Citizen science is becoming an important piece for the acquisition of scientific knowledge in the fields of natural sciences, and particularly in the inventory and monitoring of biodiversity (McKinley et al. 2017). The information generated with the collaboration of citizens has an evident importance in conservation, by providing information on the state of populations and habitats, helping in mitigation and restoration actions, and very importantly contributing to involve society in conservation (Brown and Williams 2019). An obvious advantage of these initiatives is the ability to mobilize human resources on a large territorial scale and in the medium term, which would otherwise be difficult to finance. The resulting increasing information then can be processed with advanced computational techniques (Hochachka et al 2012; Kelling et al. 2015), thus improving our interpretation of the distribution of species. Specifically, the ability to obtain information on a large territorial scale can be integrated into studies based on Species Distribution Models SDMs. One of the common problems with SDMs is that they often work from species occurrences that have been opportunistically recorded, either by professionals or amateurs. A great challenge for data obtained from non-professional citizens, however, remains to ensure its standardization and quality (Kosmala et al. 2016). This requires a clear and effective design, solid volunteer training, and a high level of coordination that turns out to be complex (Brown and Williams 2019). Finally, it is essential to perform a quality validation following scientifically recognized standards, since they are often conditioned by errors and biases in obtaining information (Bird et al. 2014). There are two basic approaches to obtain the necessary data for this validation: getting it from an external source (external validation), or allocating a part of the database itself (internal validation or cross-validation) to this function.
Matutini et al. (2020) in his work 'How citizen science could improve Species Distribution Models and their independent assessment' shows a novel application of the data generated by a citizen science initiative ('Un Dragon dans mon Jardin') by providing an external source for the validation of SDMs, as a tool to construct habitat suitability maps for nine species of amphibians in western France. Importantly, 'Un Dragon dans mon Jardin' contains standardized presence-absence data, the approximation recognized as the most robust (Guisan, et al. 2017). The SDMs to be validated, in turn, were based on opportunistic information obtained by citizens and professionals. The result shows the usefulness of this external data source by minimizing the overestimation of model accuracy that is obtained with cross-validation with the internal evaluation dataset. It also shows the importance of properly filtering the information obtained by citizens by determining the threshold of sampling effort.
The destiny of citizen science is to be integrated into the complex world of science. Supported by the increasing level of the formation of society, it is becoming a fundamental piece in the scientific system dedicated to the study of biodiversity and its conservation. After funding for scientists specialized in the recognition of biodiversity has been cut back, we are seeing a transformation of the activity of these scientists towards the design, coordination, training and verification of programs for the acquisition of field information obtained by citizens. A main goal is that a substantial part of this information will eventually get integrated into the scientific system, and rigorous verification process a fundamental element for such purpose, as shown by Matutini et al. (2020) work.

References

[1] Bird TJ et al. (2014) Statistical solutions for error and bias in global citizen science datasets. Biological Conservation 173: 144-154. doi: 10.1016/j.biocon.2013.07.037
[2] Brown ED and Williams BK (2019) The potential for citizen science to produce reliable and useful information in ecology. Conservation Biology 33: 561-569. doi: 10.1111/cobi.13223
[3] Guisan A, Thuiller W and Zimmermann N E (2017) Habitat Suitability and Distribution Models: With Applications in R. The University of Chicago Press. doi: 10.1017/9781139028271
[4] Hochachka WM, Fink D, Hutchinson RA, Sheldon D, Wong WK and Kelling S (2012) Data-intensive science applied to broad-scale citizen science. Trens Ecol Evol 27: 130-137. doi: 10.1016/j.tree.2011.11.006
[5] Kelling S, Fink D, La Sorte FA, Johnston A, Bruns NE and Hochachka WM (2015) Taking a ‘Big Data’ approach to data quality in a citizen science project. Ambio 44(Supple. 4):S601-S611. doi: 10.1007/s13280-015-0710-4
[6] Kosmala M, Wiggins A, Swanson A and Simmons B (2016) Assessing data quality in citizen science. Front Ecol Environ 14: 551–560. doi: 10.1002/fee.1436
[7] Matutini F, Baudry J, Pain G, Sineau M and Pithon J (2020) How citizen science could improve Species Distribution Models and their independent assessment. bioRxiv, 2020.06.02.129536, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/2020.06.02.129536
[8] McKinley DC et al. (2017) Citizen science can improve conservation science, natural resource management, and environmental protection. Biological Conservation 208:15-28. doi: 10.1016/j.biocon.2016.05.015

