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Levels of asynchrony for limited and unlimited dispersal metapopulation systems. (a) Semi-variogram for Callosobruchus maculatus in the 16-patch limited dispersal metapopulations. (b) Semi-variogram for C. maculatus in the 16-patch unlimited dispersal metapopulations. (c) Semi-variogram for Callosobruchus chinensis in the 16-patch limited dispersal metapopulations. (d ) Semi-variogram for C. chinensis in the 16-patch unlimited dispersal metapopulations. (e) Semi-variogram for Anisopteromalus calandrae in the 16-patch limited dispersal metapopulations. ( f ) Semi-variogram for A. calandrae in the 16- patch unlimited dispersal metapopulations. Under limited dispersal (a,c,e) the high degree of asynchrony is punctuated with periods when the systems appear to be in phase; in unlimited dispersal metapopulations (b,d,f), the systems rapidly loses asynchrony and the dynamics are completely coupled by the action of the rapidly dispersing natural enemy. Spatiotemporal semi-variograms for each metapopulation series (of length n) were calculated using gðd; tÞZ ð1=2W Þ P n
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Quantifying the role of space and spatial scale on the population dynamics of ecological assemblages is a contemporary challenge in ecology. Here, we evaluate the role of metapopulation dynamics on the persistence and dynamics of a multispecies predator-prey assemblage where two prey species shared a common natural enemy (apparent competition). By...
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... We must point out that these scenarios are only valid for specialized predators, which are highly adapted to capturing their preferred prey, such as the Eurasian lynx (Lynx lynx), specialized in small ungulates [102], or the coast horned lizard (Phrynosoma coronatum), specialized in local ants [103]. When predators are generalist they present other population dynamics because there is merely apparent competition [104,105]. We observe that two common prey of the predators remain in equilibrium despite one predator (Strong) being more efficient, due to a more uniform distribution (simulation 6 in 2SpePred.json in Supplementary Material). ...
... The habitat structure and the spatial scale play a central role in the predator-prey dynamic. These effects have been observed to operate in more complex groups of predators and preys and of multiple species [104,105]. In ECA we also see that this happens from a new perspective: the aggregate distribution prevails in the prey-predator interaction because its variability limits the effectiveness of the predator, and this means preserving the evolutionary balance of its prey and, therefore, of predators. ...
Simple Summary: ECA is similar to a cellular automaton that mimics the evolutionary dynamics of species in metapopulations, simulating the underlying mechanisms of natural selection. In this work, we carry out an in-silico study of the effects of different dispersal strategies on the evolutionary balance of interactions between digital organisms. In ECA we see how a specific type of distribution can significantly influence the dynamics, persistence, distribution, and abundance of populations of different species within a particular habitat. In the first place, we show that an aggregate distribution is more inefficient than a uniform distribution. Still, we verify that this aggregate distribution is essential in predator-prey type interactions so that the species involved do not become extinct. We also show that the aggregate distribution does not comply with the competitive exclusion principle and, for this reason, it results in a general and straightforward explanation for the paradox of plankton and the grouping of animals. Many animals and especially some planktonic species group together in specific spaces, although some travel adrift, leaving other areas or patches free where competitors or their prey can prosper, preventing them from becoming extinct. Abstract: This work analyzes the evolutionary consequences of different aggregation levels of species distribution with an Evolutionary Cellular Automaton (ECA). We have found that in habitats with the same carrying capacity, aggregated distributions preserve smaller populations than do uniform distributions, i.e., they are less efficient. Nonetheless, we have also found that aggregated distributions, among other factors, can help the evolutionary stability of some biological interactions , such as predator-prey interactions, despite their granting less individual fitness. Besides, the competitive exclusion principle does not usually stand in populations with aggregated distribution. We have applied ECA to study the effects of aggregated distribution in two notorious cases: in the so-called paradox of the plankton and in gregarious animals. In doing so, we intend to ratify long-established ecological knowledge explaining these phenomena from a new perspective. In the first case, due to aggregate distribution, large aggregations of digital organisms mimicking very abundant planktonic species, leave large patches or oceanic areas free for other less competitive organisms , which mimic rare species, to prosper. In this case, we can see how effects, such as ecological drift and the small portion, act simultaneously. In the second case of aggregation, the aggregate distribution of gregarious animals could be explained under specialized predator-prey interactions and interdemic competition. Thus, digital organisms that imitate predators reduce the competitive capacity of their prey, destabilizing their competitiveness against other species. The specialized predator also goes extinct if the prey goes extinct by natural selection. Predators that have an aggregate distribution compensate the prey and thus avoid exclusion. This way there are more predator-free patches in which the prey can prosper. However, by granting greater colonization capacity to its prey, the predator loses competitiveness. Therefore, it is a multilevel selection event in which group adaptation grows to the detriment of the predator as an individual. Citation: Falgueras-Cano, J.; Falgueras-Cano, J.A.; Moya, A.
