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26 Mar 2019
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Is behavioral flexibility linked with exploration, but not boldness, persistence, or motor diversity?

Probing behaviors correlated with behavioral flexibility

Recommended by based on reviews by 2 anonymous reviewers

Behavioral plasticity, which is a subset of phenotypic plasticity, is an important component of foraging, defense against predators, mating, and many other behaviors. More specifically, behavioral flexibility, in this study, captures how quickly individuals adapt to new circumstances. In cases where individuals disperse to new environments, which often occurs in range expansions, behavioral flexibility is likely crucial to the chance that individuals can establish in these environments. Thus, it is important to understand how best to measure behavioral flexibility and how measures of such flexibility might vary across individuals and behavioral contexts and with other measures of learning and problem solving.
In this preregistration, Logan and colleagues propose to use a long-term study of the great-tailed grackle to measure how much they can manipulate behavioral flexibility in a reversal learning task, how much behavioral flexibility in one task predicts flexibility in another task and in problem solving a new task, and how robust these patterns are within individuals and across tasks. Logan and colleagues lay out their hypotheses and predictions for each experiment in a clear and concise manner. They also are very clear about the details of their study system, such as how they determined the number of trials they use in their learning reversal experiments, and how those details have influenced their experimental design. Further, given that the preregistration uses RMarkdown and is stored on GitHub (as are other studies in the larger project), their statistical code and its history of modification are easily available. This is a crucial component of making research more reproducible, which is a recent emphasis in behavioral sciences more broadly.
Reviewers of this preregistration found the study of substantial merit. The authors have responded to the reviewers' comments and their revisions have made the preregistration much clearer and cogent. I am happy to recommend this preregistration.

Is behavioral flexibility linked with exploration, but not boldness, persistence, or motor diversity?Kelsey McCune, Carolyn Rowney, Luisa Bergeron, Corina LoganThis is a PREREGISTRATION. The DOI was issued by OSF and refers to the whole GitHub repository, which contains multiple files. The specific file we are submitting is g_exploration.Rmd, which is easily accessible at GitHub at & Ethology, Preregistrations, ZoologyJeremy Van Cleve2018-09-27 03:35:12 View
12 Oct 2019
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Investigating the use of learning mechanisms in a species that is rapidly expanding its geographic range

How would variation in environmental predictability affect the use of different learning mechanisms in a social bird?

Recommended by based on reviews by Matthew Petelle and 1 anonymous reviewer

In their pre-registered paper [1], McCune and colleagues propose a field-based study of social versus individual learning mechanisms in an avian species (great-tailed grackles) that has been expanding its geographic range. The study forms part of a longer-term project that addresses various aspects of this species’ behaviour and biology, and the experience of the team is clear from the preprint. Assessing variation in learning mechanisms in different sections of the grackles’ distribution range, the researchers will investigate how individual learning and social transmission may impact learning about novel challenges in the environment. Considering that this is a social species, the authors expect both individual learning and social transmission to occur, when groups of grackles encounter new challenges/ opportunities in the wild. This in itself is not a very unusual idea to test [2, 3], but the authors are rigorously distinguishing between imitation, emulation, local enhancement, and social enhancement. Such rigour is certainly valuable in studies of cognition in the wild.
Further, the authors predict that the contribution of individual versus social learning could vary between populations, as the core may contain fewer unfamiliar/novel stimuli than the edge, where artificial sources of water (for example) may be more common. They make an argument that the core, middle, and edge populations would experience differing levels of environmental predictability. If true, their field experiments could yield very novel results on how changes in environmental predictability affect social/individual learning in a single study species. Their data would then give unusual insights into the ecological value of individual learning and distinct forms of social learning – something that is not easy to test in wild animals. The authors consider a variety of alternative hypotheses that may ultimately explain their findings, and clarify their methods and analyses in fine detail. The authors also set out limitations clearly, and give a thorough account of their approaches and thinking.
The reviewers and I have a still-unanswered question, which is central to the study: what is the predictability or unpredictability of the core versus edge environments? Although the authors have explained similarities and distinctions between the different sections of the grackles’ range, their description feels a bit vague -- it's not as rigorous or well-defined as the rest of the paper. Such a lack of definition may be inevitable in the limitations of a preprint, but ultimately it does suggest that there may be real uncertainty about the qualitative differences between the core, edge, and middle environments. The authors do explain that a lack of variation in individual responses to the field experiments would preclude the testing of further hypothesis, but do not mention how a salient lack of variation in novelty/ predictability between the environments could impact their hypotheses.
An assessment/quantification of the rate at which the different populations of grackles encounter novel stimuli would be a cornerstone of the success of this proposed study. Certainly, the authors cannot address this in much more detail during the preprint stage, but they need to consider how to best assess/describe differences before starting the full study. Such an assessment could take the form of either a GIS desktop study (comparing, for example, rates of dam/canal construction in core versus edge sections of the distribution range), or observational/ movement data contrasting how frequently members of core versus edge populations encounter artificial sources of water/food in a given month/year. Considering the long-term nature of the larger project, it is possible that these data are already available, but I am speculating. I would highly recommend that such an assessment be undertaken, beyond the mere mention of expected differences. This would solidify the central idea that there are concrete differences between the environments.
Despite this concern, the authors attended well to the comments and recommendations of the two reviewers – both experts in cognitive ecology. It is a preprint showing clear thinking and a consideration of most of the challenges that may be encountered during the course of the study. My own opinion and the estimations of the two reviewers all underscore the originality and value of this project – this should be a very valuable and potentially novel study. I look forward to seeing the outcomes of the research.


