The importance of the vertical structure of vegetation cover for the functioning, management and conservation of ecosystems has received particular attention from ecologists in the last decades. Canopy architecture has many implications for light extinction coefficient, temperature variation reduction, self-shading which are all key parameters for the structuring and functioning of different ecosystems such as grasslands [1,2], forests [3,4], phytoplankton communities [5, 6], macroalgal populations [7] and even underwater animal forests such as octocoral communities [8].

This research topic, therefore, benefits from a large body of literature and the facilitative role of self-shadowing is no longer in question. However, it is always puzzling to note that some of the most common ecosystems turn out to be amongst the least known. This is precisely the case of the Fucus serratus communities which are widespread in Northeast Atlantic along the Atlantic coast of Europe from Svalbard to Portugal, as well as Northwest Atlantic & Gulf of St. Lawrence, easily accessible at low tide, but which have comparatively received less attention than more emblematic macro-algal communities such as Laminariales.

The lack of attention paid to these most common Fucales is particularly critical as some species such as F. serratus are proving to be particularly vulnerable to environmental change, leading to a predicted northward retreat from its current southern boundary [9].

In the present study [10], the authors showed the importance of the vegetation cover in resisting tide-induced environmental stresses. The canopy of F. serratus mitigates stress levels experienced in the lower layers during emersion, while various acclimation strategies take over to maintain the photosynthetic apparatus in optimal conditions.

They hereby highlight adaptation mechanisms to the extreme environment represented by the intertidal zone. These adaptation strategies were expected and similar mechanisms had been shown at the cellular level previously [11]. The earliest studies on the subject have shown that the structure of the bottom, the movement of water, and light availability all "influence the distribution of Fucaceae and disturb the regularity of their fine zonation, which itself is caused by the most important factor, desiccation", as Zaneveld states in his review [12]. He observed that the causes of the zonal distribution of marine algae are numerous, and identified several points of interest such as the relative period of emersion, the rapidity of desiccation, the loss of water, and the thickness of the cell walls.

The present study thus highlights the existence of facilitative mechanisms associated with F. serratus canopy and nicely confirms previous work with in situ observations. It also highlights the importance of the vegetative cover in combating desiccation and introduces the dampening effect as a facilitating mechanism.

The effect of the vegetation cover can sometimes even be felt beyond its immediate area of influence. A recent study shows that ground-level ozone is significantly reduced by the combined effects of canopy shading and turbulence [4]. Below the canopy, the light intensity becomes sufficiently low which inhibits ozone formation due to the decrease in the rates of hydroxyl radical formation and the rates of conversion of nitrogen dioxide to nitrogen oxide by photolysis. In addition, reductions in light levels associated with foliage promote ozone-destroying reactions between plant-emitted species, such as nitric oxide and/or alkenes, and ozone itself. The reduction in diffusivity slows the upward transport of surface emitted species, partially decoupling the area under the canopy from the rest of the atmosphere.

By analogy with the work of Makar et al [4], and in the light of the results provided by the authors of this study, one may wonder whether the canopy dampening of F. serratus communities (and other common fucoids widely distributed on our coasts) might not also influence atmospheric chemistry, both at the Earth's surface and in the atmospheric boundary layer. The lack of accumulation of reactive oxygen species under the canopy found by the authors is consistent with this hypothesis and suggests that the damping effect of F. serratus may well have much wider consequences than expected.