... their population-level spatial structuredetermines and constrains intra-and interspecific interactions (e.g., Skellam 1951;MacArthur and Wilson 1967;Levin 1974;Tilman and Kareiva 1997;Fagan et al. 1999). This spatial structure often arises from dispersal (e.g., Levins 1969;Gilpin and Hanski 1991;Hanski 1998Hanski , 1999Bonsall et al. 2005;Schnell et al. 2013;Gibert 2016;Yeakel et al. 2018), habitat selection and suitability (MacArthur and Levins 1964;Rosenzweig 1981;Cody 1985;Hirzel et al. 2001;Brotons et al. 2004;Abrams 2007;Hirzel and Le Lay 2008), aggregation (Hassell and May 1974;Krause and Tegeder 1994;Levin 1994;May et al. 2016) or segregation of individuals (Recher 1966;Thiebot et al. 2012;Navarro et al. 2013), local environmental conditions (Pearson and Dawson 2003;Thuiller et al. 2004), local extinction (MacArthur and Wilson 1967;Levins 1969;Gilpin and Hanski 1991) as well as interspecific ecological interactions such as competition (Tilman 1994;Tilman et al. 1994;Amarasekare 2003;Amarasekare et al. 2004), predation (Bascompte et al. 2002;McCann et al. 2005;Amarasekare 2008), and facilitation (Choler et al. 2001;Stachowicz 2001;Boulangeat et al. 2012). ...
We revisit a seminal paper by Levin (Am Nat 108:207–228, 1974), where spatially mediated coexistence and spatial pattern formation were described. We do so by reviewing and explaining the mathematical tools used to evaluate the dynamics of ecological systems in space, from the perspective of recent developments in spatial population dynamics. We stress the importance of space-mediated stability for the coexistence of competing species and explore the ecological consequences of space-induced instabilities (Turing instabilities) for spatial pattern formation in predator–prey systems. Throughout, we link existing theory to recent developments in discrete spatially structured metapopulations, such as our understanding of how ecological dynamics occurring on a network can be analyzed using the Laplacian matrix and its associated eigenvalue spectrum. We underline the validity of Levin’s message, over 40 years later, and suggest it has ever-growing implications in a changing and increasingly fragmented world.
... Alternatively, it might not have: the generalization that metapopulation persistence time is a humped function of dispersal rate rests largely on theoretical models (Yaari et al. 2012), the applicability of which to real metapopulations remains relatively little tested. The only previous replicated experiments showing that metapopulation persistence time is a humped function of dispersal rate or connectivity used only three connectivity treatments, two of which were extreme: no connectivity, or an approximately well-mixed system (Holyoak and Lawler 1996, Holyoak 2000, Ellner et al. 2001, Bonsall et al. 2005. Other experiments testing effects of dispersal rate or connectivity on metapopulation dynamics were unreplicated (Huffaker 1958, Janssen et al. 1997, did not explore the full range of possible dispersal rates or connectivity (Burkey 1997, Janssen et al. 1997, Kerr et al. 2006, Bull et al. 2007), or did not find that metapopulation persistence is a humped function of dispersal rate or connectivity (Dey and Joshi 2006, Griffen and Drake 2009, Vogwill et al. 2009). ...
