The evolution of thermal performance can constrain dispersal during range shifting

Organisms can cope with changing temperature under climate change by either adapting to the temperature at which they perform best and/or by dispersing to more benign locations. The evolution of a new thermal niche during range shifting is, however, expected to be strongly constrained by genetic load because spatial sorting is known to induce fast evolution of dispersal. To broaden our understanding of this interaction, we studied the joint evolution of dispersal and thermal performance curves (TPCs) of a population during range shifting by applying an individual-based spatially explicit model. Always, TPCs adapted to the local thermal conditions. Remarkably, this adaptation coincided with an evolution of dispersal at the shifting range front being equally high or lower than at the trailing edge. This optimal strategy reduces genetic load and highlights that evolutionary dynamics during range shifting change when crucial traits such as dispersal and thermal performance jointly evolve.     

Exploring stable pattern formation in models of tussock moth populations

1. The western tussock moth ( Orgyia vetusta ) at the University of California Bodega Marine Reserve (Sonoma County, California, USA) exhibits dense, localized populations in the midst of extensive habitats where variation in host plant quality or predator abundance is unable to explain the restricted extent of the outbreaks.
2. Two primary features suggest that the host patterning is intrinsically generated: (i) female tussock moths are wingless, producing a low effective dispersal distance for the hosts; and (ii) the tussock moth population is attacked by several species of widely dispersing wasp and fly parasitoids.
3. We consider a set of spatially explicit host–parasitoid models to examine whether intrinsically generated patterns are possible within this system. These models include a spatially extended Nicholson–Bailey model to examine general features of pattern formation in host–parasitoid systems, and two system-specific models, an individual-based simulation and a population-level analytic model, to examine the details of this empirical system.
4. Both stable patterning and rapid extinction of the host population are initial-condition dependent outcomes of the general and specific models, implying that an intrinsically generated stable host pattern is feasible within the tussock moth system.
5. Stable patterning is enhanced by a large parasitoid-to-host dispersal ratio, local host resource limitation, and increased parasitism at the host patch's edge.     

On the evolution of dispersal via heterogeneity in spatial connectivity

Dispersal has long been recognized as a mechanism that shapes many observed ecological and evolutionary processes. Thus, understanding the factors that promote its evolution remains a major goal in evolutionary ecology. Landscape connectivity may mediate the trade-off between the forces in favour of dispersal propensity (e.g. kin-competition, local extinction probability) and those against it (e.g. energetic or survival costs of dispersal). It remains, however, an open question how differing degrees of landscape connectivity may select for different dispersal strategies. We implemented an individual-based model to study the evolution of dispersal on landscapes that differed in the variance of connectivity across patches ranging from networks with all patches equally connected to highly heterogeneous networks. The parthenogenetic individuals dispersed based on a flexible logistic function of local abundance. Our results suggest, all else being equal, that landscapes differing in their connectivity patterns will select for different dispersal strategies and that these strategies confer a long-term fitness advantage to individuals at the regional scale. The strength of the selection will, however, vary across network types, being stronger on heterogeneous landscapes compared with the ones where all patches have equal connectivity. Our findings highlight how landscape connectivity can determine the evolution of dispersal strategies, which in turn affects how we think about important ecological dynamics such as metapopulation persistence and range expansion.     

Individual variation and population dynamics: lessons from a simple system

The mapping of environment, through variation in individuals' life histories, to dynamics can be complex and often poorly known. Consequently, it is not clear how important it is dynamically. To explore this, I incorporated lessons from an empirical system, a soil mite, into an individual-based model. Individuals compete for resource and allocate this according to eight 'genetic' rules that specify investment in growth or reserves (which influences survival or fecundity), size at maturation and reproductive allocation. Density dependence, therefore, emerges from competition for food, limiting individual's growth and fecundity. We use this model to examine the role that genetic and phenotypically plastic variation plays in dynamics, by fixing phenotypes, by allowing phenotypes to vary plastically and by creating genetic variation between individuals. Variation, and how it arises, influences short- and long-run dynamics in a way comparable in magnitude with halving food supply. In particular, by switching variation on and off, it is possible to identify a range of processes necessary to capture the dynamics of the 'full model'. Exercises like this can help identify key processes and parameters, but a concerted effort is needed across many different systems to search for shared understanding of both process and modelling.     

Dynamics of range margins for metapopulations under climate change

We link spatially explicit climate change predictions to a dynamic metapopulation model. Predictions of species'' responses to climate change, incorporating metapopulation dynamics and elements of dispersal, allow us to explore the range margin dynamics for two lagomorphs of conservation concern. Although the lagomorphs have very different distribution patterns, shifts at the edge of the range were more pronounced than shifts in the overall metapopulation. For Romerolagus diazi (volcano rabbit), the lower elevation range limit shifted upslope by approximately 700 m. This reduced the area occupied by the metapopulation, as the mountain peak currently lacks suitable vegetation. For Lepus timidus (European mountain hare), we modelled the British metapopulation. Increasing the dispersive estimate caused the metapopulation to shift faster on the northern range margin (leading edge). By contrast, it caused the metapopulation to respond to climate change slower, rather than faster, on the southern range margin (trailing edge). The differential responses of the leading and trailing range margins and the relative sensitivity of range limits to climate change compared with that of the metapopulation centroid have important implications for where conservation monitoring should be targeted. Our study demonstrates the importance and possibility of moving from simple bioclimatic envelope models to second-generation models that incorporate both dynamic climate change and metapopulation dynamics.     

