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01 Feb 2020
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Evidence of tool use in a seabird?

Touchy matter: the delicate balance between Morgan’s canon and open-minded description of advanced cognitive skills in the animal

Recommended by ORCID_LOGO based on reviews by Valérie Dufour and Alex Taylor

In a recent paper published in PNAS, Fayet et al. [1] reported scarce field observations of two Atlantic puffins (four years apart) apparently scratching their bodies using sticks, which was interpreted by the authors as evidence of tool use in this species. In a short response, Benjamin Farrar [2] raises serious concerns about this interpretation and proposes simpler, more parsimonious, mechanisms explaining the observed behaviour: a textbook case of Morgan's canon.
In virtually all introductory lectures on animal behaviour, students are advised to exercise caution when interpreting empirical data and weighting alternative explanations. We are sometimes prisoner of our assumptions: our desire of beliefs in advanced cognitive skills in non-human species make us more receptive to facts confirming our preconceptions than to simpler, less exciting, interpretations (a phenomenon known as "confirmation bias" in psychology). We must resist the temptation to accept appealing explanations without enough critical thinking. Our students are thus taught to apply the Lloyd Morgan's canon, a variant of one of the most important heuristics in Science, the principle of parsimony or Occam's razor, rephrased by Morgan [3, page 53] in the context of animal behaviour: "In no case may we interpret an action as the outcome of a higher psychical faculty, if it can be interpreted as the outcome of the exercise of one that stands lower in the psychological scale". In absence of evidence to the contrary, one should postulate the simplest cognitive skill consistent with the observed behaviour. While sometimes criticized from an epistemological point of view [4-6], it remains an essential and largely accepted framework of animal cognition. It has repeatedly proved to be a useful guide in the minefield of comparative psychology. Classical ethology questions related to the existence of, for instance, meta-cognition [7], intentionality or problem solving [8] have been convincingly investigated using this principle.
Yet, there is a downside to this conservative approach. Blind reference to Morgan's canon may narrow our theoretical thinking about animal cognition [7,9]. It could be counter-productive to systematically deny advanced cognitive skills in animals. On the contrary, keeping our mind open to unplanned observations, unexpected discoveries, or serendipity [10], and being prepared to accept new hypotheses, sometimes fairly remote from the dominant paradigm, may be a fruitful research strategy. To quote Darwin's famous letter to Alfred Wallace: "I am a firm believer, that without speculation there is no good and original observation" [11]. Brief notes in specialized scientific journals, or even in grey literature (by enthusiast amateur ornithologists, ichthyologists, or entomologists), constitutes a rich array of anecdotal observations. For instance, Sol et al. [12] convincingly compared the innovation propensity across bird species by screening ornithology literature using keywords like 'never reported', 'not seen before', 'first report', 'unusual' or 'novel'. Even if "the plural of anecdote is not data" as the saying goes, such descriptions of novel behaviours, even single-subject observations, are indisputably precious: taxonomic ubiquity of a behaviour is a powerful argument in favour of evolutionary convergence. Of course, a race to the bottom, amplified by the inevitable media hypes around scientific articles questioning human exceptionalism, is another possible scientific trap for behavioural biologists in search of skills characteristic of so-called advanced species, but never described so far in supposedly cognitively simpler organisms. As stated by Franz de Waal [9]: "I have nothing against anecdotes, especially if they have been caught on camera or come from reputable observers who know their animals; but I do view them as a starting point of research, never an end point".
In the case of the two video observations of puffins apparently using sticks as scratching tool, it must be considered as a mere anecdote unless scientists systematically investigate this behaviour. In his constructive criticism of Fayet et al.'s paper, Benjamin Farrar [2] proposes interesting directions of research and testable predictions. A correlation between the background rate of stick picking and the rate of stick preening would indicate that this behaviour was more likely explained by fluke than genuine innovation in this species.


[1] Fayet, A. L., Hansen, E. S., and Biro, D. (2020). Evidence of tool use in a seabird. Proceedings of the National Academy of Sciences, 117(3), 1277–1279. doi: 10.1073/pnas.1918060117
[2] Farrar, B. G. (2020). Evidence of tool use in a seabird? PsyArXiv, 463hk, ver. 5 recommended and peer-reviewed by Peer Community In Ecology. doi: 10.31234/
[3] Morgan, C. L. (1894). An introduction to comparative psychology. London, UK: Walter Scott, Ltd. Retrieved from
[4] Meketa, I. (2014). A critique of the principle of cognitive simplicity in comparative cognition. Biology and Philosophy, 29(5), 731–745. doi: 10.1007/s10539-014-9429-z
[5] Fitzpatrick, S. (2017). Against Morgan's Canon. In K. Andrews and J. Beck (Eds.), The Routledge handbook of philosophy of animal minds (pp. 437–447). London, UK: Routledge, Taylor and Francis Group. doi: 10.4324/9781315742250.ch42
[6] Starzak, T. (2017). Interpretations without justification: a general argument against Morgan's Canon. Synthese, 194(5), 1681–1701. doi: 10.1007/s11229-016-1013-4
[7] Arbilly, M., and Lotem, A. (2017). Constructive anthropomorphism: a functional evolutionary approach to the study of human-like cognitive mechanisms in animals. Proceedings of the Royal Society B: Biological Sciences, 284(1865), 20171616. doi: 10.1098/rspb.2017.1616
[8] Taylor, A. H., Knaebe, B., and Gray, R. D. (2012). An end to insight? New Caledonian crows can spontaneously solve problems without planning their actions. Proceedings of the Royal Society B: Biological Sciences, 279(1749), 4977–4981. doi: 10.1098/rspb.2012.1998
[9] de Waal, F. (2016). Are we smart enough to know how smart animals are? New-York, USA: W. W. Norton and Company.
[10] Scheffer, M. (2014). The forgotten half of scientific thinking. Proceedings of the National Academy of Sciences, 111(17), 6119–6119. doi: 10.1073/pnas.1404649111
[11] Darwin, C. R. (1857). Letter to A. R. Wallace, 22 December 1857. Retrieved 30 January 2020, from
[12] Sol, D., Lefebvre, L., and Rodríguez-Teijeiro, J. D. (2005). Brain size, innovative propensity and migratory behaviour in temperate Palaearctic birds. Proceedings of the Royal Society B: Biological Sciences, 272(1571), 1433–1441. doi: 10.1098/rspb.2005.3099

