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14 Jul 2023
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Field margins as substitute habitat for the conservation of birds in agricultural wetlands

Searching for conservation opportunities at the margins

Recommended by based on reviews by Scott Wilson and Elena D Concepción

In a progressively human-dominated planet (Venter et al., 2016), the fate of many species will depend on the extent to which they can persist in anthropogenic landscapes. In Western Europe, where only small areas of primary habitat remain (e.g. Sabatini et al., 2018), semi-natural areas are crucial habitats to many native species, yet they are threatened by the expansion of human activities, including agricultural expansion and intensification (Rigal et al., 2023). 

A new study by Mallet and colleagues (Mallet et al., 2023) investigates the extent to which bird species in the Camargue region are able to use the margins of agricultural fields as substitutes for their preferred semi-natural habitats. Located in the delta of the Rhône River in Southern France, the Camargue is internationally recognized for its biodiversity value, classified as a Biosphere Reserve by UNESCO and as a Wetland of International Importance under the Ramsar Convention (IUCN & UN-WCMC, 2023). Mallet and colleagues tested three specific hypotheses: that grass strips (grassy field boundaries, including grassy tracks or dirt roads used for moving agricultural machinery) can function as substitute habitats for grassland species; that reed strips along drainage ditches (common in the rice paddy landscapes of the Camargue) can function as substitute habitats to wetland species; and that hedgerows can function as substitute habitats to species that favour woodland edges. They did so by measuring how the local abundances of 14 bird species (nine typical of forest edges, 3 of grasslands, and two of reedbeds) respond to increasing coverage of either the three types of field margins or of the three types of semi-natural habitat. 

This is an elegant study design, yet – as is often the case with real field data – results are not as simple as expected. Indeed, for most species (11 out of 14) local abundances did not increase significantly with the area of their supposed primary habitat, undermining the assumption that they are strongly associated with (or dependent on) those habitats. Among the three species that did respond positively to the area of their primary habitat, one (a forest edge species) responded positively but not significantly to the area of field margins (hedgerows), providing weak evidence to the habitat compensation hypothesis. For the other two (grassland and a wetland species), abundance responded even more strongly to the area of field margins (grass and reed strips, respectively) than to the primary habitat, suggesting that the field margins are not so much a substitute but valuable habitats in their own right. 

It would have been good conservation news if field margins were found to be suitable habitat substitutes to semi-natural habitats, or at least reasonable approximations, to most species. Given that these margins have functional roles in agricultural landscapes (marking boundaries, access areas, water drainage), they could constitute good win-win solutions for reconciling biodiversity conservation with agricultural production. Alas, the results are more complicated than that, with wide variation in species responses that could not have been predicted from presumed habitat affinities. These results illustrate the challenges of conservation practice in complex landscapes formed by mosaics of variable land use types. With species not necessarily falling neatly into habitat guilds, it becomes even more challenging to plan strategically how to manage landscapes to optimize their conservation. The results presented here suggest that species’ abundances may be responding to landscape variables not taken into account in the analyses, such as connectivity between habitat patches, or maybe positive and negative edge effects between land use types. That such uncertainties remain even in a well-studied region as the Camargue, and for such a well-studied taxon such as birds, only demonstrates the continued importance of rigorous field studies testing explicit hypotheses such as this one by Mallet and colleagues. 


IUCN, & UN-WCMC (2023). Protected Planet. Protected Planet. 

Mallet, P., Béchet, A., Sirami, C., Mesléard, F., Blanchon, T., Calatayud, F., Dagonet, T., Gaget, E., Leray, C., & Galewski, T. (2023). Field margins as substitute habitat for the conservation of birds in agricultural wetlands. bioRxiv, 2022.05.05.490780, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. 

Rigal, S., Dakos, V., Alonso, H., Auniņš, A., Benkő, Z., Brotons, L., Chodkiewicz, T., Chylarecki, P., de Carli, E., del Moral, J. C. et al. (2023). Farmland practices are driving bird population decline across Europe. Proceedings of the National Academy of Sciences, 120, e2216573120. 

Sabatini, F. M., Burrascano, S., Keeton, W. S., Levers, C., Lindner, M., Pötzschner, F., Verkerk, P. J., Bauhus, J., Buchwald, E., Chaskovsky, O., Debaive, N. et al. (2018). Where are Europe’s last primary forests? Diversity and Distributions, 24, 1426–1439. 

Venter, O., Sanderson, E. W., Magrach, A., Allan, J. R., Beher, J., Jones, K. R., Possingham, H. P., Laurance, W. F., Wood, P., Fekete, B. M., Levy, M. A., & Watson, J. E. M. (2016). Sixteen years of change in the global terrestrial human footprint and implications for biodiversity conservation. Nature Communications, 7, 12558. 

Field margins as substitute habitat for the conservation of birds in agricultural wetlandsMallet Pierre, Béchet Arnaud, Sirami Clélia, Mesléard François, Blanchon Thomas, Calatayud François, Dagonet Thomas, Gaget Elie, Leray Carole, Galewski Thomas<p style="text-align: justify;">Breeding birds in agricultural landscapes have declined considerably since the 1950s and the beginning of agricultural intensification in Europe. Given the increasing pressure on agricultural land, it is necessary t...Agroecology, Biodiversity, Conservation biology, Landscape ecologyAna S. L. Rodrigues2022-05-09 10:48:49 View
13 Jul 2023
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Parasites make hosts more profitable but less available to predators

Indirect effects of parasitism include increased profitability of prey to optimal foragers

Recommended by based on reviews by Thierry DE MEEUS and Eglantine Mathieu-Bégné

Even though all living organisms are, at the same time, involved in host-parasite interactions and embedded in complex food webs, the indirect effects of parasitism are only beginning to be unveiled.

Prosnier et al. investigated the direct and indirect effects of parasitism making use of a very interesting biological system comprising the freshwater zooplankton Daphnia magna and its highly specific parasite, the iridovirus DIV-1 (Daphnia-iridescent virus 1). Daphnia are typically semitransparent, but once infected develop a white phenotype with a characteristic iridescent shine due to the enlargement of white fat cells.