Shifting dispersal modes at an expanding species’ range margin

While it is generally recognized that noncontiguous (long‐distance) dispersal of small numbers of individuals is important for range expansion over large geographic areas, it is often assumed that colonization on more local scales proceeds by population expansion and diffusion dispersal (larger numbers of individuals colonizing adjacent sites). There are few empirical studies of dispersal modes at the front of expanding ranges, and very little information is available on dispersal dynamics at smaller geographic scales where we expect contiguous (diffusion) dispersal to be prevalent. We used highly polymorphic genetic markers to characterize dispersal modes at a local geographic scale for populations at the edge of the range of a newly invasive grass species (Brachypodium sylvaticum) that is undergoing rapid range expansion in the Pacific Northwest of North America. Comparisons of Bayesian clustering of populations, patterns of genetic diversity, and gametic disequilibrium indicate that new populations are colonized ahead of the invasion front by noncontiguous dispersal from source populations, with admixture occurring as populations age. This pattern of noncontiguous colonization was maintained even at a local scale. Absence of evidence for dispersal among adjacent pioneer sites at the edge of the expanding range of this species suggests that pioneer populations undergo an establishment phase during which they do not contribute emigrants for colonization of neighbouring sites. Our data indicate that dispersal modes change as the invasion matures: initial colonization processes appear to be dominated by noncontiguous dispersal from only a few sources, while contiguous dispersal may play a greater role once populations become established.     

Trait evolution in an individual-based model of herbaceous vegetation

Many theoretical studies of evolution are based upon the concepts of the evolutionary stable strategy and optimal life-history solutions. An individual based model of vegetation is used to simulate life-history evolution under two different sets of environmental conditions. At one level the results suggest that optimal life-history solutions do appear to evolve. At the end of the simulations the vegetation that evolved in a fertile and uncut environment was taller, thinner and germinated later than that which developed in a less fertile and cut habitat. However, between simulation variation was observed to be high, particularly for the parameter regulating the timing of reproduction, and it showed no indication of reaching fixation. When this trait was prevented from mutating, the variances of other traits were seen to increase. Although at the population level between simulation variation was high, some traits achieved a degree of stability within simulations, suggesting that multiple adaptive peaks may be being approached. However, there was little evidence of trait fixation occurring within the most abundant genotype. It is considered that frequency dependent selection/Red Queen dynamics may be acting to prevent the most abundant genotype from reaching fixation. It is argued that if such processes prevent optimal genetic solutions from being achieved then the search for evolutionary stable strategies within the evolution of life-histories may be over simplistic.     

Waldfragmentierung, Konkurrenz und klimatische Abh?ngigkeit beim Wespenbussard (Pernis apivorus)

Zusammenfassung Zwischen 1992 und 1997 betrug der Wespenbussard-Bestand eines 110 km² großen, waldarmen Gebietes am nördlichen Alpenrand 7 bis 10 territoriale Paare. Von 10 Waldfragmenten wurden die kleineren signifikant häufiger besetzt. Die Besetzung durch Mäusebussard-Brutpaare beeinflußte die Besetzungsrate von 10 Waldfragmenten nicht signifikant, auch nicht im Zusammenhang mit der Größe der Fragmente. Eine höhere Anzahl von Regentagen im Mai hatte keine Auswirkungen auf die Dichte, und im Juni/Juli bewirkten sie während des Untersuchungszeitraumes keine Einbrüche der Reproduktion. Es wird vorgeschlagen, daß den Faktoren Waldfragmentierung, Konkurrenz und Klima das Nahrungsangebot übergeordnet ist.

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Forest fragmentation, competition and climatic dependence in the Honey Buzzard (Pernis apivorus)

Summary Between 1992 and 1997 a population of Honey Buzzards varied between 7 and 10 territorial pairs on the northern fringe of the Alps. The amount of forest cover in the study plot of 110 km² was only 10 %. For 10 forest fragments the frequency of occupation increased significantly with the decreasing size of the fragments. The presence of breeding pairs of Common Buzzards did not significantly affect the frequency of occupation of 10 fragments of <27 ha. Additionally, for fragments occupied by both species, there was no significant increase in the occupation by Honey Buzzards with inreasing patch size. Both species used to breed successfully alongside each other in small patches. The number of rain days in May had no significant impact on the density during the study period. Similarly, I confirmed at least 4–6 successful broods each year. I conclude that forest fragmentation does not carry any threat to the Honey Buzzard population. Mechanisms like competition, habitat fragmentation and climate seem to act through food supply.