Evidence of tool use in a seabird?Benjamin G. FarrarFayet, Hansen and Biro (1) provide two observations of Atlantic puffins, *Fratercula arctica*, performing self-directed actions while holding a stick in their beaks. The authors interpret this as evidence of tool use as they suggest that the stick...Behaviour & EthologyFrancois-Xavier Dechaume-Moncharmont2020-01-22 11:55:27 View
28 Sep 2020
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The dynamics of spawning acts by a semelparous fish and its associated energetic costs

Extreme weight loss: when accelerometer could reveal reproductive investment in a semelparous fish species

Recommended by ORCID_LOGO based on reviews by Aidan Jonathan Mark Hewison, Loïc Teulier and 1 anonymous reviewer

Continuous observation of animal behaviour could be quite a challenge in the field, and the situation becomes even more complicated with aquatic species mostly active at night. In such cases, biologging techniques are real game changers in ecology, behavioural ecology or eco-physiology. An accelerating number of methodological applications of these tools in natural condition are thus published each year [1]. Biologging is not limited to movement ecology. For instance, fine grain information about energy expenditure can be inferred from body acceleration [2], and accelerometers has already proven useful in monitoring reproductive costs in some fish species [3,4]. The first part of the study by Tentelier et al. [5] is in line with this growing literature. It describes measurements of energy expenditure during reproduction in a fish species, Allis shad (Alosa Alosa), based on tail beat frequency and occurrence of spawning acts. The study has been convincingly conducted, and the results are important for fish biologists. But this is not the whole story: the authors added to this otherwise classical study a very original and insightful analysis which deserves closer interest.
Tentelier et al. propose to use static accelerometer to monitor change in body roundness through the reproductive season. These semelparous fish first mature and built up reserves in the Atlantic Ocean and migrate into fresh water to reproduce. Contrary to iteroparous species, female shads do not have to strategically preserve energy for future reproduction. The females die few days after spawning having exhausted their energetic reserves: they typically lose almost half of their body mass during the spawning season. The beautiful idea in this study was to track down information about this dramatic slimming in the accelerometer data. Indeed, the accelerometer was attached on the side of the fish (close to the dorsal fin). A change in its angle with the vertical plane could be correlated with the change in roundness, the angle declining with the female thinning. Accelerometers have already been used to record body posture [6] but, in the present study, the novelty was to monitor the change in body shape.
Unfortunately, the data by Tentelier et al. are inconclusive so far. Broadly speaking, the accelerometer angle recorded declined through the spawning season, indicating an average slimming of the females, but there was no correlation between the change in angle and the mass loss at the individual level. This was partly due to the fact that the dorsal position of the accelerometer was not optimized to measures egg laying whose effects are mostly observable on ventral side.
Yet, this nice idea deserves more scrutiny. The method seems to be sensitive enough to detect inflation of swim bladder, the gas-filled organ helping the fish to control their position in the water column, as the accelerometer angle increased when the fish stayed close to the water surface. Additional works and proper calibration are certainly needed to validate the use of accelerometer angle as a proxy for body roundness. The actual data were not strong enough to justify a standalone publication on the subject, but it would have been shame to lose traces of such analysis and keep it in the file drawer. This is why I strongly support its report as a side question in a broader study. Science progresses not only with neat conclusive studies but also when unexpected (apparently anecdotal) observations stimulate new researches.


[1] Börger L, Bijleveld AI, Fayet AL, Machovsky‐Capuska GE, Patrick SC, Street GM and Vander Wal E. (2020) Biologging special feature. J. Anim. Ecol. 89, 6–15. 10.1111/1365-2656.13163
[2] Wilson RP et al. (2020) Estimates for energy expenditure in free‐living animals using acceleration proxies: A reappraisal. J. Anim. Ecol. 89, 161–172. 10.1111/1365-2656.13040
[3] Tsuda Y, Kawabe R, Tanaka H, Mitsunaga Y, Hiraishi T, Yamamoto K and Nashimoto K. (2006) Monitoring the spawning behaviour of chum salmon with an acceleration data logger. Ecol. Freshw. Fish 15, 264–274. 10.1111/j.1600-0633.2006.00147.x
[4] Sakaji H, Hamada K, Naito Y. 2018 Identifying spawning events of greater amberjack using accelerometers. Mar. Biol. Res. 14, 637–641. 10.1080/17451000.2018.1492140
[5] Tentelier C, Bouchard C, Bernardin A, Tauzin A, Aymes J-C, Lange F, Récapet C, Rives J (2020) The dynamics of spawning acts by a semelparous fish and its associated energetic costs. bioRxiv, 436295. doi: 10.1101/436295 ver. 7 peer-reviewed and recommended by PCI Ecology. 10.1101/436295
[6] Brown DD, Kays R, Wikelski M, Wilson R, Klimley AP. 2013 Observing the unwatchable through acceleration logging of animal behavior. Anim. Biotelemetry 1, 20. 10.1186/2050-3385-1-20

