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    Pairwise interact-and-imitate dynamics

    The modelConsider a unit-mass population of agents who repeatedly interact in pairs to play a symmetric stage game. The set of strategies available to each agent is finite and denoted by (S equiv {1, ldots , n}). A population state is a vector (x in X equiv {x in {mathbb{R}}^n_+: sum _{i in S} x_i = 1}), with (x_i) the fraction of the population playing strategy (i in S). Payoffs are described by a function (F: S times S rightarrow {mathbb{R}}), where F(i, j) is the payoff received by an agent playing strategy i when the opponent plays strategy j. As a shorthand, we refer to an undirected pair of individuals, one playing i and the other playing j, as an ij pair. The set of all possible undirected pairs is denoted by (mathscr {P}).The interaction structure is modeled as a function (p : X times mathscr {P} rightarrow left[ 0, 1/2 right] ) subject to (sum _{ij in mathscr {P}} p_{ij}(x)=1/2) (since the mass of pairs is half the mass of agents), with (p_{ij}(x)) indicating the mass of ij pairs formed in state x. Note that the mass of ij pairs can never exceed (min {x_i,x_j}), that is, (p_{ij}(x) le min {x_i,x_j}) for all x. We assume that p is continuous in X, and that (p_{ij}(x) > 0) if and only if (x_i > 0) and (x_j > 0 )—meaning that the probability of an ij pair being formed is strictly positive if and only if strategies i and j are played by someone. In the case of uniform random matching, (p_{ii} = x_i^2/2) and (p_{ij} = x_i x_j) for any i and (j ne i).The revision protocol is modeled as a function (phi : X times S times S rightarrow [-1,1]), where (phi _{ij}(x) in [-1,1]) is the probability that an ij pair will turn into an ii pair minus the probability that it will turn into a jj pair, conditional on the population state being x and an ij pair being formed. We assume that (phi ) is continuous in X. We note that by construction (phi _{ij}=-phi _{ji}) for all (i,j in S), and hence (phi _{ii}=0) for all (i in S). Our main assumption on the revision protocol is the following, which is met, among others, by pairwise proportional imitative and imitate-if-better rules22.
    Assumption 1

    For every (x in X), (phi _{ij}(x) > 0) if (F(i,j) > F(j,i)).
    In what follows we consider a dynamical system in continuous time with state space X, characterized by the following equation of motion.

    Definition 1

    (Pairwise interact-and-imitate dynamics—PIID) For every (x in X) and every (i in S):$$begin{aligned} dot{x}_i = sum _{j in S} p_{ij}(x) phi _{ij}(x). end{aligned}$$
    (1)

    Main findingsGlobal asymptotic convergenceIn any purely imitative dynamics, if (x_i(t)=0), then (x_i(t^{prime})=0) for every (t^{prime} > t). This implies that we cannot hope for global asymptotic convergence in a strict sense. Thus, to assess convergence towards a certain state x in a meaningful way, we restrict our attention to those states where all strategies that have positive frequency in x have positive frequency as well. We denote by (X_x) the set of states whose support contains the support of x.

    Definition 2

    (Supremacy) Strategy (iin S) is supreme if (F(i,j) >F(j,i)) for every (j in S setminus {i}).
    We note that under PIID, the concept of supremacy is closely related to that of asymmetry33,34, in that (F(i,j) > F(j,i)) implies that agents can only switch from strategy j to strategy i.

    Proposition 1

    If (i in S) is a supreme strategy, then state (x^* equiv left{ x in X : x_i = 1 right} ) is globally asymptotically stable for the dynamical system with state space (X_{x^*}) and PIID as equation of motion.
    Relation to replicator dynamicsTo further characterize the dynamics induced by the pairwise interact-and-imitate protocol, we make two additional assumptions. First, matching is uniformly random, meaning that everyone in the population has the same probability of interacting with everyone else; formally, (p_{ii} = x_i^2/2) and (p_{ij} = x_i x_j) for all i and (j ne i). Second, the probability that an agent has to imitate the opponent is proportional to the difference in their payoffs if the opponent’s payoff exceeds her own, and is zero otherwise. As a consequence, (phi _{ij} = F(i,j) – F(j,i)) up to a proportionality factor. Let

    (F left( i, x right) :=sum _j x_j F left( i, j right) ),

    (F left( x, i right) :=sum _j x_j F left( j, i right) ), and

    ( F left( x, x right) :=sum _i sum _j x_i x_j F left( i, j right) ).

