Morand et al. (2024) designed convolutional neural networks to predict the occurrences of 38 marine animals worldwide. The environmental predictors were sea surface temperature, chlorophyll concentration, salinity and fifteen others. The time of some of the predictors was chosen to be as close as possible to the time of the observed occurrence.

This approach has previously only been applied to the analysis of the distribution of terrestrial plant species (Botella et al. 2018, Deneu et al. 2021), so the application here to very different marine ecosystems and organisms is a novelty worth highlighting and discussing.

A very interesting feature of PCI Ecology is that reviews are provided with the final manuscript and the present recommendation text.

In the case of the Morand et al. article, the reviewers provided very detailed and insightful comments that deserve to be published and read alongside the article.

The reviewers' comments question the ecological significance and implications of choosing fine temporal and spatial scales in CNN distribution modelling in order to obtain species distribution modelling (SDM).

The main question debated during the review process was whether the CNN modeling approach used here can be defined as a kind of niche modeling.

The fact is that most of the organisms studied here are mobile, and the authors have taken into account precise environmental information at dates close to those of species appearance (for example, "Temperature and chlorophyll values were also included 15 and 5 days before the occurrences"). In doing so, they took into account the fine spatial and temporal scales of species occurrences and environmental conditions, which can be influenced by both environmental preferences and the movement behaviors of individuals. The question then arises: does this approach really represent the ecological niches of the marine organisms selected? Given that most selected organisms may have specific seasonal movement dynamics, the CNN model also learns the individual movement behaviors of organisms over seasons and years. The ecological niche is a broader concept that takes into account all the environmental conditions that enable species to persist over the course of their lives and over generations. This differs from the case of sessile land plants, which must respond to the environmental context only at the points of appearance.

This is not a shortcoming of the methodology proposed here but rather an interesting conceptual issue to be considered and discussed. Modelling the occurrence of individuals at a given time and position can characterize not only the species' niche but also the dynamics of organisms' temporal movements. As a result, the model predicts the position of individuals at a given time, while the niche should also represent the role of environmental conditions faced by individuals at other times in their lives.
A relevant perspective would then be to analyze whether and how the neural network can help disentangle the ranges of environmental conditions defining the niche from those influencing the movement dynamics of individuals.

Another interesting point is that the CNN model is used here as a multi-species classifier, meaning that it provides the ranked probability that a given observation corresponds to one of the 38 species considered in the study, depending on the environmental conditions at the location and time of the observation. In other words, the model provides the relative chance of choosing each of the 38 species at a given time and place. Imagine that you are only studying two species that have exactly the same niche, a standard SDM approach should provide a high probability of occurrence close to 1 in localities where environmental conditions are very and equally suited to both species, while the CNN classifier would provide a value close to 0.5 for both species, meaning that we have an equal chance of choosing one or the other. Consequently, the fact that the probability given by the classifier is higher for a species at a given point than at another point does not (necessarily) mean that the first point presents better environmental conditions for that species but rather that we are more likely to choose it over one of the other species at this point than at another. In fact, the classification task also reflects whether the other 37 species are more or less likely to be found at each point. The classifier, therefore, does not provide the relative probability of occurrence of a species in space but rather a relative chance of finding it instead of one of the other 37 species at each point of space and time.

It is important that an ecologist designing a multi-species classifier for species distribution modelling is well aware of this point and does not interpret the variation of probabilities for a species in space as an indication of more or less suitable habitat for that specific species. On the other hand, predicting the relative probabilities of finding species to a given point at a given time gives an indication of the dynamics of their local co-occurrence. In this respect, the CNN approach is closer to a joint species distribution model (jSDM). As Ovaskainen et al. (2017) mention, "By simultaneously drawing on the information from multiple species, these (jSDM) models allow one to seek community-level patterns in how species respond to their environment". Let's return to the two species example we used above. The fact that the probabilities are 0.5 for both species actually suggests that both species can coexist at the same abundance at this location. In this respect, the CNN multi-species classifier offers promising prospects for the prediction of assemblages and habitats thanks to the relative importance of the most characteristic/dominant species from a species pool. The species pool comprises all classified species and must be sufficiently representative of the ecological diversity of species niches in the area.