Urban ecology, the study of ecological systems in our increasingly urbanized world, is crucial to planning and redesigning cities to enhance ecosystem services (Kremer et al. 2016), human health and well-being and further conservation goals (Dallimer et al. 2012). Urban trees are a crucial component of urban streets and parks that provide shade and cooling through evapotranspiration (Fung and Jim 2019), improve air quality (Lai and Kontokosta 2019), help control storm water (Johnson and Handel 2016), and conserve wildlife (Herrmann et al. 2012; de Andrade et al. 2020).
Ideally, management of urban forests strikes a balance between maintaining the health of urban trees while retaining those organisms, such as herbivores, that connect a tree to the urban ecosystem. Herbivory by arthropods can substantially affect tree growth and reproduction (Whittaker and Warrington 1985), and so understanding factors that influence herbivory in urban forests is important to effective management. At the same time, herbivorous arthropods are important as key components of urban bird diets (Airola and Greco 2019) and provide a backyard glimpse at forest ecosystems in an increasingly built environment (Pearse 2019). Maintenance of arthropod predators may be one way to retain arthropods in urban forests while keeping detrimental outbreaks of herbivores in check. In “Insect herbivory on urban trees: Complementary effects of tree neighbors and predation” Stemmelen and colleagues (Stemmelen et al. 2020) use a clever sampling design to show that insect herbivory decreases as the diversity of neighboring trees increased. By placing artificial larvae out on trees, they provide evidence that increased predation in higher diversity urban forest patches might drive patterns in herbivory. The paper also demonstrates the importance of tree species identity in determining leaf herbivory.
The implications of this research for urban foresters is that deliberately planting diverse urban forests will help manage insect herbivores and should thus improve tree health. Potential knock-on effects could be seen for the ecosystem services provided by urban forests. While it might be tempting to simply plant more of the species that are subject to low current rates of herbivory, other research on the long-term vulnerability of monocultures to attack by specialist pathogens and herbivores (Tooker and Frank 2012) cautions against such an approach. Furthermore, the importance of urban forest insects to birds, including migrating birds, argues for managing urban forests more holistically (Greco and Airola 2018).
Stemmelen et al. (2020) used an observational approach focused on urban forests in Montreal, Canada in their research. Their findings suggest follow-up research focused on a broader cross-section of urban forests across latitudes, as well as experimental research. Experiments could, for example, exclude avian predators with netting (e.g. (Marquis and Whelan 1994)) to evaluate the relative importance of birds to managing urban insects on trees, as well as the flip side of that equation, the important to birds of insects on urban trees.
In summary, Stemmelen and colleague’s manuscript illustrates clever sampling and use of observational data to infer broader ecological patterns. It is worth reading to better understand the role of diversity in driving plant-insect community interactions and given the implications of the findings for sustainable long-term management of urban forests.

