Guiding and monitoring the efficiency of conservation efforts needs robust scientific background information, of which one key element is estimating wildlife abundance and its spatial and temporal variation. As raw counts are by nature incomplete counts of a population, correcting for detectability is required (Clobert, 1995; Turlure et al., 2018). This can be done with Capture-Mark-Recapture protocols (Iijima, 2020). Techniques for marking individuals are diverse, e.g. writing on butterfly wings, banding birds, or using natural specific patterns in the individual’s body such as leopard fur or whale tail. Advancement in technology opens new opportunities for developing marking techniques, including strategies to limit mark identification errors (Burchill & Pavlic, 2019), and for using active marks that can transmit data remotely or be read automatically.

The details of such methodological developments frequently remain unpublished, the method being briefly described in studies that use it. For a few years, there has been however a renewed interest in proper publishing of methods for ecology and evolution. This study by Folk & Mennerat (2023) fits in this context, offering a nice example of detailed description and testing of a method to mark salmon ectoparasites using RFID tags. Such tags are extremely small, yet easy to use, even with automatic recording procedure. The study provides a very good basis protocol that should help researchers working for small species, in particular invertebrates. The study is complemented by a video illustrating the placement of the tag so the reader who would like to replicate the procedure can get a very precise idea of it.

References

Burchill, A. T., & Pavlic, T. P. (2019). Dude, where’s my mark? Creating robust animal identification schemes informed by communication theory. Animal Behaviour, 154, 203–208. https://doi.org/10.1016/j.anbehav.2019.05.013

Clobert, J. (1995). Capture-recapture and evolutionary ecology: A difficult wedding ? Journal of Applied Statistics, 22(5–6), 989–1008.

Folk, A., & Mennerat, A. (2023). Methods for tagging an ectoparasite, the salmon louse Lepeophtheirus salmonis (p. 2023.08.31.555695). bioRxiv, ver. 2 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2023.08.31.555695

Iijima, H. (2020). A Review of Wildlife Abundance Estimation Models: Comparison of Models for Correct Application. Mammal Study, 45(3), 177–188. https://doi.org/10.3106/ms2019-0082

Turlure, C., Pe’er, G., Baguette, M., & Schtickzelle, N. (2018). A simplified mark–release–recapture protocol to improve the cost effectiveness of repeated population size quantification. Methods in Ecology and Evolution, 9(3), 645–656. https://doi.org/10.1111/2041-210X.12900

Humans are exploiting biological resources since thousands of years. Exploitation of biological resources has become particularly intense since the beginning of the 20th century and the steep increase in the worldwide human population size. Marine and freshwater fishes are not exception to that rule, and they have been (and continue to be) strongly harvested as a source of proteins for humans. For some species, fishery has been so intense that natural stocks have virtually collapsed in only a few decades. The worst example begin that of the Northwest Atlantic cod that has declined by more than 95% of its historical biomasses in only 20-30 years of intensive exploitation (Frank et al. 2005). These rapid and steep changes in biomasses have huge impacts on the entire ecosystems since species targeted by fisheries are often at the top of trophic chains (Frank et al. 2005). 

Beyond demographic impacts, fisheries also have evolutionary impacts on populations, which can also indirectly alter ecosystems (Uusi-Heikkilä et al. 2015; Palkovacs et al. 2018). Fishermen generally focus on the largest specimens, and hence exert a strong selective pressure against these largest fish (which is called “harvest selection”). There is now ample evidence that harvest selection can lead to rapid evolutionary changes in natural populations toward small individuals (Kuparinen & Festa-Bianchet 2017). These evolutionary changes are of course undesirable from a human perspective, and have attracted many scientific questions. Nonetheless, the consequence of harvest selection is not always observable in natural populations, and there are cases in which no phenotypic change (or on the contrary an increase in mean body size) has been observed after intense harvest pressures. In a conceptual Essay, Edeline and Loeuille (Edeline & Loeuille 2020) propose novel ideas to explain why the evolutionary consequences of harvest selection can be so diverse, and how a cross talk between ecological and evolutionary dynamics can explain patterns observed in natural stocks.