References

[1] Jurik TW, Kliebenstein H (2000) Canopy Architecture, Light Extinction and Self-Shading of a Prairie Grass, Andropogon Gerardii. The American Midland Naturalist, 144, 51–65. http://www.jstor.org/stable/3083010

[2] Mitchley J, Willems JH (1995) Vertical canopy structure of Dutch chalk grasslands in relation to their management. Vegetatio, 117, 17–27. https://doi.org/10.1007/BF00033256

[3] Kane VR, Gillespie AR, McGaughey R, Lutz JA, Ceder K, Franklin JF (2008) Interpretation and topographic compensation of conifer canopy self-shadowing. Remote Sensing of Environment, 112, 3820–3832. https://doi.org/10.1016/j.rse.2008.06.001

[4] Makar PA, Staebler RM, Akingunola A, Zhang J, McLinden C, Kharol SK, Pabla B, Cheung P, Zheng Q (2017) The effects of forest canopy shading and turbulence on boundary layer ozone. Nature Communications, 8, 15243. https://doi.org/10.1038/ncomms15243

[5] Shigesada N, Okubo A (1981) Analysis of the self-shading effect on algal vertical distribution in natural waters. Journal of Mathematical Biology, 12, 311–326. https://doi.org/10.1007/BF00276919

[6] Barros MP, Pedersén M, Colepicolo P, Snoeijs P (2003) Self-shading protects phytoplankton communities against H2O2-induced oxidative damage. Aquatic Microbial Ecology, 30, 275–282. https://doi.org/10.3354/ame030275

[7] Ørberg SB, Krause-Jensen D, Mouritsen KN, Olesen B, Marbà N, Larsen MH, Blicher ME, Sejr MK (2018) Canopy-Forming Macroalgae Facilitate Recolonization of Sub-Arctic Intertidal Fauna and Reduce Temperature Extremes. Frontiers in Marine Science, 5. https://doi.org/10.3389/fmars.2018.00332

[8] Nelson H, Bramanti L (2020) From Trees to Octocorals: The Role of Self-Thinning and Shading in Underwater Animal Forests. In: Perspectives on the Marine Animal Forests of the World (eds Rossi S, Bramanti L), pp. 401–417. Springer International Publishing, Cham. https://doi.org/10.1007/978-3-030-57054-5_12

[9] Jueterbock A, Kollias S, Smolina I, Fernandes JMO, Coyer JA, Olsen JL, Hoarau G (2014) Thermal stress resistance of the brown alga Fucus serratus along the North-Atlantic coast: Acclimatization potential to climate change. Marine Genomics, 13, 27–36. https://doi.org/10.1016/j.margen.2013.12.008

[10] Migné A, Duong G, Menu D, Davoult D, Gévaert F (2021) Dynamics of Fucus serratus thallus photosynthesis and community primary production during emersion across seasons: canopy dampening and biochemical acclimation. HAL, hal-03079617, ver. 4 peer-reviewed and recommended by Peer community in Ecology. https://hal.archives-ouvertes.fr/hal-03079617

[11] Lichtenberg M, Kühl M (2015) Pronounced gradients of light, photosynthesis and O2 consumption in the tissue of the brown alga Fucus serratus. New Phytologist, 207, 559–569. https://doi.org/10.1111/nph.13396

[12] Zaneveld JS (1937) The Littoral Zonation of Some Fucaceae in Relation to Desiccation. Journal of Ecology, 25, 431–468. https://doi.org/10.2307/2256204

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
 
Morel-Journel T, Assa CR, Mailleret L, Vercken E (2019) Its all about connections: hubs and invasion in habitat networks. Ecology Letters, 22, 313–321. https://doi.org/10.1111/ele.13192

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
 
Skellam JG (1951) Random Dispersal in Theoretical Populations. Biometrika, 38, 196–218. https://doi.org/10.2307/2332328
 
Zelnik YR, Arnoldi J-F, Loreau M (2019) The three regimes of spatial recovery. Ecology, 100, e02586. https://doi.org/10.1002/ecy.2586

The slippersnail (Crepidula fornicata), originally from the eastern coast of North America, has invaded European coastlines from Norway to the Mediterranean Sea [1]. This species is capable of achieving incredibly high densities (up to several thousand individuals per square meter) and likely has major impacts on a variety of community- and ecosystem-level processes, including alteration of carbon and nitrogen fluxes and competition with native suspension feeders [2].