In a combination of infection trials and comparison of white and non-white phenotypes collected in natural ponds, the authors demonstrated increased mortality and reduced lifetime fitness in infected Daphnia. Furthermore, white phenotypes had lower mobility, increased reflectance, larger body sizes and higher protein content than non-white phenotypes. As a consequence, total energy content was effectively doubled in white Daphnia when compared to non-white broodless Daphnia

Next the authors conducted foraging trials with Daphnia predators Notonecta (the backswimmer) and Phoxinus (the European minnow). Focusing on Notonecta, unchanged search time and increased handling time were more than compensated by the increased energy content of white Daphnia. White Daphnia were 24% more profitable and consistently preferred by Notonecta, as the optimal foraging theory would predict. The authors argue that menu decisions of optimal foragers in the field might be different, however, as the prevalence – and therefore availability - of white phenotypes in natural populations is very low.

The study therefore contributes to our understanding of the trophic context of parasitism. One shortcoming of the study is that the authors rely exclusively on phenotypic signs for determining infection. On their side, DIV-1 is currently known to be highly specific to Daphnia, their study site is well within DIV-1 distributional range, and the symptoms of infection are very conspicuous. Furthermore, the infection trial – in which non-white Daphnia were exposed to white Daphnia homogenates - effectively caused several lethal and sublethal effects associated with DIV-1 infection, including iridescence. However, the infection trial also demonstrated that part of the exposed individuals developed intermediate traits while still keeping the non-white, non-iridescent phenotype. Thus, there may be more subtleties to the association of DIV-1 infection of Daphnia with ecological and evolutionary consequences, such as costs to resistance or covert infection, that the authors acknowledge, and that would be benefitted by coupling experimental and observational studies with the determination of actual infection and viral loads.​​​


Prosnier L., N. Loeuille, F.D. Hulot, D. Renault, C. Piscart, B. Bicocchi, M, Deparis, M. Lam, & V. Médoc. (2023). Parasites make hosts more profitable but less available to predators. BioRxiv, ver. 4 peer-reviewed and recommended by Peer Community in Ecology.

Parasites make hosts more profitable but less available to predatorsLoïc Prosnier, Nicolas Loeuille, Florence D. Hulot, David Renault, Christophe Piscart, Baptiste Bicocchi, Muriel Deparis, Matthieu Lam, Vincent Médoc<p>Parasites are omnipresent, and their eco-evolutionary significance has aroused much interest from scientists. Parasites may affect their hosts in many ways by altering host density, vulnerability to predation, and energy content, thus modifying...Community ecology, Eco-evolutionary dynamics, Epidemiology, Experimental ecology, Food webs, Foraging, Freshwater ecology, Host-parasite interactions, Life history, Parasitology, Statistical ecologyLuis Schiesari2022-05-20 10:15:41 View
03 Feb 2023
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The role of climate change and niche shifts in divergent range dynamics of a sister-species pair

Drivers of range expansion in a pair of sister grackle species

Recommended by ORCID_LOGO based on reviews by 2 anonymous reviewers

The spatial distribution of a species is driven by both biotic and abiotic factors that may change over time (Soberón & Nakamura, 2009; Paquette & Hargreaves, 2021).  Therefore, species ranges are dynamic, especially in humanized landscapes where changes occur at high speeds (Sirén & Morelli, 2020). The distribution of many species is being reduced because of human impacts; however, some species are expanding their distributions, even over their niche (Lustenhouwer & Parker, 2022). One of the factors that may lead to a geographic niche expansion is behavioral flexibility (Mikhalevich et al., 2017), but the mechanisms determining range expansion through behavioral changes are not fully understood. 

The PCI Ecology study by Summers et al. (2023) uses a very large database on the current and historic distribution of two species of grackles that have shown different trends in their distribution. The great-tailed grackle has largely expanded its range over the 20th century, while the range of the boat-tailed grackle has remained very similar. They take advantage of this differential response in the distribution of the two species and run several analyses to test whether it was a change in habitat availability, in the realized niche, in habitat connectivity or in in the other traits or conditions that previously limited the species range, what is driving the observed distribution of the species. The study finds a change in the niche of great-tailed grackle, consistent with the high behavioral flexibility of the species.

The two reviewers and I have seen a lot of value in this study because 1) it addresses a very timely question, especially in the current changing world; 2) it is a first step to better understanding if behavioral attributes may affect species’ ability to change their niche; 3) it contrasts the results using several complementary statistical analyses, reinforcing their conclusions; 4) it is based on the preregistration Logan et al (2021), and any deviations from it are carefully explained and justified in the text and 5) the limitations of the study have been carefully discussed. It remains to know if the boat-tailed grackle has more limited behavioral flexibility than the great-tailed grackle, further confirming the results of this study.

Logan CJ, McCune KB, Chen N, Lukas D (2021) Implementing a rapid geographic range expansion - the role of behavior and habitat changes.

Lustenhouwer N, Parker IM (2022) Beyond tracking climate: Niche shifts during native range expansion and their implications for novel invasions. Journal of Biogeography, 49, 1481–1493.

Mikhalevich I, Powell R, Logan C (2017) Is behavioural flexibility evidence of cognitive complexity? How evolution can inform comparative cognition. Interface Focus, 7, 20160121.

Paquette A, Hargreaves AL (2021) Biotic interactions are more often important at species’ warm versus cool range edges. Ecology Letters, 24, 2427–2438.

Sirén APK, Morelli TL (2020) Interactive range-limit theory (iRLT): An extension for predicting range shifts. Journal of Animal Ecology, 89, 940–954.

Soberón J, Nakamura M (2009) Niches and distributional areas: Concepts, methods, and assumptions. Proceedings of the National Academy of Sciences, 106, 19644–19650.

Summers JT, Lukas D, Logan CJ, Chen N (2022) The role of climate change and niche shifts in divergent range dynamics of a sister-species pair. EcoEvoRxiv, ver. 3 peer-reviewed and recommended by Peer Community in Ecology.

