivoltine butterfly (Pieris oleracea). Using species-specific temperature- and photoperiod-sensitive vital rates, we estimated the number of generations per year and population growth rate over the set of climate conditions experienced during the past 38 years. We predicted that populations in the southern portion of its range have added a fourth generation in recent years, resulting in higher annual population growth rates (demographic bonanzas). We predicted that populations in the Northeast United States have experienced developmental traps, where increases in the thermal window initially caused mortality of the final generation and reduced growth rates. These populations may recover if more growing degree days are added to the year. Our framework for incorporating detailed demography into commonly used photothermal models demonstrates the importance of using both demography and phenology to predict consequences of phenological shifts., (© 2019 John Wiley & Sons Ltd.)

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Understanding the factors associated with declines of at-risk species is an important first step in setting management and recovery targets. This step can be challenging when multiple aspects of climate and land use are changing simultaneously, and any or all could be contributing to population declines. We analyzed population trends of monarch butterflies in western North America in relation to likely environmental drivers. Unlike the larger eastern monarch population, past analyses of western monarchs have only evaluated the importance of climate (i.e., not land use) factors as drivers of abundance. We used partial least squares regression (PLSR) to evaluate the potential importance of changes in land use and climate variables. Trends in western monarch abundance were more strongly associated with land use variables than climate variables. Conclusions about importance of climate and land use variables were robust to changes in PLSR model structure. However, individual variables were too collinear to unambiguously separate their effects. We compared these conclusions to the more widely used technique of multiple regression, followed by multi-model inference (MRMI). Naïve interpretation of MRMI results could be misleading, if collinearity were not taken into account. MRMI was also highly sensitive to variation in model construction. Our results suggest a two-pronged approach to monarch conservation, specifically, starting efforts now to restore habitat, while also using experiments to more clearly delineate separate effects of climate and land use factors. They also demonstrate the utility of PLSR, a technique that is growing in use but is still relatively under-appreciated in conservation biology., (© 2019 by the Ecological Society of America.)

Synchronized and variable reproduction by perennial plants, called mast seeding, is a major reproductive strategy of trees. The need to accumulate sufficient resources after depletion following fruiting (resource budget), the efficiency of mass flowering for outcross pollination (pollen coupling), or the external factors preventing reproduction (environmental veto) could all synchronize masting. We used seed production data for four species ( Quercus ilex , Quercus humilis , Sorbus aucuparia , and Pinus albicaulis ) to parametrize resource budget models of masting. Based on species life-history characteristics, we hypothesized that pollen coupling should synchronize reproduction in S. aucuparia and P. albicaulis , while in Q. ilex and Q. humilis , environmental veto should be a major factor. Pollen coupling was stronger in S. aucuparia and P. albicaulis than in oaks, while veto was more frequent in the latter. Yet in all species, costs of reproduction were too small to impose a replenishment period. A synchronous environmental veto, in the presence of environmental stochasticity, was sufficient to produce observed variability and synchrony in reproduction. In the past, vetoes like frost events that prevent reproduction have been perceived as negative for plants. In fact, they could be selectively favored as a way to create mast seeding.

Ecologists often assume that range expansion will be fastest in landscapes composed entirely of the highest-quality habitat. Theoretical models, however, show that range expansion depends on both habitat quality and habitat-specific movement rates. Using data from 78 species in 70 studies, we find that animals typically have faster movement through lower-quality environments (73% of published cases). Therefore, if we want to manage landscapes for range expansion, there is a trade-off between promoting movement with nonhostile matrix, and promoting population growth with high-quality habitat. We illustrate how this trade-off plays out with the use of an exemplar species, the Baltimore checkerspot butterfly. For this species, we calculate that the expected rate of range expansion is fastest in landscapes with ~15% high-quality habitat. Behavioral responses to nonhabitat matrix have often been documented in animal populations, but rarely included in empirical predictions of range expansion. Considering movement behavior could change land-planning priorities from focus on high-quality habitat only to integrating high- and low-quality land cover types, and evaluating the costs and benefits of different matrix land covers for range expansion., (© 2019 by the Ecological Society of America.)