References

Airola, D. and Greco, S. (2019). Birds and oaks in California’s urban forest. Int. Oaks, 30, 109–116.
de Andrade, A.C., Medeiros, S. and Chiarello, A.G. (2020). City sloths and marmosets in Atlantic forest fragments with contrasting levels of anthropogenic disturbance. Mammal Res., 65, 481–491. doi: https://doi.org/10.1007/s13364-020-00492-0
Dallimer, M., Irvine, K.N., Skinner, A.M.J., Davies, Z.G., Rouquette, J.R., Maltby, L.L., et al. (2012). Biodiversity and the Feel-Good Factor: Understanding Associations between Self-Reported Human Well-being and Species Richness. Bioscience, 62, 47–55. doi: https://doi.org/10.1525/bio.2012.62.1.9
Fung, C.K.W. and Jim, C.Y. (2019). Microclimatic resilience of subtropical woodlands and urban-forest benefits. Urban For. Urban Green., 42, 100–112. doi: https://doi.org/10.1016/j.ufug.2019.05.014
Greco, S.E. and Airola, D.A. (2018). The importance of native valley oaks (Quercus lobata) as stopover habitat for migratory songbirds in urban Sacramento, California, USA. Urban For. Urban Green., 29, 303–311. doi: https://doi.org/10.1016/j.ufug.2018.01.005
Herrmann, D.L., Pearse, I.S. and Baty, J.H. (2012). Drivers of specialist herbivore diversity across 10 cities. Landsc. Urban Plan., 108, 123–130. doi: https://doi.org/10.1016/j.landurbplan.2012.08.007
Johnson, L.R. and Handel, S.N. (2016). Restoration treatments in urban park forests drive long-term changes in vegetation trajectories. Ecol. Appl., 26, 940–956. doi: https://doi.org/10.1890/14-2063
Kremer, P., Hamstead, Z., Haase, D., McPhearson, T., Frantzeskaki, N., Andersson, E., et al. (2016). Key insights for the future of urban ecosystem services research. Ecol. Soc., 21: 29. doi: http://doi.org/10.5751/ES-08445-210229
Lai, Y. and Kontokosta, C.E. (2019). The impact of urban street tree species on air quality and respiratory illness: A spatial analysis of large-scale, high-resolution urban data. Heal. Place, 56, 80–87. doi: https://doi.org/10.1016/j.healthplace.2019.01.016
Marquis, R.J. and Whelan, C.J. (1994). Insectivorous birds increase growth of white oak through consumption of leaf-chewing insects. Ecology, 75, 2007–2014. doi: https://doi.org/10.2307/1941605
Pearse, I.S. (2019). Insect herbivores on urban native oak trees. Int. Oaks, 30, 101–108.
Stemmelen, A., Paquette, A., Benot, M.-L., Kadiri, Y., Jactel, H. and Castagneyrol, B. (2020) Insect herbivory on urban trees: Complementary effects of tree neighbours and predation. bioRxiv, 2020.04.15.042317, ver. 5 peer-reviewed and recommended by PCI Ecology. doi: https://doi.org/10.1101/2020.04.15.042317
Tooker, J. F., and Frank, S. D. (2012). Genotypically diverse cultivar mixtures for insect pest management and increased crop yields. J. Appl. Ecol., 49(5), 974-985. doi: https://doi.org/10.1111/j.1365-2664.2012.02173.x
Whittaker, J.B. and Warrington, S. (1985). An experimental field study of different levels of insect herbivory induced By Formica rufa predation on Sycamore (Acer pseudoplatanus) III. Effects on Tree Growth. J. Appl. Ecol., 22, 797. doi: https://doi.org/10.2307/2403230

In this article, Saade et al. (2021) investigate how the rate of local extinctions and their spatial distribution affect recolonization dynamics in metacommunities. They use an elegant combination of microcosm experiments with metacommunities of freshwater ciliates and mathematical modelling mirroring their experimental system. Their main findings are (i) that local patch extinctions increase both local (α-) and inter-patch (β-) diversity in a transient way during the recolonization process, (ii) that these effects depend more on the spatial distribution of extinctions (dispersed or clustered) than on their amount, and (iii) that they may spread regionally.
Microcosm experiments are already quite cool just by themselves and have contributed largely to conceptual advances in community ecology (see Fraser and Keddy 1997, or Jessup et al. 2004 for reviews on this topic), but they are here exploited to a whole further level by the fitting of a metapopulation dynamics model. The model allows both to identify the underlying mechanisms most likely to generate the patterns observed (here, competitive interactions) and to assess the robustness of these patterns when considering larger spatial or temporal scales. This release of experimental limitations allows here for the analysis of quantitative metrics of spatial structure, like the distance to the closest patch, which gives an interesting insight into the functional basis of the effect of the spatial distribution of extinctions.

