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Understanding movement in heterogeneous environments is central to predicting how landscape changes affect animal populations. Several recent studies point out an intriguing and distinctive looping behavior by butterflies at habitat patch edges and hypothesize that this behavior requires a new framework for analyzing animal movement. We show that this looping behavior could be caused by a longstanding movement model, biased correlated random walk, with bias toward habitat patches. The ability of this longstanding model to explain recent observations reinforces the point that butterflies respond to habitat heterogeneity and do not move randomly through heterogeneous environments. We discuss the implications of different movement models for predicting butterfly responses to landscape change, and our rationale for retaining longstanding movement models, rather than developing new modeling frameworks for looping behavior at patch edges.

Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60-90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.

Negative impacts of invasive plants on natives have been well documented, but much less is known about whether invasive plants can cause population level declines. We used demographic models to investigate the effects of two invasive grasses on the demography and population growth of Anemone patens, a long-lived native perennial of North American grasslands. Demographic data of A. patens growing in patches characterized by Bromus inermis, Poa pratensis, or native grasses were used to parameterize integral projection models. Models based on both average conditions and those allowing for environmental stochasticity indicate that A. patens is slowly increasing in patches of native grass (lambda = 1.02) and declining in patches of invasive grasses, particularly those dominated by B. inermis (lambda = 0.93). Extinction probabilities indicate that A. patens should persist in native grass patches, but has a much higher probability of extinction in Bromus patches compared to Poa patches. While sensitivity analyses showed that survival had the biggest effect on population growth rates in all habitats, results of a Life Table Response Experiment (LTRE) revealed that slower individual growth rates in patches of invasive grasses contributed the most to the observed reduction in population growth. These results suggest that invasive grasses may cause slow declines in A. patens, despite short-term coexistence, and that controlling B. inermis only would not be sufficient to ensure A. patens persistence.

Mast seeding is common in plant populations, but its causes have rarely been tested experimentally. We tested mechanisms of alternate-year flowering and fruit set in an iteroparous, bee-pollinated, herbaceous plant, Astragalus scaphoides, in semi-arid sagebrush steppe. Patterns of reproduction from 1986 to 1999 indicated that spring precipitation was a cue for synchronous flowering, and that increased pollination in high-flowering years was a fitness advantage of synchrony. We tested these patterns by adding supplemental water and pollen to plants in high- and low-flowering sites and years. Supplemental water had no effect on flowering or seed set, so water is not a proximate cue for reproduction, though it could be important over longer (>3 year) time scales. Supplemental pollination increased fruit set in low- but not high-flowering years, indicating that synchronous flowering increases pollination success. Many shorter-term studies also report increased fruit set after pollen supplementation, but not after resource addition. This pattern may reflect the fact that plants can store and reallocate resources, but not pollen, across multiple years. For animal-pollinated herbs such as these, uniting theories about pollination ecology and mast seeding may promote an understanding of the mechanisms that determine patterns of reproduction over time.

Broad patterns in distribution and abundance can elucidate processes of evolution. A positive association between local abundance and the size of the geographic range has been demonstrated for closely related species across many taxa. This pattern is usually explained by assuming that species with smaller ranges are ecologically inferior (e.g., poor competitors or dispersers). Many areas of high endemism support local species that have evolved recently. The distribution of these neoendemics may reflect historical processes not accounted for by ecological, equilibrium hypotheses. We asked whether such traditional macroecological hypotheses also applied to the local abundance of seven narrowly endemic species and ecologically similar widespread congeners in the northern Rocky Mountains. For each of the 14 species, we estimated abundance of five randomly chosen populations by counting plants in 10 randomly located plots. The association between range size and local abundance was not positive. Instead, all seven narrow endemics were more abundant than their widespread congeners. Ecological specialization or differences in dispersal ability are not likely explanations for our results. We believe the local abundance of narrowly endemic species may be a sign of recent speciation. Most or all of our narrowly distributed species have probably not yet had time to spread to their full potential. Furthermore, theory predicts that speciation is more likely to occur in locally abundant populations. Our results suggest that strictly ecological mechanisms cannot explain abundance and distribution in regions with high neoendemism.

