Novel host plant use by a specialist insect depends on geographic variation in both the host and herbivore species.

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.

Ecological traits explain long-term phenological trends in solitary bees.

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.

Global gene flow releases invasive plants from environmental constraints on genetic diversity.

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.

The contribution of plant spatial arrangement to bumble bee flower constancy.

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.

Resilience or Catastrophe? A possible state change for monarch butterflies in western North America.

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.

Changes in flight period predict trends in abundance of Massachusetts butterflies.

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.

Phenotypic plasticity masks range-wide genetic differentiation for vegetative but not reproductive traits in a short-lived plant.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

Using the right tool for the job: the difference between unsupervised and supervised analyses of multivariate ecological data.

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.

Flowering synchrony drives reproductive success in a wind-pollinated tree.

Synchronised and quasi-periodic production of seeds by plant populations, known as masting, is implicated in many ecological processes, but how it arises remains poorly understood. Flowering and pollination dynamics are hypothesised to provide the mechanistic link for the observed relationship between weather and population-level seed production. We report the first experimental test of the phenological synchrony hypotheses as a driver of pollen limitation in mast seeding oaks (Quercus ilex). Higher flowering synchrony yielded greater pollination efficiency, which resulted in 2-fold greater seed set in highly synchronised oaks compared to asynchronous individuals. Pollen addition removed the negative effect of asynchronous flowering on seed set. Because phenological synchrony operates through environmental variation, this result suggests that oak masting is synchronised by exogenous rather than endogenous factors. It also points to a mechanism by which changes in flowering phenology can affect plant reproduction of mast-seeding plants, with subsequent implications for community dynamics.

Developmental trap or demographic bonanza? Opposing consequences of earlier phenology in a changing climate for a multivoltine butterfly.

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.

Does masting scale with plant size? High reproductive variability and low synchrony in small and unproductive individuals.

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.

Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation.

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.

Environmental Veto Synchronizes Mast Seeding in Four Contrasting Tree Species.

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.

Why are monarch butterflies declining in the West? Understanding the importance of multiple correlated drivers.

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.

Correlated seed failure as an environmental veto to synchronize reproduction of masting plants.

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.

Time-lagged effects of weather on plant demography: drought and Astragalus scaphoides.

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).

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.

Arctic and boreal plant species decline at their southern range limits in the Rocky Mountains.

Climate change is predicted to cause a decline in warm-margin plant populations, but this hypothesis has rarely been tested. Understanding which species and habitats are most likely to be affected is critical for adaptive management and conservation. We monitored the density of 46 populations representing 28 species of arctic-alpine or boreal plants at the southern margin of their ranges in the Rocky Mountains of Montana, USA, between 1988 and 2014 and analysed population trends and relationships to phylogeny and habitat. Marginal populations declined overall during the past two decades; however, the mean trend for 18 dicot populations was -5.8% per year, but only -0.4% per year for the 28 populations of monocots and pteridophytes. Declines in the size of peripheral populations did not differ significantly among tundra, fen and forest habitats. Results of our study support predicted effects of climate change and suggest that vulnerability may depend on phylogeny or associated anatomical/physiological attributes.

Does movement behaviour predict population densities? A test with 25 butterfly species.

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.

Losing a battle but winning the war: moving past preference-performance to understand native herbivore-novel host plant interactions.

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.