Given this potential for competition, it is important to understand the diet of C. fornicata and its potential overlap with native species. However, previous research on the diet of C. fornicata and related species suggests that the types of food consumed may change with age [3, 4]. This species has an unusual reproductive strategy. It is a sequential hermaphrodite, which begins life as a somewhat mobile male but eventually slows down to become sessile. Sessile individuals form stacks of up to 10 or more individuals, with larger individuals on the bottom of the stack, and decreasingly smaller individuals piled on top. Snails at the bottom of the stack are female, whereas snails at the top of the stack are male; when the females die, the largest males become female [5]. Thus, understanding these potential ontogenetic dietary shifts has implications for both intraspecific (juvenile vs. male vs. female) and interspecific competition associated with an abundant, invasive species.

To this end, Androuin and colleagues evaluated the stable-isotope (d13C and d15N) and fatty-acid profiles of food sources and different life-history stages of C. fornicata [6]. Based on previous work highlighting the potential for life-history changes in the diet of this species [3,4], they hypothesized that C. fornicata would shift its diet as it aged and predicted that this shift would be reflected in changes in its stable-isotope and fatty-acid profiles. The authors found that potential food sources (biofilm, suspended particulate organic matter, and superficial sedimentary organic matter) differed substantially in both stable-isotope and fatty-acid signatures. However, whereas fatty-acid profiles changed substantially with age, there was no shift in the stable-isotope signatures. Because stable-isotope differences between food sources were not reflected in differences between life-history stages, the authors conservatively concluded that there was insufficient evidence for a diet shift with age. The ontogenetic shifts in fatty-acid profiles were intriguing, but the authors suggested that these reflected age-related physiological changes rather than changes in diet.

The authors’ work highlights the need to consider potential changes in the roles of invasive species with age, especially when evaluating interactions with native species. In this case, C. fornicata consumed a variety of food sources, including both benthic and particulate organic matter, regardless of age. The carbon stable-isotope signature of C. fornicata overlaps with those of several native suspension- and deposit-feeding species in the region [7], suggesting the possibility of resource competition, especially given the high abundances of this invader. This contribution demonstrates the potential difficulty of characterizing the impacts of an abundant invasive species with a complex life-history strategy. Like many invasive species, C. fornicata appears to be a dietary generalist, which likely contributes to its success in establishing and thriving in a variety of locations [8].

References

[1] Blanchard M (1997) Spread of the slipper limpet Crepidula fornicata (L. 1758) in Europe. Current state dans consequences. Scientia Marina, 61, 109–118. Open Access version : https://archimer.ifremer.fr/doc/00423/53398/54271.pdf

[2] Martin S, Thouzeau G, Chauvaud L, Jean F, Guérin L, Clavier J (2006) Respiration, calcification, and excretion of the invasive slipper limpet, Crepidula fornicata L.: Implications for carbon, carbonate, and nitrogen fluxes in affected areas. Limnology and Oceanography, 51, 1996–2007. https://doi.org/10.4319/lo.2006.51.5.1996

[3] Navarro JM, Chaparro OR (2002) Grazing–filtration as feeding mechanisms in motile specimens of Crepidula fecunda (Gastropoda: Calyptraeidae). Journal of Experimental Marine Biology and Ecology, 270, 111–122. https://doi.org/10.1016/S0022-0981(02)00013-8

[4] Yee AK, Padilla DK (2015) Allometric Scaling of the Radula in the Atlantic Slippersnail Crepidula fornicata. Journal of Shellfish Research, 34, 903–907. https://doi.org/10.2983/035.034.0320

[5] Collin R (1995) Sex, Size, and Position: A Test of Models Predicting Size at Sex Change in the Protandrous Gastropod Crepidula fornicata. The American Naturalist, 146, 815–831. https://doi.org/10.1086/285826

[6] Androuin T, Dubois SF, Hubas C, Lefebvre G, Grand FL, Schaal G, Carlier A (2021) Trophic niche of the invasive gregarious species Crepidula fornicata, in relation to ontogenic changes. bioRxiv, 2020.07.30.229021, ver. 4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2020.07.30.229021