     

How interactions between animal movement and landscape processes modify local range dynamics and extinction risk

Forecasts of range dynamics now incorporate many of the mechanisms and interactions that drive species distributions. However, connectivity continues to be simulated using overly simple distance-based dispersal models with little consideration of how the individual behaviour of dispersing organisms interacts with landscape structure (functional connectivity). Here, we link an individual-based model to a niche-population model to test the implications of this omission. We apply this novel approach to a turtle species inhabiting wetlands which are patchily distributed across a tropical savannah, and whose persistence is threatened by two important synergistic drivers of global change: predation by invasive species and overexploitation. We show that projections of local range dynamics in this study system change substantially when functional connectivity is modelled explicitly. Accounting for functional connectivity in model simulations causes the estimate of extinction risk to increase, and predictions of range contraction to slow. We conclude that models of range dynamics that simulate functional connectivity can reduce an important source of bias in predictions of shifts in species distributions and abundances, especially for organisms whose dispersal behaviours are strongly affected by landscape structure.     

The consequences of flooding for the dispersal and fate of poeciliid fish in an intermittent tropical stream

Summary We studied the rainy season dispersal of the fish Poecilia gillii (Poeciliidae) from pools in a steepgradient, intermittent stream in Santa Rosa National Park, Costa Rica. The stream consisted of about 20 pools separated by dry streambed except during two floods and subsequent brief periods of flow. Individually recognizable tags permitted mark-recapture estimates of population size and information on individual movements. The first flood was very severe, with pools losing an average of 75% of their populations (range 12–99%). Most of the lost fish died by becoming trapped in desiccating pools. Males and juveniles were more likely to be lost than were females. Population loss was negatively related to pool volume and positively related to streambed slope. In addition, population loss was positively related to preflood population size when the effects of pool volume and slope were removed indicating that pools with higher densities lost more fish. Of the fish recaptured after the flood, the average proportion found in the pool in which they had been tagged varied from 16%–96%, depending on the area of the stream. Of fish that moved, 92% went downstream. The second flood was less severe though stream flow lasted as long, and there was little effect on the pool populations. Involuntary flushing during the flood and voluntary departure apparently interacted to produce the observed patterns.     

Predicting differential effects of climate change at the population level with life-cycle models of spring Chinook salmon

Habitat conditions mediate the effects of climate, so neighboring populations with differing habitat conditions may differ in their responses to climate change. We have previously observed that juvenile survival in Snake River spring/summer Chinook salmon is strongly correlated with summer temperature in some populations and with fall streamflow in others. Here, we explore potential differential responses of the viability of four of these populations to changes in streamflow and temperature that might result from climate change. First, we linked predicted changes in air temperature and precipitation from several General Circulation Models to a local hydrological model to project streamflow and air temperature under two climate‐change scenarios. Then, we developed a stochastic, density‐dependent life‐cycle model with independent environmental effects in juvenile and ocean stages, and parameterized the model for each population. We found that mean abundance decreased 20–50% and the probability of quasi‐extinction increased dramatically (from 0.1–0.4 to 0.3–0.9) for all populations in both scenarios. Differences between populations were greater in the more moderate climate scenario than in the more extreme, hot/dry scenario. Model results were relatively robust to realistic uncertainty in freshwater survival parameters in all scenarios. Our results demonstrate that detailed population models can usefully incorporate climate‐change predictions, and that global warming poses a direct threat to freshwater stages in these fish, increasing their risk of extinction. Because differences in habitat may contribute to the individualistic population responses we observed, we infer that maintaining habitat diversity will help buffer some species from the impacts of climate change.     

Fragmentation of clones: how does it influence dispersal and competitive ability?

We applied individual-based simulations to study the effect of physiological integration among ramets in clonal species that live in patchy habitats. Three strategies were compared: (1) Splitter, in which the genet was fragmented into independent ramets; (2) Transient Integrator, where only groups of ramets were connected; and (3) Permanent Integrator, in which fragmentation did not occur, and the whole genet was integrated. We studied the dynamics of spatial spreading and population growth in these strategies separately and in competition. Various habitat types were modeled by changing the density of favorable habitat patches. We found that the spatial pattern of good patches significantly influenced the growth of the populations. When the resource patches were scarce, a large proportion of the carrying capacity of the habitat was not utilized by any of the strategies. It was the Splitter that proved to be the most severely dispersal-limited. But at the same time, it could compete for the good patches most efficiently. The balance between these two contradictory effects was largely determined by the proportion of favorable to unfavorable areas. When this proportion was low or intermediate (up to ca. 50% good), integration was more advantageous. At higher proportions, fragmentation became beneficial. Fragmentation into groups of ramets (Transient Integration) was not sufficient, only radical splitting could ensure a significant selective advantage. Transient Integrators got fragmented according to the spatial pattern of ramet mortality. It was interesting that the enrichment of the area in good sites did not lead to larger fragment sizes. It merely raised the number of fragments. Nevertheless, these small fragments were more similar to integrated genets (in the Permanent Integrator) than to solitary ramets (in the Splitter) in terms of dispersal and competitive ability. This suggests that even a slightly integrated clonal species can be ecologically considered as an integrator.