    Under these assumptions, at any point in time, the motion of (x_i) is described by:$$begin{aligned} dot{x}_i&= sum _{j ne i} x_j x_i left[ F left( i, j right) – F left( j, i right) right] = x_i sum _{j} x_j left[ F left( i, j right) – F left( j, i right) right] nonumber \&= x_i left[ F left( i, x right) – F left( x, i right) right] , end{aligned}$$
    (2)
    which is a modified replicator equation. According to (2), for every strategy i chosen by one or more agents in the population, the rate of growth of the fraction of i-players, (dot{x}_i / x_i), equals the difference between the expected payoff from playing i in state x and the average payoff received by those who are matched against an agent playing i. In contrast, under standard replicator dynamics35, the fraction of agents playing i varies depending on the excess payoff of i with respect to the current average payoff in the whole population, i.e., (dot{x}_i = x_i left[ F left( i, x right) – F left( x, x right) right] ).A noteworthy feature of replicator dynamics is that they are always payoff monotone: for any (i,j in S), the proportions of agents playing i and j grow at rates that are ordered in the same way as the expected payoffs from the two strategies36. In the case of PIID, this result fails.

    Proposition 2

    Pairwise-Interact-and-Imitate dynamics need not satisfy payoff monotonicity.
    To verify this, it is sufficient to consider any symmetric (2 times 2) game where (F left( i, j right) > F left( j, i right) ) but (F left( j, x right) > F left( i, x right) ) for some (x in X), meaning that i is the supreme strategy but j yields a higher expected payoff in state x. See Fig. 1 for an example where, in the case of uniform random matching, the above inequalities hold for any x; if strategies are updated according to the interact-and-imitate protocol, then this game only admits switches from i to j, therefore violating payoff monotonicity. Proposition 2 can have important consequences, including the survival of pure strategies that are strictly dominated.Survival of strictly dominated strategiesAn recurring topic in evolutionary game theory is to what extent does support exist for the idea that strictly dominated strategies will not be played. It has been shown that if strategy i does not survive the iterated elimination of pure strategies strictly dominated by other pure strategies, then the fraction of the population playing i will converge to zero in all payoff monotone dynamics37,38. This result does not hold in our case, as PIID is not payoff monotone.More precisely, under PIID, a strictly dominated strategy may be supreme and, therefore, not only survive but even end up being adopted by the whole population. This suggests that from an evolutionary perspective, support for the elimination of dominated strategies may be weaker than is often thought. Our result contributes to the literature on the conditions under which evolutionary dynamics fail to eliminate strictly dominated strategies in some games, examining a case which has not yet been studied39.To see that a strictly dominated strategy may be supreme, consider the simple example shown in Fig. 1. Here each agent has a strictly dominant strategy to play A; however, since the payoff from playing B against A exceeds that from playing A against B, strategy B is supreme. Thus, by Proposition 1, the population state in which all agents choose B is globally asymptotically stable.Figure 1A game where the supreme strategy is strictly dominated.Full size imageFigure 1 can also be used to comment on the relation between a supreme strategy and an evolutionary stable strategy, which is a widely used concept in evolutionary game theory40,41. Indeed, while B is the supreme strategy, A is the unique evolutionary stable strategy because it is strictly dominant. However, if F(B, A) were reduced below 2, holding everything else constant, then B would become both supreme and evolutionary stable. We therefore conclude that no particular relation holds between evolutionary stability and supremacy: neither one property implies the other, nor are they incompatible.ApplicationsHaving obtained general results for the class of finite symmetric games, we now restrict the discussion to the evolution of behavior in social dilemmas. We show that if the conditions of Proposition 1 are met, then inefficient conventions emerge in the Prisoner’s Dilemma, Stag Hunt, Minimum Effort, and Hawk–Dove games. Furthermore, this result holds both without and with the assumption that agents interact assortatively.Ineffectiveness of assortmentConsider the (2 times 2) game represented in Fig. 2. If (c > a > d > b), then mutual cooperation is Pareto superior to mutual defection but agents have a dominant strategy to defect. The resulting stage game is the Prisoner’s Dilemma, whose unique Nash equilibrium is (B, B). Moreover, since (F (B,A) > F(A,B)), B is the supreme strategy and the population state in which all agents defect is globally asymptotically stable.We stress that defection emerges in the long run for every matching rule satisfying our assumptions, and therefore also in the case of assortative interactions. Assortment reflects the tendency of similar people to clump together, and can play an important role in the evolution of cooperation42,43,44,45. Intuitively, when agents meet assortatively, the risk of cooperating in a social dilemma may be offset by a higher probability of playing against other cooperators. However, under PIID, this is not