[7] Dauby P, Khomsi A, Bouquegneau J-M (1998) Trophic Relationships within Intertidal Communities of the Brittany Coasts: A Stable Carbon Isotope Analysis. Journal of Coastal Research, 14, 1202–1212. Retrieved May 4, 2021, from http://www.jstor.org/stable/4298880

[8] Machovsky-Capuska GE, Senior AM, Simpson SJ, Raubenheimer D (2016) The Multidimensional Nutritional Niche. Trends in Ecology & Evolution, 31, 355–365. https://doi.org/10.1016/j.tree.2016.02.009

Understanding why and how species coexist in local communities is one of the central questions in ecology. There is general agreement that species distribution and coexistence are determined by a number of key mechanisms, including the environmental requirements of species, dispersal, evolutionary constraints, resource availability and selection, metapopulation dynamics, and biotic interactions (e.g. Soberón & Nakamura 2009; Colwell & Rangel 2009; Ricklefs 2015). These factors are however intricately intertwined in a scale-structured fashion (Hortal et al. 2010; D’Amen et al. 2017), making it particularly difficult to tease apart the effects of each one of them. This could be addressed by the novel field of Joint Species Distribution Modelling (JSDM; Okasvainen & Abrego 2020), as it allows assessing the effects of several sets of factors and the co-occurrence and/or covariation in abundances of potentially interacting species at the same time (Pollock et al. 2014; Ovaskainen et al. 2016; Dormann et al. 2018). However, the development of JSDM has been hampered by the general lack of good-quality detailed data on species co-occurrences and abundances (see Hortal et al. 2015).

Facon et al. (2021) use a particularly large compilation of field surveys to study the abundance and co-occurrence of Tephritidae fruit flies in c. 400 orchards, gardens and natural areas throughout the island of Réunion. Further, they combine such information with lab data on their host-selection fundamental niche (i.e. in the absence of competitors), codifying traits of female choice and larval performances in 21 host species. They use Poisson Log-Normal models, a type of mixed model that allows one to jointly model the random effects associated with all species, and retrieve the covariations in abundance that are not explained by environmental conditions or differences in sampling effort. Then, they use a series of models to evaluate the effects on these matrices of ecological covariates (date, elevation, habitat, climate and host plant), species interactions (by comparing with a constrained residual variance-covariance matrix) and the species’ host-selection fundamental niches (through separate models for each fly species).

The eight Tephritidae species inhabiting Réunion include both generalists and specialists in Solanaceae and Cucurbitaceae with a known history of interspecific competition. Facon et al. (2021) use a comprehensive JSDM approach to assess the effects of different factors separately and altogether. This allows them to identify large effects of plant hosts and the fundamental host-selection niche on species co-occurrence, but also to show that ecological covariates and weak –though not negligible– species interactions are necessary to account for all residual variance in the matrix of joint species abundances per site. Further, they also find evidence that the fitness per host measured in the lab has a strong influence on the abundances in each host plant in the field for specialist species, but not for generalists. Indeed, the stronger effects of competitive exclusion were found in pairs of Cucurbitaceae specialist species. However, these analyses fail to provide solid grounds to assess why generalists are rarely found in Cucurbitaceae and Solanaceae. Although they argue that this may be due to Connell’s (1980) ghost of competition past (past competition that led to current niche differentiation), further data on the evolutionary history of these fruit flies is needed to assess this hypothesis.