 The general and novel concept proposed by Edeline and Loeuille is actually as old as Darwin’s book; The Origin of Species (Darwin 1859). It is based on the simple idea that natural selection acting on harvested populations can actually be strong, and counter-balance (or on the contrary reinforce) the evolutionary consequence of harvest selection. Although simple, the idea that natural and harvest selection are jointly shaping contemporary evolution of exploited populations lead to various and sometimes complex scenarios that can (i) explain unresolved empirical patterns and (ii) refine predictions regarding the long-term viability of exploited populations. 

The Edeline and Loeuille’s crafty inspiration is that natural selection acting on exploited populations is itself an indirect consequence of harvest (Edeline & Loeuille 2020). They suggest that, by modifying the size structure of populations (a key parameter for ecological interactions), harvest indirectly alters interactions between populations and their biotic environment through competition and predation, which changes the ecological theatre and hence the selective pressures acting back to populations. They named this process “size-dependent eco-evolutionary feedback loops” and develop several scenarios in which these feedback loops ultimately deviate the evolutionary outcome of harvest selection from expectation. The scenarios they explore are based on strong theoretical knowledge, and range from simple ones in which a single species (the harvest species) is evolving to more complex (and realistic) ones in which multiple (e.g. the harvest species and its prey) species are co-evolving.

I will not come into the details of each scenario here, and I will let the readers (re-)discovering the complex beauty of biological life and natural selection. Nonetheless, I will emphasize the importance of considering these eco-evolutionary processes altogether to fully grasp the response of exploited populations. Edeline and Loeuille convincingly demonstrate that reduced body size due to harvest selection is obviously not the only response of exploited fish populations when natural selection is jointly considered (Edeline & Loeuille 2020). On the contrary, they show that –under some realistic ecological circumstances relaxing exploitative competition due to reduced population densities- natural selection can act antagonistically, and hence favour stable body size in exploited populations. Although this seems further desirable from a human perspective than a downsizing of exploited populations, it is actually mere window dressing as Edeline and Loeuille further showed that this response is accompanied by an erosion of the evolvability –and hence a lowest probability of long-term persistence- of these exploited populations.

Humans, by exploiting biological resources, are breaking the relative equilibrium of complex entities, and the response of populations to this disturbance is itself often complex and heterogeneous. In this Essay, Edeline and Loeuille provide –under simple terms- the theoretical and conceptual bases required to improve predictions regarding the evolutionary responses of natural populations to exploitation by humans (Edeline & Loeuille 2020). An important next step will be to generate data and methods allowing confronting the empirical reality to these novel concepts (e.g. (Monk et al. 2021), so as to identify the most likely evolutionary scenarios sustaining biological responses of exploited populations, and hence to set the best management plans for the long-term sustainability of these populations.

References

Darwin, C. (1859). On the Origin of Species by Means of Natural Selection. John Murray, London.

Edeline, E. & Loeuille, N. (2021) Size-dependent eco-evolutionary feedbacks in fisheries. bioRxiv, 2020.04.03.022905, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: https://doi.org/10.1101/2020.04.03.022905

Frank, K.T., Petrie, B., Choi, J. S. & Leggett, W.C. (2005). Trophic Cascades in a Formerly Cod-Dominated Ecosystem. Science, 308, 1621–1623. doi: https://doi.org/10.1126/science.1113075

Kuparinen, A. & Festa-Bianchet, M. (2017). Harvest-induced evolution: insights from aquatic and terrestrial systems. Philos. Trans. R. Soc. B Biol. Sci., 372, 20160036. doi: https://doi.org/10.1098/rstb.2016.0036

Monk, C.T., Bekkevold, D., Klefoth, T., Pagel, T., Palmer, M. & Arlinghaus, R. (2021). The battle between harvest and natural selection creates small and shy fish. Proc. Natl. Acad. Sci., 118, e2009451118. doi: https://doi.org/10.1073/pnas.2009451118 

Palkovacs, E.P., Moritsch, M.M., Contolini, G.M. & Pelletier, F. (2018). Ecology of harvest-driven trait changes and implications for ecosystem management. Front. Ecol. Environ., 16, 20–28. doi: https://doi.org/10.1002/fee.1743