The role of climate change and niche shifts in divergent range dynamics of a sister-species pairJeremy Summers, Dieter Lukas, Corina J. Logan, Nancy Chen<p>---This is a POST-STUDY manuscript for the PREREGISTRATION, which received in principle acceptance in 2020 from Dr. Sebastián González (reviewed by Caroline Nieberding, Tim Parker, and Pizza Ka Yee Chow; <a href=" & Ethology, Biogeography, Dispersal & Migration, Human impact, Landscape ecology, Preregistrations, Species distributionsEsther Sebastián González2022-05-26 20:07:33 View
20 Feb 2023
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Best organic farming deployment scenarios for pest control: a modeling approach

Towards model-guided organic farming expansion for crop pest management

Recommended by based on reviews by Julia Astegiano, Lionel Hertzog and Sylvain Bart

Reduce the impact the intensification of human activities has on the environmental is the challenge the humanity faces today, a major challenge that could be compared to climbing Everest without an oxygen supply. Indeed, over-population, pollution, burning fossil fuels, and deforestation are all evils which have had hugely detrimental effects on the environment such as climate change, soil erosion, poor air quality, and scarcity of drinking water to name but a few. In response to the ever-growing consumer demand, agriculture has intensified massively along with a drastic increase in the use of chemicals to ensure an adequate food supply while controlling crop pests. In this context, to address the disastrous effects of the intensive usage of pesticides on both human health and biodiversity, organic farming (OF) revealed as a miracle remedy with multiple benefits. Delattre et al. (2023) present a powerful modelling approach to decipher the crossed effects of the landscape structure and the OF expansion scenario on the pest abundance, both in organic and conventional (CF) crop fields. To this end, the authors ingeniously combined a grid-based landscape model with a spatially explicit predator-pest model. Based on an extensive in silico simulation process, they explore a diversity of landscape structures differing in their amount of semi-natural habitats (SHN) and in their fragmentation, to finally propose a ranking of various expansion scenarios according to the pest control methods in organic farming as well as to the pest and predators’ dissemination capacities. In total, 9 landscape structures (3 proportions of SHN x 3 fragmentation levels) were crossed with 3 expansion scenarios (RD = a random distribution of OF and CF in the grid; IP = isolated CF are converted; GP = CF within aggregates are converted), 4 pest management practices, 3 initial densities and 36 biological parameter combinations driving the predator’ and pest’s population dynamics. This exhaustive exploration of possible combinations of landscape and farming practices highlighted the main drivers of the various OF expansion scenarios, such as increased spillover of predators in isolated OF/CF fields, increased pest management efficiency in large patches of CF and the importance of the distance between OF and CF. In the end, this study brings to light the crucial role that landscape planning plays when OF practices have limited efficiency on pests. It also provides convincing arguments to the fact that converting to organic isolated CF as a priority seems to be the most promising scenario to limit pest densities in CF crops while improving predator to pest ratios (considered as a proxy of conservation biological control) in OF ones without increasing pest densities. Once further completed with model calibration validation based on observed life history traits data for both predators and pests, this work should be very helpful in sustaining policy makers to convince farmers of engaging in organic farming.


Delattre T, Memah M-M, Franck P, Valsesia P, Lavigne C (2023) Best organic farming deployment scenarios for pest control: a modeling approach. bioRxiv, 2022.05.31.494006, ver. 2 peer-reviewed and recommended by Peer Community in Ecology.

Best organic farming deployment scenarios for pest control: a modeling approachThomas Delattre, Mohamed-Mahmoud Memah, Pierre Franck, Pierre Valsesia, Claire Lavigne<p style="text-align: justify;">Organic Farming (OF) has been expanding recently around the world in response to growing consumer demand and as a response to environmental concerns. Its share of agricultural landscapes is expected to increase in t...Agroecology, Biological control, Landscape ecologySandrine Charles2022-06-03 11:41:14 View
15 Jul 2023
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Evolution of dispersal and the maintenance of fragmented metapopulations

The spatial dynamics of habitat fragmentation drives the evolution of dispersal and metapopulation persistence

Recommended by based on reviews by Eva Kisdi, David Murray-Stoker, Shripad Tuljapurkar and 1 anonymous reviewer

​​​​​The persistence of populations facing the destruction of their habitat is a multifaceted question that has mobilized theoreticians and empiricists alike for decades. As an ecological question, persistence has been studied as the spatial rescue of populations via dispersal into remaining suitable habitats. The spatial aggregation of habitat destruction has been a key component of these studies, and it has been applied to the problem of coexistence by integrating competition-colonization tradeoffs. There is a rich ecological literature on this topic, both from theoretical and field studies (Fahrig 2003). The relationship between life-history strategies of species and their resilience to spatially structured habitat fragmentation is also an important component of conservation strategies through the management of land use, networks of protected areas, and the creation of corridors. In the context of environmental change, the ability of species to adapt to changes in landscape configuration and availability can be treated as an eco-evolutionary process by considering the possibility of evolutionary rescue (Heino and Hanski 2001; Bell 2017). However, eco-evolutionary dynamics considering spatially structured changes in landscapes and life-history tradeoffs remains an outstanding question. Finand et al. (2023) formulate the problem of persistence in fragmented landscapes over evolutionary time scales by studying models for the evolution of dispersal in relation to habitat fragmentation and spatial aggregation. Their simulations were conducted on a spatial grid where individuals can colonize suitable patch as a function of their competitive rank that decreases as a function of their (ii) dispersal distance trait. Simulations were run under fixed habitat fragmentation (proportion of unsuitable habitat) and aggregation, and with an explicit rate of habitat destruction to study evolutionary rescue.

