The timing of biological events (phenology) is an important aspect of both a species' life cycle and how it interacts with other species and its environment. Patterns of phenological change have been given much scientific attention, particularly recently in relation to climate change. For pairs of interacting species, if their rates of phenological change differ, then this may lead to asynchrony between them and disruption of their ecological interactions. However it is often difficult to interpret differential rates of phenological change and to predict their ecological and evolutionary consequences. We review theoretical results regarding this topic, with special emphasis on those arising from life history theory, evolutionary game theory and population dynamic models. Much ecological research on phenological change builds upon the concept of match/mismatch, so we start by putting forward a simple but general model that captures essential elements of this concept. We then systematically compare the predictions of this baseline model with expectations from theory in which additional ecological mechanisms and features of species life cycles are taken into account. We discuss the ways in which the fitness consequences of interspecific phenological asynchrony may be weak, strong, or idiosyncratic. We discuss theory showing that synchrony is not necessarily an expected evolutionary outcome, and how population densities are not necessarily maximized by adaptation, and the implications of these findings. By bringing together theoretical developments regarding the eco-evolutionary consequences of phenological asynchrony, we provide an overview of available alternative hypotheses for interpreting empirical patterns as well as the starting point for the next generation of theory in this field.Global warming has caused phenological changes in all major ecosystems and taxa (Peñuelas and Filella 2001, Fitter and Fitter 2002, Sparks and Menzel 2002, Parmesan and Yohe 2003. Many spring and summer events have advanced but there is also a considerable variation in the rate of phenological change among species and phenophases (Schwartz et al. 2006, Thackeray et al. 2010. Such variation may reflect differential effects of warming on development rates of different organisms, and on cues that affect the seasonal scheduling of species activities (Yang and Rudolf 2010). It may also reflect variation in either genetic variance affecting the rate of adaptation through microevolution, or constraints on phenotypic plasticity (Gienapp et al. 2007).Many interspecific ecological interactions are dependent upon the temporal coordination of the seasonal activities of the interacting species. For example, Thomson (2010) has shown that pollen limitation in a subalpine lily (Erythronium grandiflorum) has increased over a period of 17 years, and the suggested mechanism is that climate change has decreased the temporal match between plants and pollinators. Spawning phenology in the marine intertidal bivalve Macoma balthica is under selection by te...
In migratory birds, arrival date and hatching date are two key phenological markers that have responded to global warming. A body of knowledge exists relating these traits to evolutionary pressures. In this study, we formalize this knowledge into general mathematical assumptions, and use them in an ecoevolutionary model. In contrast to previous models, this study novelty accounts for both traits-arrival date and hatching date-and the interdependence between them, revealing when one, the other or both will respond to climate. For all models sharing the assumptions, the following phenological responses will occur. First, if the nestling-prey peak is late enough, hatching is synchronous with, and arrival date evolves independently of, prey phenology. Second, when resource availability constrains the length of the pre-laying period, hatching is adaptively asynchronous with prey phenology. Predictions for both traits compare well with empirical observations. In response to advancing prey phenology, arrival date may advance, remain unchanged, or even become delayed; the latter occurring when egg-laying resources are only available relatively late in the season. The model shows that asynchronous hatching and unresponsive arrival date are not sufficient evidence that phenological adaptation is constrained. The work provides a framework for exploring microevolution of interdependent phenological traits.
1. Local food webs result from a sequence of colonisations and extinctions by species from the regional pool or metaweb, that is, the assembly process. Assembly is theorised to be a selective process: whether or not certain species or network structures can persist is partly determined by local processes including habitat filtering and dynamical constraints. Consequently, local food web structure should reflect these processes.2. The goal of this study was to test evidence for these selective processes by comparing the structural properties of real food webs to the expected distribution given the metaweb. We were particularly interested in ecological dynamics; if the network properties commonly associated with dynamical stability are indeed the result of stability constraints, then they should deviate from expectation in the direction predicted by theory.3. To create a null expectation, we used the novel approach of randomly assembling model webs by drawing species and interactions from the empirical metaweb.The assembly model permitted colonisation and extinction, and required a consumer species to have at least one prey, but had no habitat type nor population dynamical constraints. Three datasets were used: (a) the marine Antarctic metaweb, with two local food webs; (b) the 50 lakes of the Adirondacks; and (c) the arthropod community from Florida Keys' classic defaunation experiment.4. Contrary to our expectations, we found that there were almost no differences between empirical webs and those resulting from the null assembly model. Few empirical food webs showed significant differences with network properties, motif representations and topological roles. Network properties associated with stability did not deviate from expectation in the direction predicted by theory. 5. Our results suggest that-for the commonly used metrics we considered-local food web structure is not strongly influenced by dynamical nor habitat restrictions. Instead, the structure is inherited from the metaweb. This suggests that the network properties typically attributed as causes or consequences of ecological stability are instead a by-product of the assembly process (i.e. spandrels), and may potentially be too coarse to detect the true signal of dynamical constraint.
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