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20 Sep 2018
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When higher carrying capacities lead to faster propagation

When the dispersal of the many outruns the dispersal of the few

Recommended by based on reviews by Yuval Zelnik and 1 anonymous reviewer

Are biological invasions driven by a few pioneers, running ahead of their conspecifics? Or are these pioneers constantly being caught up by, and folded into, the larger flux of propagules from the established populations behind them?
In ecology and beyond, these two scenarios are known as "pulled" and "pushed" fronts, and they come with different expectations. In a pushed front, invasion speed is not just a matter of how good individuals are at dispersing and settling new locations. It becomes a collective, density-dependent property of population fluxes. And in particular, it can depend on the equilibrium abundance of the established populations inside the range, i.e. the species’ carrying capacity K, factoring in its abiotic environment and biotic interactions.
This realization is especially important because it can flip around our expectations about which species expand fast, and how to manage them. We tend to think of initial colonization and long-term abundance as two independent axes of variation among species or indeed as two ends of a spectrum, in the classic competition-colonization tradeoff [1]. When both play into invasion speed, good dispersers might not outrun good competitors. This is useful knowledge, whether we want to contain an invasion or secure a reintroduction.
In their study "When higher carrying capacities lead to faster propagation", Haond et al [2] combine mathematical analysis, Individual-Based simulations and experiments to show that various mechanisms can cause pushed fronts, whose speed increases with the carrying capacity K of the species. Rather than focus on one particular angle, the authors endeavor to demonstrate that this qualitative effect appears again and again in a variety of settings.
It is perhaps surprising that this notable and general connection between K and invasion speed has managed to garner so little fame in ecology. A large fraction of the literature employs the venerable Fisher-KPP reaction-diffusion model, which combines local logistic growth with linear diffusion in space. This model has prompted both considerable mathematical developments [3] and many applications to modelling real invasions [4]. But it only allows pulled fronts, driven by the small populations at the edge of a species range, with a speed that depends only on their initial growth rate r.
This classic setup is, however, singular in many ways. Haond et al [2] use it as a null model, and introduce three mechanisms or factors that each ensure a role of K in invasion speed, while giving less importance to the pioneers at the border.
Two factors, the Allee effect and demographic stochasticity, make small edge populations slower to grow or less likely to survive. These two factors are studied theoretically, and to make their claims stronger, the authors stack the deck against K. When generalizing equations or simulations beyond the null case, it is easy to obtain functional forms where the parameter K does not only play the role of equilibrium carrying capacity, but also affects dynamical properties such as the maximum or mean growth rate. In that case, it can trivially change the propagation speed, without it meaning anything about the role of established populations behind the front. Haond et al [2] avoid this pitfall by disentangling these effects, at the cost of slightly more peculiar expressions, and show that varying essentially nothing but the carrying capacity can still impact the speed of the invasion front.
The third factor, density-dependent dispersal, makes small populations less prone to disperse. It is well established empirically and theoretically that various biological mechanisms, from collective organization to behavioral switches, can prompt organisms in denser populations to disperse more, e.g. in such a way as to escape competition [5]. The authors demonstrate how this effect induces a link between carrying capacity and invasion speed, both theoretically and in a dispersal experiment on the parasitoid wasp, Trichogramma chilonis.
Overall, this study carries a simple and clear message, supported by valuable contributions from different angles. Although some sections are clearly written for the theoretical ecology crowd, this article has something for everyone, from the stray physicist to the open-minded manager. The collaboration between theoreticians and experimentalists, while not central, is worthy of note. Because the narrative of this study is the variety of mechanisms that can lead to the same qualitative effect, the inclusion of various approaches is not a gimmick, but helps drive home its main message. The work is fairly self-contained, although one could always wish for further developments, especially in the direction of more quantitative testing of these mechanisms.
In conclusion, Haond et al [2] effectively convey the widely relevant message that, for some species, invading is not just about the destination, it is about the many offspring one makes along the way.


[1] Levins, R., & Culver, D. (1971). Regional Coexistence of Species and Competition between Rare Species. Proceedings of the National Academy of Sciences, 68(6), 1246–1248. doi: 10.1073/pnas.68.6.1246
[2] Haond, M., Morel-Journel, T., Lombaert, E., Vercken, E., Mailleret, L., & Roques, L. (2018). When higher carrying capacities lead to faster propagation. BioRxiv, 307322. doi: 10.1101/307322
[3] Crooks, E. C. M., Dancer, E. N., Hilhorst, D., Mimura, M., & Ninomiya, H. (2004). Spatial segregation limit of a competition-diffusion system with Dirichlet boundary conditions. Nonlinear Analysis: Real World Applications, 5(4), 645–665. doi: 10.1016/j.nonrwa.2004.01.004
[4] Shigesada, N., & Kawasaki, K. (1997). Biological Invasions: Theory and Practice. Oxford University Press, UK.
[5] Matthysen, E. (2005). Density-dependent dispersal in birds and mammals. Ecography, 28(3), 403–416. doi: 10.1111/j.0906-7590.2005.04073.x