Their results reveal a balance between the selection for high dispersal under increasing habitat fragmentation and selection for lower dispersal in response to habitat aggregation. This balance leads to the coexistence of polymorphic dispersal strategies in highly aggregated landscapes with low fragmentation where high dispersers inhabit aggregated habitats while low dispersers are found in isolated habitats. The authors then integrate the spatial rescue mechanism to the problem of evolutionary rescue in response to temporally increasing fragmentation. There they show how rapid evolution allows for evolutionary rescue through the evolution of high dispersal. They also show the limits to this evolutionary rescue to cases where both aggregation and fragmentation are not too high. Interestingly, habitat aggregation prevents evolutionary rescue by directly affecting the evolutionary potential of dispersal. The study is based on simple scenarios that ignore the complexity of relationships between dispersal, landscape properties, and species interactions. This simplicity is the strength of the study, revealing basic mechanisms that can now be tested against other life-history tradeoffs and species interactions. Finand et al. (2023) provide a novel foundation for the study of eco-evolutionary dynamics in metacommunities exposed to spatially structured habitat destruction. They point to important assumptions that must be made along the way, including the relationships between dispersal distance and fecundity (they assume a positive relationship), and the nature of life-history tradeoffs between dispersal rate and local competitive abilities. 


Bell, G. 2017. Evolutionary Rescue. Annual Review of Ecology, Evolution, and Systematics 48:605–627. 
Fahrig, L. 2003. Effects of Habitat Fragmentation on Biodiversity. Annual Review of Ecology, Evolution, and Systematics 34:487–515. 
Finand, B., T. Monnin, and N. Loeuille. 2023. Evolution of dispersal and the maintenance of fragmented metapopulations. bioRxiv, 2022.06.08.495260, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. 
Heino, M., and I. Hanski. 2001. Evolution of Migration Rate in a Spatially Realistic Metapopulation Model. The American Naturalist 157:495–511.

Evolution of dispersal and the maintenance of fragmented metapopulationsBasile Finand, Thibaud Monnin, Nicolas Loeuille<p>Because it affects dispersal risk and modifies competition levels, habitat fragmentation directly constrains dispersal evolution. When dispersal is traded-off against competitive ability, increased fragmentation is often expected to select high...Colonization, Competition, Dispersal & Migration, Eco-evolutionary dynamics, Spatial ecology, Metacommunities & Metapopulations, Theoretical ecologyFrédéric Guichard2022-06-10 13:51:15 View
12 Mar 2023
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Different approaches to processing environmental DNA samples in turbid waters have distinct effects for fish, bacterial and archaea communities.

Processing environmental DNA samples in turbid waters from coastal lagoons

Recommended by based on reviews by David Murray-Stoker and Rutger De Wit

Coastal lagoons are among the most productive natural ecosystems on Earth. These relatively closed basins are important habitats and nursery for endemic and endangered species and are extremely vulnerable to nutrient input from the surrounding catchment; therefore, they are highly susceptible to anthropogenic influence, pollution and invasion (Pérez-Ruzafa et al., 2019). In general, coastal lagoons exhibit great spatial and temporal variability in their physicochemical water characteristics due to the sporadic mixing of freshwater with marine influx. One of the alternatives for monitoring communities or target species in aquatic ecosystems is the environmental DNA (eDNA), since overcomes some limitations from traditional methods and enables the investigation of multiple species from a single sample (Thomsen and Willerslev, 2015). In coastal lagoons, where the water turbidity is highly variable, there is a major challenge for monitoring the eDNA because filtering turbid water to obtain the eDNA is problematic (filters get rapidly clogged, there is organic and inorganic matter accumulation, etc.). 

The study by Turba et al. (2023) analyzes different ways of dealing with eDNA sampling and processing in turbid waters and sediments of coastal lagoons, and offers guidelines to obtain unbiased results from the subsequent sequencing using 12S (fish) and 16S (Bacteria and Archaea) universal primers. They analyzed the effect on taxa detection of: i) freezing or not prior to filtering; ii) freezing prior to centrifugation to obtain a sample pellet; and iii) using frozen sediment samples as a proxy of what happens in the water. The authors propose these different alternatives (freeze, do not freeze, sediment sampling) because they consider that they are the easiest to carry out. They found that freezing before filtering using a 3 µm pore size filter had no effects on community composition for either primer, and therefore it is a worthwhile approach for comparison of fish, bacteria and archaea in this kind of system. However, significantly different bacterial community composition was found for sediment compared to water samples. Also, in sediment samples the replicates showed to be more heterogeneous, so the authors suggest increasing the number of replicates when using sediment samples. Something that could be a concern with the study is that the rarefaction curves based on sequencing effort for each protocol did not saturate in any case, this being especially evident in sediment samples. The authors were aware of this, used the slopes obtained from each curve as a measure of comparison between samples and considering that the sequencing depth did not meet their expectations, they managed to achieve their goal and to determine which protocol is the most promising for eDNA monitoring in coastal lagoons. Although there are details that could be adjusted in relation to this item, I consider that the approach is promising for this type of turbid system.


Pérez-Ruzafa A, Campillo S, Fernández-Palacios JM, García-Lacunza A, García-Oliva M, Ibañez H, Navarro-Martínez PC, Pérez-Marcos M, Pérez-Ruzafa IM, Quispe-Becerra JI, Sala-Mirete A, Sánchez O, Marcos C (2019) Long-Term Dynamic in Nutrients, Chlorophyll a, and Water Quality Parameters in a Coastal Lagoon During a Process of Eutrophication for Decades, a Sudden Break and a Relatively Rapid Recovery. Frontiers in Marine Science, 6.

Thomsen PF, Willerslev E (2015) Environmental DNA – An emerging tool in conservation for monitoring past and present biodiversity. Biological Conservation, 183, 4–18.

Turba R, Thai GH, Jacobs DK (2023) Different approaches to processing environmental DNA samples in turbid waters have distinct effects for fish, bacterial and archaea communities. bioRxiv, 2022.06.17.495388, ver. 2 peer-reviewed and recommended by Peer Community in Ecology.