Finding evidence for the effects of competitive interactions on species’ occurrences and spatial distributions is often difficult, perhaps because these effects occur over longer time scales than the ones usually studied by ecologists (Yackulic 2017). The work by Facon and colleagues shows that weak effects of competition can be detected also at the short ecological timescales that determine coexistence in local communities, under the virtuous combination of good-quality data and sound analytical designs that account for several aspects of species’ niches, their biotopes and their joint population responses. This adds a new dimension to the application of Hutchinson’s (1978) niche framework to understand the spatial dynamics of species and communities (see also Colwell & Rangel 2009), although further advances to incorporate dispersal-driven metacommunity dynamics (see, e.g., Ovaskainen et al. 2016; Leibold et al. 2017) are certainly needed. Nonetheless, this work shows the potential value of in-depth analyses of species coexistence based on combining good-quality field data with well-thought out JSDM applications. If many studies like this are conducted, it is likely that the uprising field of Joint Species Distribution Modelling will improve our understanding of the hierarchical relationships between the different factors affecting species coexistence in ecological communities in the near future.

References

Colwell RK, Rangel TF (2009) Hutchinson’s duality: The once and future niche. Proceedings of the National Academy of Sciences, 106, 19651–19658. https://doi.org/10.1073/pnas.0901650106

Connell JH (1980) Diversity and the Coevolution of Competitors, or the Ghost of Competition Past. Oikos, 35, 131–138. https://doi.org/10.2307/3544421

D’Amen M, Rahbek C, Zimmermann NE, Guisan A (2017) Spatial predictions at the community level: from current approaches to future frameworks. Biological Reviews, 92, 169–187. https://doi.org/10.1111/brv.12222

Dormann CF, Bobrowski M, Dehling DM, Harris DJ, Hartig F, Lischke H, Moretti MD, Pagel J, Pinkert S, Schleuning M, Schmidt SI, Sheppard CS, Steinbauer MJ, Zeuss D, Kraan C (2018) Biotic interactions in species distribution modelling: 10 questions to guide interpretation and avoid false conclusions. Global Ecology and Biogeography, 27, 1004–1016. https://doi.org/10.1111/geb.12759

Facon B, Hafsi A, Masselière MC de la, Robin S, Massol F, Dubart M, Chiquet J, Frago E, Chiroleu F, Duyck P-F, Ravigné V (2021) Joint species distributions reveal the combined effects of host plants, abiotic factors and species competition as drivers of community structure in fruit flies. bioRxiv, 2020.12.07.414326. ver. 4 peer-reviewed and recommended by Peer community in Ecology. https://doi.org/10.1101/2020.12.07.414326

Hortal J, de Bello F, Diniz-Filho JAF, Lewinsohn TM, Lobo JM, Ladle RJ (2015) Seven Shortfalls that Beset Large-Scale Knowledge of Biodiversity. Annual Review of Ecology, Evolution, and Systematics, 46, 523–549. https://doi.org/10.1146/annurev-ecolsys-112414-054400

Hortal J, Roura‐Pascual N, Sanders NJ, Rahbek C (2010) Understanding (insect) species distributions across spatial scales. Ecography, 33, 51–53. https://doi.org/10.1111/j.1600-0587.2009.06428.x

Hutchinson, G.E. (1978) An introduction to population biology. Yale University Press, New Haven, CT.

Leibold MA, Chase JM, Ernest SKM (2017) Community assembly and the functioning of ecosystems: how metacommunity processes alter ecosystems attributes. Ecology, 98, 909–919. https://doi.org/10.1002/ecy.1697

Ovaskainen O, Abrego N (2020) Joint Species Distribution Modelling: With Applications in R. Cambridge University Press, Cambridge. https://doi.org/10.1017/9781108591720

Ovaskainen O, Roy DB, Fox R, Anderson BJ (2016) Uncovering hidden spatial structure in species communities with spatially explicit joint species distribution models. Methods in Ecology and Evolution, 7, 428–436. https://doi.org/10.1111/2041-210X.12502

Pollock LJ, Tingley R, Morris WK, Golding N, O’Hara RB, Parris KM, Vesk PA, McCarthy MA (2014) Understanding co-occurrence by modelling species simultaneously with a Joint Species Distribution Model (JSDM). Methods in Ecology and Evolution, 5, 397–406. https://doi.org/10.1111/2041-210X.12180