The dynamics of spawning acts by a semelparous fish and its associated energetic costsCédric Tentelier, Colin Bouchard, Anaïs Bernardin, Amandine Tauzin, Jean-Christophe Aymes, Frédéric Lange, Charlotte Recapet, Jacques Rives<p>1. During the reproductive season, animals have to manage both their energetic budget and gamete stock. In particular, for semelparous capital breeders with determinate fecundity and no parental care other than gametic investment, the depletion...Behaviour & Ethology, Freshwater ecology, Life historyFrancois-Xavier Dechaume-Moncharmont2020-06-04 15:18:56 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
16 Nov 2020
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Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functions

Hidden diversity: how genetic richness affects species diversity and ecosystem processes in freshwater ponds

Recommended by based on reviews by Andrew Barnes and Jes Hines

Biodiversity loss can have important consequences for ecosystem functions, as exemplified by a large body of literature spanning at least three decades [1–3]. While connections between species diversity and ecosystem functions are now well-defined and understood, the importance of diversity within species is more elusive. Despite a surge in theoretical work on how intraspecific diversity can affect coexistence in simple community types [4,5], not much is known about how intraspecific diversity drives ecosystem processes in more complex community types. One particular challenge is that intraspecific diversity can be expressed as observable variation of functional traits, or instead subsist as genetic variation of which the consequences for ecosystem processes are harder to grasp.
Raffard et al. [6] examined how intraspecific biodiversity loss in a consumer fish changes species diversity at lower trophic levels and ecosystem processes in pond mesocosms. An interesting feature of this experiment is that it crosses functional and genetic intraspecific diversity. To do so, Raffard and colleagues measured and genotyped European minnow (P. phoxinus) individuals sampled from streams across southern France. Combining these collected specimens into experimental ponds allowed them to control functional (population variance of body size) and genetic intraspecific richness (number of genotypes).
Effects on minnow biomass production were mostly small; biomass was significantly reduced only when lowering both functional and genetic richness. However, the consequences for lower trophic levels (zooplankton and macroinvertebrates) were more pronounced and – importantly – not intuitive. For instance, the macroinvertebrate community was less species-diverse at higher minnow functional richness. If minnows with different body sizes would be the main regulator factors [7] explaining macroinvertebrate interactions, one would expect a more diverse set of minnow body sizes (i.e. higher functional minnow richness) to permit higher instead of lower macroinvertebrate richness. At the same time, the macroinvertebrate community was more species-diverse at higher minnow genotype richness, which could indicate unobserved minnow traits determining macroinvertebrate diversity more than the usual suspects (functional consumer richness). Such unobserved traits could be behavioral traits, allowing for resource partitioning among fish.
The consequences of functional minnow diversity loss on zooplankton diversity were negative, as expected in case body size differences among fish would facilitate coexistence of their zooplankton prey, as explained above. However, this was only the case when genetic diversity was high, suggesting nonstraightforward interactive effects of observed and non-observed traits on prey diversity.
The effects of functional and genetic minnow diversity loss on invertebrate (macroinvertebrates and zooplankton) abundance were more consistent than for invertebrate diversity. This suggests again nonstraightforward relationships in this experimental ecosystem, but now between invertebrate diversity and abundance. When using abundance as a proxy for an ecosystem process (which the authors did not), this result illustrates that biodiversity loss in multitrophic communities can have consequences that are challenging to interpret, let alone predict [8,9]. Path analyses showed how the observed changes of invertebrate diversity and abundance co-determined decomposition, a key ecosystem function. These path analyses had highest explanatory power show when including both kinds of intraspecific diversity.
Taken together, the results by Raffard and colleagues suggest that genetic consumer richness can drive species diversity of connected trophic levels and ecosystem processes with similar magnitude as functional diversity. Indeed, the effects of genetic consumer richness were shown to be so strong as to compensate or exacerbate the loss of observed functional richness. The exact mechanisms explaining these effects remain to be identified, however. The possibility that fish grazing by fish with different (observed or not observed) traits regulates coexistence among invertebrate prey, for instance, would depend on how strong fish consumption feeds back on prey growth during a 30-week experiment. As the authors indicate, detailed studies on resource partitioning among consumers (e.g. using stable isotope labelling) can shed light on these matters. Doing so may address a more fundamental question, which is if the mechanisms linking intraspecific diversity to function are different from those linking interspecific diversity to function, and at what time scales.


[1] Tilman D, Downing JA (1994) Biodiversity and stability in grasslands. Nature, 367, 363–365.
[2] Cardinale BJ, Duffy JE, Gonzalez A, Hooper DU, Perrings C, Venail P, Narwani A, Mace GM, Tilman D, Wardle DA, Kinzig AP, Daily GC, Loreau M, Grace JB, Larigauderie A, Srivastava DS, Naeem S (2012) Biodiversity loss and its impact on humanity. Nature, 486, 59–67.
[3] De Laender F, Rohr JR, Ashauer R, Baird DJ, Berger U, Eisenhauer N, Grimm V, Hommen U, Maltby L, Meliàn CJ, Pomati F, Roessink I, Radchuk V, Brink PJV den (2016) Reintroducing Environmental Change Drivers in Biodiversity–Ecosystem Functioning Research. Trends in Ecology & Evolution, 31, 905–915.
[4] Hart SP, Schreiber SJ, Levine JM (2016) How variation between individuals affects species coexistence. Ecology Letters, 19, 825–838.
[5] Barabás G, D’Andrea R (2016) The effect of intraspecific variation and heritability on community pattern and robustness. Ecology Letters, 19, 977–986.
[6] Raffard A, Cucherousset J, Montoya JM, Richard M, Acoca-Pidolle S, Poésy C, Garreau A, Santoul F, Blanchet S (2020) Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functions. bioRxiv, 2020.06.10.144337, ver. 3 peer-reviewed and recommended by PCI Ecology.
[7] Pásztor L, Botta-Dukát Z, Magyar G, Czárán T, Meszéna G. Theory-Based Ecology: A Darwinian approach. Oxford University Press.
[8] Binzer A, Guill C, Rall BC, Brose U (2016) Interactive effects of warming, eutrophication and size structure: impacts on biodiversity and food-web structure. Global Change Biology, 22, 220–227.
[9] Schwarz B, Barnes AD, Thakur MP, Brose U, Ciobanu M, Reich PB, Rich RL, Rosenbaum B, Stefanski A, Eisenhauer N (2017) Warming alters energetic structure and function but not resilience of soil food webs. Nature Climate Change, 7, 895–900.

Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functionsAllan Raffard, Julien Cucherousset, José M. Montoya, Murielle Richard, Samson Acoca-Pidolle, Camille Poésy, Alexandre Garreau, Frédéric Santoul & Simon Blanchet.<p>Loss in intraspecific diversity can alter ecosystem functions, but the underlying mechanisms are still elusive, and intraspecific biodiversity-ecosystem function relationships (iBEF) have been restrained to primary producers. Here, we manipulat...Community ecology, Ecosystem functioning, Experimental ecology, Food webs, Freshwater ecologyFrederik De Laender Andrew Barnes2020-06-15 09:04:53 View
10 Aug 2023
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Coexistence of many species under a random competition-colonization trade-off

Assembly in metacommunities driven by a competition-colonization tradeoff: more species in, more species out

Recommended by based on reviews by Canan Karakoç and 1 anonymous reviewer

The output of a community model depends on how you set its parameters. Thus, analyses of specific parameter settings hardwire the results to specific ecological scenarios. Because more general answers are often of interest, one tradition is to give models a statistical treatment: one summarizes how model parameters vary across species, and then predicts how changing the summary, instead of the individual parameters themselves, would change model output. Arguably the best-known example is the work initiated by May, showing that the properties of a community matrix, encoding effects species have on each other near their equilibrium, determine stability (1,2). More recently, this statistical treatment has also been applied to one of community ecology’s more prickly and slippery subjects: community assembly, which deals with the question “Given some regional species pool, which species will be able to persist together at some local ecosystem?”. Summaries of how species grow and interact in this regional pool predict the fraction of survivors and their relative abundances, the kind of dynamics, and various kinds of stability (3,4). One common characteristic of such statistical treatments is the assumption of disorder: if species do not interact in too structured ways, simple and therefore powerful predictions ensue that often stand up to scrutiny in relatively ordered systems. 
In their recent preprint, Miller, Clenet, et al. (5) subscribe to this tradition and consider tractable assembly scenarios (6) to study the outcome of assembly in a metacommunity. They recover a result of remarkable simplicity: roughly half of the species pool makes it into the final assemblage. Their vehicle is Tilman’s classic metacommunity model (7), where colonization rates are traded off with competitive ability. More precisely, in this model, one ranks species according to their colonization rate and attributes a greater competitive strength to lower-ranked species, which makes competition strictly hierarchical and thus departs from the disorder usually imposed by statistical approaches. The authors then leverage the simplicity of the species interaction network implied by this recursive setting to analytically probe how many species survive assembly. This turns out to be a fixed fraction that is distributed according to a Binomial with a mean of 0.5. While these results should not be extrapolated beyond the system at hand (4), they are important for two reasons. First, they imply that, within the framework of metacommunities driven by competition-colonization tradeoffs, richer species pools will produce richer communities: there is no upper bound on species richness, other than the one set by the raw material available for assembly. Second, this conclusion does not rely on simulation or equation solving and is, therefore, a hopeful sign of the palatability of the problem, if formalized in the right way. Their paper then shows that varying some of the settings does not change the main conclusion: changing how colonization rates distribute across species, and therefore the nature of the tradeoff, or the order with which species invade seems not to disrupt the big picture. Only when invaders are created “de novo” during assembly, a scenario akin to “de novo” mutation, a smaller fraction of species will survive assembly. 
As always, logical extensions of this study involve complicating the model and then looking if the results stay on par. The manuscript cites switching to other kinds of competition-colonization tradeoffs, and the addition of spatial heterogeneity as two potential avenues for further research. While certainly of merit, alternative albeit more bumpy roads would encompass models with radically different behavior. Most notably, one wonders how priority effects would play out. The current analysis shows that different invasion orders always lead to the same final composition, and therefore the same final species richness, confirming earlier results from models with similar structures (6). In models with priority effects, different invasion orders will surely lead to different compositions at the end. However, if one only cares about how many (and not which) species survive, it is unsure how much priority effects will qualitatively affect assembly. Because priority effects are varied in their topological manifestation (8), an important first step will be to evaluate which kinds of priority effects are compliant with formal analysis. 
1. May, R. M. (1972). Will a Large Complex System be Stable? Nature 238, 413–414.

2. Allesina, S. & Tang, S. (2015). The stability–complexity relationship at age 40: a random matrix perspective. Population Ecology, 57, 63–75.

3. Bunin, G. (2016). Interaction patterns and diversity in assembled ecological communities. Preprint at

4. Barbier, M., Arnoldi, J.-F., Bunin, G. & Loreau, M. (2018). Generic assembly patterns in complex ecological communities. Proceeding of the National Academy of Sciences, 115, 2156–2161.

5. Miller, Z. R., Clenet, M., Libera, K. D., Massol, F. & Allesina, S. (2023). Coexistence of many species under a random competition-colonization trade-off. bioRxiv 2023.03.23.533867, ver 3 peer-reviewed and recommended by PCI Ecology.