Different approaches to processing environmental DNA samples in turbid waters have distinct effects for fish, bacterial and archaea communities.Rachel Turba, Glory H. Thai, and David K Jacobs<p style="text-align: justify;">Coastal lagoons are an important habitat for endemic and threatened species in California that have suffered impacts from urbanization and increased drought. Environmental DNA has been promoted as a way to aid in th...Biodiversity, Community genetics, Conservation biology, Freshwater ecology, Marine ecology, Molecular ecologyClaudia Piccini David Murray-Stoker2022-06-20 20:31:51 View
14 Dec 2022
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The contrasted impacts of grasshoppers on soil microbial activities in function of primary production and herbivore diet

Complex interactions between ecosystem productivity and herbivore diets lead to non-predicted effects on nutrient cycling

Recommended by based on reviews by Manuel Blouin and Tord Ranheim Sveen

The authors present a study typical of the field of belowground-aboveground interactions [1]. This framework has been extremely fruitful since the beginning of 2000s [2]. It has also contributed to bridge the gap between soil ecology and the rest of ecology [3]. The study also pertains to the rich field on the impacts of herbivores on soil functioning [4].

The study more precisely tested during two years the effect on nutrient cycling of the interaction between the type of grassland (along a gradient of biomass productivity) and the diet of the community of insect herbivores (5 treatments manipulating the grasshopper community on 1 m2 plots, with a gradient from no grasshopper to grasshoppers either specialized on forbs or grasses). What seems extremely interesting is that the study is based on a rigorous hypothesis-testing approach. They compare the predictions of two frameworks: (1) The “productivity model” predicts that in productive ecosystems herbivores consume a high percentage of the net primary production thus accelerating nutrient cycling. (2) The “diet model” distinguishes herbivores consuming exploitative plants from those eating conservative plants. The former (later) type of herbivores favours conservative (exploitative) plants therefore decelerating (accelerating) nutrient cycling. Interestingly, the two frameworks have similar predictions (and symmetrically opposite predictions) in two cases out of four combinations between ecosystem productivities and types of diet (see Table 1). An other merit of the study is to combine in a rather comprehensive way all the necessary measurements to test these frameworks in combination: grasshopper diet, soil properties, characteristics of the soil microbial community, plant traits, vegetation survey and plant biomass.

The results were in contradiction with the ‘‘diet model’’: microbial properties and nitrogen cycling did not depend on grasshopper diet. The productivity of the grasslands did impact nutrient cycling but not in the direction predicted by the “productivity model”: productive grasslands hosted exploitative plants that depleted N resources in the soil and microbes producing few extracellular enzymes, which led to a lower potential N mineralization and a deceleration of nutrient cycling. Because, the authors stuck to their original hypotheses (that were not confirmed), they were able to discuss in a very relevant way their results and to propose some interpretations, at least partially based on the time scales involved by the productivity and diet models.

Beyond all the merits of this article, I think that two issues remain largely open in relation with the dynamics of the studied systems, and would deserve future research efforts. First, on the ‘‘short’’ term (up to several decades), can we predict how the communities of plants, soil microbes, and herbivores interact to drive the dynamics of the ecosystems? Second, at the evolutionary time scale, can we understand and predict the interactions between the evolution of plant, microbe and herbivore strategies and the consequences for the functioning of the grasslands? The two issues are difficult because of the multiple feedbacks involved. One way to go further would be to complement the empirical approach with models along existing research avenues [5, 6]. 


[1] Ibanez S, Foulquier A, Brun C, Colace M-P, Piton G, Bernard L, Gallet C, Clément J-C (2022) The contrasted impacts of grasshoppers on soil microbial activities in function of primary production and herbivore diet. bioRxiv, 2022.07.04.497718, ver. 2 peer-reviewed and recommended by Peer Community in Ecology.

[2] Hooper, D. U., Bignell, D. E., Brown, V. K., Brussaard, L., Dangerfield, J. M., Wall, D. H., Wardle, D. A., Coleman, D. C., Giller, K. E., Lavelle, P., Van der Putten, W. H., De Ruiter, P. C., et al. 2000. Interactions between aboveground and belowground biodiversity in terretrial ecosystems: patterns, mechanisms, and feedbacks. BioScience, 50, 1049-1061.[1049:IBAABB]2.0.CO;2

[3] Barot, S., Blouin, M., Fontaine, S., Jouquet, P., Lata, J.-C., and Mathieu, J. 2007. A tale of four stories: soil ecology, theory, evolution and the publication system. PLoS ONE, 2, e1248.

[4] Bardgett, R. D., and Wardle, D. A. 2003. Herbivore-mediated linkages between aboveground and belowground communities. Ecology, 84, 2258-2268.

[5] Barot, S., Bornhofen, S., Loeuille, N., Perveen, N., Shahzad, T., and Fontaine, S. 2014. Nutrient enrichment and local competition influence the evolution of plant mineralization strategy, a modelling approach. J. Ecol., 102, 357-366.

[6] Schweitzer, J. A., Juric, I., van de Voorde, T. F. J., Clay, K., van der Putten, W. H., Bailey, J. K., and Fox, C. 2014. Are there evolutionary consequences of plant-soil feedbacks along soil gradients? Func. Ecol., 28, 55-64.


The contrasted impacts of grasshoppers on soil microbial activities in function of primary production and herbivore dietSébastien Ibanez, Arnaud Foulquier, Charles Brun, Marie-Pascale Colace, Gabin Piton, Lionel Bernard, Christiane Gallet, Jean-Christophe Clément<p style="text-align: justify;">Herbivory can have contrasted impacts on soil microbes and nutrient cycling, which has stimulated the development of conceptual frameworks exploring the links between below- and aboveground processes. The "productiv...Ecosystem functioning, Herbivory, Soil ecology, Terrestrial ecologySébastien Barot2022-07-14 09:06:13 View
17 May 2023
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Distinct impacts of food restriction and warming on life history traits affect population fitness in vertebrate ectotherms

Effect of food conditions on the Temperature-Size Rule

Recommended by based on reviews by Wolf Blanckenhorn and Wilco Verberk

Temperature-size rule (TSR) is a phenomenon of plastic changes in body size in response to temperature, originally observed in more than 80% of ectothermic organisms representing various groups (Atkinson 1994). In particular, ectotherms were observed to grow faster and reach smaller size at higher temperature and grow slower and achieve larger size at lower temperature. This response has fired the imagination of researchers since its invention, due to its counterintuitive pattern from an evolutionary perspective (Berrigan and Charnov 1994). The main question to be resolved is: why do organisms grow fast and achieve smaller sizes under more favourable conditions (= relatively higher temperature), while they grow longer and achieve larger sizes under less favourable conditions (relatively lower temperature), if larger size means higher fitness, while longer development may be risky? 