the case: the decision whether to adopt a strategy or not is independent of expected payoffs, and like-with-like interactions have no effect except to reduce the frequency of switches from A to B.Figure 2A (2 times 2) stage game.Full size imageEmergence of the maximin conventionIf (a > c > b), (a > d) and (d > b), then the game in Fig. 2 becomes a Stag Hunt game, which contrasts risky cooperation and safe individualism. The payoffs are such that both (left( A, Aright) ) and (left( B, Bright) ) are strict Nash equilibria, that (left( A, Aright) ) is Pareto superior to (left( B, Bright) ), and that B is the maximin strategy, i.e., the strategy which maximizes the minimum payoff an agent could possibly receive. We also assume that (a + c ne c + d), so that one of A and B is risk dominant46. If (a + b > c + d), then A (Stag) is both payoff and risk dominant. When the opposite inequality holds, the risk dominant strategy is B (Hare).Since (F (B,A) > F(A,B)), B is supreme independently of whether or not it is risk dominant to cooperate. This can result in large inefficiencies because, in the long run, the process will converge to the state in which all agents play the riskless strategy regardless of how rewarding social coordination is. As in the case of the Prisoner’s Dilemma, this holds for all matching rules satisfying our assumptions.Evolution of effort exertionIn a minimum effort game, agents simultaneously choose a strategy i, usually interpreted as a costly effort level, from a finite subset S of ({mathbb{R}}). An agent’s payoff depends on her own effort and on the minimum effort in the pair:$$begin{aligned} F left( i, j right) = alpha min left{ i, j right} – beta i , end{aligned}$$where (beta > 0) and (alpha > beta ) are the cost and benefit of effort, respectively. From a strategic viewpoint, this game can be seen as an extension of the Stag Hunt to cases where there are more than two actions. The best response to a choice of j by the opponent is to choose j as well, and coordinating on any common effort level gives a Nash equilibrium. Nash outcomes can be Pareto-ranked, with the highest-effort equilibrium being the best possible outcome for all agents. Thus, choosing a high i is rationalizable and potentially rewarding but may also result in a waste of effort.Under PIID, any (i > j) implies (phi _{ij} < 0) by Assumption 1, meaning that agents will tend to imitate the opponent when the opponent’s effort is lower than their own. The supreme strategy is therefore to exert as little effort as possible, and the population state in which all agents choose the minimum effort level is the unique globally asymptotically stable state.Emergence of aggressive behaviorConsider again the payoff matrix shown in Fig. 2. If (c > a > b > d), then the stage game is a Hawk–Dove game, which is often used to model the evolution of aggressive and sharing behaviors. Interactions can be framed as disputes over a contested resource. When two Doves (who play A) meet, they share the resource equally, whereas two Hawks (who play B) engage in a fight and suffer a cost. Moreover, when a Dove meets a Hawk, the latter takes the entire prize. Again we have that (F (A,B) < F(B,A)), implying that B is the supreme strategy and that the state where all agents play Hawk is the sole asymptotically stable state.The inefficiency that characterizes the (B, B) equilibrium in the Hawk–Dove game arises from the cost that Hawks impose on one another. This can be viewed as stemming from the fact that neither agent owns the resource prior to the interaction or cares about property. A way to overcome this problem may be to introduce a strategy associated with respect for ownership rights, the Bourgeois, who behaves as a Dove or Hawk depending on whether or not the opponent owns the resource41. If we make the standard assumption that each member of a pair has a probability of 1/2 to be an owner, then in all interactions where a Bourgeois is involved there is a 50 percent chance that she will behave hawkishly (i.e., fight for control over the resource) and a 50 percent chance that she will act as a Dove.Let R and C denote the agent chosen as row and column player, respectively, and let (omega _R) and (omega _C) be the states of the world in which R and C owns the resource. The payoffs of the resulting Hawk–Dove–Bourgeois game are shown in Fig. 3. If agents behave as expected payoff maximizers, then All Bourgeois can be singled out as the unique asymptotically stable state. Under PIID, this is not so; depending on who owns the resource, an agent playing C against an opponent playing B may either fight or avoid conflict and let the opponent have the prize. It is easy to see that (F left( C, B mid omega _R right) = F left( B,C mid omega _C right) = d), meaning that the payoff from playing C against B, conditional on owning the resource, equals the payoff from playing B against C conditional on not being an owner. In contrast, the payoff from playing C against B, conditional on not owning the resource, is always worse than that of the opponent, i.e., (F left( C, B mid omega _C right) = b < c = F left( B, C mid omega _R right) ). Thus, in every state of the world, B (Hawk) yields a payoff that is greater or equal to that from C (Bourgeois). Moreover, since (F left( B,A right) > F left( A, B right) ) in both states of the world, strategy B is weakly supreme by Definition 4, and play unfolds as an escalation of hawkishness and fights.Figure 3The Hawk–Dove–Bourgeois game.Full size image More