Uusi-Heikkilä, S., Whiteley, A.R., Kuparinen, A., Matsumura, S., Venturelli, P.A., Wolter, C., et al. (2015). The evolutionary legacy of size-selective harvesting extends from genes to populations. Evol. Appl., 8, 597–620. doi: https://doi.org/10.1111/eva.12268

In this study, the authors report a new method for estimating the abundance of the Pyrenean brown bear population. Precisely, the methodology involved aims to apply Pollock's closed robust design (PCRD) capture-recapture models to estimate population abundance and trends over time. Overall, the results encourage the use of PCRD to study populations' demographic rates, while minimizing biases due to inter-individual heterogeneity in detection probabilities.

Estimating the size and trends of animal population over time is essential for informing conservation status and management decision-making (Nichols & Williams 2006). This is particularly the case when the population is small, geographically scattered, and threatened. Although several methods can be used to estimate population abundance, they may be difficult to implement when individuals are rare, elusive, solitary, largely nocturnal, highly mobile, and/or occupy large home ranges in remote and/or rugged habitats. Moreover, in such standard methods,

• the population is assumed to be closed both geographically (no immigration nor emigration) and demographically (no births nor deaths) and
• all individuals are assumed to have identical detection probabilities regardless of their individual attributes (e.g., age, body mass, social status) and habitat features (home-range location and composition) (Otis et al. 1978).

However, these conditions are rarely met in real populations, such as wild mammals (e.g., Bellemain et al. 2005; Solbert et al. 2006), and therefore the risk of underestimating population size can rapidly increase because the assumption of perfect detection of all individuals in the population is violated.

Focusing on the critically endangered Pyrenean brown bear that was close to extinction in the mid-1990s, the study by Vanpe et al. (2022), uses protocol and opportunistic data to describe a statistical modeling exercise to construct mark-recapture histories from 2008 to 2020. Among the data, the authors collected non-invasive samples such as a mixture of hair and scat samples used for genetic identification, as well as photographic trap data of recognized individuals. These data are then analyzed in RMark to provide detection and survival estimates. The final model (i.e. PCRD capture-recapture) is then used to provide Bayesian population estimates. Results show a five-fold increase in population size between 2008 and 2020, from 13 to 66 individuals. Thus, this study represents the first published annual abundance and temporal trend estimates of the Pyrenean brown bear population since 2008.

Then, although the results emphasize that the PCRD estimates were broadly close to the MRS counts and had reasonably narrow associated 95% Credibility Intervals, they also highlight that the sampling effort is different according to individuals. Indeed, as expected, the detection of an individual depends on

• the intraspecific home range size variation that results in individuals that move the most being most likely to be detected and
• the mortality rate which is higher on cubs than on adults and subadults (due to infanticide by males, predation, death of the mother, or abandonment).

Overall, the PCRD capture-recapture modelling approach, involved in this study, provides robust estimates of abundance and demographic rates of the Pyrenean brown bear population (with associated uncertainty) while minimizing and considering bias due to inter-individual heterogeneity in detection probabilities.

The authors conclude that mark-recapture provides useful population estimates and urge wildlife ecologists and managers to use robust approaches, such as the RDPC capture-recapture model, when studying large mammal populations. This information is essential to inform management decisions and assess the conservation status of populations.

References

Bellemain, E.V.A., Swenson, J.E., Tallmon, D., Brunberg, S. and Taberlet, P. (2005). Estimating population size of elusive animals with DNA from hunter-collected feces: four methods for brown bears. Cons. Biol. 19(1), 150-161. https://doi.org/10.1111/j.1523-1739.2005.00549.x

Nichols, J.D. and Williams, B.K. (2006). Monitoring for conservation. Trends Ecol. Evol. 21(12), 668-673. https://doi.org/10.1016/j.tree.2006.08.007

Otis, D.L., Burnham, K.P., White, G.C. and Anderson, D.R. (1978). Statistical inference from capture data on closed animal populations. Wildlife Monographs (62), 3-135.