Many species exhibit high‐amplitude, extinction‐prone cycles, for instance due to resource enrichment (called the paradox of enrichment). How do such species manage to persist? One possibility is metapopulation dynamics, but it is unclear if these can mitigate the paradox of enrichment. The paradox of enrichment might increase local population extinction rates, and might also increase the spatial synchrony of population fluctuations because population cycles are easily synchronized by even low rates of dispersal. Spatially‐synchronous population fluctuations would leave no scope for rescue effects, and spatially‐synchronous local extinctions would leave no source for recolonization. We conducted a protist microcosm experiment to test how dispersal and enrichment affect the persistence of predator‐prey metapopulations by altering spatial synchrony. We assembled 54, 4‐patch metapopulations of the protist predator Euplotes patella and its protist prey Tetrahymena pyriformis. Each metapopulation experienced one of nine dispersal rates, crossed with one of six enrichment levels. We tested the effects of enrichment and dispersal on predator‐prey cycle amplitude, predator population extinction risk, spatial synchrony of predator population fluctuations, and predator metapopulation persistence time. Enrichment increased predator‐prey cycle amplitude but had no detectable effect on population extinction risk. Dispersal had no effect on predator‐prey cycle amplitude. Spatial synchrony increased with enrichment and dispersal. Metapopulation persistence decreased with enrichment and spatial synchrony. We conclude that increasing enrichment reduced metapopulation persistence primarily by increasing spatial synchrony of local extinctions, rather than by increasing their frequency. These results highlight the importance of synchrony‐preventing mechanisms for metapopulation persistence. This article is protected by copyright. All rights reserved.
... The presence of a shared predator can release competition between prey species and promote their coexistence in productive open vegetation structures with high risk, and in landscapes that contain spatial variation in habitat riskiness (Bonsall, Bull, Pickup, & Hassell, 2005;Bonsall & Hassell, 2000;DeCesare et al., 2010;Grand & Dill, 1999;Gurevitch, Morrison, & Hedges, 2000). Nevertheless, it seemed that even though foxes were present, the negative association between hares and rabbits in tall shrubs with low risk persisted (see Table 2 dataset D), possibly because rabbit as a stronger competitor prefers burrows in edge habitat with protective shrub cover (Bakker et al., 2005). ...
Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.
... Nevertheless, it seemed that the avoidance in tall shrubs with low risk was not released, possibly because of the thermoregulatory benefits for the hare ) and presence of rabbit burrows . At our study sites, hares and rabbits seem to coexist, probably because of a shared generalist predator (Bonsall & Hassell, 2000), apparent competition, spatial variation in risk factors (i.e., refuges, Bonsall et al., 2005) and segregation in time. ...
... Metacommunity persistence is difficult to study in larger organisms due to longer generation times (but see Bonsall et al. 2002Bonsall et al. , 2005Bull et al. 2006). However, persistence may be studied by quantifying the movements of predators and prey among habitat patches and their effects on prey survival. ...
... This provides insights into mechanisms of metacommunity dynamics operating within generations. Individual-based explanations for short-term metacommunity dynamics include processes, such as predation (Bonsall et al. 2002(Bonsall et al. , 2005Bull et al. 2006), herbivory (Matthiessen et al. 2007), and habitat selection (Resetarits 2005, Binckley andResetarits 2007). The factors influencing dispersal among habitat patches are important to understand as dispersal among habitat patches is thought to be required for long-term metacommunity persistence (Holyoak et al. 2005). ...
... Most experimental metacommunity studies use prototozoans to assess persistence due to their short generation times (Holyoak and Lawler 1996a,b;Forbes and Chase 2002;Kneitel and Miller 2003;Cadotte and Fukami 2005;Cadotte 2006;Hauzy et al. 2007). However, it is important to understand the short-term behaviors that influence dispersal among local patches, such as predation risk (Resetarits 2005), habitat quality (Binckley and Resetarits 2007), patch arrangement (Bull et al. 2006), number of habitat patches (Bonsall et al. 2002(Bonsall et al. , 2005, and connectivity (Matthiessen et al. 2007, Start andGilbert 2016). This study is one of the first experiments to manipulate dispersal rates of predators and prey on prey survival in insects. ...
