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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. 

References

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. https://doi.org/10.1073/pnas.0706174104

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. https://doi.org/10.1017/CBO9781139939492.004

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. https://doi.org/10.32942/X2N30J

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. https://doi.org/10.1016/S0006-3207(03)00099-5

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. https://doi.org/10.1111/j.1472-4642.2006.00272.x

Sebastián González, E. (2020) The role of behavior and habitat availability on species geographic expansion. Peer Community in Ecology, 100062. https://doi.org/10.24072/pci.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
30 Sep 2020
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How citizen science could improve Species Distribution Models and their independent assessment

Citizen science contributes to SDM validation

Recommended by based on reviews by Maria Angeles Perez-Navarro and 1 anonymous reviewer

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

References

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

How citizen science could improve Species Distribution Models and their independent assessmentFlorence Matutini, Jacques Baudry, Guillaume Pain, Morgane Sineau, Josephine Pithon<p>Species distribution models (SDM) have been increasingly developed in recent years but their validity is questioned. Their assessment can be improved by the use of independent data but this can be difficult to obtain and prohibitive to collect....Biodiversity, Biogeography, Conservation biology, Habitat selection, Spatial ecology, Metacommunities & Metapopulations, Species distributions, Statistical ecologyFrancisco Lloret2020-06-03 09:36:34 View
29 Dec 2018
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The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web

From deserts to avocado orchards - understanding realized trophic interactions in communities

Recommended by based on reviews by Owen Petchey and 2 anonymous reviewers

The late eminent ecologist Gary Polis once stated that “most catalogued food-webs are oversimplified caricatures of actual communities” and are “grossly incomplete representations of communities in terms of both diversity and trophic connections.” Not content with that damning indictment, he went further by railing that “theorists are trying to explain phenomena that do not exist” [1]. The latter critique might have been push back for Robert May´s ground-breaking but ultimately flawed research on the relationship between food-web complexity and stability [2]. Polis was a brilliant ecologist, and his thinking was clearly influenced by his experiences researching desert food webs. Those food webs possess an uncommon combination of properties, such as frequent omnivory, cannibalism, and looping; high linkage density (L/S); and a nearly complete absence of apex consumers, since few species completely lack predators or parasites [3]. During my PhD studies, I was lucky enough to visit Joshua Tree National Park on the way to a conference in New England, and I could immediately see the problems posed by desert ecosystems. At the time, I was ruminating on the “harsh-benign” hypothesis [4], which predicts that the relative importance of abiotic and biotic forces should vary with changes in local environmental conditions (from harsh to benign). Specifically, in more “harsh” environments, abiotic factors should determine community composition whilst weakening the influence of biotic interactions. However, in the harsh desert environment I saw first-hand evidence that species interactions were not diminished; if anything, they were strengthened. Teddy-bear chollas possessed murderously sharp defenses to protect precious water, creosote bushes engaged in belowground “chemical warfare” (allelopathy) to deter potential competitors, and rampant cannibalism amongst scorpions drove temporal and spatial ontogenetic niche partitioning. Life in the desert was hard, but you couldn´t expect your competition to go easy on you.
If that experience colored my thinking about nature’s reaction to a capricious environment, then the seminal work by Robert Paine on the marine rocky shore helped further cement the importance of biotic interactions. The insights provided by Paine [5] brings us closer to the research reported in the preprint “The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web” [6], given that the authors in that study hold the environment constant and test the interactions between different permutations of a simple community. Paine [5] was able to elegantly demonstrate using the chief protagonist Pisaster ochraceus (a predatory echinoderm also known as the purple sea star) that a keystone consumer could exert strong top-down control that radically reshaped the interactions amongst other community members. What was special about this study was that the presence of Pisaster promoted species diversity by altering competition for space by sedentary species, providing a rare example of an ecological network experiment combining trophic and non-trophic interactions. Whilst there are increasing efforts to describe these interactions (e.g., competition and facilitation) in multiplex networks [7], the authors of “The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web” [6] have avoided strictly competitive interactions for the sake of simplicity. They do focus on two trophic forms of competition, namely intraguild predation and apparent competition. These two interaction motifs, along with prey switching are relevant to my own research on the influence of cross-ecosystem prey subsidies to receiving food webs [8]. In particular, the apparent competition motif may be particularly important in the context of emergent adult aquatic insects as prey subsidies to terrestrial consumers. This was demonstrated by Henschel et al. [9] where the abundances of emergent adult aquatic midges in riparian fields adjacent to a large river helped stimulate higher abundances of spiders and lower abundances of herbivorous leafhoppers, leading to a trophic cascade. The aquatic insects had a bottom-up effect on spiders and this subsidy facilitated a top-down effect that cascaded from spiders to leafhoppers to plants. The apparent competition motif becomes relevant because the aquatic midges exerted a negative indirect effect on leafhoppers mediated through their common arachnid predators.
In the preprint “The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web” [6], the authors have described different permutations of a simple mite community present in avocado orchards (Persea americana). This community comprises of two predators (Euseius stipulatus and Neoseiulus californicus), one herbivore as shared prey (Oligonychus perseae), and pollen of Carpobrotus edulis as alternative food resource, with the potential for the intraguild predation and apparent competition interaction motifs to be expressed. The authors determined that these motifs should be realized based off pairwise feeding trials. It is common for food-web researchers to depict potential food webs, which contain all species sampled and all potential trophic links based on laboratory feeding trials (as demonstrated here) or from observational data and literature reviews [10]. In reality, not all these potential feeding links are realized because species may partition space and time, thus driving alternative food-web architectures. In “The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web” [6], the authors are able to show that placing species in combinations that should yield more complex interaction motifs based off pairwise feeding trials fails to deliver – the predators revert to their preferred prey resulting in modular and simple trophic chains to be expressed. Whilst these realized interaction motifs may be stable, there might also be a tradeoff with function by yielding less top-down control than desirable when considering the potential for ecosystem services such as pest management. These are valuable insights, although it should be noted that here the fundamental niche is described in a strictly Eltonian sense as a trophic role [11]. Adding additional niche dimensions (sensu [12]), such as a thermal gradient could alter the observed interactions, although it might be possible to explain these contingencies through metabolic and optimal foraging theory combined with species traits. Nonetheless, the results of these experiments further demonstrate the need for ecologists to cross-validate theory with empirical approaches to develop more realistic and predictive food-web models, lest they invoke the wrath of Gary Polis´ ghost by “trying to explain phenomena that do not exist”.