Finally, CNN-based species distribution modelling is a powerful and promising tool for studying the distributions of multi-species assemblages as a function of local environmental features but also of the spatial heterogeneity of each feature around the observation point in space and time (Deneu et al. 2021). It allows acknowledging the complex effects of environmental predictors and the roles of their spatial and temporal heterogeneity through the convolution operations performed in the neural network. As more and more computationally intensive tools become available, and as more and more environmental data becomes available at finer and finer temporal and spatial scales, the CNN approach is likely to be increasingly used to study biodiversity patterns across spatial and temporal scales.

References

Botella, C., Joly, A., Bonnet, P., Monestiez, P., and Munoz, F. (2018). Species distribution modeling based on the automated identification of citizen observations. Applications in Plant Sciences, 6(2), e1029. https://doi.org/10.1002/aps3.1029

Deneu, B., Servajean, M., Bonnet, P., Botella, C., Munoz, F., and Joly, A. (2021). Convolutional neural networks improve species distribution modelling by capturing the spatial structure of the environment. PLoS Computational Biology, 17(4), e1008856. https://doi.org/10.1371/journal.pcbi.1008856

Morand, G., Joly, A., Rouyer, T., Lorieul, T., and Barde, J. (2024) Predicting species distributions in the open ocean with convolutional neural networks. bioRxiv, ver.3 peer-reviewed and recommended by PCI Ecology https://doi.org/10.1101/2023.08.11.551418

Ovaskainen, O., Tikhonov, G., Norberg, A., Guillaume Blanchet, F., Duan, L., Dunson, D., ... and Abrego, N. (2017). How to make more out of community data? A conceptual framework and its implementation as models and software. Ecology letters, 20(5), 561-576. https://doi.org/10.1111/ele.12757

Ivimey Cook et al. (2023) provide a concise and useful “How to” review code for researchers in the fields of ecology and evolutionary biology, where the systematic review of code is not yet standard practice during the peer review of articles. Consequently, this article is full of tips for authors on how to make their code easier to review. This handy article applies not only to ecology and evolutionary biology, but to many fields that are learning how to make code more reproducible and shareable. Taking this step toward transparency is key to improving research rigor (Brito et al. 2020) and is a necessary step in helping make research trustable by the public (Rosman et al. 2022).

References

Brito, J. J., Li, J., Moore, J. H., Greene, C. S., Nogoy, N. A., Garmire, L. X., & Mangul, S. (2020). Recommendations to enhance rigor and reproducibility in biomedical research. GigaScience, 9(6), giaa056. https://doi.org/10.1093/gigascience/giaa056

Ivimey-Cook, E. R., Pick, J. L., Bairos-Novak, K., Culina, A., Gould, E., Grainger, M., Marshall, B., Moreau, D., Paquet, M., Royauté, R., Sanchez-Tojar, A., Silva, I., Windecker, S. (2023). Implementing Code Review in the Scientific Workflow: Insights from Ecology and Evolutionary Biology. EcoEvoRxiv, ver 5 peer-reviewed and recommended by Peer Community In Ecology. https://doi.org/10.32942/X2CG64

Rosman, T., Bosnjak, M., Silber, H., Koßmann, J., & Heycke, T. (2022). Open science and public trust in science: Results from two studies. Public Understanding of Science, 31(8), 1046-1062. https://doi.org/10.1177/09636625221100686

In the recent years, morphometrics, the quantitative description of shape and its covariation [1] gained considerable momentum in evolutionary ecology. Using the form of organisms to describe, classify and try to understand their diversity can be traced back at least to Aristotle. More recently, two successive revolutions rejuvenated this idea [1–3]: first, a proper mathematical refoundation of the theory of shape, then a technical revolution in the apparatus able to acquire raw data. By using a feature extraction method and planning its massive use on data acquired by aerial drones, the study by Lacaux and colleagues [4] retraces this curse of events.
The radial symmetry of Aurelia spp. jelly fish, a common species complex, is affected by stress and more largely by environmental variations, such as pollution exposition. Aurelia spp. normally present four gonads so that the proportion of non-tetramerous individuals in a population has been proposed as a biomarker [5,6].
In this study, the authors implemented the Hough transform to largely automate the detection of the gonads in Aurelia spp. Such use of the Hough transform, a long-used approach to identify shapes through edge detection, is new to morphometrics. Here, the Aurelia spp. gonads are identified as ellipses from which aspect descriptors can be derived, and primarily counted and thus can be used to quantify the proportion of individuals presenting body plans disorders.