6. Serván, C. A. & Allesina, S. (2021). Tractable models of ecological assembly. Ecology Letters, 24, 1029–1037.

7. Tilman, D. (1994). Competition and Biodiversity in Spatially Structured Habitats. Ecology, 75, 2–16.

8. Song, C., Fukami, T. & Saavedra, S. (2021). Untangling the complexity of priority effects in multispecies communities. Ecolygy Letters, 24, 2301–2313.

Coexistence of many species under a random competition-colonization trade-offZachary R. Miller, Maxime Clenet, Katja Della Libera, François Massol, Stefano Allesina<p>The competition-colonization trade-off is a well-studied coexistence mechanism for metacommunities. In this setting, it is believed that coexistence of all species requires their traits to satisfy restrictive conditions limiting their similarit...Biodiversity, Coexistence, Colonization, Community ecology, Competition, Population ecology, Spatial ecology, Metacommunities & Metapopulations, Theoretical ecologyFrederik De Laender2023-03-30 20:42:48 View
05 Nov 2019
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Crown defoliation decreases reproduction and wood growth in a marginal European beech population.

Defoliation induces a trade-off between reproduction and growth in a southern population of Beech

Recommended by based on reviews by 3 anonymous reviewers

Individuals ability to withstand abiotic and biotic stresses is crucial to the maintenance of populations at climate edge of tree species distribution. We start to have a detailed understanding of tree growth response and to a lesser extent mortality response in these populations. In contrast, our understanding of the response of tree fecundity and recruitment remains limited because of the difficulty to monitor it at the individual tree level in the field. Tree recruitment limitation is, however, crucial for tree population dynamics [1-2].
In their study Oddou-Muratorio et al. [3] use a new method that they recently developed that jointly estimate male and female effective fecundity in natural populations using naturally established seedlings [4]. Their method uses a spatially explicit Bayesian analysis based on molecular markers and parentage analyses (MEMM program [4]). They apply this method to an unmanaged Beech forest at the southern edge of Beech distribution, where tree defoliation – taken as an integrative indicator of tree abiotic and biotic stress – and growth have been monitored for all adult trees.
This allows the authors to explore alternative hypothesis about tree fecundity response to stress. In one hand, biotic and abiotic stresses are thought to negatively impact tree fecundity. In the other hand, management and studies of orchard fruit tree support the idea that stress could trigger a compensatory increase of fecundity at the cost of other performances such as growth and survival.
They show that both growth and female fecundity are negatively affected by defoliation. There was no evidence that stresses trigger a compensatory increase of fecundity. Yet, they also found that, for large highly defoliated trees, there was a trade-off between growth and female fecundity. Some individuals are able to mitigate stress impact on fecundity by decreasing their growth. It is difficult to understand with available data what is driving such divergent responses between defoliated individuals. This could be related to differences in micro-environmental conditions or genetic background of individual trees. Such individual-level difference in response to stress could be crucial to understand tree populations response to ongoing climate change. This study clearly opens exciting new perspectives to apply such new methods to understand the role of fecundity on tree population dynamics. For instance, could we apply this method across the species distribution to understand how effective fecundity and its response to abiotic stress change between southern edge populations, core populations, and northern edge populations? Using time-series retrieved from such analysis can we disentangle the effect of different climatic drivers? It would also be interesting to see how such results can contribute to analyses covering the full tree life cycle to understand the contribution of fecundity response to population and evolutionary.


[1] Clark, J. S. et al. (1999). Interpreting recruitment limitation in forests. American Journal of Botany, 86(1), 1-16. doi: 10.2307/2656950
[2] Petit, R. J., and Hampe, A. (2006). Some evolutionary consequences of being a tree. Annu. Rev. Ecol. Evol. Syst., 37, 187-214. doi: 10.1146/annurev.ecolsys.37.091305.110215
[3] Oddou-Muratorio, S., Petit, C., Journe, V., Lingrand, M., Magdalou, J. A., Hurson, C., Garrigue, J., Davi, H. and Magnanou, E. (2019). Crown defoliation decreases reproduction and wood growth in a marginal European beech population. bioRxiv, 474874, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/474874
[4] Oddou‐Muratorio, S. and Klein, E. K. (2008). Comparing direct vs. indirect estimates of gene flow within a population of a scattered tree species. Molecular Ecology, 17(11), 2743-2754. doi: 10.1111/j.1365-294X.2008.03783.x

Crown defoliation decreases reproduction and wood growth in a marginal European beech population.Sylvie Oddou-Muratorio, Cathleen Petit-Cailleux, Valentin Journé, Matthieu Lingrand, Jean-André Magdalou, Christophe Hurson, Joseph Garrigue, Hendrik Davi, Elodie Magnanou.<p>1. Although droughts and heatwaves have been associated to increased crown defoliation, decreased growth and a higher risk of mortality in many forest tree species, their impact on tree reproduction and forest regeneration still remains underst...Climate change, Eco-evolutionary dynamics, Molecular ecology, Physiology, Population ecologyGeorges Kunstler2018-11-20 13:29:42 View
21 Dec 2020
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Influence of local landscape and time of year on bat-road collision risks

Assessing bat-vehicle collision risks using acoustic 3D tracking

Recommended by ORCID_LOGO based on reviews by Mark Brigham and ?