References

[1] Polis, G. A. (1991). Complex trophic interactions in deserts: an empirical critique of food-web theory. The American Naturalist, 138(1), 123-155. doi: 10.1086/285208
[2] May, R. M. (1973). Stability and complexity in model ecosystems. Princeton University Press, Princeton, NJ, USA
[3] Dunne, J. A. (2006). The network structure of food webs. In Pascual, M., & Dunne, J. A. (eds) Ecological Networks: Linking Structure to Dynamics in Food Webs. Oxford University Press, New York, USA, 27-86
[4] Menge, B. A., & Sutherland, J. P. (1976). Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. The American Naturalist, 110(973), 351-369. doi: 10.1086/283073
[5] Paine, R. T. (1966). Food web complexity and species diversity. The American Naturalist, 100(910), 65-75. doi: 10.1086/282400
[6] Torres-Campos, I., Magalhães, S., Moya-Laraño, J., & Montserrat, M. (2018). The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food web. bioRxiv, 324178, ver. 5 peer-reviewed and recommended by PCI Ecol. doi: 10.1101/324178
[7] Kéfi, S., Berlow, E. L., Wieters, E. A., Joppa, L. N., Wood, S. A., Brose, U., & Navarrete, S. A. (2015). Network structure beyond food webs: mapping non‐trophic and trophic interactions on Chilean rocky shores. Ecology, 96(1), 291-303. doi: 10.1890/13-1424.1
[8] Burdon, F. J., & Harding, J. S. (2008). The linkage between riparian predators and aquatic insects across a stream‐resource spectrum. Freshwater Biology, 53(2), 330-346. doi: 10.1111/j.1365-2427.2007.01897.x
[9] Henschel, J. R., Mahsberg, D., & Stumpf, H. (2001). Allochthonous aquatic insects increase predation and decrease herbivory in river shore food webs. Oikos, 93(3), 429-438. doi: 10.1034/j.1600-0706.2001.930308.x
[10] Brose, U., Pavao-Zuckerman, M., Eklöf, A., Bengtsson, J., Berg, M. P., Cousins, S. H., Mulder, C., Verhoef, H. A., & Wolters, V. (2005). Spatial aspects of food webs. In de Ruiter, P., Wolters, V., Moore, J. C., & Melville-Smith, K. (eds) Dynamic Food Webs. vol 3. Academic Press, Burlington, 463-469
[11] Elton, C. (1927). Animal Ecology. Sidgwick and Jackson, London, UK
[12] Hutchinson, G. E. (1957). Concluding Remarks. Cold Spring Harbor Symposia on Quantitative Biology, 22, 415-427. doi: 10.1101/sqb.1957.022.01.039

The return of the trophic chain: fundamental vs realized interactions in a simple arthropod food webInmaculada Torres-Campos, Sara Magalhães, Jordi Moya-Laraño, Marta Montserrat<p>The mathematical theory describing small assemblages of interacting species (community modules or motifs) has proved to be essential in understanding the emergent properties of ecological communities. These models use differential equations to ...Community ecology, Experimental ecologyFrancis John Burdon2018-05-16 19:34:10 View
03 Jan 2024
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Diagnosis of planktonic trophic network dynamics with sharp qualitative changes

A new approach to describe qualitative changes of complex trophic networks

Recommended by based on reviews by Tim Coulson and 1 anonymous reviewer

Modelling the temporal dynamics of trophic networks has been a key challenge for community ecologists for decades, especially when anthropogenic and natural forces drive changes in species composition, abundance, and interactions over time. So far, most modelling methods fail to incorporate the inherent complexity of such systems, and its variability, to adequately describe and predict temporal changes in the topology of trophic networks. 

Taking benefit from theoretical computer science advances, Gaucherel and colleagues (2024) propose a new methodological framework to tackle this challenge based on discrete-event Petri net methodology. To introduce the concept to naïve readers the authors provide a useful example using a simplistic predator-prey model.

The core biological system of the article is a freshwater trophic network of western France in the Charente-Maritime marshes of the French Atlantic coast. A directed graph describing this system was constructed to incorporate different functional groups (phytoplankton, zooplankton, resources, microbes, and abiotic components of the environment) and their interactions. Rules and constraints were then defined to, respectively, represent physiochemical, biological, or ecological processes linking network components, and prevent the model from simulating unrealistic trajectories. Then the full range of possible trajectories of this mechanistic and qualitative model was computed.

The model performed well enough to successfully predict a theoretical trajectory plus two trajectories of the trophic network observed in the field at two different stations, therefore validating the new methodology introduced here. The authors conclude their paper by presenting the power and drawbacks of such a new approach to qualitatively model trophic networks dynamics.