Trophic interactions are often studied within habitat patches, but among-patch dispersal of individuals may influence local patch dynamics. Metacommunity concepts incorporate the effects of dispersal on local and community dynamics. There are few experimental tests of metacommunity theory using insects compared to those conducted in microbial microcosms. Using connected experimental mesocosms, we varied the density of the leafhopper Agallia constricta Van Duzee (Homoptera: Cicadellidae) and a generalist insect predator, the damsel bug (Nabis spp., Heteroptera: Nabidae), to determine the effects of conspecific and predator density and varying the time available to dispersal among mesocosms on predation rates, dispersal rates, and leafhopper survival. Conspecific and damsel bug density did not affect dispersal rates in leafhoppers, but this may be due to leafhoppers' aversion to leaving the host plants or the connecting tubes between mesocosms hindering leafhopper movement. Leafhopper dispersal was higher in high-dispersal treatments. Survival rates of A. constricta were also lowest in treatments where dispersal was not limited. This is one of the first experimental studies to vary predator density and the time available to dispersal. Our results indicate that dispersal is the key to understanding short-term processes such as prey survival in predator-prey metacommunities. Further work is needed to determine how dispersal rates influence persistence of communities in multigenerational studies.
... As first suggested by Nicholson & Bailey (1935), spatially structured habitats where dispersal between sites is limited may substantially increase the duration of coexistence. This was demonstrated by the classic experiments by Huffaker (1958) and in various other experiments thereafter (Holyoak & Lawler 1996;Kerr et al. 2002Kerr et al. , 2006Bonsall et al. 2005;Cooper et al. 2012), as well as in theoretical studies including agent-based simulations in spatial habitats (Crowley 1981;Hassell et al. 1991;Nowak & May 1992;Durrett & Levin 1994) and metapopulation models (Hastings 1977(Hastings , 1980Hanski & Gilpin 1991;Tilman 1994). There are, however, several distinct spatial mechanisms for coexistence, which may lead to different spatial-temporal patterns. ...
... We analysed data from several spatially subdivided microcosm experiments, including beetles and wasps (Bonsall et al. 2005) and micro-organisms (Holyoak 2000) (Fig. 4ad), as well as from long term in natura monitoring projects (Miller & terHorst 2012;Brooks, A. of Moorea Coral Reef LTER, 2014) (Fig. 4e,f). The two-dimensional histograms in Fig. 4 demonstrate the number of sites for which the numbers of both prey (or hosts) and predators (or parasites) are within a given range. ...
... Finally, our analyses of empirical data from spatially structured microcosms reveal both unimodal and bimodal patterns of spatial abundance distributions (Fig. 4a-d). Specifically, analysing data from experiments by Bonsall et al. (2005) with ...
Explaining the coexistence and distribution of species in time and space remains a fundamental challenge. While species coexistence depends on both local and regional mechanisms, it is sometimes unclear which role each mechanism takes in a given ecosystem. Consequently, it is very hard to predict the response of the ecosystem to environmental changes. Here, we develop a model to study spatial patterns of coexistence, focusing on predator–prey and host–parasite populations. We show, both theoretically and empirically, that these systems may exhibit both local and regional patterns and mechanisms of coexistence. Changes in environmental parameters, such as spatial connectivity, may lead to a transition from regional to local coexistence or it may lead directly to extinction, depending on demographic parameters. This demonstrates the importance of simultaneously analysing interacting mechanisms that act at different spatial scales to understand the response of ecosystems to environmental changes.
... Other experiments have shown that predators and prey persist longest in meta-communities with more patches (Bonsall (Holyoak 2000a). However, in highly connected patches that could be considered a single homogenous habitat, persistence of predators and prey declined due to synchrony of population dynamics (Bonsall et al. 2005(Bonsall et al. , 2002Holyoak 2000b;Vogwill et al. 2009). The importance of asynchrony among patches for maintaining the stability of meta-communities has been supported by several empirical studies. ...