The loss of biodiversity is an issue of great concern, especially if the extinction of species or the loss of a large number of individuals within populations results in a loss of critical ecosystem services. We know that the most important threat to most species is habitat loss and degradation (Keil et al., 2015; Pimm et al., 2014); the latter can be caused by multiple anthropogenic activities, including pollution, introduction of invasive species and fragmentation (Brook et al., 2008; Scanes, 2018). Roads are a major cause of habitat fragmentation, isolating previously connected populations and being a direct source of mortality for animals that attempt to cross them (Spellberg, 1998).
While most studies have focused on the effect of roads on larger mammals (Bartonička et al., 2018; Litvaitis and Tash, 2008), in recent years many researchers have grown increasingly concerned about the risk of collision between bats and vehicles (Fensome and Mathews, 2016). For example, a recent publication by Medinas et al. (2021) found 509 bat casualties along a 51-km-long transect during a period of 3 years. Their study provides extremely valuable information to asses which factors primarily drive bat mortality on roads, yet it required a substantial investment of time coupled with the difficulty of detecting bat carcasses. Other studies have used acoustic monitoring as a proxy to gauge risk of collision based on estimates of bat density along roads (reviewed in Fensome and Mathews 2016); while the results of such studies are valuable, the number of passes recorded does not necessarily equal collision risk, as many species may simply avoid crossing the roads. Understanding the risk of collisions is of vital importance for adequate planning of road construction, particularly for key sites that harbor threatened bat species or unusually large populations, especially if these are already greatly impacted by other anthropogenic activities (e.g. wind turbines; Kunz et al. 2007) or unusually deadly pathogens (e.g. white-nose syndrome; Blehert et al. 2009).
The study by Roemer et al. (2020) titled “Influence of local landscape and time of year on bat-road collision risks”, is a welcome addition to our understanding of bat collision risk as it employs a more accurate assessment of bat collision risk based on acoustic monitoring and tracking of flight paths. The goal of the study of Roemer and collaborators, which was conducted at 66 study sites in the Mediterranean region, is to provide an assessment of collision risk based on bat activity near roads. They collected a substantial amount of information for several species: more than 30,000 estimated flight trajectories for 21+ species, including Barbastella barbastellus, Myotis spp., Plecotus sp., Rhinolophus ferrumequinum, Miniopterus schreibersii, Pipistrellus spp., Nyctalus leisleri, and others. They assess risk based on estimates of 1) species abundance from acoustic monitoring, 2) direction of flight paths along roads, and 3) bat-vehicle co-occurrence.
Their findings suggest that risk is habitat, species, guild, and season-specific. Roads within forested habitats posed the largest threats for most species, particularly since most flights within these habitats occurred at the zone of collision risk. They also found that bats typically fly parallel to the road axis regardless of habitat type, which they argue supports the idea that bats may use roads as corridors. The results of their study, as expected, also show that the majority of bat passes were detected during summer or autumn, depending on species, yet they provide novel findings of an increase in risky behaviors during autumn, when the number of passes at the zone of collision risk increased significantly. Their results also suggest that mid-range echolocators, a classification that is based on call design and parameters (Frey-Ehrenbold et al., 2013), had a larger portion of flights in the zone at risk, thus potentially making them more susceptible than short and long-range echolocators to collisions with vehicles.
The methods employed by Roemer et al. (2020) could further help us determine how roads pose species and site-specific threats in a diversity of places without the need to invest a significant amount of time locating bat carcasses. Their findings are also important as they could provide valuable information for deciding where new roads should be constructed, particularly if the most vulnerable species are abundant, perhaps due to the presence of important roost sites. They also show how habitats near larger roads could increase threats, providing an important first step for recommendations regarding road construction and maintenance. As pointed out by one reviewer, one possible limitation of the study is that the results are not supported by the identification of carcasses. For example, does an increase in the number of identified flights at the zone of risk really translate into an increase in the number of collisions? Regardless of the latter, the paper’s methods and results are very valuable and provide an important step towards developing additional tools to assess bat-vehicle collision risks.


[1] Bartonička T, Andrášik R, Duľa M, Sedoník J, Bíl M (2018) Identification of local factors causing clustering of animal-vehicle collisions. The Journal of Wildlife Management, 82, 940–947.
[2] Blehert DS, Hicks AC, Behr M, Meteyer CU, Berlowski-Zier BM, Buckles EL, Coleman JTH, Darling SR, Gargas A, Niver R, Okoniewski JC, Rudd RJ, Stone WB (2009) Bat White-Nose Syndrome: An Emerging Fungal Pathogen? Science, 323, 227–227.
[3] Brook BW, Sodhi NS, Bradshaw CJA (2008) Synergies among extinction drivers under global change. Trends in Ecology & Evolution, 23, 453–460.
[4] Fensome AG, Mathews F (2016) Roads and bats: a meta-analysis and review of the evidence on vehicle collisions and barrier effects. Mammal Review, 46, 311–323.
[5] Frey‐Ehrenbold A, Bontadina F, Arlettaz R, Obrist MK (2013) Landscape connectivity, habitat structure and activity of bat guilds in farmland-dominated matrices. Journal of Applied Ecology, 50, 252–261.
[6] Keil P, Storch D, Jetz W (2015) On the decline of biodiversity due to area loss. Nature Communications, 6, 8837.
[7] Kunz TH, Arnett EB, Erickson WP, Hoar AR, Johnson GD, Larkin RP, Strickland MD, Thresher RW, Tuttle MD (2007) Ecological impacts of wind energy development on bats: questions, research needs, and hypotheses. Frontiers in Ecology and the Environment, 5, 315–324.[315:EIOWED]2.0.CO;2
[8] Litvaitis JA, Tash JP (2008) An Approach Toward Understanding Wildlife-Vehicle Collisions. Environmental Management, 42, 688–697.
[9] Medinas D, Marques JT, Costa P, Santos S, Rebelo H, Barbosa AM, Mira A (2021) Spatiotemporal persistence of bat roadkill hotspots in response to dynamics of habitat suitability and activity patterns. Journal of Environmental Management, 277, 111412.
[10] Pimm SL, Jenkins CN, Abell R, Brooks TM, Gittleman JL, Joppa LN, Raven PH, Roberts CM, Sexton JO (2014) The biodiversity of species and their rates of extinction, distribution, and protection. Science, 344.
[11] Roemer C, Coulon A, Disca T, Bas Y (2020) Influence of local landscape and time of year on bat-road collision risks. bioRxiv, 2020.07.15.204115, ver. 3 peer-reviewed and recommended by Peer Community in Ecology.
[12] Scanes CG (2018) Chapter 19 - Human Activity and Habitat Loss: Destruction, Fragmentation, and Degradation. In: Animals and Human Society (eds Scanes CG, Toukhsati SR), pp. 451–482. Academic Press.
[13] Spellerberg I (1998) Ecological effects of roads and traffic: a literature review. Global Ecology & Biogeography Letters, 7, 317–333.