Reference

Cedric Gaucherel, Stolian Fayolle, Raphael Savelli, Olivier Philippine, Franck Pommereau, Christine Dupuy (2024) Diagnosis of planktonic trophic network dynamics with sharp qualitative changes. bioRxiv 2023.06.29.547055, ver. 2 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2023.06.29.547055

Diagnosis of planktonic trophic network dynamics with sharp qualitative changesCedric Gaucherel, Stolian Fayolle, Raphael Savelli, Olivier Philippine, Franck Pommereau, Christine Dupuy<p>Trophic interaction networks are notoriously difficult to understand and to diagnose (i.e., to identify contrasted network functioning regimes). Such ecological networks have many direct and indirect connections between species, and these conne...Community ecology, Ecosystem functioning, Food webs, Freshwater ecology, Interaction networks, Microbial ecology & microbiologyFrancis Raoul Tim Coulson2023-07-03 10:42:34 View
21 Nov 2023
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Pathogen community composition and co-infection patterns in a wild community of rodents

Reservoirs of pestilence: what pathogen and rodent community analyses can tell us about transmission risk

Recommended by ORCID_LOGO based on reviews by Adrian Diaz, Romain Pigeault and 1 anonymous reviewer

Rodents are well known as one of the main animal groups responsible for human-transmitted pathogens. As such, it seems logical to try and survey what kinds of pathogenic microbes might be harboured by wild rodents, in order to establish some baseline surveillance and prevent future zoonotic outbreaks (Bernstein et al., 2022). This is exactly what Abbate et al. (2023) endeavoured and their findings are intimidating. Based on quite a large sampling effort, they collected more than 700 rodents of seven species around two villages in northeastern France. They looked for molecular markers indicative of viral and bacterial infections and proceeded to analyze their pathogen communities using multivariate techniques.

Variation in the prevalence of the different pathogens was found among host species, with e.g. signs of CPXV more prevalent in Cricetidae while some Mycoplasma strains were more prevalent in Muridae. Co-circulation of pathogens was found in all species, with some evidencing signs of up to 12 different pathogen taxa. The diversity of co-circulating pathogens was markedly different between host species and higher in adult hosts, but not affected by sex. The dataset also evinced some slight differences between habitats, with meadows harbouring a little more diversity of rodent pathogens than forests. Less intuitively, some pathogen associations seemed quite repeatable, such as the positive association of Bartonella spp. with CPXV in the montane water vole. The study allowed the authors to test several associations already described in the literature, including associations between different hemotropic Mycoplasma species.

I strongly invite colleagues interested in zoonoses, emerging pandemics and more generally One Health to read the paper of Abbate et al. (2023) and try to replicate them across the world. To prevent the next sanitary crises, monitoring rodents, and more generally vertebrates, population demographics is a necessary and enlightening step (Johnson et al., 2020), but insufficient. Following the lead of colleagues working on rodent ectoparasites (Krasnov et al., 2014), we need more surveys like the one described by Abbate et al. (2023) to understand the importance of the dilution effect in the prevalence and transmission of microbial pathogens (Andreazzi et al., 2023) and the formation of epidemics. We also need other similar studies to assess the potential of different rodent species to carry pathogens more or less capable of infecting other mammalian species (Morand et al., 2015), in other places in the world.

References

Abbate, J. L., Galan, M., Razzauti, M., Sironen, T., Voutilainen, L., Henttonen, H., Gasqui, P., Cosson, J.-F. & Charbonnel, N. (2023) Pathogen community composition and co-infection patterns in a wild community of rodents. BioRxiv, ver.4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2020.02.09.940494 

Andreazzi, C. S., Martinez-Vaquero, L. A., Winck, G. R., Cardoso, T. S., Teixeira, B. R., Xavier, S. C. C., Gentile, R., Jansen, A. M. & D'Andrea, P. S. (2023) Vegetation cover and biodiversity reduce parasite infection in wild hosts across ecological levels and scales. Ecography, 2023, e06579.
https://doi.org/10.1111/ecog.06579
 
Bernstein, A. S., Ando, A. W., Loch-Temzelides, T., Vale, M. M., Li, B. V., Li, H., Busch, J., Chapman, C. A., Kinnaird, M., Nowak, K., Castro, M. C., Zambrana-Torrelio, C., Ahumada, J. A., Xiao, L., Roehrdanz, P., Kaufman, L., Hannah, L., Daszak, P., Pimm, S. L. & Dobson, A. P. (2022) The costs and benefits of primary prevention of zoonotic pandemics. Science Advances, 8, eabl4183.
https://doi.org/10.1126/sciadv.abl4183
 
Johnson, C. K., Hitchens, P. L., Pandit, P. S., Rushmore, J., Evans, T. S., Young, C. C. W. & Doyle, M. M. (2020) Global shifts in mammalian population trends reveal key predictors of virus spillover risk. Proceedings of the Royal Society B: Biological Sciences, 287, 20192736.
https://doi.org/10.1098/rspb.2019.2736
 
Krasnov, B. R., Pilosof, S., Stanko, M., Morand, S., Korallo-Vinarskaya, N. P., Vinarski, M. V. & Poulin, R. (2014) Co-occurrence and phylogenetic distance in communities of mammalian ectoparasites: limiting similarity versus environmental filtering. Oikos, 123, 63-70.
https://doi.org/10.1111/j.1600-0706.2013.00646.x
 
Morand, S., Bordes, F., Chen, H.-W., Claude, J., Cosson, J.-F., Galan, M., Czirjak, G. Á., Greenwood, A. D., Latinne, A., Michaux, J. & Ribas, A. (2015) Global parasite and Rattus rodent invasions: The consequences for rodent-borne diseases. Integrative Zoology, 10, 409-423.
https://doi.org/10.1111/1749-4877.12143