A major strength of this study is that it highlights the importance of considering the spatial structure explicitly. Recent work on ecological networks has shown repeatedly that network structure affects the propagation of pathogens (Badham and Stocker 2010), invaders (Morel-Journel et al. 2019), or perturbation events (Gilarranz et al. 2017). Here, the spatial structure of the metacommunity is a regular grid of patches, but the distribution of extinction events may be either regularly dispersed (i.e., extinct patches are distributed evenly over the grid and are all surrounded by non-extinct patches only) or clustered (all extinct patches are neighbours). This has a direct effect on the neighbourhood of perturbed patches, and because perturbations have mostly local effects, their recovery dynamics are dominated by the composition of this immediate neighbourhood. In landscapes with dispersed extinctions, the neighbourhood of a perturbed patch is not affected by the amount of extinctions, and neither is its recovery time. In contrast, in landscapes with clustered extinctions, the amount of extinctions affects the depth of the perturbed area, which takes longer to recover when it is larger. Interestingly, the spatial distribution of extinctions here is functionally equivalent to differences in connectivity between perturbed and unperturbed patches, which results in contrasted “rescue recovery” and “mixing recovery” regimes as described by Zelnick et al. (2019).
 
Furthermore, this study focuses on local dynamics of competition and short-term, transient patterns that may have been overlooked by more classical, equilibrium-based approaches of dynamical systems of metacommunities. Indeed, in a metacommunity composed of several competitors, early theoretical work demonstrated that species coexistence is possible at the regional scale only, provided that spatial heterogeneity creates spatial variance in fitness or precludes the superior competitor from accessing certain habitat patches (Skellam 1951, Levins 1969). In the spatially homogeneous experimental system of Saade et al., one of the three ciliate species ends up dominating the community at equilibrium. However, following local, one-time extinction events, the community endures a recolonization process in which differences in dispersal may provide temporary spatial niches for inferior competitors. These transient patterns might prove essential to understand and anticipate the resilience of natural systems that are under increasing pressure, and enduring ever more frequent and intense perturbations (IPBES 2019). Spatial autocorrelation in extinction events was previously identified as a risk for stability and persistence of metacommunities (Ruokolainen 2013, Kahilainen et al. 2018). These new results show that autocorrelated perturbations also have longer-lasting effects, which is likely to increase their overall impact on metacommunity dynamics. As spatial and temporal autocorrelation of temperature and extreme climatic events are expected to increase (Di Cecco and Gouthier 2018), studies that investigate how metacommunities respond to the structure of the distribution of perturbations are more necessary than ever.
 
References

Badham J, Stocker R (2010) The impact of network clustering and assortativity on epidemic behaviour. Theoretical Population Biology, 77, 71–75. https://doi.org/10.1016/j.tpb.2009.11.003
 
Di Cecco GJ, Gouhier TC (2018) Increased spatial and temporal autocorrelation of temperature under climate change. Scientific Reports, 8, 14850. https://doi.org/10.1038/s41598-018-33217-0
 
Fraser LH, Keddy P (1997) The role of experimental microcosms in ecological research. Trends in Ecology & Evolution, 12, 478–481. https://doi.org/10.1016/S0169-5347(97)01220-2
 
Gilarranz LJ, Rayfield B, Liñán-Cembrano G, Bascompte J, Gonzalez A (2017) Effects of network modularity on the spread of perturbation impact in experimental metapopulations. Science, 357, 199–201. https://doi.org/10.1126/science.aal4122
 
IPBES (2019) Summary for policymakers of the global assessment report on biodiversity and ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. S. Díaz, J. Settele, E. S. Brondízio E.S., H. T. Ngo, M. Guèze, J. Agard, A. Arneth, P. Balvanera, K. A. Brauman, S. H. M. Butchart, K. M. A. Chan, L. A. Garibaldi, K. Ichii, J. Liu, S. M. Subramanian, G. F. Midgley, P. Miloslavich, Z. Molnár, D. Obura, A. Pfaff, S. Polasky, A. Purvis, J. Razzaque, B. Reyers, R. Roy Chowdhury, Y. J. Shin, I. J. Visseren-Hamakers, K. J. Willis, and C. N. Zayas (eds.). IPBES secretariat, Bonn, Germany. 56 pages. https://doi.org/10.5281/zenodo.3553579 
 
Jessup CM, Kassen R, Forde SE, Kerr B, Buckling A, Rainey PB, Bohannan BJM (2004) Big questions, small worlds: microbial model systems in ecology. Trends in Ecology & Evolution, 19, 189–197. https://doi.org/10.1016/j.tree.2004.01.008
 
