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In this paper, we review mate-finding Allee effects from ecological and evolutionary points of view. We define 'mate-finding' as mate searching in mobile animals, and also as the meeting of gametes for sessile animals and plants (pollination). We consider related issues such as mate quality and choice, sperm limitation and physiological stimulation of reproduction by conspecifics, as well as discussing the role of demographic stochasticity in generating mate-finding Allee effects. We consider the role of component Allee effects due to mate-finding in generating demographic Allee effects (at the population level). Compelling evidence for demographic Allee effects due to matefinding (as well as via other mechanisms) is still limited, due to difficulties in censusing rare populations or a failure to identify underlying mechanisms, but also because of fitness trade-offs, population spatial structure and metapopulation dynamics, and because the strength of component Allee effects may vary in time and space. Mate-finding Allee effects act on individual fitness and are thus susceptible to change via natural selection. We believe it is useful to distinguish two routes by which evolution can act to mitigate mate-finding Allee effects. The first is evolution of characteristics such as calls, pheromones, hermaphroditism, etc.which make mate-finding more efficient at low density, thus eliminating the Allee effect. Such adaptations are very abundant in the natural world, and may have arisen to avoid Allee effects, although other hypotheses are also possible. The second route is to avoid low density via adaptations such as permanent or periodic aggregation. In this case, the Allee effect is still present, but its effects are avoided. These two strategies may have different consequences in a world where many populations are being artificially reduced to low density: in the first case, population growth rate can be maintained, while in the second case, the mechanism to avoid Allee effects has been destroyed. It is therefore in these latter populations that we predict the greatest evidence for mate-finding Allee effects and associated demographic consequences. This idea is supported by the existing empirical evidence for demographic Allee effects. Given a strong effect that mate-finding appears to have on individual fitness, we support the continuing quest to find connections between component mate-finding Allee effects (individual reproductive fitness) and the demographic consequences. There are many reasons why such studies are difficult, but it is important, particularly given the increasing number of populations and species of conservation concern, that the ecological community understands more about how widespread demographic Allee effects really are, and why.

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.

Summary1. In a population with Allee effects a positive relationship exists between fitness and population size or density. Allee effects may result in extinction thresholds and are therefore crucial in conservation and management. It has been shown theoretically that Allee effects can be driven by predation; however, there are few empirical data. Previous empirical work on Allee effects has emphasized that taxa with life-history characteristics such as co-operative breeding may be prone to such effects. Because predation is a general ecological mechanism, Allee effects may be more widespread than previously thought. 2. We used a series of simple heuristic models to develop a theoretical framework for understanding predation-driven Allee effects as a function of predator functional and aggregative responses. 3. Predators can create an Allee effect if they have a type I (linear) or type II (saturating) functional response without a type III (sigmoid) aggregative response, or vice versa. In addition, predation must be the main driver of prey dynamics, and prey must have little spatial or temporal refuge from predation. 4. We highlighted several, mainly unrecognized, examples of predation-driven Allee effects from the literature, the majority of which came from systems that had been perturbed by exploitation or introduced predators. 5. Synthesis and applications . Allee effects can arise from a general ecological process under a variety of different combinations of functional and aggregative responses. Allee effects may thus be present in a broad spectrum of different taxa with different types of life history, not only those taxa, such as broadcast spawners and co-operative breeders, on which empirical work has focused thus far. Conservation biologists and managers working with heavily exploited or threatened populations, or attempting reintroductions, should be aware of the possibility of a threshold population size or density below which extinction is likely. These thresholds can occur regardless of species life history, if predation is a major source of mortality and spatial and temporal predation refuges are limited.

The fitness or population growth rate of populations with Allee effects increases with increasing population size or density up to a certain threshold. Allee effects are possible in marine populations, as they are less open than has been assumed and may have a metapopulation structure. We modelled the population consequences of Allee effects and show that increases in mortality interact with critical Allee thresholds, such that an Allee effect with no population consequences at low mortality can drive a population to extinction when mortality is increased. In heavily fished species, populations with strong Allee effects go extinct at lower levels of fishing mortality, or conversely as fishing mortality increases, weaker Allee effects can drive population to extinction. We found little empirical evidence in the literature for widespread Allee effects in marine populations, although we found some suggestive observations, particularly for broadcast spawners and in exploited populations This might be due to methodological problems or long time lags. Many marine species have components of their life history or ecology which could in theory generate Allee effects; however the population level consequences of these potential mechanisms remains virtually unexplored. We suggest that including Allee effects in models of vulnerable populations may be critical for the precautionary management of exploited and threatened marine species.KEY WORDS: Allee effects · Positive density dependence · Inverse density dependence · DepensationResale or republication not permitted without written consent of the publisher

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.

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.

Feeding behavior of mussels (Mytilus edulis) was measured in situ using a video camera and expressed as the mean percentage of valve gape aperture (VA), concomitant with environmental and biological parameters over two tidal cycles. Mussel feeding behavior and the physical parameters responded to three primary tidal components, of which semidiurnal was dominant (12.42 h). VA was synchronized with chlorophyll a (Chl a) concentration (proxy for food) with a strong positive correlation (r 5 0.72, p , 0.001). Chl a and suspended particulate matter (SPM) were dependent on tidal advection. The combination of the reconstructed tidal constituents derived from harmonic analysis were used to successfully model mussel feeding behavior (r 5 0.90, p , 0.001). In this concentration range (0.6 to 2.5 mg L 21 ), Chl a, measured at 1 m above the mussel bed, regulates mussel feeding behavior irrespective of the presence of predators, changes in SPM, or flow velocity.