[1] McCune, K. B., McElreath, R., and Logan, C. J. (2019). Investigating the use of learning mechanisms in a species that is rapidly expanding its geographic range. In principle recommendation by Peer Community In Ecology.
[2] Benson-Amram, S. and Holekamp, K. E. (2012). Innovative problem solving by wild spotted hyenas. Proceedings of the Royal Society B: Biological Sciences, 279(1744), 4087–4095. doi: 10.1098/rspb.2012.1450
[3] Federspiel, I. G., Boeckle, M., von Bayern, A. M. P. and Emery, N. J. (2019). Exploring individual and social learning in jackdaws (Corvus monedula). Learning & Behavior, 47(3), 258–270. doi: 10.3758/s13420-019-00383-8

Investigating the use of learning mechanisms in a species that is rapidly expanding its geographic rangeKelsey McCune, Richard McElreath, Corina LoganThis is one of many studies planned for our long-term research on the role of behavior and learning in rapid geographic range expansions. Project background: Behavioral flexibility, the ability to change behavior when circumstances change based on...Behaviour & Ethology, Eco-evolutionary dynamics, Foraging, Preregistrations, Social structure, Spatial ecology, Metacommunities & Metapopulations, ZoologyAliza le Roux2019-07-23 18:45:20 View
08 Jan 2020
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Studies of NH4+ and NO3- uptake ability of subalpine plants and resource-use strategy identified by their functional traits

Nitrate or not nitrate. That is the question

Recommended by based on reviews by Vincent Maire and 1 anonymous reviewer

The article by Legay et al. [1] addresses two main issues: the links between belowground and aboveground plant traits and the links between plant strategies (as defined by these traits) and the capacity to absorb nitrate and ammonium. I recommend this work because these are important and current issues. The literature on plant traits is extremely rich and the existence of a leaf economic spectrum linked to a gradient between conservative and acquisitive plants is now extremely well established [2-3]. Many teams are now working on belowground traits and possible links with the aboveground gradients [4-5]. It seems indeed that there is a root economic spectrum but this spectrum is apparently less pronounced than the leaf economic spectrum. The existence of links between the two spectrums are still controversial and are likely not universal as suggested by discrepant results and after all a plant could have a conservative strategy aboveground and an acquisitive strategy belowground (or vice-versa) because, indeed, constraints are different belowground and aboveground (for example because in given ecosystem/vegetation type light may be abundant but not water or mineral nutrients). The various results obtained also suggest that we do not full understand the diversity of belowground strategies, what is at stake with these strategies, and the links with root characteristics.
Each time I give a conference on the work we are carrying out on African grasses that likely absorb ammonium preferentially because they inhibit nitrification [6-7], somebody asks me a question about the fact that plant essentially absorb nitrate because ammonium is toxic and nitrate more available in the soil. The present article confirms that this is not the case and that, though there are currently some teams working on the subject, we do not really know for the moment whether plants absorb nitrate or ammonium, in which proportion, how plastic this proportion is within individuals and within species. This subject seems to me crucial because it is linked to (1) the capacity of ecosystems to conserve nitrogen [8], because nitrate, much more than ammonium, goes out of ecosystems through leaching and denitrification, (2) to carbon cycling and plant energy budget because absorbing nitrate requires spending mucho more energy than absorbing ammonium because nitrate must be reduced before being incorporated in plant biomass, which is very energy costly. These two issues are naturally very relevant to develop efficient cropping systems in terms of carbon and nitrogen.
Interestingly, the present article, comparing three grass species in different sites, suggests that there is no trade-off between the absorption of nitrate and ammonium: more acquisitive individuals tend to absorb more ammonium and nitrate. This is contrary to hypotheses we made to predict the outcome of competition between plants absorbing nitrate and ammonium in different proportions [9] but should be tested in the future comparing many different types of plants. The results also suggest that more conservative plants absorb relatively more ammonium, which makes sense because this allows them to spare the energy necessary to reduce nitrate. This leads to the question of the effect of these strategies on nitrogen retention within the ecosystem. If nitrification is high (low), absorbing ammonium is not efficient and likely leads to high (low) nitrogen losses. This should be tested in the future. Moreover, the authors have measured the absorption of nitrate and ammonium through measurements at the root scale on cut roots. This should be complemented by measurements at the whole plant scale.


