Double Allee effects and extinction in the island fox.

An Allee effect (AE) occurs in populations when individuals suffer a decrease in fitness at low densities. If a fitness component is reduced (component AE), per capita population growth rates may decline as a consequence (demographic AE) and extinction risk is increased. The island fox (Urocyon littoralis) is endemic to six of the eight California Channel Islands. Population crashes have coincided with an increase in predation by Golden Eagles (Aquila chrysaetos). We propose that AEs could render fox populations more sensitive and may be a likely explanation for their sharp decline. We analyzed demographic data collected between 1988 and 2000 to test whether fox density (1) influences survival and reproductive rates; (2) interacts with eagle presence and affects fox fitness parameters; and (3) influences per capita fox population trends. A double component AE simultaneously influenced survival (of adults and pups) and proportion of breeding adult females. The adult survival AE was driven by predation by eagles. These component AEs led to a demographic AE. Multiple-component AEs, a predation-driven AE, and the simultaneous occurrence of both component and demographic AEs in a mammal are all previously unreported processes. Populations below 7 foxes/km(2) could have suboptimal population growth rates due to the demographic AE, and AEs may have contributed to the dramatic declines in three fox populations. Because fox densities in critically endangered populations are well below this level, removing Golden Eagles appears necessary to prevent a predation-driven AE. Conservationists should also be aware of AEs when planning the release of captive foxes. More generally, our findings highlight the danger of overlooking AEs in the conservation of populations of rare or threatened species.

Experimental evidence for spatial self-organization and its emergent effects in mussel bed ecosystems.

Spatial self-organization is the main theoretical explanation for the global occurrence of regular or otherwise coherent spatial patterns in ecosystems. Using mussel beds as a model ecosystem, we provide an experimental demonstration of spatial self-organization. Under homogeneous laboratory conditions, mussels developed regular patterns, similar to those in the field. An individual-based model derived from our experiments showed that interactions between individuals explained the observed patterns. Furthermore, a field study showed that pattern formation affected ecosystem-level processes in terms of improved growth and resistance to wave action. Our results imply that spatial self-organization is an important determinant of the structure and functioning of ecosystems, and it needs to be considered in their conservation.

Periodic dynamics in a two-stage Allee effect model are driven by tension between stage equilibria.

Single species difference population models can show complex dynamics such as periodicity and chaos under certain circumstances, but usually only when rates of intrinsic population growth or other life history parameter are unrealistically high. Single species models with Allee effects (positive density dependence at low density) have also been shown to exhibit complex dynamics when combined with over-compensatory density dependence or a narrow fertility window. Here we present a simple two-stage model with Allee effects which shows large amplitude periodic fluctuations for some initial conditions, without these requirements. Periodicity arises out of a tension between the critical equilibrium of each stage, i.e. when the initial population vector is such that the adult stage is above the critical value, while the juvenile stage is below the critical value. Within this area of parameter space, the range of initial conditions giving rise to periodic dynamics is driven mainly by adult mortality rates. Periodic dynamics become more important as adult mortality increases up to a certain point, after which periodic dynamics are replaced by extinction. This model has more realistic life history parameter values than most 'chaotic' models. Conditions for periodic dynamics might arise in some marine species which are exploited (high adult mortality) leading to recruitment limitation (low juvenile density) and might be an additional source of extinction risk.

Density dependence, spatial scale and patterning in sessile biota.

Sessile biota can compete with or facilitate each other, and the interaction of facilitation and competition at different spatial scales is key to developing spatial patchiness and patterning. We examined density and scale dependence in a patterned, soft sediment mussel bed. We followed mussel growth and density at two spatial scales separated by four orders of magnitude. In summer, competition was important at both scales. In winter, there was net facilitation at the small scale with no evidence of density dependence at the large scale. The mechanism for facilitation is probably density dependent protection from wave dislodgement. Intraspecific interactions in soft sediment mussel beds thus vary both temporally and spatially. Our data support the idea that pattern formation in ecological systems arises from competition at large scales and facilitation at smaller scales, so far only shown in vegetation systems. The data, and a simple, heuristic model, also suggest that facilitative interactions in sessile biota are mediated by physical stress, and that interactions change in strength and sign along a spatial or temporal gradient of physical stress.