The sample sizes studied here were too low to allow finer-grained ecophysiological investigations. That being said, the proof-of-concept is convincing and this paper paths the way for an operational and innovative approach to the ecological monitoring of sensible aquatic ecosystems.

References

[1] Kendall, D. G. (1989). A survey of the statistical theory of shape. Statistical Science, 87-99. doi: https://doi.org/10.1214/ss/1177012589
[2] Rohlf, F. J., and Marcus, L. F. (1993). A revolution morphometrics. Trends in ecology & evolution, 8(4), 129-132. doi: https://doi.org/10.1016/0169-5347(93)90024-J
[3] Adams, D. C., Rohlf, F. J., and Slice, D. E. (2004). Geometric morphometrics: ten years of progress following the ‘revolution’. Italian Journal of Zoology, 71(1), 5-16. doi: https://doi.org/10.1080/11250000409356545
[4] Lacaux, C., Desolneux, A., Gadreaud, J., Martin-Garin, B. and Thiéry, A. (2020) Hough transform implementation to evaluate the morphological variability of the moon jellyfish (Aurelia spp.). bioRxiv, 2020.03.11.986984, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. doi: https://doi.org/10.1101/2020.03.11.986984
[5] Gershwin, L. A. (1999). Clonal and population variation in jellyfish symmetry. Journal of the Marine Biological Association of the United Kingdom, 79(6), 993-1000. doi: https://doi.org/10.1017/S0025315499001228
[6] Gadreaud, J., Martin-Garin, B., Artells, E., Levard, C., Auffan, M., Barkate, A.-L. and Thiéry, A. (2017) The moon jellyfish as a new bioindicator: impact of silver nanoparticles on the morphogenesis. In: Mariottini GL, editor. Jellyfish: ecology, distribution patterns and human interactions. Nova Science Publishers; 2017. pp. 277–292.

In 2018 Blaisdell and colleagues set out to study how causal cognition may impact large scale macroecological patterns, more specifically range dynamics, in the great-tailed grackle (Fronhofer 2019). This line of research is at the forefront of current thought in macroecology, a field that has started to recognize the importance of animal behaviour more generally (see e.g. Keith and Bull (2017)). Importantly, the authors were pioneering the use of preregistrations in ecology and evolution with the aim of improving the quality of academic research.

Now, nearly 3 years later, it is thanks to their endeavour of making research better that we learn that the authors are “[...] unable to speculate about the potential role of causal cognition in a species that is rapidly expanding its geographic range.” (Blaisdell et al. 2021; page 2). Is this a success or a failure? Every reader will have to find an answer to this question individually and there will certainly be variation in these answers as becomes clear from the referees’ comments. In my opinion, this is a success story of a more stringent and transparent approach to doing research which will help us move forward, both methodologically and conceptually.

References

Fronhofer (2019) From cognition to range dynamics: advancing our understanding of macroe-
cological patterns. Peer Community in Ecology, 100014. doi: https://doi.org/10.24072/pci.ecology.100014

Keith, S. A. and Bull, J. W. (2017) Animal culture impacts species' capacity to realise climate-driven range shifts. Ecography, 40: 296-304. doi: https://doi.org/10.1111/ecog.02481

Blaisdell, A., Seitz, B., Rowney, C., Folsom, M., MacPherson, M., Deffner, D., and Logan, C. J. (2021) Do the more flexible individuals rely more on causal cognition? Observation versus intervention in causal inference in great-tailed grackles. PsyArXiv, ver. 5 peer-reviewed and recommended by Peer community in Ecology. doi: https://doi.org/10.31234/osf.io/z4p6s

A core challenge facing ecologists is to work through an ever-increasing amount of data. The accelerating decline in biodiversity worldwide, mounting pressure of anthropogenic impacts, and increasing demand for actionable indicators to guide effective policy means that monitoring will only intensify, and rely on tools that can generate even more information (Gonzalez et al., 2023). How, then, do we handle this new volume and diversity of data?