[1] Legay, N., Grassein, F., Arnoldi, C., Segura, R., Laîné, P., Lavorel, S. and Clément, J.-C. (2020). Studies of NH4+ and NO3- uptake ability of subalpine plants and resource-use strategy identified by their functional traits. bioRxiv, 372235, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/372235
[2] Shipley, B., Lechowicz, M.J., Wright, I. & Reich, P.B. (2006) Fundamental trade-offs generating the worldwide leaf economics spectrum. Ecology, 87, 535-541. doi: 10.1890/05-1051
[3] Reich, P.B. (2014) The world-wide ‘fast-slow’ plant economics spectrum: a traits manifesto. J. Ecol., 102, 275-301. doi: 10.1111/1365-2745.12211
[4] Maire, V., Gross, N., Pontes, L.D.S., Picon-Cochard, C. & Soussana, J.F. (2009) Trade-off between root nitrogen acquisition and shoot nitrogen utilization across 13 co-occurring pasture grass species. Func. Ecol., 23, 668-679. doi: 10.1111/j.1365-2435.2009.01557.x
[5] Roumet, C., Birouste, M., Picon-Cochard, C., Ghestem, M., Osman, N., Vrignon-Brenas, S., Cao, K.F. & Stokes, A. (2016) Root structure-function relationships in 74 species: evidence of a root economics spectrum related to carbon economy. New. Phytol., 210, 815-826. doi: 10.1111/nph.13828
[6] Lata, J.-C., Degrange, V., Raynaud, X., Maron, P.-A., Lensi, R. & Abbadie, L. (2004) Grass populations control nitrification in savanna soils. Funct. Ecol., 18, 605-611. doi: 10.1111/j.0269-8463.2004.00880.x
[7] Srikanthasamy, T., Leloup, J., N’Dri, A.B., Barot, S., Gervaix, J., Koné, A.W., Koffi, K.F., Le Roux, X., Raynaud, X. & Lata, J.-C. (2018) Contrasting effects of grasses and trees on microbial N-cycling in an African humid savanna. Soil Biol. Biochem., 117, 153-163. doi: 10.1016/j.soilbio.2017.11.016
[8] Boudsocq, S., Lata, J.C., Mathieu, J., Abbadie, L. & Barot, S. (2009) Modelling approach to analyze the effects of nitrification inhibition on primary production. Func. Ecol., 23, 220-230. doi: 10.1111/j.1365-2435.2008.01476.x
[9] Boudsocq, S., Niboyet, A., Lata, J.-C., Raynaud, X., Loeuille, N., Mathieu, J., Blouin, M., Abbadie, L. & Barot, S. (2012) Plant preference for ammonium versus nitrate: a neglected determinant of ecosystem functioning? Am. Nat., 180, 60-69. doi: 10.1086/665997

Studies of NH4+ and NO3- uptake ability of subalpine plants and resource-use strategy identified by their functional traitsLegay Nicolas, Grassein Fabrice, Arnoldi Cindy, Segura Raphaël, Laîné Philippe, Lavorel Sandra, Clément Jean-Christophe<p>The leaf economics spectrum (LES) is based on a suite of leaf traits related to plant functioning and ranges from resource-conservative to resource-acquisitive strategies. However, the relationships with root traits, and the associated belowgro...Community ecology, Physiology, Terrestrial ecologySébastien Barot2018-07-19 14:22:28 View
15 May 2023
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Behavioral flexibility is manipulable and it improves flexibility and innovativeness in a new context

An experiment to improve our understanding of the link between behavioral flexibility and innovativeness

Recommended by ORCID_LOGO based on reviews by Maxime Dahirel, Andrea Griffin, Aliza le Roux and 1 anonymous reviewer