Influence of local landscape and time of year on bat-road collision risksCharlotte Roemer, Aurélie Coulon, Thierry Disca, and Yves Bas<p>Roads impact bat populations through habitat loss and collisions. High quality habitats particularly increase bat mortalities on roads, yet many questions remain concerning how local landscape features may influence bat behaviour and lead to hi...Behaviour & Ethology, Biodiversity, Conservation biology, Human impact, Landscape ecologyGloriana Chaverri2020-07-20 10:56:29 View
12 Apr 2023
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Feeding and growth variations affect δ13C and δ15N budgets during ontogeny in a lepidopteran larva

Refining our understanding how nutritional conditions affect 13C and 15N isotopic fractionation during ontogeny in a herbivorous insect

Recommended by based on reviews by Anton Potapov and 1 anonymous reviewer

Using stable isotope fractionation to disentangle and understand the trophic positions of animals within the food webs they are embedded within has a long tradition in ecology (Post, 2002; Scheu, 2002). Recent years have seen increasing application of the method with several recent reviews summarizing past advancements in this field (e.g. Potapov et al., 2019; Quinby et al., 2020).

In their new manuscript, Charberet and colleagues (2023) set out to refine our understanding of the processes that lead to nitrogen and carbon stable isotope fractionation by investigating how herbivorous insect larvae (specifically, the noctuid moth Spodoptera littoralis) respond to varying nutritional conditions (from starving to ad libitum feeding) in terms of stable isotopes enrichment. Though the underlying mechanisms have been experimentally investigated before in terrestrial invertebrates (e.g. in wolf spiders; Oelbermann & Scheu, 2002), the elegantly designed and adequately replicated experiments by Charberet and colleagues add new insights into this topic. Particularly, the authors provide support for the hypotheses that (A) 15N is disproportionately accumulated under fast growth rates (i.e. when fed ad libitum) and that (B) 13C is accumulated under low growth rates and starvation due to depletion of 13C-poor fat tissues. Applying this knowledge to field samples where feeding conditions are usually not known in detail is not straightforward, but the new findings could still help better interpretation of field data under specific conditions that make starvation for herbivores much more likely (e.g. droughts).

Overall this study provides important methodological advancements for a better understanding of plant-herbivore interactions in a changing world.


Charberet, S., Maria, A., Siaussat, D., Gounand, I., & Mathieu, J. (2023). Feeding and growth variations affect δ13C and δ15N budgets during ontogeny in a lepidopteran larva. bioRxiv, ver. 3 peer-reviewed and recommended by Peer Community in Ecology.

Oelbermann, K., & Scheu, S. (2002). Stable Isotope Enrichment (δ 15N and δ 13C) in a Generalist Predator (Pardosa lugubris, Araneae: Lycosidae): Effects of Prey Quality. Oecologia, 130(3), 337–344.

Post, D. M. (2002). Using stable isotopes to estimate trophic position: Models, methods, and assumptions. Ecology, 83(3), 703–718.[0703:USITET]2.0.CO;2

Potapov, A. M., Tiunov, A. V., & Scheu, S. (2019). Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition. Biological Reviews, 94(1), 37–59.

Quinby, B. M., Creighton, J. C., & Flaherty, E. A. (2020). Stable isotope ecology in insects: A review. Ecological Entomology, 45(6), 1231–1246.

Scheu, S. (2002). The soil food web: Structure and perspectives. European Journal of Soil Biology, 38(1), 11–20.

Feeding and growth variations affect δ13C and δ15N budgets during ontogeny in a lepidopteran larvaSamuel M. Charberet, Annick Maria, David Siaussat, Isabelle Gounand, Jérôme Mathieu<p style="text-align: justify;">Isotopes are widely used in ecology to study food webs and physiology. The fractionation observed between trophic levels in nitrogen and carbon isotopes, explained by isotopic biochemical selectivity, is subject to ...Experimental ecology, Food webs, PhysiologyGregor Kalinkat2022-11-16 15:23:31 View
07 Oct 2019
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Which pitfall traps and sampling efforts should be used to evaluate the effects of cropping systems on the taxonomic and functional composition of arthropod communities?

On the importance of experimental design: pitfall traps and arthropod communities

Recommended by ORCID_LOGO based on reviews by Cécile ALBERT and Matthias Foellmer