Kahilainen A, van Nouhuys S, Schulz T, Saastamoinen M (2018) Metapopulation dynamics in a changing climate: Increasing spatial synchrony in weather conditions drives metapopulation synchrony of a butterfly inhabiting a fragmented landscape. Global Change Biology, 24, 4316–4329. https://doi.org/10.1111/gcb.14280

Levins R (1969) Some Demographic and Genetic Consequences of Environmental Heterogeneity for Biological Control1. Bulletin of the Entomological Society of America, 15, 237–240. https://doi.org/10.1093/besa/15.3.237
 
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Ruokolainen L (2013) Spatio-Temporal Environmental Correlation and Population Variability in Simple Metacommunities. PLOS ONE, 8, e72325. https://doi.org/10.1371/journal.pone.0072325

Saade C, Kefi S, Gougat-Barbera C, Rosenbaum B, Fronhofer EA (2021) Spatial distribution of local patch extinctions drives recovery dynamics in metacommunities. bioRxiv, 2020.12.03.409524, ver. 4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2020.12.03.409524
 
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As far back as Clements [1] and Gleason [2], the historical schism between deterministic and stochastic perspectives has divided ecologists. Deterministic theories tend to emphasize niche-based processes such as environmental filtering and species interactions as the main drivers of species distribution in nature, while stochastic theories mainly focus on chance colonization, random extinctions and ecological drift [3]. Although the old days when ecologists were fighting fiercely over null models and their adequacy to capture niche-based processes is over [4], the ghost of that debate between deterministic and stochastic perspectives came back to haunt ecologists in the form of the ‘environment versus space’ debate with the development of metacommunity theory [5]. While interest in that question led to meaningful syntheses of metacommunity dynamics in natural systems [6], it also illustrated how context-dependant the answer was [7]. One of the next frontiers in metacommunity ecology is to identify the underlying drivers of this observed context-dependency in the relative importance of ecological processus [7, 8].
Reflecting on seminal work by Robert MacArthur emphasizing different processes at different spatial scales [9, 10] (the so-called ‘MacArthur paradox’), Chase and Myers proposed in 2011 that a key in solving the deterministic versus stochastic debate was probably to turn our attention to how the relative importance of local processes changes across spatial scales [3]. Scale-dependance is a well-acknowledged challenge in ecology, hampering empirical syntheses and comparisons between studies [11-14]. Embracing the scale-dependance of ecological processes would not only lead to stronger syntheses and consolidation of current knowledge, it could also help resolve many current debates or apparent contradictions [11, 15, 16].
The timely study by Siqueira et al. [17] fits well within this historical context by exploring the relative importance of ecological drift and selection across a gradient of community size (number of individuals in a given community). More specifically, they tested the hypothesis that small communities are more dissimilar among each other because of ecological drift compared to large communities, which are mainly structured by niche selection [17]. That smaller populations or communities should be more affected by drift is a mathematical given [18], but the main questions are i) for a given community size how important is ecological drift relative to other processes, and ii) how small does a community have to be before random assembly dominates? The authors answer these questions using an extensive stream dataset with a community size gradient sampled from 200 streams in two climatic regions (Brazil and Finland). Combining linear models with recent null model approaches to measure deviations from random expectations [19], they show that, as expected based on theory and recent experimental work, smaller communities tend to have higher β-diversity, and that those β-diversity patterns could not be distinguished from random assembly processes [17]. Spatial turnover among larger communities is mainly driven by niche-based processes related to species sorting or dispersal dynamics [17]. Given the current environmental context, with many anthropogenic perturbations leading to reduced community size, it is legitimate to wonder, as the authors do, whether we are moving toward a more stochastic and thus less predictable world with obvious implications for the conservation of biodiversity [17].
The real strength of the study by Siqueira et al. [17], in my opinion, is in the inclusion of stream data from boreal and tropical regions. Interestingly and most importantly, the largest communities in the tropical streams are as large as the smallest communities in the boreal streams. This is where the study should really have us reflect on the notions of context-dependency in observed patterns because the negative relationship between community size and β-diversity was only observed in the tropical streams, but not in the boreal streams [17]. This interesting nonlinearity in the response means that a study that would have investigated the drift versus niche-based question only in Finland would have found very different results from the same study in Brazil. Only by integrating such a large scale gradient of community sizes together could the authors show the actual shape of the relationship, which is the first step toward building a comprehensive synthesis on a debate that has challenged ecologists for almost a century.