Whether individuals are able to cope with new environmental conditions, and whether this ability can be improved, is certainly of great interest in our changing world. One way to cope with new conditions is through behavioral flexibility, which can be defined as “the ability to adapt behavior to new circumstances through packaging information and making it available to other cognitive processes” (Logan et al. 2023). Flexibility is predicted to be positively correlated with innovativeness, the ability to create a new behavior or use an existing behavior in a few situations (Griffin & Guez 2014). 
The post-study manuscript by Logan et al. (2023) proposes to test flexibility manipulability, and the relationship between flexibility and innovativeness. The authors did so with an experimental study on great-tailed grackles (Quiscalus mexicanus), an expanding species in the US, known to be flexible. 
The authors used serial reversal learning to investigate (1) whether behavioral flexibility, as measured by reversal learning using tubes of different shades, is manipulable; (2) whether manipulating (improving/training) behavioral flexibility improves flexibility and innovativeness in new contexts; (3) the type of learning strategy used by the individuals throughout the serial reversals.
The study described in this manuscript was pre-registered in Logan et al. (2019) and received in-principle recommendation on 26 Mar 2019 (Coulon 2019). One hypothesis from this original preregistration will be treated in a separate manuscript.
Among several interesting results, what I found most striking is that flexibility, in this species, seems to be a trait that is acquired by experience (vs. inherent to the individual). This opens exciting interrogations on the role of social learning, and on the impact of rapid environmental changes (which may force the individuals to experiment new ways to access to resources, for example), on individual flexibility and adaptability to new conditions. 

Coulon A (2019) Can context changes improve behavioral flexibility? Towards a better understanding of species adaptability to environmental changes. Peer Community in Ecology, 100019.

Griffin, A. S., & Guez, D. (2014). Innovation and problem solving: A review of common mechanisms. Behavioural Processes, 109, 121–134.

Logan C, Rowney C, Bergeron L, Seitz B, Blaisdell A, Johnson-Ulrich Z, McCune K (2019)
Is behavioral flexibility manipulatable and, if so, does it improve flexibility and problem solving in a new context? In Principle Recommendation 2019. PCI Ecology.

Logan CJ, Lukas D, Blaisdell AP, Johnson-Ulrich Z, MacPherson M, Seitz B, Sevchik A, McCune KB (2023) Behavioral flexibility is manipulable and it improves flexibility and innovativeness in a new context. EcoEcoRxiv, version 5 peer-reviewed and recommended by Peer Community in Ecology.

Behavioral flexibility is manipulable and it improves flexibility and innovativeness in a new contextLogan CJ, Lukas D, Blaisdell AP, Johnson-Ulrich Z, MacPherson M, Seitz BM, Sevchik A, McCune KB<p style="text-align: justify;">Behavioral flexibility, the ability to adapt behavior to new circumstances, is thought to play an important role in a species’ ability to successfully adapt to new environments and expand its geographic range. Howev...Behaviour & Ethology, Preregistrations, ZoologyAurélie Coulon2022-01-13 19:08:52 View
06 Oct 2020
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Implementing a rapid geographic range expansion - the role of behavior and habitat changes

The role of behavior and habitat availability on species geographic expansion

Recommended by ORCID_LOGO based on reviews by Caroline Marie Jeanne Yvonne Nieberding, Pizza Ka Yee Chow, Tim Parker and 1 anonymous reviewer

Understanding the relative importance of species-specific traits and environmental factors in modulating species distributions is an intriguing question in ecology [1]. Both behavioral flexibility (i.e., the ability to change the behavior in changing circumstances) and habitat availability are known to influence the ability of a species to expand its geographic range [2,3]. However, the role of each factor is context and species dependent and more information is needed to understand how these two factors interact. In this pre-registration, Logan et al. [4] explain how they will use Great-tailed grackles (Quiscalus mexicanus), a species with a flexible behavior and a rapid geographic range expansion, to evaluate the relative role of habitat and behavior as drivers of the species’ expansion [4]. The authors present very clear hypotheses, predicted results and also include alternative predictions. The rationales for all the hypotheses are clearly stated, and the methodology (data and analyses plans) are described with detail. The large amount of information already collected by the authors for the studied species during previous projects warrants the success of this study. It is also remarkable that the authors will make all their data available in a public repository, and that the pre-registration in already stored in GitHub, supporting open access and reproducible science. I agree with the three reviewers of this pre-registration about its value and I think its quality has largely improved during the review process. Thus, I am happy to recommend it and I am looking forward to seeing the results.


[1] Gaston KJ. 2003. The structure and dynamics of geographic ranges. Oxford series in Ecology and Evolution. Oxford University Press, New York.

[2] Sol D, Lefebvre L. 2000. Behavioural flexibility predicts invasion success in birds introduced to new zealand. Oikos. 90(3): 599–605.

[3] Hanski I, Gilpin M. 1991. Metapopulation dynamics: Brief history and conceptual domain. Biological journal of the Linnean Society. 42(1-2): 3–16.