This evolutionary conundrum still awaits an ultimate explanation (Angilletta Jr et al. 2004; Angilletta and Dunham 2003; Verberk et al. 2021). Although theoretical modelling has shown that such a growth pattern can be achieved as a response to temperature alone, with a specific combination of energetic parameters and external mortality (Kozłowski et al. 2004), it has been suggested that other temperature-dependent environmental variables may be the actual drivers of this pattern. One of the most frequently invoked variable is the relative oxygen availability in the environment (e.g., Atkinson et al. 2006; Audzijonyte et al. 2019; Verberk et al. 2021; Woods 1999), which decreases with temperature increase. Importantly, this effect is more pronounced in aquatic systems (Forster et al. 2012). However, other temperature-dependent parameters are also being examined in the context of their possible effect on TSR induction and strength.

Food availability is among the interfering factors in this regard. In aquatic systems, nutritional conditions are generally better at higher temperature, while a range of relatively mild thermal conditions is considered. However, there are no conclusive results so far on how nutritional conditions affect the plastic body size response to acute temperature changes. A study by Bazin et al. (2023) examined this particular issue, the effects of food and temperature on TSR, in medaka fish. An important value of the study was to relate the patterns found to fitness. This is a rare and highly desirable approach since evolutionary significance of any results cannot be reliably interpreted unless shown as expressed in light of fitness. 

The authors compared the body size of fish kept at 20°C and 30°C under conditions of food abundance or food restriction. The results showed that the TSR (smaller body size at 30°C compared to 20°C) was observed in both food treatments, but the effect was delayed during fish development under food restriction. Regarding the relevance to fitness, increased temperature resulted in more eggs laid but higher mortality, while food restriction increased survival but decreased the number of eggs laid in both thermal treatments. Overall, food restriction seemed to have a more severe effect on development at 20°C than at 30°C, contrary to the authors’ expectations. 

I found this result particularly interesting. One possible interpretation, also suggested by the authors, is that the relative oxygen availability, which was not controlled for in this study, could have affected this pattern. According to theoretical predictions confirmed in quite many empirical studies so far, oxygen restriction is more severe at higher temperatures. Perhaps for these particular two thermal treatments and in the case of the particular species studied, this restriction was more severe for organismal performance than the food restriction. This result is an example that all three variables, temperature, food and oxygen, should be taken into account in future studies if the interrelationship between them is to be understood in the context of TSR. It also shows that the reasons for growing smaller in warm may be different from those for growing larger in cold, as suggested, directly or indirectly, in some previous studies (Hessen et al. 2010; Leiva et al. 2019). 

Since medaka fish represent predatory vertebrates, the results of the study contribute to the issue of global warming effect on food webs, as the authors rightly point out. This is an important issue because the general decrease in the size or organisms in the aquatic environment with global warming is a fact (e.g., Daufresne et al. 2009), while the question of how this might affect entire communities is not trivial to resolve (Ohlberger 2013). 


Angilletta Jr, M. J., T. D. Steury & M. W. Sears, 2004. Temperature, growth rate, and body size in ectotherms: fitting pieces of a life–history puzzle. Integrative and Comparative Biology 44:498-509.

Angilletta, M. J. & A. E. Dunham, 2003. The temperature-size rule in ectotherms: Simple evolutionary explanations may not be general. American Naturalist 162(3):332-342.

Atkinson, D., 1994. Temperature and organism size – a biological law for ectotherms. Advances in Ecological Research 25:1-58.

Atkinson, D., S. A. Morley & R. N. Hughes, 2006. From cells to colonies: at what levels of body organization does the 'temperature-size rule' apply? Evolution & Development 8(2):202-214

Audzijonyte, A., D. R. Barneche, A. R. Baudron, J. Belmaker, T. D. Clark, C. T. Marshall, J. R. Morrongiello & I. van Rijn, 2019. Is oxygen limitation in warming waters a valid mechanism to explain decreased body sizes in aquatic ectotherms? Global Ecology and Biogeography 28(2):64-77

Bazin, S., Hemmer-Brepson, C., Logez, M., Sentis, A. & Daufresne, M. 2023. Distinct impacts of food restriction and warming on life history traits affect population fitness in vertebrate ectotherms. HAL, ver.3  peer-reviewed and recommended by PCI Ecology.

Berrigan, D. & E. L. Charnov, 1994. Reaction norms for age and size at maturity in response to temperature – a puzzle for life historians. Oikos 70:474-478.

Daufresne, M., K. Lengfellner & U. Sommer, 2009. Global warming benefits the small in aquatic ecosystems. Proceedings of the National Academy of Sciences USA 106(31):12788-93

Forster, J., A. G. Hirst & D. Atkinson, 2012. Warming-induced reductions in body size are greater in aquatic than terrestrial species. Proceedings of the National Academy of Sciences of the United States of America 109(47):19310-19314.

Hessen, D. O., P. D. Jeyasingh, M. Neiman & L. J. Weider, 2010. Genome streamlining and the elemental costs of growth. Trends in Ecology & Evolution 25(2):75-80.

Kozłowski, J., M. Czarnoleski & M. Dańko, 2004. Can optimal resource allocation models explain why ectotherms grow larger in cold? Integrative and Comparative Biology 44(6):480-493.

Leiva, F. P., P. Calosi & W. C. E. P. Verberk, 2019. Scaling of thermal tolerance with body mass and genome size in ectotherms: a comparison between water- and air-breathers. Philosophical Transactions of the Royal Society B 374:20190035.

Ohlberger, J., 2013. Climate warming and ectotherm body szie - from individual physiology to community ecology. Functional Ecology 27:991-1001.