The number of insect species at risk of population decline and extinction is increasing rapidly. Yet we know almost nothing about the ecology of these species, except for at-risk butterflies. A growing body of literature shows how butterfly vital rates, including demography and movement, are essential for guiding conservation and recovery. History has shown us that without these data, conservation decisions often weaken, rather than enhance, population viability. This is especially true in changing landscapes. We review knowledge of vital rates across all at-risk butterflies. We have information on movement for 17 of 283 butterfly species and information on demography for 19 species. We find that habitat-specific movement behavior is key to understanding how to connect populations, and habitat-specific demography is central to managing habitats. Methods and analyses worked out for butterflies can provide a scaffold around which to build studies for the conservation of other at-risk insects.

Variable, synchronized seed production, called masting, is a widespread reproductive strategy in plants. Resource dynamics, pollination success, and, as described here, environmental veto are possible proximate mechanisms driving masting. We explored the environmental veto hypothesis, which assumes that reproductive synchrony is driven by external factors preventing reproduction in some years, by extending the resource budget model of masting with correlated reproductive failure. We ran this model across its parameter space to explore how key parameters interact to drive seeding dynamics. Next, we parameterized the model based on 16 yr of seed production data for populations of red (Quercus rubra) and white (Quercus alba) oaks. We used these empirical models to simulate seeding dynamics, and compared simulated time series with patterns observed in the field. Simulations showed that resource dynamics and reproduction failure can produce masting even in the absence of pollen coupling. In concordance with this, in both oaks, among-year variation in resource gain and correlated reproductive failure were necessary and sufficient to reproduce masting, whereas pollen coupling, although present, was not necessary. Reproductive failure caused by environmental veto may drive large-scale synchronization without density-dependent pollen limitation. Reproduction-inhibiting weather events are prevalent in ecosystems, making described mechanisms likely to operate in many systems., (© 2018 The Authors New Phytologist © 2018 New Phytologist Trust.)

Temperature and precipitation determine the conditions where plant species can occur. Despite their significance, to date, surprisingly few demographic field studies have considered the effects of abiotic drivers. This is problematic because anticipating the effect of global climate change on plant population viability requires understanding how weather variables affect population dynamics. One possible reason for omitting the effect of weather variables in demographic studies is the difficulty in detecting tight associations between vital rates and environmental drivers. In this paper, we applied Functional Linear Models (FLMs) to long-term demographic data of the perennial wildflower, Astragalus scaphoides, and explored sensitivity of the results to reduced amounts of data. We compared models of the effect of average temperature, total precipitation, or an integrated measure of drought intensity (standardized precipitation evapotranspiration index, SPEI), on plant vital rates. We found that transitions to flowering and recruitment in year t were highest if winter/spring of year t was wet (positive effect of SPEI). Counterintuitively, if the preceding spring of year t - 1 was wet, flowering probabilities were decreased (negative effect of SPEI). Survival of vegetative plants from t - 1 to t was also negatively affected by wet weather in the spring of year t - 1 and, for large plants, even wet weather in the spring of t - 2 had a negative effect. We assessed the integrated effect of all vital rates on life history performance by fitting FLMs to the asymptotic growth rate, log(λt). Log(λt) was highest if dry conditions in year t - 1 were followed by wet conditions in the year t. Overall, the positive effects of wet years exceeded their negative effects, suggesting that increasing frequency of drought conditions would reduce population viability of A. scaphoides. The drought signal weakened when reducing the number of monitoring years. Substituting space for time did not recover the weather signal, probably because the weather variables varied little between sites. We detected the SPEI signal when the analysis included data from two sites monitored over 20 yr (2 × 20 observations), but not when analyzing data from four sites monitored over 10 yr (4 × 10 observations)., (© 2018 by the Ecological Society of America.)

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., (© 2017 by the Ecological Society of America.)

Diffusion, which approximates a correlated random walk, has been used by ecologists to describe movement, and forms the basis for many theoretical models. However, it is often criticized as too simple a model to describe animal movement in real populations. We test a key prediction of diffusion models, namely, that animals should be more abundant in land cover classes through which they move more slowly. This relationship between density and diffusion has rarely been tested across multiple species within a given landscape. We estimated diffusion rates and corresponding densities of 25 Israeli butterfly species from flight path data and visual surveys. The data were collected across 19 sites in heterogeneous landscapes with four land cover classes: semi-natural habitat, olive groves, wheat fields and field margins. As expected from theory, species tended to have higher densities in land cover classes through which they moved more slowly and lower densities in land cover classes through which they moved more quickly. Two components of movement (move length and turning angle) were not associated with density, nor was expected net squared displacement. Move time, however, was associated with density, and animals spent more time per move step in areas with higher density. The broad association we document between movement behaviour and density suggests that diffusion is a good first approximation of movement in butterflies. Moreover, our analyses demonstrate that dispersal is not a species-invariant trait, but rather one that depends on landscape context. Thus, land cover classes with high diffusion rates are likely to have low densities and be effective conduits for movement., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