When higher carrying capacities lead to faster propagationMarjorie Haond, Thibaut Morel-Journel, Eric Lombaert, Elodie Vercken, Ludovic Mailleret & Lionel Roques<p>This preprint has been reviewed and recommended by Peer Community In Ecology ( Finding general patterns in the expansion of natural populations is a major challenge in ecology and invasion biology...Biological invasions, Colonization, Dispersal & Migration, Experimental ecology, Population ecology, Spatial ecology, Metacommunities & Metapopulations, Theoretical ecologyMatthieu Barbier Yuval Zelnik2018-04-25 10:18:48 View
10 Jun 2018
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A reply to “Ranging Behavior Drives Parasite Richness: A More Parsimonious Hypothesis”

Does elevated parasite richness in the environment affect daily path length of animals or is it the converse? An answer bringing some new elements of discussion

Recommended by based on reviews by 2 anonymous reviewers

In 2015, Brockmeyer et al. [1] suggested that mandrills (Mandrillus sphinx) may accept additional ranging costs to avoid heavily parasitized areas. Following this paper, Bicca-Marques and Calegaro-Marques [2] questioned this interpretation and presented other hypotheses. To summarize, whilst Brockmeyer et al. [1] proposed that elevated daily path length may be a consequence of elevated parasite richness, Bicca-Marques and Calegaro-Marques [2] viewed it as a cause. In this current paper, Charpentier and Kappeler [3] respond to some of the criticisms by Bicca-Marques and Calegaro-Marques and discuss the putative parsimony of the two competing scenarios. The manuscript is interesting and focuses on an important question concerning the discussion about the social organization and home range use in wild mandrills. This answer helps to move this debate forward and should stimulate more empirical studies of the role of environmentally-transmitted parasites in shaping ranging and movement patterns of wild vertebrates. Given the elements this paper brings to the topics, it should have been published in American Journal of Primatology, the journal that published the two previous articles.


[1] Brockmeyer, T., Kappeler, P. M., Willaume, E., Benoit, L., Mboumba, S., & Charpentier, M. J. E. (2015). Social organization and space use of a wild mandrill (Mandrillus sphinx) group. American Journal of Primatology, 77(10), 1036–1048. doi: 10.1002/ajp.22439
[2] Bicca-Marques, J. C., & Calegaro-Marques, C. (2016). Ranging behavior drives parasite richness: A more parsimonious hypothesis. American Journal of Primatology, 78(9), 923–927. doi: 10.1002/ajp.22561
[3] Charpentier, M. J., & Kappeler, P. M. (2018). A reply to “Ranging Behavior Drives Parasite Richness: A More Parsimonious Hypothesis.” ArXiv:1805.08151v2 [q-Bio]. Retrieved from

A reply to “Ranging Behavior Drives Parasite Richness: A More Parsimonious Hypothesis”Charpentier MJE, Kappeler PMIn a recent article, Bicca-Marques and Calegaro-Marques [2016] discussed the putative assumptions related to an interpretation we provided regarding an observed positive relationship between weekly averaged parasite richness of a group of mandrill...Behaviour & Ethology, Evolutionary ecology, Foraging, Host-parasite interactions, Spatial ecology, Metacommunities & Metapopulations, ZoologyCédric Sueur2018-05-22 10:59:33 View
01 Jun 2018
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Data-based, synthesis-driven: setting the agenda for computational ecology

Some thoughts on computational ecology from people who I’m sure use different passwords for each of their accounts

Recommended by based on reviews by Matthieu Barbier and 1 anonymous reviewer

Are you an ecologist who uses a computer or know someone that does? Even if your research doesn’t rely heavily on advanced computational techniques, it likely hasn’t escaped your attention that computers are increasingly being used to analyse field data and make predictions about the consequences of environmental change. So before artificial intelligence and robots take over from scientists, now is great time to read about how experts think computers could make your life easier and lead to innovations in ecological research. In “Data-based, synthesis-driven: setting the agenda for computational ecology”, Poisot and colleagues [1] provide a brief history of computational ecology and offer their thoughts on how computational thinking can help to bridge different types of ecological knowledge. In this wide-ranging article, the authors share practical strategies for realising three main goals: (i) tighter integration of data and models to make predictions that motivate action by practitioners and policy-makers; (ii) closer interaction between data-collectors and data-users; and (iii) enthusiasm and aptitude for computational techniques in future generations of ecologists. The key, Poisot and colleagues argue, is for ecologists to “engage in meaningful dialogue across disciplines, and recognize the currencies of their collaborations.” Yes, this is easier said than done. However, the journey is much easier with a guide and when everyone involved serves to benefit not only from the eventual outcome, but also the process.


[1] Poisot, T., Labrie, R., Larson, E., & Rahlin, A. (2018). Data-based, synthesis-driven: setting the agenda for computational ecology. BioRxiv, 150128, ver. 4 recommended and peer-reviewed by PCI Ecology. doi: 10.1101/150128

Data-based, synthesis-driven: setting the agenda for computational ecologyTimothée Poisot, Richard Labrie, Erin Larson, Anastasia RahlinComputational ecology, defined as the application of computational thinking to ecological problems, has the potential to transform the way ecologists think about the integration of data and models. As the practice is gaining prominence as a way to...Meta-analyses, Statistical ecology, Theoretical ecologyPhillip P.A. Staniczenko2018-02-05 20:51:41 View