Despite the increasing refinement of statistical methods, a robust experimental design is still one of the most important cornerstones to answer ecological and evolutionary questions. However, there is a strong trade-off between a perfect design and its feasibility. A common mantra is that more data is always better, but how much is enough is complex to answer, specially when we want to capture the spatial and temporal variability of a given process. Gardarin and Valantin-Morison [1] make an effort to answer these questions for a practical case: How many pitfalls traps, of which type, and over which extent, do we need to detect shifts in arthropod community composition in agricultural landscapes. There is extense literature on how to approach these challenges using preliminary data in combination with simulation methods [e.g. 2], but practical cases are always welcomed to illustrate the complexity of the decisions to be made. A key challenge in this situation is the nature of simplified and patchy agricultural arthropod communities. In this context, small effect sizes are expected, but those small effects are relevant from an ecological point of view because small increases at low biodiversity may produce large gains in ecosystem functioning [3].
The paper shows that some variables are not important, such as the type of fluid used to fill the pitfall traps. This is good news for potential comparisons among studies using slightly different protocols. However, the bad news are that the sampling effort needed for detecting community changes is larger than the average effort currently implemented. A potential solution is to focus on Community Weighed Mean metrics (CWM; i.e. a functional descriptor of the community body size distribution) rather than on classic metrics such as species richness, as detecting changes on CWM requires a lower sampling effort and it has a clear ecological interpretation linked to ecosystem functioning.
Beyond the scope of the data presented, which is limited to a single region over two years, and hence it is hard to extrapolate to other regions and years, the big message of the paper is the need to incorporate statistical power simulations as a central piece of the ecologist's toolbox. This is challenging, especially when you face questions such as: Should I replicate over space, or over time? The recommended paper is accompanied by the statistical code used, which should facilitate this task to other researchers. Furthermore, we should be aware that some important questions in ecology are highly variable in space and time, and hence, larger sampling effort across space and time is needed to detect patterns. Larger and longer monitoring schemes require a large effort (and funding), but if we want to make relevant ecology, nobody said it would be easy.


[1] Gardarin, A. and Valantin-Morison, M. (2019). Which pitfall traps and sampling efforts should be used to evaluate the effects of cropping systems on the taxonomic and functional composition of arthropod communities? Zenodo, 3468920, ver. 3 peer-reviewed and recommended by PCI Ecology. doi: 10.5281/zenodo.3468920
[2] Johnson, P. C., Barry, S. J., Ferguson, H. M., and Müller, P. (2015). Power analysis for generalized linear mixed models in ecology and evolution. Methods in ecology and evolution, 6(2), 133-142. doi: 10.1111/2041-210X.12306
[3] Cardinale, B. J. et al. (2012). Biodiversity loss and its impact on humanity. Nature, 486(7401), 59-67. doi: 10.1038/nature11148

Which pitfall traps and sampling efforts should be used to evaluate the effects of cropping systems on the taxonomic and functional composition of arthropod communities?Antoine Gardarin and Muriel Valantin-Morison<p>1. Ground dwelling arthropods are affected by agricultural practices, and analyses of their responses to different crop management are required. The sampling efficiency of pitfall traps has been widely studied in natural ecosystems. In arable a...Agroecology, Biodiversity, Biological control, Community ecologyIgnasi Bartomeus2019-01-08 09:40:14 View
07 Aug 2023
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Heather pollen is not necessarily a healthy diet for bumble bees

The importance of understanding bee nutrition

Recommended by ORCID_LOGO based on reviews by Cristina Botías and 1 anonymous reviewer

​​Contrasting with the great alarm on bee declines, it is astonishing how little basic biology we know about bees, including on abundant and widespread species that are becoming model species. Plant-pollinator relationships are one of the cornerstones of bee ecology, and researchers are increasingly documenting bees' diets. However, we rarely know which effects feeding on different flowers has on bees' health. This paper (Tourbez et al. 2023) uses an elegant experimental setting to test the effect of heather pollen on bumblebees' (Bombus terrestris) reproductive success. This is a timely question as heather is frequently used by bumblebees, and its nectar has been reported to reduce parasite infections. In fact, it has been suggested that bumblebees can medicate themselves when infected (Richardson et al. 2014), and the pollen of some Asteraceae has been shown to help them fight parasites (Gekière​ et al. 2022). The starting hypothesis is that heather pollen contains flavonoids that might have a similar effect. Unfortunately, Tourbez​ and collaborators do not support this hypothesis, showing a negative effect of heather pollen, in particular its flavonoids, in bumblebees offspring, and an increase in parasite loads when fed on flavonoids. This is important because it challenges the idea that many pollen and nectar chemical compounds might have a medicinal use, and force us to critically analyze the effect of chemical compounds in each particular case. The results open several questions, such as why bumblebees collect heather pollen, or in which concentrations or pollen mixes it is deleterious. A limitation of the study is that it uses micro-colonies, and extrapolating this to real-world conditions is always complex. Understanding bee declines require a holistic approach starting with bee physiology and scaling up to multispecies population dynamics.  


Gekière, A., Semay, I., Gérard, M., Michez, D., Gerbaux, P., & Vanderplanck, M. 2022. Poison or Potion: Effects of Sunflower Phenolamides on Bumble Bees and Their Gut Parasite. Biology, 11(4), 545.​

Richardson, L.L., Adler, L.S., Leonard, A.S., Andicoechea, J., Regan, K.H., Anthony, W.E., Manson, J.S., &​ Irwin, R.E. 2015. Secondary metabolites in floral nectar reduce parasite infections in bumblebees. Proceedings of the Royal Society of London B: Biological Sciences 282 (1803), 20142471.

Tourbez, C., Semay, I., Michel, A., Michez, D., Gerbaux, P., Gekière A. & Vanderplanck, M. 2023. Heather pollen is not necessarily a healthy diet for bumble bees. Zenodo, ver 3, reviewed and recommended by PCI Ecology.​​

Heather pollen is not necessarily a healthy diet for bumble bees Clément Tourbez, Irène Semay, Apolline Michel, Denis Michez, Pascal Gerbaux, Antoine Gekière, Maryse Vanderplanck<p>There is evidence that specialised metabolites of flowering plants occur in both vegetative parts and floral resources (i.e., pollen and nectar), exposing pollinators to their biological activities. While such metabolites may be toxic to bees, ...Botany, Chemical ecology, Host-parasite interactions, Pollination, ZoologyIgnasi Bartomeus2023-04-10 21:22:34 View