The autocatalytic maturation of the chromophore in green fluorescent protein (GFP) was thought to require the precise positioning of the side chains surrounding it in the core of the protein, many of which are strongly conserved among homologous fluorescent proteins. In this study, we screened for green fluorescence in an exhaustive set of point mutations of seven residues that make up the chromophore microenvironment, excluding R96 and E222 because mutations at these positions have been previously characterized. Contrary to expectations, nearly all amino acids were tolerated at all seven positions. Only four point mutations knocked out fluorescence entirely. However, chromophore maturation was found to be slower and/or fluorescence reduced in several cases. Selected combinations of mutations showed nonadditive effects, including cooperativity and rescue. The results provide guidelines for the computational engineering of GFPs.

Introduced plants can positively affect population viability by augmenting the diet of native herbivores, but can negatively affect populations if they are subpar or toxic resources. In organisms with complex life histories, such as insects specializing on host plants, the impacts of a novel host may differ across life stages, with divergent effects on population persistence. Most research on effects of novel hosts has focused on adult oviposition preference and larval performance, but adult preference may not optimize offspring performance, nor be indicative of host quality from a demographic perspective. We compared population growth rates of the Baltimore checkerspot butterfly, Euphydryas phaeton, on an introduced host, Plantago lanceolata (English plantain), and the native host Chelone glabra (white turtlehead). Contrary to the previous findings suggesting that P. lanceolata could be a population sink, we found higher population growth rates (λ) on the introduced than the native host, even though some component parameters of λ were higher on the native host. Our findings illustrate the importance of moving beyond preference-performance studies to integrate vital rates across all life stages for evaluating herbivore-host plant relationships. Single measures of preference or performance are not sufficient proxies for overall host quality nor do they provide insights into longer term consequences of novel host plant use. In our system, in particular, P. lanceolata may buffer checkerspot populations when the native host is limiting, but high growth rates could lead to crashes over longer time scales.

Home range size generally decreases with increasing population density, but testing how this relationship is influenced by other factors (e.g., food availability, kin structure) is a difficult task. We used spatially explicit capture-recapture models to examine how home range size varies with population density in the yellow-necked mouse ( Apodemus flavicollis ). The relationship between population density and home range size was studied at two distinct phases of population fluctuations induced by beech ( Fagus sylvatica ) masting: post-mast peak in abundance (first summer after mast, n  = 2) and subsequent crash (second summer after mast, n  = 2). We live-trapped mice from June to September to avoid the confounding effects of autumn seedfall on home range size. In accordance with general predictions, we found that home range size was negatively associated with population density. However, after controlling for the effect of density, home ranges of mice were larger in post-mast years than during the crash phase. This indicates a higher spatial overlap among neighbors in post-mast years. We suggest that the increased spatial overlap is caused by negative density-dependent dispersal that leads to high relatedness of individuals within population in the peak phase of the cycle.

The predator satiation and predator dispersal hypotheses provide alternative explanations for masting. Both assume satiation of seed-eating vertebrates. They differ in whether satiation occurs before or after seed removal and caching by granivores (predator satiation and predator dispersal, respectively). This difference is largely unrecognized, but it is demographically important because cached seeds are dispersed and often have a microsite advantage over nondispersed seeds. We conducted rodent exclosure experiments in two mast and two nonmast years to test predictions of the predator dispersal hypothesis in our study system of yellow-necked mice (Apodemus flavicollis) and European beech (Fagus sylvatica). Specifically, we tested whether the fraction of seeds removed from the forest floor is similar during mast and nonmast years (i.e., lack of satiation before seed caching), whether masting decreases the removal of cached seeds (i.e., satiation after seed storage), and whether seed caching increases the probability of seedling emergence. We found that masting did not result in satiation at the seed removal stage. However, masting decreased the removal of cached seeds, and seed caching dramatically increased the probability of seedling emergence relative to noncached seeds. European beech thus benefits from masting through the satiation of scatterhoarders that occurs only after seeds are removed and cached. Although these findings do not exclude other evolutionary advantages of beech masting, they indicate that fitness benefits of masting extend beyond the most commonly considered advantages of predator satiation and increased pollination efficiency.