References

[1] Clements, F. E. (1936). Nature and structure of the climax. Journal of ecology, 24(1), 252-284. doi: 10.2307/2256278
[2] Gleason, H. A. (1917). The structure and development of the plant association. Bulletin of the Torrey Botanical Club, 44(10), 463-481. doi: 10.2307/2479596
[3] Chase, J. M., and Myers, J. A. (2011). Disentangling the importance of ecological niches from stochastic processes across scales. Philosophical transactions of the Royal Society B: Biological sciences, 366(1576), 2351-2363. doi: 10.1098/rstb.2011.0063
[4] Diamond, J. M., and Gilpin, M. E. (1982). Examination of the “null” model of Connor and Simberloff for species co-occurrences on islands. Oecologia, 52(1), 64-74. doi: 10.1007/BF00349013
[5] Leibold M. A., et al. (2004). The metacommunity concept: a framework for multi‐scale community ecology. Ecology letters, 7(7), 601-613. doi: 10.1111/j.1461-0248.2004.00608.x
[6] Cottenie, K. (2005). Integrating environmental and spatial processes in ecological community dynamics. Ecology letters, 8(11), 1175-1182. doi: 10.1111/j.1461-0248.2005.00820.x
[7] Leibold, M. A. and Chase, J. M. (2018). Metacommunity Ecology. Monographs in Population Biology, vol. 59. Princeton University Press. [8] Vellend, M. (2010). Conceptual synthesis in community ecology. The Quarterly review of biology, 85(2), 183-206. doi: 10.1086/652373
[9] MacArthur, R. H., and Wilson, E. O. (1963). An equilibrium theory of insular zoogeography. Evolution, 17(4), 373-387. doi: 10.1111/j.1558-5646.1963.tb03295.x
[10] MacArthur, R. H., and Levins, R. (1967). The limiting similarity, convergence, and divergence of coexisting species. The American Naturalist, 101(921), 377-385. doi: 10.1086/282505
[11] Viana, D. S., and Chase, J. M. (2019). Spatial scale modulates the inference of metacommunity assembly processes. Ecology, 100(2), e02576. doi: 10.1002/ecy.2576
[12] Chave, J. (2013). The problem of pattern and scale in ecology: what have we learned in 20 years?. Ecology letters, 16, 4-16. doi: 10.1111/ele.12048
[13] Patrick, C. J., and Yuan, L. L. (2019). The challenges that spatial context present for synthesizing community ecology across scales. Oikos, 128(3), 297-308. doi: 10.1111/oik.05802
[14] Chase, J. M., and Knight, T. M. (2013). Scale‐dependent effect sizes of ecological drivers on biodiversity: why standardised sampling is not enough. Ecology letters, 16, 17-26. doi: 10.1111/ele.12112
[15] Horváth, Z., Ptacnik, R., Vad, C. F., and Chase, J. M. (2019). Habitat loss over six decades accelerates regional and local biodiversity loss via changing landscape connectance. Ecology letters, 22(6), 1019-1027. doi: 10.1111/ele.13260
[16] Chase, J. M, Gooriah, L., May, F., Ryberg, W. A, Schuler, M. S, Craven, D., and Knight, T. M. (2019). A framework for disentangling ecological mechanisms underlying the island species–area relationship. Frontiers of Biogeography, 11(1). doi: 10.21425/F5FBG40844.
[17] Siqueira T., Saito V. S., Bini L. M., Melo A. S., Petsch D. K. , Landeiro V. L., Tolonen K. T., Jyrkänkallio-Mikkola J., Soininen J. and Heino J. (2019). Community size affects the signals of ecological drift and niche selection on biodiversity. bioRxiv 515098, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/515098
[18] Hastings A., Gross L. J. eds. (2012). Encyclopedia of theoretical ecology (University of California Press, Berkeley).
[19] Chase, J. M., Kraft, N. J., Smith, K. G., Vellend, M., and Inouye, B. D. (2011). Using null models to disentangle variation in community dissimilarity from variation in α‐diversity. Ecosphere, 2(2), 1-11. doi: 10.1890/ES10-00117.1