Pathogen community composition and co-infection patterns in a wild community of rodentsJessica Lee Abbate, Maxime Galan, Maria Razzauti, Tarja Sironen, Liina Voutilainen, Heikki Henttonen, Patrick Gasqui, Jean-François Cosson, Nathalie Charbonnel<p style="text-align: justify;">Rodents are major reservoirs of pathogens that can cause disease in humans and livestock. It is therefore important to know what pathogens naturally circulate in rodent populations, and to understand the factors tha...Biodiversity, Coexistence, Community ecology, Eco-immunology & Immunity, Epidemiology, Host-parasite interactions, Population ecology, Species distributionsFrancois Massol2020-02-11 12:42:28 View
19 Feb 2020
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Soil variation response is mediated by growth trajectories rather than functional traits in a widespread pioneer Neotropical tree

Growth trajectories, better than organ-level functional traits, reveal intraspecific response to environmental variation

Recommended by ORCID_LOGO based on reviews by Georges Kunstler and François Munoz

Functional traits are “morpho-physio-phenological traits which impact fitness indirectly via their effects on growth, reproduction and survival” [1]. Most functional traits are defined at organ level, e.g. for leaves, roots and stems, and reflect key aspects of resource acquisition and resource use by organisms for their development and reproduction [2]. More rarely, some functional traits can be related to spatial development, such as vegetative height and lateral spread in plants.
Organ-level traits are especially popular because they can be measured in a standard way and easily compared over many plants. But these traits can broadly vary during the life of an organism. For instance, Roggy et al. [3] found that Leaf Mass Area can vary from 30 to 140 g.m^(-2) between seedling and adult stages for the canopy tree Dicorynia guianensis in French Guiana. Fortunel et al. [4] have also showed that developmental stages much contribute to functional trait variation within several Micropholis tree species in lowland Amazonia.
The way plants grow and invest resources into organs is variable during life and allows defining specific developmental sequences and architectural models [5,6]. There is clear ontogenic variation in leaf number, leaf properties and ramification patterns. Ontogenic variations reflect changing adaptation of an individual over its life, depending on the changing environmental conditions.
In this regard, measuring a single functional trait at organ level in adult trees should miss the variation of resource acquisition and use strategies over time. Thus we should built a more integrative approach of ecological development, also called “eco-devo” approach [7].
Although the ecological significance of ontogeny and developmental strategies is now well known, the extent to which it contributes to explain species survival and coexistence in communities is still broadly ignored in functional ecology. Levionnois et al. [8] investigated intraspecific variation of functional traits and growth trajectories in a typical, early-successional tree species in French Guiana, Amazonia. This species, Cecropia obtusa, is generalist regarding soil type and can be found on both white sand and ferralitic soil. The study examines whether there in intraspecific variation in functional traits and growth trajectories of C. obtusa in response to the contrasted soil types.
The tree communities observed on the two types of soils include species with distinctive functional trait values, that is, there are changes in species composition related to different species strategies along the classical wood and leaf economic spectra. The populations of C. obtusa found on the two soils showed some difference in functional traits, but it did not concern traits related to the main economic spectra. Conversely, the populations showed different growth strategies, in terms of spatial and temporal development.
The major lessons we can learn from the study are:
(i) Functional traits measured at organ level cannot reflect well how long-lived plants collect and invest resources during their life. The results show the potential of considering architectural and developmental traits together with organ-level functional traits, to better acknowledge the variation in ecological strategies over plant life, and thus to better understand community assembly processes.
(ii) What makes functional changes between communities differs when considering interspecific and intraspecific variation. Species turnover should encompass different corteges of soil specialists. These specialists are sorted along economic spectra, as shown in tropical rainforests and globally [2]. Conversely, a generalist species such as C. obtusa does occur on contrasted soil, which entails that it can accommodate the contrasted ecological conditions. However, the phenotypic adjustment is not related to how leaves and wood ensure photosynthesis, water and nutrient acquisition, but regards the way the resources are allocated to growth and reproduction over time.
The results of the study stress the need to better integrate growth strategies and ontogeny in the research agenda of functional ecology. We can anticipate that organ-level functional traits and growth trajectories will be more often considered together in ecological studies. The integration should help better understand the temporal niche of organisms, and how organisms can coexist in space and time with other organisms during their life. Recently, Klimešová et al. [9] have proposed standardized protocols for collecting plant modularity traits. Such effort to propose easy-to-measure traits representing plant development and ontogeny, with clear functional roles, should foster the awaited development of an “eco-devo” approach.

References

[1] Violle, C., Navas, M. L., Vile, D., Kazakou, E., Fortunel, C., Hummel, I., & Garnier, E. (2007). Let the concept of trait be functional!. Oikos, 116(5), 882-892. doi: 10.1111/j.0030-1299.2007.15559.x
[2] Díaz, S. et al. (2016). The global spectrum of plant form and function. Nature, 529(7585), 167-171. doi: 10.1038/nature16489
[3] Roggy, J. C., Nicolini, E., Imbert, P., Caraglio, Y., Bosc, A., & Heuret, P. (2005). Links between tree structure and functional leaf traits in the tropical forest tree Dicorynia guianensis Amshoff (Caesalpiniaceae). Annals of forest science, 62(6), 553-564. doi: 10.1051/forest:2005048
[4] Fortunel, C., Stahl, C., Heuret, P., Nicolini, E. & Baraloto, C. (2020). Disentangling the effects of environment and ontogeny on tree functional dimensions for congeneric species in tropical forests. New Phytologist. doi: 10.1111/nph.16393
[5] Barthélémy, D., & Caraglio, Y. (2007). Plant architecture: a dynamic, multilevel and comprehensive approach to plant form, structure and ontogeny. Annals of botany, 99(3), 375-407. doi: 10.1093/aob/mcl260
[6] Hallé, F., & Oldeman, R. A. (1975). An essay on the architecture and dynamics of growth of tropical trees. Kuala Lumpur: Penerbit Universiti Malaya.
[7] Sultan, S. E. (2007). Development in context: the timely emergence of eco-devo. Trends in Ecology & Evolution, 22(11), 575-582. doi: 10.1016/j.tree.2007.06.014
[8] Levionnois, S., Tysklind, N., Nicolini, E., Ferry, B., Troispoux, V., Le Moguedec, G., Morel, H., Stahl, C., Coste, S., Caron, H. & Heuret, P. (2020). Soil variation response is mediated by growth trajectories rather than functional traits in a widespread pioneer Neotropical tree. bioRxiv, 351197, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/351197
[9] Klimešová, J. et al. (2019). Handbook of standardized protocols for collecting plant modularity traits. Perspectives in Plant Ecology, Evolution and Systematics, 40, 125485. doi: 10.1016/j.ppees.2019.125485