Solberg, K.H., Bellemain, E., Drageset, O.M., Taberlet, P. and Swenson, J.E. (2006). An evaluation of field and non-invasive genetic methods to estimate brown bear (Ursus arctos) population size. Biol. Conserv. 128(2), 158-168. https://doi.org/10.1016/j.biocon.2005.09.025

Vanpé C, Piédallu B, Quenette P-Y, Sentilles J, Queney G, Palazón S, Jordana IA, Jato R, Elósegui Irurtia MM, de la Torre JS, and Gimenez O (2022) Estimating abundance of a recovering transboundary brown bear population with capture-recapture models. bioRxiv, 2021.12.08.471719, ver. 4 recommended and peer-reviewed by PCI Ecology. https://doi.org/10.1101/2021.12.08.471719

Polar and subpolar regions are fragile environments, where the introduction of alien species may completely change ecosystem dynamics if the alien species become keystone species (e.g. Croll, 2005). The increasing number of human visits, together with climate change, are favouring the introduction and settling of new invaders to these regions, particularly in Antarctica (Hughes et al. 2015). Within this context, the joint use of Species Distribution Models (SDM) –to assess the areas potentially suitable for the aliens– with other measures of the potential to become successful invaders can inform on the need for devoting specific efforts to eradicate these new species before they become naturalized (e.g. Pertierra et al. 2016).
Bazzichetto et al. (2020) use data from a detailed inventory, SDMs and trait data altogether to assess the drivers of invasion success of six alien plants on Possession Island, in the remote sub-Antarctic archipelago of Crozet. SDMs have inherent limitations to describe different aspects of species distributions, including the fundamental niche and, with it, the areas that could host viable populations (Hortal et al. 2012). Therefore, their utility to predict future biological invasions is limited (Jiménez-Valverde et al. 2011). However, they can be powerful tools to describe species range dynamics if they are thoughtfully used by adopting conscious decisions about the techniques and data used, and interpreting carefully the actual implications of their results.
This is what Bazzichetto et al. (2020) do, using General Linear Models (GLM) –a technique well rooted in the original niche-based SDM theory (e.g. Austin 1990)– that can provide a meaningful description of the realized niche within the limits of an adequately sampled region. Further, as alien species share and are similarly affected by several steps of the invasion process (Richardson et al. 2000), these authors model the realized distribution of the six species altogether. This can be done through the recently developed joint-SDM, a group of techniques where the co-occurrence of the modelled species is explicitly taken into account during modelling (e.g. Pollock et al. 2014). Here, the addition of species traits has been identified as a key step to understand the associations of species in space (see Dormann et al. 2018). Bazzichetto et al. (2020) combine their GLM-based SDM for each species with a so-called multi-SDM approach, where they assess together the consistency in the interactions between both species and topographically-driven climate variations, and several plant traits and two key anthropic factors –accessibility from human settlements and distance to hiking paths.
This work is a good example on how a theoretically meaningful SDM approach can provide useful –though perhaps not deep– insights on biological invasions for remote landscapes threatened by biotic homogenization. By combining climate and topographic variables as proxies for the spatial variations in the abiotic conditions regulating plant growth, measures of accessibility, and traits of the plant invaders, Bazzichetto et al. (2020) are able to identify the different effects that the interactions between the potential intensity of propagule dissemination by humans, and the ecological characteristics of the invaders themselves, may have on their invasion success.
The innovation of modelling together species responses is important because it allows dissecting the spatial dynamics of spread of the invaders, which indeed vary according to a handful of their traits. For example, their results show that no all old residents have profited from the larger time of residence in the island, as Poa pratensis is seemingly as dependent of a higher intensity of human activity as the newcomer invaders in general are. According to Bazzichetto et al. trait-based analyses, these differences are apparently related with plant height, as smaller plants disperse more easily. Further, being perennial also provides an advantage for the persistence in areas with less human influence. This puts name, shame and fame to the known influence of plant life history on their dispersal success (Beckman et al. 2018), at least for the particular case of plant invasions in Possession Island.
Of course this approach has limitations, as data on the texture, chemistry and temperature of the soil are not available, and thus were not considered in the analyses. These factors may be critical for both establishment and persistence of small plants in the harsh Antarctic environments, as Bazzichetto et al. (2020) recognize. But all in all, their results provide key insights on which traits may confer alien plants with a higher likelihood of becoming successful invaders in the fragile Antarctic and sub-Antarctic ecosystems. This opens a way for rapid assessments of invasibility, which will help identifying which species in the process of naturalizing may require active contention measures to prevent them from becoming ecological game changers and cause disastrous cascade effects that shift the dynamics of native ecosystems.