Verberk, W. C. E. P., D. Atkinson, K. N. Hoefnagel, A. G. Hirst, C. R. Horne & H. Siepel, 2021. Shrinking body sizes in response to warming: explanations for the temperature-size rule with special emphasis on the role of oxygen. Biological Reviews 96:247-268.

Woods, H. A., 1999. Egg-mass size and cell size: effects of temperature on oxygen distribution. American Zoologist 39:244-252.

Distinct impacts of food restriction and warming on life history traits affect population fitness in vertebrate ectothermsSimon Bazin, Claire Hemmer-Brepson, Maxime Logez, Arnaud Sentis, Martin Daufresne<p>The reduction of body size with warming has been proposed as the third universal response to global warming, besides geographical and phenological shifts. Observed body size shifts in ectotherms are mostly attributed to the temperature size rul...Climate change, Experimental ecology, Freshwater ecology, Phenotypic plasticity, Population ecologyAleksandra Walczyńska2022-07-27 09:28:29 View
07 Jun 2023
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High intraspecific growth variability despite strong evolutionary heritage in a neotropical forest

Environmental and functional determinants of tree performance in a neotropical forest: the imprint of evolutionary legacy on growth strategies

Recommended by ORCID_LOGO based on reviews by David Murray-Stoker, Camille Girard and Jelena Pantel

The hyperdiverse tropical forests have long fascinated ecologists because the fact that so many species persist at a low density at a local scale remains hard to explain. Both niche-based and neutral hypotheses have been tested, primarily based on analyzing the taxonomic composition of tropical forest plots (Janzen 1970; Hubbell 2001). Studies of the functional and phylogenetic structure of tropical tree communities have further aimed to better assess the importance of niche-based processes. For instance, Baraloto et al. (2012) found that co-occurring species were functionally and phylogenetically more similar in a neotropical forest, suggesting a role of environmental filtering. Likewise, Schmitt et al. (2021) found the influence of environmental filtering on the functional composition of an Indian rainforest. Yet these studies evidenced non-random trait-environment association based on the composition of assemblages only (in terms of occurrences and abundances). A major challenge remains to further address whether and how tree performance varies among species and individuals in tropical forests.

Functional traits are related to components of individual fitness (Violle et al. 2007). Recently, more and more emphasis has been put on examining the relationship between functional trait values and demographic parameters (Salguero-Gómez et al. 2018), in order to better understand how functional trait values determine species population dynamics and abundances in assemblages. Fortunel et al. (2018) found an influence of functional traits on species growth variation related to topography, and less clearly to neighborhood density (crowding). Poorter et al. (2018) observed 44% of trait variation within species in a neotropical forest. Although individual trait values would be expected to be better predictors of performance than average values measured at the species level, Poorter et al still found a poor relationship.

Schmitt et al. (2023) examined how abiotic conditions and biotic interactions (considering neighborhood density) influenced the variation of individual potential tree growth, in a tropical forest plot located in French Guiana. They also considered the link between species-averaged values of growth potential and functional traits. Schmitt et al. (2023) found substantial variation in growth potential within species, that functional traits explained 40% of the variation of species-averaged growth and, noticeably, that the taxonomic structure (used as random effect in their model) explained a third of the variation in individual growth.

Although functional traits of roots, wood and leaves could predict a significant part of species growth potential, much variability of tree growth occurred within species. Intraspecific trait variation can thus be huge in response to changing abiotic and biotic contexts across individuals. The information on phylogenetic relationships can still provide a proxy of the integrated phenotypic variation that is under selection across the phylogeny, and determine a variation in growth strategies among individuals. The similarity of the phylogenetic structure suggests a joint selection of these growth strategies and related functional traits during events of convergent evolution. Baraloto et al. (2012) already noted that phylogenetic distance can be a proxy of niche overlap in tropical tree communities. Here, Schmitt et al. further demonstrate that evolutionary heritage is significantly related to individual growth variation, and plead for better acknowledging this role in future studies.

While the role of fitness differences in tropical tree community dynamics remained to be assessed, the present study provides new evidence that individual growth does vary depending on evolutionary relationships, which can reflect the roles of selection and adaptation on growth strategies. Therefore, investigating both the influence of functional traits and phylogenetic relationships on individual performance remains a promising avenue of research, for functional and community ecology in general.


Baraloto, Christopher, Olivier J. Hardy, C. E. Timothy Paine, Kyle G. Dexter, Corinne Cruaud, Luke T. Dunning, Mailyn-Adriana Gonzalez, et al. 2012. « Using functional traits and phylogenetic trees to examine the assembly of tropical tree communities ». Journal of Ecology, 100: 690‑701.
Fortunel Claire, Lasky Jesse R., Uriarte María, Valencia Renato, Wright S.Joseph, Garwood Nancy C., et Kraft Nathan J. B. 2018. « Topography and neighborhood crowding can interact to shape species growth and distribution in a diverse Amazonian forest ». Ecology, 99(10): 2272-2283.
Hubbell, S. P. 2001. The Unified Neutral Theory of Biodiversity and Biogeography. 1 vol. Princeton and Oxford: Princeton University Press.
Janzen, Daniel H. 1970. « Herbivores and the number of tree species in tropical forests ». American Naturalist, 104(940): 501-528.
Poorter, Lourens, Carolina V. Castilho, Juliana Schietti, Rafael S. Oliveira, et Flávia R. C. Costa. 2018. « Can traits predict individual growth performance? A test in a hyperdiverse tropical forest ». New Phytologist, 219 (1): 109‑21.
Salguero-Gómez, Roberto, Cyrille Violle, Olivier Gimenez, et Dylan Childs. 2018. « Delivering the promises of trait-based approaches to the needs of demographic approaches, and vice versa ». Functional Ecology, 32 (6): 1424‑35.
Schmitt, Sylvain, Valérie Raevel, Maxime Réjou‐Méchain, Narayanan Ayyappan, Natesan Balachandran, Narayanan Barathan, Gopalakrishnan Rajashekar, et François Munoz. 2021. « Canopy and understory tree guilds respond differently to the environment in an Indian rainforest ». Journal of Vegetation Science, e13075.
Sylvain Schmitt, Bruno Hérault, et Géraldine Derroire. 2023. « High intraspecific growth variability despite strong evolutionary heritage in a neotropical forest ». bioRxiv, 2022.07.27.501745, ver. 3 peer-reviewed and recommended by Peer Community in Ecology.
Violle, C., M. L. Navas, D. Vile, E. Kazakou, C. Fortunel, I. Hummel, et E. Garnier. 2007. « Let the concept of trait be functional! » Oikos, 116(5), 882-892.