Soil variation response is mediated by growth trajectories rather than functional traits in a widespread pioneer Neotropical treeSébastien Levionnois, Niklas Tysklind, Eric Nicolini, Bruno Ferry, Valérie Troispoux, Gilles Le Moguedec, Hélène Morel, Clément Stahl, Sabrina Coste, Henri Caron, Patrick Heuret<p style="text-align: justify;">1- Trait-environment relationships have been described at the community level across tree species. However, whether interspecific trait-environment relationships are consistent at the intraspecific level is yet unkn...Botany, Eco-evolutionary dynamics, Habitat selection, Ontogeny, Tropical ecologyFrançois Munoz2018-06-21 17:13:17 View
22 Nov 2021
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Beating your neighbor to the berry patch

When more competitors means less harvested resource

Recommended by ORCID_LOGO based on reviews by Francois Massol, Jeremy Van Cleve and 1 anonymous reviewer

In this paper, Alan R. Rogers (2021) examines the dynamics of foraging strategies for a resource that gains value over time (e.g., ripening fruits), while there is a fixed cost of attempting to forage the resource, and once the resource is harvested nothing is left for other harvesters. For this model, not any pure foraging strategy is evolutionary stable. A mixed equilibrium exists, i.e., with a mixture of foraging strategies within the population, which is still evolutionarily unstable. Nonetheless, Alan R. Rogers shows that for a large number of competitors and/or high harvesting cost, the mixture of strategies remains close to the mixed equilibrium when simulating the dynamics. Surprisingly, in a large population individuals will less often attempt to forage the resource and will instead “go fishing”. The paper also exposes an experiment of the game with students, which resulted in a strategy distribution somehow close to the theoretical mixture of strategies.

The economist John F. Nash Jr. (1950) gained the Nobel Prize of economy in 1994 for his game theoretical contributions. He gave his name to the “Nash equilibrium”, which represents a set of individual strategies that is reached whenever all the players have nothing to gain by changing their strategy while the strategies of others are unchanged. Alan R. Rogers shows that the mixed equilibrium in the foraging game is such a Nash equilibrium. Yet it is evolutionarily unstable insofar as a distribution close to the equilibrium can invade.

The insights of the study are twofold. First, it sheds light on the significance of Nash equilibrium in an ecological context of foraging strategies. Second, it shows that an evolutionarily unstable state can rule the composition of the ecological system. Therefore, the contribution made by the paper should be most significant to better understand the dynamics of competitive communities and their eco-evolutionary trajectories. 

References

Nash JF (1950) Equilibrium points in n-person games. Proceedings of the National Academy of Sciences, 36, 48–49. https://doi.org/10.1073/pnas.36.1.48

Rogers AR (2021) Beating your Neighbor to the Berry Patch. bioRxiv, 2020.11.12.380311, ver. 8 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2020.11.12.380311

 

Beating your neighbor to the berry patchAlan R. Rogers<p style="text-align: justify;">Foragers often compete for resources that ripen (or otherwise improve) gradually. What strategy is optimal in this situation? It turns out that there is no optimal strategy. There is no evolutionarily stable strateg...Behaviour & Ethology, Evolutionary ecology, ForagingFrançois Munoz, , Erol Akçay, , Jorge Peña, Sébastien Lion, , , , François Rousset, 2020-12-10 18:38:49 View
01 Mar 2022
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Dissimilarity of species interaction networks: quantifying the effect of turnover and rewiring

How to evaluate and interpret the contribution of species turnover and interaction rewiring when comparing ecological networks?

Recommended by ORCID_LOGO based on reviews by Ignasi Bartomeus and 1 anonymous reviewer

A network includes a set of vertices or nodes (e.g., species in an interaction network), and a set of edges or links (e.g., interactions between species). Whether and how networks vary in space and/or time are questions often addressed in ecological research. 

Two ecological networks can differ in several extents: in that species are different in the two networks and establish new interactions (species turnover), or in that species that are present in both networks establish different interactions in the two networks (rewiring). The ecological meaning of changes in network structure is quite different according to whether species turnover or interaction rewiring plays a greater role. Therefore, much attention has been devoted in recent years on quantifying and interpreting the relative changes in network structure due to species turnover and/or rewiring.

Poisot et al. (2012) proposed to partition the global variation in structure between networks, \( \beta_{WN} \) (WN = Whole Network) into two terms: \( \beta_{OS} \) (OS = Only Shared species) and \( \beta_{ST} \) (ST = Species Turnover), such as \( \beta_{WN} = \beta_{OS} + \beta_{ST} \).

The calculation lays on enumerating the interactions between species that are common or not to two networks, as illustrated on Figure 1 for a simple case. Specifically, Poisot et al. (2012) proposed to use a Sorensen type measure of network dissimilarity, i.e., \( \beta_{WN} = \frac{a+b+c}{(2a+b+c)/2} -1=\frac{b+c}{2a+b+c} \) , where \( a \) is the number of interactions shared between the networks, while \( b \) and \( c \) are interaction numbers unique to one and the other network, respectively. \( \beta_{OS} \) is calculated based on the same formula, but only for the subnetworks including the species common to the two networks, in the form \( \beta_{OS} = \frac{b_{OS}+c_{OS}}{2a_{OS}+b_{OS}+c_{OS}} \) (e.g., Fig. 1). \( \beta_{ST} \) is deduced by subtracting \( \beta_{OS} \) from \( \beta_{WN} \) and represents in essence a "dissimilarity in interaction structure introduced by dissimilarity in species composition" (Poisot et al. 2012).