Determining the minimum area required to sustain populations has a long history in theoretical and conservation biology. Correlative approaches are often used to estimate minimum area requirements (MARs) based on relationships between area and the population size required for persistence or between species' traits and distribution patterns across landscapes. Mechanistic approaches to estimating MAR facilitate prediction across space and time but are few. We used a mechanistic MAR model to determine the critical minimum patch size (CMP) for the Baltimore checkerspot butterfly (Euphydryas phaeton), a locally abundant species in decline along its southern range, and sister to several federally listed species. Our CMP is based on principles of diffusion, where individuals in smaller patches encounter edges and leave with higher probability than those in larger patches, potentially before reproducing. We estimated a CMP for the Baltimore checkerspot of 0.7-1.5 ha, in accordance with trait-based MAR estimates. The diffusion rate on which we based this CMP was broadly similar when estimated at the landscape scale (comparing flight path vs. capture-mark-recapture data), and the estimated population growth rate was consistent with observed site trends. Our mechanistic approach to estimating MAR is appropriate for species whose movement follows a correlated random walk and may be useful where landscape-scale distributions are difficult to assess, but demographic and movement data are obtainable from a single site or the literature. Just as simple estimates of lambda are often used to assess population viability, the principles of diffusion and CMP could provide a starting place for estimating MAR for conservation., (© 2015 Society for Conservation Biology.)

Individual variation in movement can have important consequences for spatial population dynamics. For instance, individual variation increases leptokurtosis in dispersal distance, such that more individuals move very short and very long distances relative to a homogeneous population. We quantified individual variation in movement of the Baltimore checkerspot butterfly (Euphydryas phaeton) to investigate its importance for two conservation-related metrics: the expected dispersal distance and the critical minimum patch size, or the smallest area within which a population can persist based on loss due to emigration. All movement parameters showed among-individual variation, with the greatest variation in move lengths and time spent resting. Correlations in among-individual movement parameters indicated that some butterflies were generally more mobile than others. We incorporated empirically estimated movement and demographic parameters into two individual-based models (IBMs), one with homogeneity in movement among individuals, and one with heterogeneity in movement. As expected, individual variation in movement increased the leptokurtosis of lifetime movement distance; the maximum difference in distance moved was substantial (-850 m vs. -5800 m) and is likely of significance for conservation. Individual variation also affected the critical minimum patch size, but the difference (-0.7 ha vs. -0.5 ha) is unlikely to be ecologically significant. Notably, populations with individual variation had higher growth rates in small patches and lower growth rates in large patches, a logical consequence of increased leptokurtosis. Individual variation in movement is fairly straightforward to quantify using mixed effects models and is important for spatial population dynamics, thus we encourage its inclusion in studies of other systems.

Masting, the highly variable and synchronous production of seeds across a population of perennial plants, is an ecologically important, but still poorly understood, phenomenon. While much is known about the fitness benefits of masting and its effects on seed consumers and trophic interactions, less is understood about the proximate mechanisms of masting. The resource budget model (RBM) posits that masting requires more resources than plants can gain in a single year. Individual plants store resources until a threshold is reached and then produce seeds, which depletes resources so that plants cannot reproduce again for 2 or more years. Individuals are synchronized by pollen coupling or environmental forcing. We review the assumptions of these models and assess the extent to which they are consistent with general patterns in plant populations. We discuss the implications of the RBM for how plants respond to changes in the external environment. Overall, the RBM is a likely cause of synchrony in many, but not all, masting species. This mechanistic hypothesis also leads to specific, but not always intuitive, expectations about how plant resources affect mast seeding., (© 2014 New York Academy of Sciences.)

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., (© 2013 Society for Conservation Biology.)