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    High-throughput 16S rRNA gene sequencing of the microbial community associated with palm oil mill effluents of two oil processing systems

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    Red light, green light: both signal ‘go’ to deadly algae

    Green and red lighting might be good for migratory birds and sea turtles, but could have undesirable effects if marine algae are present. Credit: Getty

    Ecology
    24 June 2021
    Red light, green light: both signal ‘go’ to deadly algae

    Artificial lighting thought to be more wildlife-friendly than white light could encourage algal blooms.

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    Green or red lights in seaside areas have been proposed as alternatives to white light to protect wildlife. But new experiments show that exposure to red or green light at night boosts the growth of some ocean algae — including species known to rob waters of oxygen.Little is known about the impact of artificial light on marine life, even though many brightly lit cities are coastal. To address that knowledge gap, Sofie Spatharis at the University of Glasgow, UK, and her colleagues exposed a mix of microscopic marine algae collected from Scottish waters to standard white light. They also exposed the mixture to red and green lights, which have been proposed to minimize impacts on sea turtles and migratory seabirds, respectively.The team found that all light colours enhanced growth of the microalgae mix. Red light had the most pronounced effect, doubling the number of cells produced. The proportions of species in the mixture also shifted: both red and green light especially favoured growth of harmful species in the Skeletonema genus, which form dense blooms that are deadly to fish.

    Proc. R. Soc. B (2021)

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    Coordinated gas release among the physostomous fish sprat (Sprattus sprattus)