Figure 1. Ecological networks exemplified in Fründ (2021) and discussed in Poisot (2022). a is the number of shared links (continuous lines in right figures), while b+c is the number of edges unique to one or the other network (dashed lines in right figures).

Alternatively, Fründ (2021) proposed to define \( \beta_{OS} = \frac{b_{OS}+c_{OS}}{2a+b+c} \) and \( \beta_{ST} = \frac{b_{ST}+c_{ST}}{2a+b+c} \), where \( b_{ST}=b-b_{OS} \)  and \( c_{ST}=c-c_{OS} \) , so that the components \( \beta_{OS} \) and \( \beta_{ST} \) have the same denominator. In this way, Fründ (2021) partitioned the count of unique \( b+c=b_{OS}+b_{ST}+c_{ST} \) interactions, so that \( \beta_{OS} \) and \( \beta_{ST} \) sums to \( \frac{b_{OS}+c_{OS}+b_{ST}+c_{ST}}{2a+b+c} = \frac{b+c}{2a+b+c} = \beta_{WN} \). Fründ (2021) advocated that this partition allows a more sensible comparison of \( \beta_{OS} \) and \( \beta_{ST} \), in terms of the number of links that contribute to each component.

For instance, let us consider the networks 1 and 2 in Figure 1 (left panel) such as \( a_{OS}=2 \) (continuous lines in right panel), \( b_{ST} + c_{ST} = 1 \) and \( b_{OS} + c_{OS} = 1 \) (dashed lines in right panel), and thereby \( a = 2 \), \( b+c=2 \), \( \beta_{WN} = 1/3 \). Fründ (2021) measured \( \beta_{OS}=\beta_{ST}=1/6 \) and argued that it is appropriate insofar as it reflects that the number of unique links in the OS and ST components contributing to network dissimilarity (dashed lines) are actually equal. Conversely, the formula of Poisot et al. (2012) yields \( \beta_{OS}=1/5 \), hence \( \beta_{ST} = \frac{1}{3}-\frac{1}{5}=\frac{2}{15}<\beta_{OS} \). Fründ (2021) thus argued that the method of Poisot tends to underestimate the contribution of species turnover.

To clarify and avoid misinterpretation of the calculation of \( \beta_{OS} \) and \( \beta_{ST} \) in Poisot et al. (2012), Poisot (2022) provides a new, in-depth mathematical analysis of the decomposition of \( \beta_{WN} \). Poisot et al. (2012) quantify in \( \beta_{OS} \) the actual contribution of rewiring in network structure for the subweb of common species. Poisot (2022) thus argues that \( \beta_{OS} \) relates only to the probability of rewiring in the subweb, while the definition of \( \beta_{OS} \) by Fründ (2021) is relative to the count of interactions in the global network (considered in denominator), and is thereby dependent on both rewiring probability and species turnover. Poisot (2022) further clarifies the interpretation of \( \beta_{ST} \). \( \beta_{ST} \) is obtained by subtracting \( \beta_{OS} \) from \( \beta_{WN} \) and thus represents the influence of species turnover in terms of the relative architectures of the global networks and of the subwebs of shared species. Coming back to the example of Fig.1., the Poisot et al. (2012) formula posits that \( \frac{\beta_{ST}}{\beta_{WN}}=\frac{2/15}{1/3}=2/5 \), meaning that species turnover contributes two-fifths of change in network structure, while rewiring in the subweb of common species contributed three fifths.  Conversely, the approach of Fründ (2021) does not compare the architectures of global networks and of the subwebs of shared species, but considers the relative contribution of unique links to network dissimilarity in terms of species turnover and rewiring. 

Poisot (2022) concludes that the partition proposed in Fründ (2021) does not allow unambiguous ecological interpretation of rewiring. He provides guidelines for proper interpretation of the decomposition proposed in Poisot et al. (2012).

References

Fründ J (2021) Dissimilarity of species interaction networks: how to partition rewiring and species turnover components. Ecosphere, 12, e03653. https://doi.org/10.1002/ecs2.3653

Poisot T, Canard E, Mouillot D, Mouquet N, Gravel D (2012) The dissimilarity of species interaction networks. Ecology Letters, 15, 1353–1361. https://doi.org/10.1111/ele.12002

Poisot T (2022) Dissimilarity of species interaction networks: quantifying the effect of turnover and rewiring. EcoEvoRxiv Preprints, ver. 4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.32942/osf.io/gxhu2

Dissimilarity of species interaction networks: quantifying the effect of turnover and rewiringTimothée Poisot<p style="text-align: justify;">Despite having established its usefulness in the last ten years, the decomposition of ecological networks in components allowing to measure their β-diversity retains some methodological ambiguities. Notably, how to ...Biodiversity, Interaction networks, Theoretical ecologyFrançois Munoz2021-07-31 00:18:41 View
07 Jun 2023
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High intraspecific growth variability despite strong evolutionary heritage in a neotropical forest

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

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

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

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

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

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

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

REFERENCES

Baraloto, Christopher, Olivier J. Hardy, C. E. Timothy Paine, Kyle G. Dexter, Corinne Cruaud, Luke T. Dunning, Mailyn-Adriana Gonzalez, et al. 2012. « Using functional traits and phylogenetic trees to examine the assembly of tropical tree communities ». Journal of Ecology, 100: 690‑701.
https://doi.org/10.1111/j.1365-2745.2012.01966.x
 