Pulsed food resources lead to mismatches between distribution of consumers and resources in space and time. Many studies have investigated how pollinators and floral resources covary in space, but few have looked at their covariance among years. I studied responses of two bee taxa, Bombus (a social genus) and Anthophora (a solitary genus), to variation in flowering by Astragalus scaphoides, a perennial herb that flowers in alternate years. First, I quantified the rate at which individual plants were visited by bees. Anthophora showed evidence of a demographic response to resource pulses--that is, more individuals were seen in the year after a high-flowering year--whereas Bombus did not. Second, I quantified pollinator behavior by following individual bees and recording the proportion of visits to A. scaphoides within single foraging bouts. The proportion of visits to A. scaphoides by both taxa increased with A. scaphoides's flowering density. Higher specialization in high-flowering years likely makes both taxa better pollinators in high-flowering years. If these taxa differ in effectiveness as pollinators, then these responses translate into variation in pollination services in space and time, specifically, more activity by Bombus in high-flowering years and more by Anthophora in years following high-flowering years. They also emphasize that pollinator activity depends in part on past-as well as current-floral resources.

Most populations exist in variable environments. Two sets of theory have been developed to address this variability. Stochastic dynamics focus on variation in population growth rates based on random differences in vital rates such as growth, survival, and reproduction. Transient dynamics focus on short-term, deterministic responses to changes in the stage distribution of individuals. These processes are related: demographic variation shifts stage structures, producing transient responses, which then contribute to the overall variability of population growth rate. The relative contributions of vital rates vs. transient responses to stochastic dynamics, and the implications for transient analyses, are unclear. This study explores the role of transient responses in stochastic dynamics of nine perennial plant species. Across the species, transient responses contributed more on average to variability in annual population growth rates than did variation in vital rates alone. Transient potential of an average matrix was indicative of the contribution of transient dynamics, although these metrics varied greatly across years. Transient responses were often in the opposite direction as demographic variation, suggesting that transient dynamics may at times have a buffering effect on populations. Overall, transient dynamics had an important role in modulating environmental variation, with implications for both processes in understanding stochastic dynamics.

• In masting trees, synchronized, heavy reproductive events are thought to deplete stored resources and to impose a replenishment period before subsequent masting. However, direct evidence of resource depletion in wild, masting trees is very rare. Here, we examined the timing and magnitude (local vs individual-level) of stored nutrient depletion after a heavy mast event in Pinus albicaulis. • In 2005, the mast year, we compared seasonal changes in leaf and sapwood nitrogen (N) and phosphorus (P) concentrations and leaf photosynthetic rates in cone-bearing branches, branches that never produced cones, and branches with experimentally removed cones. We also compared nutrient concentrations in cone branches and branches that had never had cones between 2005 and 2006, and measured tree ring width and new shoot growth during 2005. • During the mast year, N or P depletion occurred only in tissue fractions of reproductive branches, where photosynthetic rates were reduced. However, by the end of the following year, nutrients were depleted in all branches, indicating individual-level resource depletion. New shoot and radial growth were not affected by masting. • We provide direct evidence that mast events in wild trees deplete stored nutrients. Our results highlight the importance of evaluating reproductive costs over time and at the individual level., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

1. Two aspects of landscape composition shape the behavioural response of animals to habitat heterogeneity: physical habitat structure and abundance of key resources. In general, within-habitat movement behaviour has been investigated in relation to resources, and preference at boundaries has been quantified in response to physical structure. 2. Habitat preference studies suggest that responses to resources vs. structure should differ, e.g. between male and female animals, and effects of responses to structure and resources may also interact. However, most studies of animal movement combine various aspects of behavioural responses to 'habitat', implicitly assuming that resources and structure are broadly equivalent. 3. We conducted a large-scale experiment of the movement of Fender's blue (Icaricia icarioides fenderi), an endangered butterfly, to investigate butterfly response to physical structure of the landscape (prairie, open woods and dense woods) and to resources [presence or absence of Kincaid's lupine, Lupinus oreganus (larval hostplant patches)]. The experiment included 606 butterfly flight paths across four habitat types and nine ecotones. 4. Responses to physical structure and resource patches were not congruent. Butterflies were attracted to resource patches within both prairies and open woods and moved more slowly when in resource patches. Butterflies tended to prefer prairie at prairie-forest edges but tended to move faster in prairies than in open woods. Physical structure and resources also interacted; butterflies did not respond to physical habitat structure when resource patches spanned prairie - open woods ecotones. 5. Even dense woods were not perfect barriers, in contrast to a large body of literature that assumes insects from open habitats will not enter dense forests. 6. Movement of both males and females responded to resources and structure. However, female butterflies had stronger responses to both resources and structure in most cases. Females had strongest response to resource (hostplant) patches at patch edges, whereas the strongest preference of males was to return to prairie from open forest. 7. If other species behave like Fender's blue, then combining different definitions of 'habitat' (physical structure vs. resources), different aspects of movement (edge preference vs. within-habitat movement) and/or males and females within species could all lead to misleading conclusions. Our results highlight the importance of investigating these responses, and our study provides a framework for separating them in other systems., (© 2012 The Authors. Journal of Animal Ecology © 2012 British Ecological Society.)