    Study areaThe study was carried out in Bunnefjorden, Norway. The fjord froze over from January to April and we here analyze data from ice-free conditions in early winter (12 Nov to 2 Dec 2009). Bunnefjorden is a 150 m deep inner branch of the Oslofjord, and a 57 m deep sill at the entrance restricts water exchange with the outer part of the fjord. Klevjer and Kaartvedt24 provide a map of the study area. The fjord branch normally becomes hypoxic in the lower part of the water column. During the current study, oxygen contents were 2–3 ml l−1 between 15 and 60 m, while waters below 70–80 m were severely hypoxic and devoid of fishes9.Studies of overwintering sprat have been undertaken in Bunnefjorden during several winters, and the biology of sprat as well as the identity of the main acoustic targets in the fjord are well established9,18,24. In the winter of the current study, catches from 33 trawl samples were dominated by sprat; with ~ 40 times higher catches than the next most abundant species, herring (Clupea harengus)9.Study designSolberg and Kaartvedt9 and Solberg et al.18 provide details on methods, and we here only give a summary of the acoustic setup. In short, upward-looking Simrad EK 60 echosounders kept in pressure-proof casings were deployed at the bottom (150 m) and in buoys (80 and 30 m) for enhanced resolution in shallower part of the water column. Cables for electricity and transfer of data to a PC on shore enabled continuous operation of the systems. We here use the data from the shallowest echosounder (200 kHz) that provided superior resolution in near-surface water, though did not cover the full depth range of the population distribution. Echograms from the deeper located echosounders covering the whole (inhabited) water column and showing the full diel population behavior are given in Solberg and Kaartvedt9 and Solberg et al.18.Records of gas releaseReleased gas appeared as ascending lines in the echogram (Fig. 4). We quantified the release as explained by Solberg and Kaartvedt9. We only included ascending traces connected to the acoustic record of a fish, but without enumerating the release per individual fish. Since the same fish may release several bursts of bubbles within a short time interval, we here pooled any sequences of gas release within a 10-s period as one event. This procedure will also exclude cases with several different individuals releasing bubbles in the course of this short time interval, yet we chose this conservative approach not to generate an artificial high connection of gas releases between the fishes.Figure 4Echogram showing sprat releasing gas, with every oblique line representing one release event and lines with a different angle to the release events representing swimming sprat. Colors represent the volume-backscatter coefficient (Sv).Full size imageAnalyses of dataThe frequency of gas releases varied with time, both within a day and between the weeks. Such patterns compare to service systems like call centres and hospital emergency rooms25 that can be modelled as a Poisson process26,27. We therefore started our analysis with the statistical procedure suggested by Brown et al.28 in their influential analysis of the call dynamics in a banking call centre. The first step is to subdivide the day into time intervals, which are short enough to consider event rates as approximately constant. Here we chose to investigate alternative periods of respectively 1, 5, and 30 min, as well as 1, 2, 4, and 6 h. At the longest interval, the peaks in the gas release intensity are expected to be the result of a non-constant Poisson parameter, and therefore more likely to induce rejection of the null-hypothesis of a homogenous random process. In contrast, we expect to find higher concordance with a random process for the short intervals of 5 min. In assessing connectivity among gas bubble releases, we formulate a new model allowing for a formal test of non-randomness (summarized in Fig. 5). We name this approach the simulated connectivity test (S-CON test), which we implemented in R29, with the code being available in the Supplementary appendix.Figure 5Illustration of the steps related to the simulated connectivity test (S-CON test). Bubbles occurring within 10 s are pooled into single release events. We then determine the connectivity of each release event—aka the number of release events within the following 30 s time window. From these, we calculate the average connectivity for a specified period (1, 5, 30 min, 1-, 2-, 4- and 6-h intervals). In a final step, we compare the observed average connectivity to the critical value which is defined as the 95th percentile of 1000 simulations of random placements of the same number of releases. If the observed connectivity is larger than the critical value, we reject the null hypothesis of random gas releases for the specified time interval.Full size imageIf there is a common physiological reason or some form of communication among sprat, a burst of gas release is likely followed by subsequent releases. Thus, it is reasonable to assume that the total number of releases within a short time interval like 30 s would be effective in detecting dependencies between the releases. We therefore define the concept of connectivity as follows:Let the gas be released at times T1, T2, …, Tn and define the connectivity at each event as the numbers of records within the following 30 s. The average connectivity in any considered window of the investigated time-period (for example a window of 1 h) is defined as the average connectivity of all cases of connectivity within the considered window (see also Fig. 5).In order to test the null hypothesis of no dependency between gas bubble releases, we compare the measured average connectivity in the data set with the simulation of 1000 random placements of the total number of observations in a given time window. For example, if we consider a window of 30 min with 15 release events having an average connectivity of 2.1, we performed 1000 random placements of 15 points between 1 and 1800. In this way, we get 1000 simulated values of the average connectivity, from which we pick out the critical 95th percentile, following the common significance level of 0.05 in biology. If the observed average connectivity is larger than this critical value, we reject the null-hypothesis and conclude that the releases of gas bubbles are dependent random variables. Thus, if our example obtains a critical value of 1.7, the null-hypothesis of random arrival times of bubbles is rejected (because the observed value of 2.1 is larger than the critical value of 1.7).Since a dependency between the fish will induce a higher concentration of release events than produced by random releases, we expect the average connectivity to be quite sensitive to the alternative hypothesis of dependent arrival times. Also, note that the concept of connectivity has a combinatory nature, so we need only require that the considered window contains at least two releases of gas bubbles. In contrast, alternative approaches using Kolmogorov–Smirnov tests28 are based on the cumulative distribution function and therefore require at least five observed bubble releases.To test the dependency of the results on the chosen time interval, we also ran the analysis using connectivity intervals of 25 and 35 s, which revealed some variability to the estimates of non-random bubble release (Fig. 3) but did not influence the general pattern. We also tested whether the interval within which we consider subsequent bubbles to be part of one single release event influences our results. The more we consider sequential bubbles to be independent of each other, i.e. their own release event, the higher the proportion of non-random gas release and vice versa.Fish abundanceTo exclude the possibility that apparent connectivity would be a mere result of fluctuating fish abundance, we tested whether the number of released bubbles is a function of fish biomass. For this, we first calculated the total number of gas release events within 30-min periods. We then compared these values to the summed surface integrated acoustic scattering coefficient (SA) for the same periods and for the same depth interval (upper 30 m), assuming that the integrated scattering coefficient (SA) serves as a proxy for the total fish biomass9. We filtered the scattering data to remove noise from non-biological sources prior to use. Both variables were log-transformed prior to analysis. We then fitted a linear model of the two variables using generalized least squares. To account for temporal autocorrelation in the data, we also included a correlation structure of type 1 (corAR1). The analysis was done in R29 using the nlme package30.Ethics declarationsLive animals (fish) were not used in this study. More