Synchronous mast seeding is increasingly recognized as common in plant populations. Recent theoretical models show that synchronous mast seeding could be a consequence of resource allocation and storage within individual plants, coupled by pollen limitation in low-flowering years. We used long-term population and weather data to parameterize models of flowering based on stored resources and pollen limitation in Astragalus scaphoides, a bee-pollinated plant that flowers in alternate years. We used these models to test whether internal resource dynamics could explain mast seeding in A. scaphoides and, if so, whether synchrony was caused by pollen limitation and/or fluctuations in precipitation. We compared predictions of models that included all combinations of three factors: constant versus precipitation-dependent resource gain, uniform versus heterogeneous resource gain (among individual plants), and resource-dependent versus resource- and pollen-limited fruit set. Pollen limitation and heterogeneous resource gain were necessary and sufficient to explain alternate-year flowering, but precipitation increased the quantitative match between model predictions and flowering dynamics. Together, our results support the importance of density-dependent pollen limitation as an ultimate and proximate cause of mast seeding in A. scaphoides. Precipitation does not act as a direct cue for synchrony in this species but might affect long-term resource gain and fruiting dynamics.

Simple measures of habitat proximity made primarily on the basis of land cover are widely used in the ecological literature to infer habitat connectivity, or the potential for animal movement among resource patches. However, such indices rarely have been tested against observations of animal movement or against more detailed biological models. We developed a priori expectations as to the types of study systems and organisms for which various habitat proximity indices would be best suited. We then used data from three study systems and four species to test which, if any, of the indices were good predictors of population-level responses. Our a priori expectations about index performance were not upheld. The indices that consider both habitat area and distance from the focal patch were highly correlated with each other, suggesting that they do index similar quantities. However, none of the indices performed well in predicting population response variables. The results suggest that the pattern of habitat cover alone may be insufficient to predict the process of animal movement.

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

The presence of litter has the potential to alter the population dynamics of plants. In this paper, we explore the effects of litter on population dynamics using a simple experimental laboratory system with populations of the annual crucifer, Cardamine pensylvanica. Using a factorial experiment with four densities and three litter levels, we determined the effect of litter on biomass and plant fecundity, and the life stages responsible for these changes in yield. Although litter had significant effects on seed germination and on seedling survivorship, we show, using a population dynamics model, that these effects were not demographically significant. Rather, the potential effect of litter on population dynamics resulted almost entirely from its effect on biomass. Persistent litter suppressed plant biomass and apparently removed the direct density effect present in the absence of litter. Thus, litter changed the shape of the recruitment curve from slightly humped to asymptotic. In addition to changing the shape of the recruitment curve, litter reduced the carrying capacity of the populations. Thus, the population dynamics model indicated that not all statistically significant responses were dynamically significant. Given the potential complexity of litter effects, simple population models provide a powerful tool for understanding the potential consequences of short-term responses.

Parental environmental effects have been widely reported in plants, but these effects are often weak relative to direct effects of current environmental conditions. Few studies have asked when consideration of such effects is necessary to understand long-term plant population dynamics. In this article, I show that inclusion of effects of parental density on offspring mass fundamentally changes population dynamics models by making recruitment a function of population size in two previous generations (Nt+1 = f(Nt, Nt-1)), rather than one (Nt+1 = f(Nt,)). Models without parental density effects predict either stable population dynamics or sharp crashes from high to low population size (flip bifurcations). When parental effects are at least one-third the size of direct density effects, gradual cycles from high to low population size (Hopf bifurcations) are possible. In this study, I measured effects of parental and offspring density on offspring quality in an annual plant, Cardamine pensylvanica, by manipulating plant density independently in parent and offspring generations and by comparing the effects of parent and offspring density on offspring performance. Parental density effects were detectable but were noticeably weaker than offspring density effects. Nonetheless, the parental effect was large enough to change population dynamics predictions. Thus, parental effects may be an important component of the numerical dynamics of plant populations.