During the growing season, some individuals in perennial plant populations may remain alive belowground while others emerge. This phenomenon, known as prolonged dormancy, seems maladaptive, because prolonged dormancy delays growth and reproduction. However, prolonged dormancy may offer the benefit of safety while belowground, leading to the hypothesis that prolonged dormancy is a bet-hedging strategy. We evaluated this hypothesis using a 25-year demographic study of Astragalus scaphoides, an iteroparous perennial plant. First, we determined the relationship between prolonged dormancy and fitness using data from individuals in our population. This analysis showed that prolonged dormancy decreased arithmetic mean fitness and reduced variance in fitness. Geometric mean fitness was maximized at intermediate levels of prolonged dormancy. Empirical patterns of lifetime reproductive success confirm this relationship. We also compared fitness of plants in our population to hypothetical plants without prolonged dormancy, which generally revealed benefits of prolonged dormancy, even if plants could forgo prolonged dormancy without costs to other vital rates. Therefore, prolonged dormancy may indeed function as a bet-hedging strategy, but the benefits of remaining belowground outweigh the costs only for a subset of individuals. Bet hedging has been demonstrated in plants with simple life histories, such as annuals and monocarpic perennials; we present evidence that bet hedging may be important for plants with more complex life histories.

In plants, prolonged dormancy is often considered a response to resource depletion or environmental stress that comes at a fitness cost. However, apparent costs of dormancy could reflect the state in which plants entered dormancy, rather than effects of dormancy per se. We tested this hypothesis for a terrestrial orchid, Epipactis atrorubens, by analyzing differences in vital rates of dormant and emergent plants using generalized linear mixed models, applied to eight years of demographic data. Dormant E. atrorubens plants did not form one homogeneous stage class. Instead, the vital rates of dormant plants mirrored performance of plants in their life stage before dormancy. Plants emerging from dormancy were slightly (albeit only marginally statistically significantly) larger than plants transitioning from the matching aboveground stage class, especially for smaller and younger stage classes. Because small plants were most likely to go dormant, plants emerging from dormancy were also smaller than average, if one were to compare all previously dormant plants to all previously emergent plants. Therefore, misclassifying all dormant plants into a single stage class changes whether we view dormancy as intrinsically costly, in terms of future performance upon emergence. We suggest that prolonged dormancy may be a form of phenotypic plasticity in which plants distribute their performance and reproductive effort through time, rather than a simple stress response.

Matrix projection models are among the most widely used tools in plant ecology. However, the way in which plant ecologists use and interpret these models differs from the way in which they are presented in the broader academic literature. In contrast to calls from earlier reviews, most studies of plant populations are based on < 5 matrices and present simple metrics such as deterministic population growth rates. However, plant ecologists also cautioned against literal interpretation of model predictions. Although academic studies have emphasized testing quantitative model predictions, such forecasts are not the way in which plant ecologists find matrix models to be most useful. Improving forecasting ability would necessitate increased model complexity and longer studies. Therefore, in addition to longer term studies with better links to environmental drivers, priorities for research include critically evaluating relative/comparative uses of matrix models and asking how we can use many short-term studies to understand long-term population dynamics., (© 2010 Blackwell Publishing Ltd/CNRS.)

Prolonged dormancy is a life-history stage in which mature plants fail to resprout for one or more growing seasons and instead remain alive belowground. Prolonged dormancy is relatively common, but the proximate causes and consequences of this intriguing strategy have remained elusive. In this study we tested whether stored resources are associated with remaining belowground, and investigated the resource costs of remaining belowground during the growing season. We measured stored resources at the beginning and end of the growing season in Astragalus scaphoides, an herbaceous perennial in southwest Montana, USA. At the beginning of the growing season, dormant plants had lower concentrations of stored mobile carbon (nonstructural carbohydrates, NSC) than did emergent plants. Surprisingly, during the growing season, dormant plants gained as much NSC as photosynthetically active plants, an increase most likely due to remobilization of structural carbon. Thus, low levels of stored NSC were associated with remaining belowground, and remobilization of structural carbon may allow for dormant plants to emerge in later seasons. The dynamics of NSC in relation to dormancy highlights the ability of plants to change their own resource status somewhat independently of resource assimilation, as well as the importance of considering stored resources in understanding plant responses to the environment.