References

Austin, M. P., Nicholls, A. O., and Margules, C. R. (1990). Measurement of the realized qualitative niche: environmental niches of five Eucalyptus species. Ecological Monographs, 60(2), 161-177. doi: https://doi.org/10.2307/1943043
Bazzichetto, M., Massol, F., Carboni, M., Lenoir, J., Lembrechts, J. J. and Joly, R. (2020) Once upon a time in the far south: Influence of local drivers and functional traits on plant invasion in the harsh sub-Antarctic islands. bioRxiv, 2020.07.19.210880, ver. 3 peer-reviewed and recommended by PCI Ecology. doi: https://doi.org/10.1101/2020.07.19.210880
Beckman, N. G., Bullock, J. M., and Salguero-Gómez, R. (2018). High dispersal ability is related to fast life-history strategies. Journal of Ecology, 106(4), 1349-1362. doi: https://doi.org/10.1111/1365-2745.12989
Croll, D. A., Maron, J. L., Estes, J. A., Danner, E. M., and Byrd, G. V. (2005). Introduced predators transform subarctic islands from grassland to tundra. Science, 307(5717), 1959-1961. doi: https://doi.org/10.1126/science.1108485
Dormann, C. F., Bobrowski, M., Dehling, D. M., Harris, D. J., Hartig, F., Lischke, H., Moretti, M. D., Pagel, J., Pinkert, S., Schleuning, M., Schmidt, S. I., Sheppard, C. S., Steinbauer, M. J., Zeuss, D., and Kraan, C. (2018). Biotic interactions in species distribution modelling: 10 questions to guide interpretation and avoid false conclusions. Global Ecology and Biogeography, 27(9), 1004-1016. doi: https://doi.org/10.1111/geb.12759
Jiménez-Valverde, A., Peterson, A., Soberón, J., Overton, J., Aragón, P., and Lobo, J. (2011). Use of niche models in invasive species risk assessments. Biological Invasions, 13(12), 2785-2797. doi: https://doi.org/10.1007/s10530-011-9963-4
Hortal, J., Lobo, J. M., and Jiménez-Valverde, A. (2012). Basic questions in biogeography and the (lack of) simplicity of species distributions: Putting species distribution models in the right place. Natureza & Conservação – Brazilian Journal of Nature Conservation, 10(2), 108-118. doi: https://doi.org/10.4322/natcon.2012.029
Hughes, K. A., Pertierra, L. R., Molina-Montenegro, M. A., and Convey, P. (2015). Biological invasions in terrestrial Antarctica: what is the current status and can we respond? Biodiversity and Conservation, 24(5), 1031-1055. doi: https://doi.org/10.1007/s10531-015-0896-6
Pertierra, L. R., Baker, M., Howard, C., Vega, G. C., Olalla-Tarraga, M. A., and Scott, J. (2016). Assessing the invasive risk of two non-native Agrostis species on sub-Antarctic Macquarie Island. Polar Biology, 39(12), 2361-2371. doi: https://doi.org/10.1007/s00300-016-1912-3
Pollock, L. J., Tingley, R., Morris, W. K., Golding, N., O'Hara, R. B., Parris, K. M., Vesk, P. A., and McCarthy, M. A. (2014). Understanding co-occurrence by modelling species simultaneously with a Joint Species Distribution Model (JSDM). Methods in Ecology and Evolution, 5(5), 397-406. doi: https://doi.org/10.1111/2041-210X.12180
Richardson, D. M., Pyšek, P., Rejmánek, M., Barbour, M. G., Panetta, F. D., and West, C. J. (2000). Naturalization and invasion of alien plants: concepts and definitions. Diversity and Distributions, 6(2), 93-107. doi: https://doi.org/10.1046/j.1472-4642.2000.00083.x

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.