Fortunel Claire, Lasky Jesse R., Uriarte María, Valencia Renato, Wright S.Joseph, Garwood Nancy C., et Kraft Nathan J. B. 2018. « Topography and neighborhood crowding can interact to shape species growth and distribution in a diverse Amazonian forest ». Ecology, 99(10): 2272-2283. https://doi.org/10.1002/ecy.2441
 
Hubbell, S. P. 2001. The Unified Neutral Theory of Biodiversity and Biogeography. 1 vol. Princeton and Oxford: Princeton University Press. https://www.jstor.org/stable/j.ctt7rj8w
 
Janzen, Daniel H. 1970. « Herbivores and the number of tree species in tropical forests ». American Naturalist, 104(940): 501-528. https://doi.org/10.1086/282687
 
Poorter, Lourens, Carolina V. Castilho, Juliana Schietti, Rafael S. Oliveira, et Flávia R. C. Costa. 2018. « Can traits predict individual growth performance? A test in a hyperdiverse tropical forest ». New Phytologist, 219 (1): 109‑21. https://doi.org/10.1111/nph.15206
 
Salguero-Gómez, Roberto, Cyrille Violle, Olivier Gimenez, et Dylan Childs. 2018. « Delivering the promises of trait-based approaches to the needs of demographic approaches, and vice versa ». Functional Ecology, 32 (6): 1424‑35. https://doi.org/10.1111/1365-2435.13148
 
Schmitt, Sylvain, Valérie Raevel, Maxime Réjou‐Méchain, Narayanan Ayyappan, Natesan Balachandran, Narayanan Barathan, Gopalakrishnan Rajashekar, et François Munoz. 2021. « Canopy and understory tree guilds respond differently to the environment in an Indian rainforest ». Journal of Vegetation Science, e13075. https://doi.org/10.1111/jvs.13075
 
Sylvain Schmitt, Bruno Hérault, et Géraldine Derroire. 2023. « High intraspecific growth variability despite strong evolutionary heritage in a neotropical forest ». bioRxiv, 2022.07.27.501745, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.07.27.501745
 
Violle, C., M. L. Navas, D. Vile, E. Kazakou, C. Fortunel, I. Hummel, et E. Garnier. 2007. « Let the concept of trait be functional! » Oikos, 116(5), 882-892. https://doi.org/10.1111/j.0030-1299.2007.15559.x

High intraspecific growth variability despite strong evolutionary heritage in a neotropical forestSylvain Schmitt, Bruno Hérault, Géraldine Derroire<p style="text-align: justify;">Individual tree growth is a key determinant of species performance and a driver of forest dynamics and composition. Previous studies on tree growth unravelled the variation in species growth as a function of demogra...Community ecology, Demography, Population ecologyFrançois Munoz Jelena Pantel, David Murray-Stoker2022-08-01 14:29:04 View
02 Oct 2018
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How optimal foragers should respond to habitat changes? On the consequences of habitat conversion.

Optimal foraging in a changing world: old questions, new perspectives

Recommended by ORCID_LOGO based on reviews by Frederick Adler, Andrew Higginson and 1 anonymous reviewer

Marginal value theorem (MVT) is an archetypal model discussed in every behavioural ecology textbook. Its popularity is largely explained but the fact that it is possible to solve it graphically (at least in its simplest form) with the minimal amount of equations, which is a sensible strategy for an introductory course in behavioural ecology [1]. Apart from this heuristic value, one may be tempted to disregard it as a naive toy model. After a burst of interest in the 70's and the 80's, the once vivid literature about optimal foraging theory (OFT) has lost its momentum [2]. Yet, OFT and MVT have remained an active field of research in the parasitoidologists community, mostly because the sampling strategy of a parasitoid in patches of hosts and its resulting fitness gain are straightforward to evaluate, which eases both experimental and theoretical investigations [3].
This preprint [4] is in line with the long-established literature on OFT. It follows two theoretical articles [5,6] in which Vincent Calcagno and co-authors assessed the effect of changes in the environmental conditions on optimal foraging strategy. This time, they did not modify the shape of the gain function (describing the diminishing return of the cumulative intake as a function of the residency time in a patch) but the relative frequencies of good and bad patches. At first sight, that sounds like a minor modification of their earlier models. Actually, even the authors initially were fooled by the similarities before spotting the pitfalls. Here, they genuinely point out the erroneous verbal prediction in their previous paper in which some non-trivial effects of the change in patch frequencies have been overlooked. The present study indeed provides a striking example of ecological fallacy, and more specifically of Simpson's paradox which occurs when the aggregation of subgroups modifies the apparent pattern at the scale of the entire population [7,8]. In the case of MVT under constraints of habitat conversion, the increase of the residency times in both bad and good patches can result in a decrease of the average residency time at the level of the population. This apparently counter-intuitive property can be observed, for instance, when the proportion of bad quality patches strongly increases, which increases the probability that the individual forages on theses quickly exploited patches, and thus decreases its average residency time on the long run.
The authors thus put the model on the drawing board again. Proper assessment of the effect of change in the frequency of patch quality is more mathematically challenging than when one considers only changes in the shape of the gain function. The expected gain must be evaluated at the scale of the entire habitat instead of single patch. Overall, this study, which is based on a rigorous formalism, stands out as a warning against too rapid interpretations of theoretical outputs. It is not straightforward to generalize the predictions of previous models without careful evaluating their underlying hypotheses. The devil is in the details: some slight, seemingly minor, adjustments of the assumptions may have some major consequences.
The authors discussed the general conditions leading to changes in residency times or movement rates. Yet, it is worth pointing out again that it would be a mistake to blindly consider these theoretical results as forecasts for the foragers' behaviour in natura. OFT models has for a long time been criticized for sweeping under the carpet the key questions of the evolutionary dynamics and the maintenance of the optimal strategy in a population [9,10]. The distribution of available options is susceptible to change rapidly due to modifications of the environmental conditions or, even more simply, the presence of competitors which continuously remove the best options from the pool of available options [11]. The key point here is that the constant monitoring of available options implies cognitive (neural tissue is one of the most metabolically expensive tissues) and ecological costs: assessment and adjustment to the environmental conditions requires time, energy, and occasional mistakes (cost of naiveté, [12]). While rarely considered in optimal analyses, these costs should severely constraint the evolution of the subtle decision rules. Under rapidly fluctuating conditions, it could be more profitable to maintain a sub-optimal strategy (but performing reasonably well on the long run) than paying the far from negligible costs implied by the pursuit of optimal strategies [13,14]. For instance, in the analysis presented in this preprint, it is striking how close the fitness gains of the plastic and the non-plastic forager are, particularly if one remembers that the last-mentioned cognitive and ecological costs have been neglected in these calculations.
Yet, even if one can arguably question its descriptive value, such models are worth more than a cursory glance. They still have normative value insofar that they provide upper bounds for the response to modifications of the environmental conditions. Such insights are precious to design future experiments on the question. Being able to compare experimentally measured behaviours with the extremes of the null model (stubborn non-plastic forager) and the optimal strategy (only achievable by an omniscient daemon) informs about the cognitive bias or ecological costs experienced by real life foragers. I thus consider that this model, and more generally most OFT models, are still a valuable framework which deserves further examination.