Explaining variation in population growth rates is fundamental to predicting population dynamics and population responses to environmental change. In this study, we used matrix population models, which link birth, growth and survival to population growth rate, to examine how and why population growth rates vary within and among 50 terrestrial plant species. Population growth rates were more similar within species than among species; with phylogeny having a minimal influence on among-species variation. Most population growth rates decreased over the observation period and were negatively autocorrelated between years; that is, higher than average population growth rates tended to be followed by lower than average population growth rates. Population growth rates varied more through time than space; this temporal variation was due mostly to variation in post-seedling survival and for a subset of species was partly explained by response to environmental factors, such as fire and herbivory. Stochastic population growth rates departed from mean matrix population growth rate for temporally autocorrelated environments. Our findings indicate that demographic data and models of closely related plant species cannot necessarily be used to make recommendations for conservation or control, and that post-seedling survival and the sequence of environmental conditions are critical for determining plant population growth rate.

In fire-adapted ecosystems, fire is presumed to be the dominant ecological force, and little is known about how consumer interactions influence forest regeneration. Here, we investigated seed predation by deer mice (Peromyscus maniculatus) and its effects on recruitment of ponderosa pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) seedlings in unburned and recently burned fire-adapted montane forests in west-central Montana, USA. Deer mice were almost twice as abundant in burned than unburned stands. Deer mouse removal of seeds from petri dishes was two times higher in burned than in unburned stands, and seed removal levels were 8% higher for ponderosa pine than for the smaller Douglas-fir seeds. In seed-addition experiments, emergence of seedlings in deer mouse-exclusion cages was almost six times higher in burned compared to unburned forest. In both burned and unburned forest, emergence was lower for ponderosa pine than for Douglas-fir. Seedling survival to establishment did not differ between conifer species but was considerably higher in burned than in unburned forest. However, effects of seed predation on recruitment prevailed over fire effects: in cages allowing access by deer mice, emergence and establishment were extremely rare for both conifer species in both burned and unburned forest. This research suggests that consumer interactions can substantially influence recruitment even in fire-adapted forest ecosystems.

Mast-seeding is the synchronous and periodic reproduction by plant populations. This phenomenon has been widely studied from a community-level perspective, but we know extremely little about how plants are able to synchronize reproduction. Here, we present the first experimental test of proximate mechanisms of mast-seeding, by preventing reproduction in an iteroparous, mast-seeding wildflower. Through a series of experiments, we show that mobile carbohydrate stores (NSC) control alternate-year flowering by individual plants; seed set depletes NSC which prevents flowering the following year. Plants are synchronized by density-dependent pollen limitation; when plants flower asynchronously, they set fewer seeds, which prevents NSC depletion. Therefore, these individual plants flower in subsequent years and become synchronized. Because mast-seeding is a consequence of physiological controls of reproduction, differences in plant resource acquisition and allocation could dramatically change patterns of seed production, and changes in plant consumers and pollinators could change selection on physiological and developmental pathways.

Ecological restoration is a key component of biological conservation. Nevertheless, unlike protection of existing areas, restoration changes existing land use and can therefore be more controversial. Some restoration projects negatively affect surrounding landowners, creating social constraints to restoration success. Just as negative off-site impacts (i.e., negative externalities) flow from industrial areas to natural areas, restoration projects can generate negative externalities for commercial land uses, such as agriculture. Negative externalities from industry have led to government regulation to prevent human health and environmental impacts. Negative externalities from restoration projects have elicited similar legal constraint on at least one large-scale conservation project, riparian restoration in the Sacramento River Conservation Area. The negative externalities of restoration that are perceived to be the direct result of specific goals, such as endangered species management, are likely to be more contentious than externalities arising from unintended phenomena such as weed invasion. Restoration planners should give equal consideration to off-site characteristics as to on-site characteristics when choosing sites for restoration and designing projects. Efforts to control externalities can lead to off-site ecological benefits.