Phenology affects nearly all aspects of ecology and evolution. Virtually all biological phenomenafrom individual physiology to interspecific relationships to global nutrient fluxes-have annual cycles and are influenced by the timing of abiotic events. Recent years have seen a surge of interest in this topic, as an increasing number of studies document phenological responses to climate change. Much recent research has addressed the genetic controls on phenology, modelling techniques and ecosystem-level and evolutionary consequences of phenological change. To date, however, these efforts have tended to proceed independently. Here, we bring together some of these disparate lines of inquiry to clarify vocabulary, facilitate comparisons among habitat types and promote the integration of ideas and methodologies across different disciplines and scales. We discuss the relationship between phenology and life history, the distinction between organismal-and population-level perspectives on phenology and the influence of phenology on evolutionary processes, communities and ecosystems. Future work should focus on linking ecological and physiological aspects of phenology, understanding the demographic effects of phenological change and explicitly accounting for seasonality and phenology in forecasts of ecological and evolutionary responses to climate change.

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Abstract. One possible effect of climate change is the generation of a mismatch in the seasonal timing of interacting organisms, owing to species-specific shifts in phenology. Despite concerns that plants and pollinators might be at risk of such decoupling, there have been few attempts to test this hypothesis using detailed phenological data on insect emergence and flowering at the same localities. In particular, there are few data sets on pollinator flight seasons that are independent of flowering phenology, because pollinators are typically collected at flowers. To address this problem, we established standardized nesting habitat (trap nests) for solitary bees and wasps at sites along an elevational gradient in the Rocky Mountains, and monitored emergence during three growing seasons. We also recorded air temperatures and flowering phenology at each site. Using a reciprocal transplant experiment with nesting bees, we confirmed that local environmental conditions are the primary determinants of emergence phenology. We were then able to develop phenology models to describe timing of pollinator emergence or flowering, across all sites and years, as a function of accumulated degree-days. Although phenology of both plants and insects is well described by thermal models, the best models for insects suggest generally higher threshold temperatures for development or diapause termination than those required for plants. In addition, degreeday requirements for most species, both plants and insects, were lower in locations with longer winters, indicating either a chilling or vernalization requirement that is more completely fulfilled at colder sites, or a critical photoperiod before which degree-day accumulation does not contribute to development. Overall, these results suggest that phenology of plants and trap-nesting bees and wasps is regulated in similar ways by temperature, but that plants are more likely than insects to advance phenology in response to springtime warming. We discuss the implications of these results for plants and pollinators, and suggest that phenological decoupling alone is unlikely to threaten population persistence for most species in our study area.

Climate change can aff ect plant -pollinator interactions in a variety of ways, but much of the research attention has focused on whether independent shifts in phenology will alter temporal overlap between plants and pollinators. Here I review the research on plant -pollinator mismatch, assessing the potential for observational and experimental approaches to address particular aspects of the problem. Recent, primarily observational studies suggest that phenologies of co-occurring plants and pollinators tend to respond similarly to environmental cues, but that nevertheless, certain pairs of interacting species are showing independent shifts in phenology. Only in a few cases, however, have these independent shifts been shown to aff ect population vital rates (specifi cally, seed production by plants) -but this largely refl ects a lack of research. Compared to the few long-term studies of pollination in natural plant populations, experimental manipulations of phenology have yielded relatively optimistic conclusions about eff ects of phenological shifts on plant reproduction, and I discuss how issues of scale and frequency-dependence in pollinator behaviour aff ect the interpretation of these ' temporal transplant ' experiments. Comparable research on the impacts of mismatch on pollinator populations is so far lacking, but both observational studies and focused experiments have the potential to improve our forecasts of pollinator responses to changing phenologies. Finally, while there is now evidence that plant -pollinator mismatch can aff ect seed production by plants, it is still unclear whether this phenological impact will be the primary way in which climate change aff ects plant -pollinator interactions. It would be useful to test the direct eff ects of changing climate on pollinator population persistence, and to compare the importance of phenological mismatch with other threats to pollination.

Summary1. Land-use change frequently reduces local species diversity. Species losses will often result in loss of trait diversity, with likely consequences for community functioning. However, the converse need not be generally true: management approaches that succeed in retaining species richness could nevertheless fail to maintain trait diversity. We evaluated this possibility using bee communities in a California agroecosystem. 2. We examined among site patterns in bee species diversity and functional-trait diversity in a landscape composed of a mosaic of semi-natural habitat, organic farms and conventional farms. We sampled bees from all three habitat types and compiled a data base of life-history ('functional') traits for each species. 3. Although species richness was higher on organic farms than conventional farms, functional diversity was lower in both farm types than in natural habitat. Nesting location (below-ground vs. above-ground) was the primary trait contributing to differences in functional diversity between farms and natural habitat, reflecting observed differences in availability of nesting substrates among habitat types. Other traits, including phenology and sociality, were also associated with species' occurrences or dominance in particular site types. These patterns suggest that management practices common to all farms act as environmental filters that cause similarly low functional diversity in their bee communities. 4. Synthesis and applications. Although our results support the value of organic farming in maintaining abundant and species-rich bee communities, components of bee functional diversity are not well supported in farmed landscapes, regardless of farming practice. Maintenance of natural habitat, and/or the addition of natural habitat elements to farms, is therefore important for the retention of functionally diverse bee assemblages in agroecosystems.