The effects of habitat connectivity on food webs have been studied both empirically and theoretically, yet the question of whether empirical results support theoretical predictions for any food web metric other than species richness has received little attention. Our synthesis brings together theory and empirical evidence for how habitat connectivity affects both food web stability and complexity. Food web stability is often predicted to be greatest at intermediate levels of connectivity, representing a compromise between the stabilizing effects of dispersal via rescue effects and prey switching, and the destabilizing effects of dispersal via regional synchronization of population dynamics. Empirical studies of food web stability generally support both this pattern and underlying mechanisms. Food chain length has been predicted to have both increasing and unimodal relationships with connectivity as a result of predators being constrained by the patch occupancy of their prey. Although both patterns have been documented empirically, the underlying mechanisms may differ from those predicted by models. In terms of other measures of food web complexity, habitat connectivity has been empirically found to generally increase link density but either reduce or have no effect on connectance, whereas a unimodal relationship is expected. In general, there is growing concordance between empirical patterns and theoretical predictions for some effects of habitat connectivity on food webs, but many predictions remain to be tested over a full connectivity gradient, and empirical metrics of complexity are rarely modeled. Closing these gaps will allow a deeper understanding of how natural and anthropogenic changes in connectivity can affect real food webs.
... Collectively, our study highlights the role of intermediate levels of disturbance, intermediate patch connectivity, and heterogeneous resources on metapopulation dynamics and long-term persistence (Bonsall et al. 2005, Bull et al. 2007, Strevens and Bonsall 2010. We examined metapopulations subjected to fast, slow, and no patch dynamics. ...
Human-dominated landscapes often feature patches that fluctuate in suitability through space and time, but there is little experimental evidence relating the consequences of dynamic patches for species persistence. We used a spatially and temporally dynamic metapopulation model to assess and compare metapopulation capacity and persistence for red flour beetles (Tribolium castaneum) in experimental landscapes differentiated by resource structure, patch dynamics (destruction and restoration) and connectivity. High connectivity increased the colonization rate of beetles, but this effect was less pronounced in heterogeneous relative to homogeneous landscapes. Higher connectivity and faster patch dynamics increased extinction rates in landscapes. Lower connectivity promoted density-dependent emigration. Heterogeneous landscapes containing patches of different carrying capacity enhanced landscape-level occupancy probability. Highest metapopulation capacity and persistence was observed in landscapes with heterogeneous patches, low connectivity and slow patch dynamics. Control landscapes with no patch dynamics exhibited rapid declines in abundance and approached extinction due to increased adult mortality in the matrix, higher pupal cannibalism by adults and extremely low rates of exchange between remaining habitable patches. Our results highlight the role of intermediate patch dynamics, intermediate connectivity and the nature of density dependence of emigration for persistence of species in heterogeneous landscapes. Our results also demonstrate the importance of incorporating local dynamics into estimation of metapopulation capacity for conservation planning.
Read More: http://www.esajournals.org/doi/abs/10.1890/14-0044.1
... This suggests that understanding the spatial scale of host regulation, versus that of disease transmission, will be crucially important in understanding the dynamical impacts of vectors and indirect interactions. While the role of metapopulations (populations linked through limited dispersal) have recently been shown to mitigate the effects of apparent competition mediated by natural enemies (Bonsall et al., 2005;Bull et al., 2006Bull et al., , 2007, the effects of regional spatial structures on the persistence and dynamics of disease (and particular vector-borne disease) in multi-host systems still remains poorly explored. We expect this to be an important direction for future research. ...
Infectious disease influences the dynamics of host populations and the structure of species communities via impacts on host demography. Species that share infectious diseases are well-known to interact indirectly through the process of apparent competition, but there has been little attention given to the role of vectors in these indirect interactions. Here we explore how vector-borne disease and host-vector interactions can drive apparent competitive interactions. We show that different facets of the ecology associated with vector-host-host interactions affect the structure of the three-species assemblage. Crucially, the patterns associated with invasion of alternative hosts, the spread of the infectious disease by the vector, and the dynamics of the community interactions are influenced by the mode of transmission. We highlight the role of alternative hosts on disease amplification, dilution and magnification and discuss the results with reference to recent developments in apparent competition and community structure.