[4] Logan CJ, McCune KB, Chen N, Lukas D. 2020. Implementing a rapid geographic range expansion - the role of behavior and habitat changes ( In principle acceptance by PCI Ecology of the version on 16 Dec 2021

Implementing a rapid geographic range expansion - the role of behavior and habitat changesLogan CJ, McCune KB, Chen N, Lukas D<p>It is generally thought that behavioral flexibility, the ability to change behavior when circumstances change, plays an important role in the ability of a species to rapidly expand their geographic range (e.g., Lefebvre et al. (1997), Griffin a...Behaviour & Ethology, Biological invasions, Dispersal & Migration, Foraging, Habitat selection, Human impact, Phenotypic plasticity, Preregistrations, ZoologyEsther Sebastián GonzálezAnonymous, Caroline Marie Jeanne Yvonne Nieberding, Tim Parker2020-05-14 11:18:57 View
28 Aug 2023
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Implementing a rapid geographic range expansion - the role of behavior changes

Behavioral changes in the rapid geographic expansion of the great-tailed grackle

Recommended by ORCID_LOGO based on reviews by Francois-Xavier Dechaume-Moncharmont, Pizza Ka Yee Chow and 1 anonymous reviewer

While many species' populations are declining, primarily due to human-related impacts (McKnee et al., 2014), certain species have thrived by utilizing human-influenced environments, leading to their population expansion (Muñoz & Real, 2006). In this context, the capacity to adapt and modify behaviors in response to new surroundings is believed to play a crucial role in facilitating species' spread to novel areas (Duckworth & Badyaev, 2007). For example, an increase in innovative behaviors within recently established communities could aid in discovering previously untapped food resources, while a decrease in exploration might reduce the likelihood of encountering dangers in unfamiliar territories (e.g., Griffin et al., 2016). To investigate the contribution of these behaviors to rapid range expansions, it is essential to directly measure and compare behaviors in various populations of the species.

The study conducted by Logan et al. (2023) aims to comprehend the role of behavioral changes in the range expansion of great-tailed grackles (Quiscalus mexicanus). To achieve this, the researchers compared the prevalence of specific behaviors at both the expansion's edge and its middle. Great-tailed grackles were chosen as an excellent model due to their behavioral adaptability, rapid geographic expansion, and their association with human-modified environments. The authors carried out a series of experiments in captivity using wild-caught individuals, following a detailed protocol. The study successfully identified differences in two of the studied behavioral traits: persistence (individuals participated in a larger proportion of trials) and flexibility variance (a component of the species' behavioral flexibility, indicating a higher chance that at least some individuals in the population could be more flexible). Notably, individuals at the edge of the population exhibited higher values of persistence and flexibility, suggesting that these behavioral traits might be contributing factors to the species' expansion. Overall, the study by Logan et al. (2023) is an excellent example of the importance of behavioral flexibility and other related behaviors in the process of species' range expansion and the significance of studying these behaviors across different populations to gain a better understanding of their role in the expansion process.

Finally, it is important to underline that this study is part of a pre-registration that received an In Principle Recommendation in PCI Ecology (Sebastián-González 2020) where objectives, methodology, and expected results were described in detail. The authors have identified any deviation from the original pre-registration and thoroughly explained the reasons for their deviations, which were very clear. 


Duckworth, R. A., & Badyaev, A. V. (2007). Coupling of dispersal and aggression facilitates the rapid range expansion of a passerine bird. Proceedings of the National Academy of Sciences, 104(38), 15017-15022.

Griffin, A.S., Guez, D., Federspiel, I., Diquelou, M., Lermite, F. (2016). Invading new environments: A mechanistic framework linking motor diversity and cognition to establishment success. Biological Invasions and Animal Behaviour, 26e46.

Logan, C. J., McCune, K., LeGrande-Rolls, C., Marfori, Z., Hubbard, J., Lukas, D. 2023. Implementing a rapid geographic range expansion - the role of behavior changes. EcoEvoRxiv, ver. 3 peer-reviewed and recommended by PCI Ecology.

McKee, J. K., Sciulli, P. W., Fooce, C. D., & Waite, T. A. (2004). Forecasting global biodiversity threats associated with human population growth. Biological Conservation, 115(1), 161-164.

Muñoz, A. R., & Real, R. (2006). Assessing the potential range expansion of the exotic monk parakeet in Spain. Diversity and Distributions, 12(6), 656-665.

Sebastián González, E. (2020) The role of behavior and habitat availability on species geographic expansion. Peer Community in Ecology, 100062. Reviewers: Caroline Nieberding, Tim Parker, and Pizza Ka Yee Chow.