In his 1972 paper “Will a Large Complex System Be Stable?” [1], May challenges the idea that large communities are more stable than small ones. This was the beginning of a fundamental debate that still structures an entire research area in ecology: the diversity-stability debate [2]. The most salient strength of May’s work was to use a mathematical argument to refute an idea based on the observations that simple communities are less stable than large ones. Using the formalism of dynamical systems and a major results on the distribution of the eigen values for random matrices, May demonstrated that the addition of random interactions destabilizes ecological communities and thus, rich communities with a higher number of interactions should be less stable. But May also noted that his mathematical argument holds true only if ecological interactions are randomly distributed and thus concluded that this must not be true! This is how the contradiction between mathematics and empirical observations led to new developments in the study of ecological networks.
Since 1972, the theoretical corpus of ecology has advanced, building on the formalism of dynamical systems, ecologists have revealed that ecological interactions are indeed not randomly distributed [3,4], but general rules are still missing and we are far from understanding what determine the exact network topology of a given community. One promising avenue is to understand the relationship between different facets of the concept of stability [5,6]. Indeed, the classical approach to determine whether a system is stable is qualitative: if a system returns to its equilibrium when it is slightly moved away from it, then the system is considered stable. But there are several other aspects that are worth scrutinizing. For instance, when a system returns to its equilibrium, one can characterize the corresponding transient dynamics [7,8], that is asking fundamental questions such as: what is the trajectory of return? How long does it take to return to the equilibrium? Another fundamental question is whether the system remains qualitatively stable when the distributions of interactions strengths change? From a biological standpoint, all of these questions matter as all these aspects of stability may partially explain the actual structure of ecological networks, and hence, frameworks that integrate several facets of stability are much needed.
The study by Arnoldi et al. [9] is a significant step towards such a framework. The strength of their formalism is threefold. First, instead of considering separately the system and its perturbations, they considering the fluctuations of a perturbed ecological systems and thus, perturbations are parts of the ecological system. Second, they use of a broad definition of perturbation that encompasses the types of perturbations (whether the individual respond synchronously or not), their intensity and their direction (how the perturbations are correlated across species). Third, they quantify the instability of the system using variability which integrates the consequences of perturbations over the whole set of species of a community: such a measure is comparable across communities and accounts for the trivial effect of the perturbations on the system dynamics.
Using this framework, the authors show that interactions within a stable community leads to a general relationship between variability and the abundance of individually perturbed species: if individuals of species respond in synchrony to a perturbation, then the more abundant the species perturbed the higher the variability of the system, but the relationship is reverse when individual respond asynchronously. A direct implications of these results for the classical debate is that the diversity-stability relationship is negative for the former type of perturbations (as in May’s seminal paper) but positive for the latter type. Hence, the rigorous work of Arnoldi and colleagues sheds a new light upon the classical debate: the nature of the perturbation regime prevailing within a community affects the slope of the diversity-stability relationships and given the vast diversity of ecological communities, this may very well be one of the reasons why the debate still endures.
From a historical perspective, it is interesting that ecologists have gone from looking at random webs to structured webs and now, in a sense, Arnoldi et al. are unpacking the role of differentially structured perturbations. The work they achieved will doubtlessly be followed by further theoretical investigations. One natural research avenue is to revisit the role of the topology of ecological networks with this framework: how the distribution of interactions and their strength affect the general relationship they unravel? Finally, this study demonstrate that the impact of the abundance of a species on the variability of the system depends on the nature of the perturbation regime and so the distribution of species abundances within a community should be determined by the prevailing perturbation regime which is a prediction that remains to be tested.

References

[1] May, Robert M (1972). Will a Large Complex System Be Stable? Nature 238, 413–414. doi: 10.1038/238413a0
[2] McCann, Kevin Shear (2000). The Diversity–Stability Debate. Nature 405, 228–233. doi: 10.1038/35012234
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