High intraspecific growth variability despite strong evolutionary heritage in a neotropical forestSylvain Schmitt, Bruno Hérault, Géraldine Derroire<p style="text-align: justify;">Individual tree growth is a key determinant of species performance and a driver of forest dynamics and composition. Previous studies on tree growth unravelled the variation in species growth as a function of demogra...Community ecology, Demography, Population ecologyFrançois Munoz Jelena Pantel, David Murray-Stoker2022-08-01 14:29:04 View
10 Jan 2024
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Beyond variance: simple random distributions are not a good proxy for intraspecific variability in systems with environmental structure

Two paradigms for intraspecific variability

Recommended by ORCID_LOGO based on reviews by Simon Blanchet and Bart Haegeman

Community ecology usually concerns itself with understanding the causes and consequences of diversity at a given taxonomic resolution, most classically at the species level. Yet there is no doubt that diversity exists at all scales, and phenotypic variability within a taxon can be comparable to differences between taxa, as observed from bacteria to fish and trees. The question that motivates an active and growing body of work (e.g. Raffard et al 2019) is not so much whether intraspecific variability matters, but what we get wrong by ignoring it and how to incorporate it into our understanding of communities. There is no established way to think about diversity at multiple nested taxonomic levels, and it is tempting to summarize intraspecific variability simply by measuring species mean and variance in any trait and metric.

In this study, Girard-Tercieux et al (2023a) propose that, to understand its impact on community-level outcomes and in particular on species coexistence, we should carefully distinguish between two ways of thinking about intraspecific variability:

-"unstructured" variation, where every individual's features are like an independent random draw from a species-specific distribution, for instance, due to genetic lottery and developmental accidents

-"structured" variation that is due to each individual encountering a different but enduring microenvironment.

The latter type of variability may still appear complex and random-like when the environment is high-dimensional (i.e. multifaceted, with many different factors contributing to each individual's performance and development). Thus, it is not necessarily "structured" in the sense of being easily understood -- we may need to measure more aspects of the environment than is practical if we want to fully predict these variations.

What distinguishes this "structured" variability is that it is, in a loose sense, inheritable: individuals from the same species that grow in the same microenvironment will have the same performance, in a repeatable fashion. Thus, if each species is best at exploiting at least a fraction of environmental conditions, it is likely to avoid extinction by competition, except in the unlucky case of no propagule reaching any of the favorable sites.
By contrast, drawing each individual's preferences and performance randomly at each generation (from its own species distribution, but independently from other and past individuals) leads to stochastic dynamics, so-called ecological drift, that easily induce a large number of species extinctions.

The core intuition, that the complex spatial structure and high-dimensional nature of the environment plays a key explanatory role in species coexistence, is a running thread through several of the authors' work (e.g. Clark et al 2010), clearly inspired by their focus on tropical forests. This study, by tackling the question of intraspecific determinants of interspecific outcomes, makes a compelling addition to this line of investigation, coming as a theoretical companion to a more data-oriented study (Girard-Tercieux et al 2023b). But I believe it raises a question that is even broader in scope.

This kind of intraspecific variability, due to different individuals growing in different microenvironments, is perhaps most relevant for trees and other sessile organisms, but the distinction made here between "unstructured" and "structured" variability can likely be extended to many other ecological settings.

In my understanding, what matters most in "structured" variability is not so much it stemming from a fixed environment, but rather it being maintained across generations, rather than possibly lost by drift. This difference between variability in the form of "frozen" randomness and in the form of stochastic drift over time is highly relevant in other theoretical fields (e.g. in physics, where it is the difference between a disordered solid and a liquid), and thus, I expect that it is a meaningful distinction to make throughout community ecology.


James S. Clark, David Bell, Chengjin Chu, Benoit Courbaud, Michael Dietze, Michelle Hersh, Janneke HilleRisLambers et al. (2010) "High‐dimensional coexistence based on individual variation: a synthesis of evidence." Ecological Monographs 80, no. 4 : 569-608.

Camille Girard-Tercieux, Ghislain Vieilledent, Adam Clark, James S. Clark, Benoît Courbaud, Claire Fortunel, Georges Kunstler, Raphaël Pélissier, Nadja Rüger, Isabelle Maréchaux (2023a) "Beyond variance: simple random distributions are not a good proxy for intraspecific variability in systems with environmental structure." bioRxiv, ver. 4 peer-reviewed and recommended by Peer Community in Ecology.

Camille Girard‐Tercieux, Isabelle Maréchaux, Adam T. Clark, James S. Clark, Benoît Courbaud, Claire Fortunel, Joannès Guillemot et al. (2023b) "Rethinking the nature of intraspecific variability and its consequences on species coexistence." Ecology and Evolution 13, no. 3 : e9860.

Allan Raffard, Frédéric Santoul, Julien Cucherousset, and Simon Blanchet. (2019) "The community and ecosystem consequences of intraspecific diversity: A meta‐analysis." Biological Reviews 94, no. 2: 648-661.

Beyond variance: simple random distributions are not a good proxy for intraspecific variability in systems with environmental structureCamille Girard-Tercieux, Ghislain Vieilledent, Adam Clark, James S. Clark, Benoit Courbaud, Claire Fortunel, Georges Kunstler, Raphaël Pélissier, Nadja Rüger, Isabelle Maréchaux<p>The role of intraspecific variability (IV) in shaping community dynamics and species coexistence has been intensively discussed over the past decade and modelling studies have played an important role in that respect. However, these studies oft...Biodiversity, Coexistence, Community ecology, Competition, Theoretical ecologyMatthieu Barbier2022-08-07 12:51:30 View