Implementing a rapid geographic range expansion - the role of behavior changesLogan CJ, McCune KB, LeGrande-Rolls C, Marfori Z, Hubbard J, Lukas D<p>It is generally thought that behavioral flexibility, the ability to change behavior when circumstances change, plays an important role in the ability of species to rapidly expand their geographic range. Great-tailed grackles (<em>Quiscalus mexi...Behaviour & Ethology, Preregistrations, ZoologyEsther Sebastián González2023-04-12 11:00:42 View
04 May 2021
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Are the more flexible great-tailed grackles also better at behavioral inhibition?

Great-tailed grackle research reveals need for researchers to consider their own flexibility and test limitations in cognitive test batteries.

Recommended by based on reviews by Pizza Ka Yee Chow and Alex DeCasian

In the article, "Are the more flexible great-tailed grackles also better at behavioral inhibition?", Logan and colleagues (2021) are setting an excellent standard for cognitive research on wild-caught animals. Using a decent sample (N=18) of wild-caught birds, they set out to test the ambiguous link between behavioral flexibility and behavioral inhibition, which is supported by some studies but rejected by others. Where this study is more thorough and therefore also more revealing than most extant research, the authors ran a battery of tests, examining both flexibility (reversal learning and solution switching) and inhibition (go/no go task; detour task; delay of gratification) through multiple different test series. They also -- somewhat accidentally -- performed their experiments and analyses with and without different criteria for correctness (85%, 100%). Their mistakes, assumptions and amendments of plans made during preregistration are clearly stated and this demonstrates the thought-process of the researchers very clearly.

Logan et al. (2021) show that inhibition in great-tailed grackles is a multi-faceted construct, and demonstrate that the traditional go/no go task likely tests a very different aspect of inhibition than the detour task, which was never linked to any of their flexibility measures. Their comprehensive Bayesian analyses held up the results of some of the frequentist statistics, indicating a consistent relationship between flexibility and inhibition, with more flexible individuals also showing better inhibition (in the go/no go task). This same model, combined with inconsistencies in the GLM analyses (depending on the inclusion or exclusion of an outlier), led them to recommend caution in the creation of arbitrary thresholds for "success" in any cognitive tasks. Their accidental longer-term data collection also hinted at patterns of behaviour that shorter-term data collection did not. Of course, researchers have to decide on success criteria in order to conduct experiments, but in the same way that frequentist statistics are acknowledged to have flaws, the setting of success criteria must be acknowledged as inherently arbitrary. Where possible, researchers could reveal novel, biologically salient patterns by continuing beyond the point where a convenient success criterion has been reached. This research also underscores that tests may not be examining the features we expected them to measure, and are highly sensitive to biological and ecological variation between species as well as individual variation within populations.

To me, this study is an excellent argument for pre-registration of research (registered as Logan et al. 2019 and accepted by Vogel 2019), as the authors did not end up cherry-picking only those results or methods that worked. The fact that some of the tests did not "work", but was still examined, added much value to the study. The current paper is a bit densely written because of the comprehensiveness of the research. Some editorial polishing would likely make for more elegant writing. However, the arguments are clear, the results novel, and the questions thoroughly examined. The results are important not only for cognitive research on birds, but are potentially valuable to any cognitive scientist. I recommend this article as excellent food for thought.


Logan CJ, McCune K, Johnson-Ulrich Z, Bergeron L, Seitz B, Blaisdell AP, Wascher CAF. (2019) Are the more flexible individuals also better at inhibition?  In principle acceptance by PCI Ecology of the version on 6 Mar 2019

Logan CJ, McCune KB, MacPherson M, Johnson-Ulrich Z, Rowney C, Seitz B, Blaisdell AP, Deffner D, Wascher CAF (2021) Are the more flexible great-tailed grackles also better at behavioral inhibition? PsyArXiv, ver. 7 peer-reviewed and recommended by Peer community in Ecology.

Vogel E (2019) Adapting to a changing environment: advancing our understanding of the mechanisms that lead to behavioral flexibility. Peer Community in Ecology, 100016. 

Are the more flexible great-tailed grackles also better at behavioral inhibition?Logan CJ, McCune KB, MacPherson M, Johnson-Ulrich Z, Rowney C, Seitz B, Blaisdell AP, Deffner D, Wascher CAF<p style="text-align: justify;">Behavioral flexibility (hereafter, flexibility) should theoretically be positively related to behavioral inhibition (hereafter, inhibition) because one should need to inhibit a previously learned behavior to change ...PreregistrationsAliza le Roux2020-12-04 13:57:07 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
24 May 2023
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Evolutionary determinants of reproductive seasonality: a theoretical approach

When does seasonal reproduction evolve?

Recommended by ORCID_LOGO based on reviews by Francois-Xavier Dechaume-Moncharmont, Nigel Yoccoz and 1 anonymous reviewer

Have you ever wondered why some species breed seasonally while others do not? You might think it is all down to lattitude and the harshness of winters but it turns out it is quite a bit more complicated than that. A consequence of this is that climate change may result in the evolution of the degree of seasonal reproduction, with some species perhaps becoming less seasonal and others more so even in the same habitat. 