References

[1] Fawcett, T. W. & Higginson, A. D. 2012 Heavy use of equations impedes communication among biologists. Proc. Natl. Acad. Sci. 109, 11735–11739. doi: 10.1073/pnas.1205259109
[2] Owens, I. P. F. 2006 Where is behavioural ecology going? Trends Ecol. Evol. 21, 356–361. doi: 10.1016/j.tree.2006.03.014
[3] Louâpre, P., Fauvergue, X., van Baaren, J. & Martel, V. 2015 The male mate search: an optimal foraging issue? Curr. Opin. Insect Sci. 9, 91–95. doi: 10.1016/j.cois.2015.02.012
[4] Calcagno, V., Hamelin, F., Mailleret, L., & Grognard, F. (2018). How optimal foragers should respond to habitat changes? On the consequences of habitat conversion. bioRxiv, 273557, ver. 4 peer-reviewed and recommended by PCI Ecol. doi: 10.1101/273557
[5] Calcagno, V., Grognard, F., Hamelin, F. M., Wajnberg, É. & Mailleret, L. 2014 The functional response predicts the effect of resource distribution on the optimal movement rate of consumers. Ecol. Lett. 17, 1570–1579. doi: 10.1111/ele.12379
[6] Calcagno, V., Mailleret, L., Wajnberg, É. & Grognard, F. 2013 How optimal foragers should respond to habitat changes: a reanalysis of the Marginal Value Theorem. J. Math. Biol. 69, 1237–1265. doi: 10.1007/s00285-013-0734-y
[7] Galipaud, M., Bollache, L., Wattier, R., Dechaume-Moncharmont, F.-X. & Lagrue, C. 2015 Overestimation of the strength of size-assortative pairing in taxa with cryptic diversity: a case of Simpson's paradox. Anim. Behav. 102, 217–221. doi: 10.1016/j.anbehav.2015.01.032
[8] Kievit, R. A., Frankenhuis, W. E., Waldorp, L. J. & Borsboom, D. 2013 Simpson's paradox in psychological science: a practical guide. Front. Psychol. 4, 513. doi: 10.3389/fpsyg.2013.00513
[9] Bolduc, J.-S. & Cézilly, F. 2012 Optimality modelling in the real world. Biol. Philos. 27, 851–869. doi: 10.1007/s10539-012-9333-3
[10] Pierce, G. J. & Ollason, J. G. 1987 Eight reasons why optimal foraging theory is a complete waste of time. Oikos 49, 111–118. doi: 10.2307/3565560
[11] Dechaume-Moncharmont, F.-X., Brom, T. & Cézilly, F. 2016 Opportunity costs resulting from scramble competition within the choosy sex severely impair mate choosiness. Anim. Behav. 114, 249–260. doi: 10.1016/j.anbehav.2016.02.019
[12] Snell-Rood, E. C. 2013 An overview of the evolutionary causes and consequences of behavioural plasticity. Anim. Behav. 85, 1004–1011. doi: 10.1016/j.anbehav.2012.12.031
[13] Fawcett, T. W., Fallenstein, B., Higginson, A. D., Houston, A. I., Mallpress, D. E. W., Trimmer, P. C. & McNamara, J. M. 2014 The evolution of decision rules in complex environments. Trends Cogn. Sci. 18, 153–161. doi: 10.1016/j.tics.2013.12.012
[14] Marshall, J. A. R., Trimmer, P. C., Houston, A. I. & McNamara, J. M. 2013 On evolutionary explanations of cognitive biases. Trends Ecol. Evol. 28, 469-473. doi: 10.1016/j.tree.2013.05.013

How optimal foragers should respond to habitat changes? On the consequences of habitat conversion.Vincent Calcagno, Frederic Hamelin, Ludovic Mailleret, Frederic GrognardThe Marginal Value Theorem (MVT) provides a framework to predict how habitat modifications related to the distribution of resources over patches should impact the realized fitness of individuals and their optimal rate of movement (or patch residen...Behaviour & Ethology, Dispersal & Migration, Foraging, Landscape ecology, Spatial ecology, Metacommunities & Metapopulations, Theoretical ecologyFrancois-Xavier Dechaume-Moncharmont2018-03-05 10:42:11 View