Ricklefs RE (2015) Intrinsic dynamics of the regional community. Ecology Letters, 18, 497–503. https://doi.org/10.1111/ele.12431

Soberón J, Nakamura M (2009) Niches and distributional areas: Concepts, methods, and assumptions. Proceedings of the National Academy of Sciences, 106, 19644–19650. https://doi.org/10.1073/pnas.0901637106

Yackulic CB (2017) Competitive exclusion over broad spatial extents is a slow process: evidence and implications for species distribution modeling. Ecography, 40, 305–313. https://doi.org/10.1111/ecog.02836

For decades, the effect of population density on individual performance has been studied by ecologists using both theoretical, observational, and experimental approaches. The generally accepted definition of the Allee effect is a positive correlation between population density and average individual fitness that occurs at low population densities, while individual fitness is typically decreased through intraspecific competition for resources at high population densities.  Allee effects are very relevant in conservation biology because species at low population densities would then be subjected to much higher extinction risks. 

However, due to all kinds of stochasticity, low population numbers are always more vulnerable to extinction than larger population sizes. This effect by itself cannot be necessarily ascribed to lower individual performance at low densities, i.e, Allee effects. Vercken and colleagues (2021) address this challenging question and measure the extent to which average individual fitness is affected by population density analyzing 30 experimental populations. As a model system, they use populations of parasitoid wasps of the genus Trichogramma. They report Allee effect in 8 out 30 experimental populations. Vercken and colleagues's work has several strengths. 

First of all, it is nice to see that they put theory at work. This is a very productive way of using theory in ecology. As a starting point, they look at what simple theoretical population models say about Allee effects (Lewis and Kareiva 1993; Amarasekare 1998; Boukal and Berec 2002). These models invariably predict a one-humped relation between population-density and per-capita growth rate. It is important to remark that pure logistic growth, the paradigm of density-dependence, would never predict such qualitative behavior. It is only when there is a depression of per-capita growth rates at low densities that true Allee effects arise. Second, these authors manage to not only experimentally test this main prediction but also report additional demographic traits that are consistently affected by population density. 

In these wasps, individual performance can be measured in terms of the average number of individuals every adult is able to put into the next generation ---the lambda parameter in their analysis. The first panel in figure 3 shows that the per-capita growth rates are lower in populations presenting Allee effects, the ones showing a one-humped behavior in the relation between per-capita growth rates and population densities (see figure 2). Also other population traits, such maximum population size and exitinction probability, change in a correlated and consistent manner. 

In sum, Vercken and colleagues's results are experimentally solid and based on theory expectations. However, they are very intriguing. They find the signature of Allee effects in only 8 out 30 populations, all from the same genus Trichogramma, and some populations belonging to the same species (from different sampling sites) do not show consistently Allee effects. Where does this population variability comes from? What are the reasons underlying this within- and between-species variability? What are the individual mechanisms driving Allee effects in these populations? Good enough, this piece of work generates more intriguing questions than the question is able to clearly answer. Science is not a collection of final answers but instead good questions are the ones that make science progress. 

References

Amarasekare P (1998) Allee Effects in Metapopulation Dynamics. The American Naturalist, 152, 298–302. https://doi.org/10.1086/286169

Boukal DS, Berec L (2002) Single-species Models of the Allee Effect: Extinction Boundaries, Sex Ratios and Mate Encounters. Journal of Theoretical Biology, 218, 375–394. https://doi.org/10.1006/jtbi.2002.3084

Lewis MA, Kareiva P (1993) Allee Dynamics and the Spread of Invading Organisms. Theoretical Population Biology, 43, 141–158. https://doi.org/10.1006/tpbi.1993.1007

Vercken E, Groussier G, Lamy L, Mailleret L (2021) The hidden side of the Allee effect: correlated demographic traits and extinction risk in experimental populations. HAL, hal-02570868, ver. 4 peer-reviewed and recommended by Peer community in Ecology. https://hal.archives-ouvertes.fr/hal-02570868