Burtschell et al. (2023) investigated how various factors influence seasonal breeding by building an individual-based model of a baboon population from which they calculated the degree of seasonality for the fittest reproductive strategy. They then altered key aspects of their model to examine how these changes impacted the degree of seasonality in the reproductive strategy. What they found is fascinating. 

The degree of seasonality in reproductive strategy is expected to increase with increased seasonality in the environment, decreased food availability, increased energy expenditure, and how predictable resource availability is. Interestingly, neither female cycle length nor extrinsic infant mortality influenced the degree of seasonality in reproduction.

What this means in reality for seasonal species is more challenging to understand. Some environments appear to be becoming more seasonal yet less predictable, and some species appear to be altering their daily energy budgets in response to changing climate in quite complex ways. As with pretty much everything in biology, Burtschell et al.'s work reveals much nuance and complexity, and that predicting how species might alter their reproductive timing is fraught with challenges.

The paper is very well written. With a simpler model it may have proven possible to achieve analytical solutions, but this is a very minor gripe. The reviewers were positive about the paper, and I have little doubt it will be well-cited. 


Burtschell L, Dezeure J, Huchard E, Godelle B (2023) Evolutionary determinants of reproductive seasonality: a theoretical approach. bioRxiv, 2022.08.22.504761, ver. 2 peer-reviewed and recommended by Peer Community in Ecology.

Evolutionary determinants of reproductive seasonality: a theoretical approachLugdiwine Burtschell, Jules Dezeure, Elise Huchard, Bernard Godelle<p style="text-align: justify;">Reproductive seasonality is a major adaptation to seasonal cycles and varies substantially among organisms. This variation, which was long thought to reflect a simple latitudinal gradient, remains poorly understood ...Evolutionary ecology, Life history, Theoretical ecologyTim Coulson Nigel Yoccoz2022-08-23 21:37:28 View
05 Apr 2022
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Late-acting self-incompatible system, preferential allogamy and delayed selfing in the heterostylous invasive populations of Ludwigia grandiflora subsp. hexapetala

Water primerose (Ludwigia grandiflora subsp. hexapetala) auto- and allogamy: an ecological perspective

Recommended by ORCID_LOGO based on reviews by Juan Arroyo, Emiliano Mora-Carrera and 1 anonymous reviewer

Invasive plant species are widely studied by the ecologist community, especially in wetlands. Indeed, alien plants are considered one of the major threats to wetland biodiversity (Reid et al., 2019). Ludwigia grandiflora subsp. hexapetala (Hook. & Arn.) G.L.Nesom & Kartesz, 2000 (Lgh) is one of them and has received particular attention for a long time (Hieda et al., 2020; Thouvenot, Haury, & Thiebaut, 2013). The ecology of this invasive species and its effect on its biotic and abiotic environment has been studied in previous works. Different processes were demonstrated to explain their invasibility such as allelopathic interference (Dandelot et al., 2008), resource competition (Gérard et al., 2014), and high phenotypic plasticity (Thouvenot, Haury, & Thiébaut, 2013), to cite a few of them. However, although vegetative reproduction is a well-known invasive process for alien plants like Lgh (Glover et al., 2015), the sexual reproduction of this species is still unclear and may help to understand the Lgh population dynamics.

Portillo Lemus et al. (2021) showed that two floral morphs of Lgh co-exist in natura, involving self-compatibility for short-styled phenotype and self-incompatibility for long-styled phenotype processes. This new article (Portillo Lemus et al., 2022) goes further and details the underlying mechanisms of the sexual reproduction of the two floral morphs.

Complementing their previous study, the authors have described a late self-incompatible process associated with the long-styled morph, which authorized a small proportion of autogamy. Although this represents a small fraction of the L-morph reproduction, it may have a considerable impact on the L-morph population dynamics. Indeed, authors report that “floral morphs are mostly found in allopatric monomorphic populations (i.e., exclusively S-morph or exclusively L-morph populations)” with a large proportion of L-morph populations compared to S-morph populations in the field. It may seem counterintuitive as L-morph mainly relies on cross-fecundation. 

Results show that L-morph autogamy mainly occurs in the fall, late in the reproduction season. Therefore, the reproduction may be ensured if no exogenous pollen reaches the stigma of L-morph individuals. It partly explains the large proportion of L-morph populations in the field. 