This is the question Royaux et al. (2024) are tackling with their contribution. By introducing both a conceptual ("How should we think about our work?") and an operational ("Here is a tool to do our work with") framework, they establish a series of best practices for the analysis of ecological data.

It is easy to think about best practices in ecological data analysis in its most proximal form: is it good statistical practice? Is the experimental design correct? These have formed the basis of many recommendations over the years (see e.g. Popovic et al., 2024, for a recent example). But the contribution of Royaux et al. focuses on a different part of the analysis pipeline: the computer science (and software engineering) aspect of it.

As data grows in volume and complexity, the code needed to handle it follows the same trend. It is not a surprise, therefore, to see that the demand for programming skills in ecologists has doubled recently (Feng et al., 2020), prompting calls to make computational literacy a core component of undergraduate education (Farrell & Carrey, 2018). But beyond training, an obvious way to make computational analysis ecological data more reliable and effective is to build better tools. This is precisely what Royaux et al. have achieved.

They illustrate their approach through their experience building Galaxy-Ecology, a computing environment for ecological analysis: by introducing a clear taxonomy of computing concepts (data exploration, pre-processing, analysis, representation), with a hierarchy between them (formatting, data correction, anonymization), they show that we can think about the pipeline going from data to results in a way that is more systematized, and therefore more prone to generalization.

We may buckle at the idea of yet another ontology, or yet another framework, for our work, but I am convinced that the work of Royaux et al. is precisely what our field needs. Because their levels of atomization (their term for the splitting of complex pipelines into small, single-purpose tasks) are easy to understand, and map naturally onto tasks that we already perform, it is likely to see wide adoption. Solving the big, existential challenges of monitoring and managing biodiversity at the global scale requires the adoption of good practices, and a tool like Galaxy-Ecology goes a long way towards this goal.

References

Farrell, K.J., and Carey, C.C. (2018). Power, pitfalls, and potential for integrating computational literacy into undergraduate ecology courses. Ecol. Evol. 8, 7744-7751.
https://doi.org/10.1002/ece3.4363

Feng, X., Qiao, H., and Enquist, B. (2020). Doubling demands in programming skills call for ecoinformatics education. Frontiers in Ecology and the Environment 18, 123-124.
https://doi.org/10.1002/fee.2179
 
Gonzalez, A., Vihervaara, P., Balvanera, P., Bates, A.E., Bayraktarov, E., Bellingham, P.J., Bruder, A., Campbell, J., Catchen, M.D., Cavender-Bares, J., et al. (2023). A global biodiversity observing system to unite monitoring and guide action. Nat. Ecol. Evol., 1-5. 
https://doi.org/10.1038/s41559-023-02171-0
 
Popovic, G., Mason, T.J., Drobniak, S.M., Marques, T.A., Potts, J., Joo, R., Altwegg, R., Burns, C.C.I., McCarthy, M.A., Johnston, A., et al. (2024). Four principles for improved statistical ecology. Methods Ecol. Evol. 15, 266-281.
https://doi.org/10.1111/2041-210X.14270
 
Coline Royaux, Jean-Baptiste Mihoub, Marie Jossé, Dominique Pelletier, Olivier Norvez, Yves Reecht, Anne Fouilloux, Helena Rasche, Saskia Hiltemann, Bérénice Batut, Marc Eléaume, Pauline Seguineau, Guillaume Massé, Alan Amossé, Claire Bissery, Romain Lorrilliere, Alexis Martin, Yves Bas, Thimothée Virgoulay, Valentin Chambon, Elie Arnaud, Elisa Michon, Clara Urfer, Eloïse Trigodet, Marie Delannoy, Gregoire Loïs, Romain Julliard, Björn Grüning, Yvan Le Bras (2024) Guidance framework to apply best practices in ecological data analysis: Lessons learned from building Galaxy-Ecology. EcoEvoRxiv, ver.3 peer-reviewed and recommended by PCI Ecology. 
https://doi.org/10.32942/X2G033