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    Migratory birds aid the redistribution of plants to new climates

    NEWS AND VIEWS
    23 June 2021

    Migratory birds aid the redistribution of plants to new climates

    Birds that travel long distances can disperse seeds far and wide. An assessment of the timing and direction of European bird migration reveals how these patterns might affect seed dispersal as the planet warms.

    Barnabas H. Daru

     ORCID: http://orcid.org/0000-0002-2115-0257

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    Barnabas H. Daru

    Barnabas H. Daru is in the Department of Life Sciences, Texas A&M University-Corpus Christi, Corpus Christi, Texas 78412, USA.

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    The rapid pace of global warming and its effects on habitats raise the question of whether species are able to keep up so that they remain in suitable living conditions. Some animals can move fast to adjust to a swiftly changing climate. Plants, being less mobile, rely on means such as seed dispersal by animals, wind or water to move to new areas, but this redistribution typically occurs within one kilometre of the original plant1. Writing in Nature, González-Varo et al.2 shed light on the potential capacity of migratory birds to aid seed dispersal.When the climate in a plant’s usual range becomes hotter than it can tolerate, it must colonize new, cooler areas that might lie many kilometres away. It is not fully clear how plants distribute their seeds across great distances, let alone how they cross geographical barriers. One explanation for long-distance seed dispersal is through transport by migratory birds. Such birds ingest viable seeds when eating fruit (Fig. 1) and can move them tens or hundreds of kilometres outside the range of a plant species3. In this mode of dispersal, the seeds pass through the bird’s digestive tract unharmed4,5 and are deposited in faeces, which provides fertilizer that aids plant growth. In the case of European migratory birds, for example, the direction of seed dispersal will depend on whether the timing of fruit production coincides with a bird’s southward trip to warmer regions around the Equator, or northward to cooler regions. Many aspects of this process have been a mystery until now.

    Figure 1 | A young blackcap bird (Sylvia atricapilla) eating elderberries.Credit: Getty