Beyond the description of late-acting self-incompatibility, which makes the Onagraceae a third family of Myrtales with this reproductive adaptation, the study raises several ecological questions linked to the results presented in the article. First, it seems that even if autogamy is possible, Lgh would favour allogamy, even in S-morph, through the faster development of pollen tubes from other individuals. This may confer an adaptative and evolutive advantage for the Lgh, increasing its invasive potential. The article shows this faster pollen tube development in S-morph but does not test the evolutive consequences. It is an interesting perspective for future research. It would also be interesting to describe cellular processes which recognize and then influence the speed of the pollen tube. Second, the importance of sexual reproduction vs vegetative reproduction would also provide information on the benefits of sexual dimorphism within populations. For instance, how fruit production increases the dispersal potential of Lgh would help to understand Lgh population dynamics and to propose adapted management practices (Delbart et al., 2013; Meisler, 2009).

To conclude, the study proposes a morphological, reproductive and physiological description of the Lgh sexual reproduction process. However, underlying ecological questions are well included in the article and the ecophysiological results enlighten some questions about the role of sexual reproduction in the invasiveness of Lgh. I advise the reader to pay attention to the reviewers’ comments; the debates were very constructive and, thanks to the great collaboration with the authorship, lead to an interesting paper about Lgh reproduction and with promising perspectives in ecology and invasion ecology.


Dandelot S, Robles C, Pech N, Cazaubon A, Verlaque R (2008) Allelopathic potential of two invasive alien Ludwigia spp. Aquatic Botany, 88, 311–316.

Delbart E, Mahy G, Monty A (2013) Efficacité des méthodes de lutte contre le développement de cinq espèces de plantes invasives amphibies : Crassula helmsii, Hydrocotyle ranunculoides, Ludwigia grandiflora, Ludwigia peploides et Myriophyllum aquaticum (synthèse bibliographique). BASE, 17, 87–102.

Gérard J, Brion N, Triest L (2014) Effect of water column phosphorus reduction on competitive outcome and traits of Ludwigia grandiflora and L. peploides, invasive species in Europe. Aquatic Invasions, 9, 157–166.

Glover R, Drenovsky RE, Futrell CJ, Grewell BJ (2015) Clonal integration in Ludwigia hexapetala under different light regimes. Aquatic Botany, 122, 40–46.

Hieda S, Kaneko Y, Nakagawa M, Noma N (2020) Ludwigia grandiflora (Michx.) Greuter & Burdet subsp. hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, an Invasive Aquatic Plant in Lake Biwa, the Largest Lake in Japan. Acta Phytotaxonomica et Geobotanica, 71, 65–71.

Meisler J (2009) Controlling Ludwigia hexaplata in Northern California. Wetland Science and Practice, 26, 15–19.

Portillo Lemus LO, Harang M, Bozec M, Haury J, Stoeckel S, Barloy D (2022) Late-acting self-incompatible system, preferential allogamy and delayed selfing in the heteromorphic invasive populations of Ludwigia grandiflora subsp. hexapetala. bioRxiv, 2021.07.15.452457, ver. 4 peer-reviewed and recommended by Peer Community in Ecology.

Portillo Lemus LO, Bozec M, Harang M, Coudreuse J, Haury J, Stoeckel S, Barloy D (2021) Self-incompatibility limits sexual reproduction rather than environmental conditions in an invasive water primrose. Plant-Environment Interactions, 2, 74–86.

Reid AJ, Carlson AK, Creed IF, Eliason EJ, Gell PA, Johnson PTJ, Kidd KA, MacCormack TJ, Olden JD, Ormerod SJ, Smol JP, Taylor WW, Tockner K, Vermaire JC, Dudgeon D, Cooke SJ (2019) Emerging threats and persistent conservation challenges for freshwater biodiversity. Biological Reviews, 94, 849–873.

Thouvenot L, Haury J, Thiebaut G (2013) A success story: water primroses, aquatic plant pests. Aquatic Conservation: Marine and Freshwater Ecosystems, 23, 790–803.

Thouvenot L, Haury J, Thiébaut G (2013) Seasonal plasticity of Ludwigia grandiflora under light and water depth gradients: An outdoor mesocosm experiment. Flora - Morphology, Distribution, Functional Ecology of Plants, 208, 430–437.

Late-acting self-incompatible system, preferential allogamy and delayed selfing in the heterostylous invasive populations of Ludwigia grandiflora subsp. hexapetalaLuis O. Portillo Lemus, Maryline Harang, Michel Bozec, Jacques Haury, Solenn Stoeckel, Dominique Barloy<p style="text-align: justify;">Breeding system influences local population genetic structure, effective size, offspring fitness and functional variation. Determining the respective importance of self- and cross-fertilization in hermaphroditic flo...Biological invasions, Botany, Freshwater ecology, PollinationAntoine Vernay2021-07-16 09:53:50 View