    González-Varo and colleagues report how plants might be able to keep pace with rapid climate change through the help of migrating birds. The authors analysed the fruiting times of plants, patterns of bird migration and the interactions between fruit-eating birds and fleshy-fruited plants across Europe. Plants with fleshy fruits were chosen for this study because most of their seed transport is by migratory birds6, and because fleshy-fruited plants are an important component of the woody-plant community in Europe. The common approach until now has been to predict plant dispersal and colonization using models fitted to abiotic factors, such as the current climate. González-Varo et al. instead analysed an impressive data set of 949 different seed-dispersal interactions between bird and plant communities, together with data on entire fruiting times and migratory patterns of birds across Europe. The researchers also analysed DNA traces from bird faeces to identify the plants and birds responsible for seed dispersal.
    Read the paper: Limited potential for bird migration to disperse plants to cooler latitudes
    The authors hypothesized that the direction of seed migration depends on how the plants interact with migratory birds, the frequency of these interactions or the number of bird species that might transport seeds from each plant species. González-Varo and colleagues found that 86% of plant species studied might have seeds dispersed by birds during their southward trip towards drier and hotter equatorial regions in autumn, whereas only about one-third of the plant species might be dispersed by birds migrating north in spring. This dispersal trend was more pronounced in temperate plants than in the Mediterranean plant communities examined. These results are in general agreement with well-known patterns of fruiting times and bird migrations. For example, the fruit of most fleshy-fruited plants in Europe ripens at a time that coincides with when birds migrate south towards the Equator7.Perhaps the most striking feature of these inferred seed movements is the observation that 35% of plant species across European communities, which are closely related on the evolutionary tree (phylogenetically related), might benefit from long-distance dispersal by the northward journey of migratory birds. This particular subset of plants tends to fruit over a long period of time, or has fruits that persist over the winter. This means that the ability of plants to keep up with climate change could be shaped by their evolutionary history — implying that future plant communities in the Northern Hemisphere will probably come from plant species that are phylogenetically closely related and that have migrated from the south. Or, to put it another way, the overwhelming majority of plant species that are dispersed south towards drier and hotter regions at the Equator will probably be less able to keep pace with rapid climate change in their new locations than will the few ‘winners’ that are instead dispersed north to cooler climates. This has implications for understanding how plants will respond to climate change, and for assessing ecosystem functions and community assembly at higher levels of the food chain. However, for seeds of a given plant species, more evidence is needed to assess whether passing through the guts of birds affects germination success.To determine which birds might be responsible for the plant redistributions to cooler climates in the north, the authors categorized European bird migrants into Palaearctic (those that fly to southern Europe and northern Africa during their non-breeding season) and Afro-Palaearctic (those that winter in sub-Saharan Africa). Only a few common Palaearctic migrants, such as the blackcap (Sylvia atricapilla; Fig. 1) or blackbird (Turdus merula), provide most of this crucial dispersal service northwards to cooler regions across Europe. Because migratory birds are able to relocate a small, non-random subset of plants, this could well have a strong influence on the types of plant community that will form under climate-change conditions.
    A bird’s migration decoded
    A major problem, however, is that the role of these birds in dispersing seeds over long distances is already at risk from human pressures and environmental changes8. Understanding these large-scale seed-dispersal interactions offers a way for targeted conservation actions to protect the areas that are most vulnerable to climate change. This could include boosting protection efforts in and around the wintering grounds of migratory birds — locations that are already experiencing a rise in human pressures, such as illegal bird hunting.González-Varo and colleagues’ focus on seed dispersal across a Northern Hemisphere region means that, as with most ecological analyses, the results are dependent on scale, which can cause issues when interpreting data9. Because the Northern Hemisphere has more land area and steeper seasonal temperature gradients than the Southern Hemisphere does, seed-dispersal interactions might have different patterns from those occurring in the Southern Hemisphere or in aquatic systems.For example, seed-eating birds from the genus Quelea migrate from the Southern Hemisphere to spend the dry season in equatorial West Africa, then move southwards again when the rains arrive. Their arrival in southern Africa usually coincides with the end of the wet season in this region, when annual grass seeds are in abundance. It will be worth investigating whether migratory birds in the Southern Hemisphere also influence the redistribution of plant communities during global warming. Likewise, exploring the long-distance dispersal of seeds of aquatic plants, such as seagrasses10 by water birds, is another area for future research that might benefit from González-Varo and colleagues’ methods.This study provides a great example of how migratory birds might assist plant redistribution to new locations that would normally be difficult for them to reach on their own, and which might offer a suitable climate. As the planet warms, understanding how such biological mechanisms reorganize plant communities complements the information available from climate-projection models, which offer predictions of future species distributions.

    doi: https://doi.org/10.1038/d41586-021-01547-1

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    Competing Interests
    The author declares no competing interests.

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