The mean trophic level of catches in world fisheries has generally declined over the 20th century, a phenomenon called "fishing down the food web" (Pauly et al. 1998). Several mechanisms have been proposed to explain this decline including the collapse of, or decline in, higher trophic level stocks leading to the inclusion of lower trophic level stocks in the fishery. Fishing down the food web may lead to a reduction in the resilience, i.e., the capacity to rebound from change, of the fished community, which is concerning given the necessity of resilience in the face of climate change. 

The practice of adaptive harvesting, which involves fishing stocks based on their availability, can also result in a reduction in the average trophic level of a fishery (Branch et al. 2010). Adaptive harvesting, similar to adaptive foraging, can affect the resilience of fisheries. Generally, adaptive foraging acts as a stabilizing force in communities (Valdovinos et al. 2010), however it is not clear how including harvesters as the adaptive foragers will affect the resilience of the system.

Tromeur and Loeuille (2023) analyze the effects of adaptively harvesting a trophic community. Using a system of ordinary differential equations representing a predator-prey model where both species are harvested, the researchers mathematically analyze the impact of increasing fishing effort and adaptive harvesting on the mean trophic level and resilience of the fished community. This is achieved by computing the equilibrium densities and equilibrium allocation of harvest effort.  In addition, the researchers numerically evaluate adaptive harvesting in a tri-trophic system (predator, prey, and resource). The study focuses on the effect of adaptively distributing harvest across trophic levels on the mean trophic level of catches, the propensity for regime shifts to occur, the ability to return to equilibrium after a disturbance, and the speed of this return. 

The results indicate that adaptive harvesting leads to a decline in the mean trophic level of catches, resulting in “fishing down the food web”. Furthermore, the study shows that adaptive harvesting may harm the overall resilience of the system. Similar results were observed numerically in a tri-trophic community.

While adaptive foraging is generally a stabilizing force on communities, the researchers found that adaptive harvesting can destabilize the harvested community. One of the key differences between adaptive foraging models and the model presented here, is that the harvesters do not exhibit population dynamics. This lack of a numerical response by the harvesters to decreasing population sizes of their stocks leads to regime shifts. The realism of a fishery that does not respond numerically to declining stock is debatable, however it is very likely that there will a least be significant delays due to social and economic barriers to leaving the fishery, that will lead to similar results.

This study is not unique in demonstrating the ability of adaptive harvesting to result in “fishing down the food web”. As pointed out by the researchers, the same results have been shown with several different model formulations (e.g., age and size structured models). Similarly, this study is not unique to showing that increasing adaptation speeds decreases the resilience of non-linear predator-prey systems by inducing oscillatory behaviours. Much of this can be explained by the destabilising effect of increasing interaction strengths on food webs (McCann et al. 1998). 

By employing a straightforward model, the researchers were able to demonstrate that adaptive harvesting, a common strategy employed by fishermen, can result in a decline in the average trophic level of catches, regime shifts, and reduced resilience in the fished community. While previous studies have observed some of these effects, the fact that the current study was able to capture them all with a simple model is notable. This modeling approach can offer insight into the role of human behavior on the complex dynamics observed in fisheries worldwide.

References

Branch, T. A., R. Watson, E. A. Fulton, S. Jennings, C. R. McGilliard, G. T. Pablico, D. Ricard, et al. 2010. The trophic fingerprint of marine fisheries. Nature 468:431–435. https://doi.org/10.1038/nature09528

Tromeur, E., and N. Loeuille. 2023. Effects of adaptive harvesting on fishing down processes and resilience changes in predator-prey and tritrophic systems. bioRxiv 290460, ver 5 peer-reviewed and recommended by PCI Ecology. https://doi.org/10.1101/290460

McCann, K., A. Hastings, and G.R. Huxel. 1998. Weak trophic interactions and the balance of nature. Nature 395: 794-798. https://doi.org/10.1038/27427

Pauly, D., V. Christensen, J. Dalsgaard, R. Froese, and F. Torres Jr. 1998. Fishing down marine food webs. Science 279:860–86. https://doi.org/10.1126/science.279.5352.860

Valdovinos, F.S., R. Ramos-Jiliberto, L. Garay-Naravez, P. Urbani, and J.A. Dunne. 2010. Consequences of adaptive behaviour for the structure and dynamics of food webs. Ecology Letters 13: 1546-1559. https://doi.org/10.1111/j.1461-0248.2010.01535.x

In a progressively human-dominated planet (Venter et al., 2016), the fate of many species will depend on the extent to which they can persist in anthropogenic landscapes. In Western Europe, where only small areas of primary habitat remain (e.g. Sabatini et al., 2018), semi-natural areas are crucial habitats to many native species, yet they are threatened by the expansion of human activities, including agricultural expansion and intensification (Rigal et al., 2023). 

A new study by Mallet and colleagues (Mallet et al., 2023) investigates the extent to which bird species in the Camargue region are able to use the margins of agricultural fields as substitutes for their preferred semi-natural habitats. Located in the delta of the Rhône River in Southern France, the Camargue is internationally recognized for its biodiversity value, classified as a Biosphere Reserve by UNESCO and as a Wetland of International Importance under the Ramsar Convention (IUCN & UN-WCMC, 2023). Mallet and colleagues tested three specific hypotheses: that grass strips (grassy field boundaries, including grassy tracks or dirt roads used for moving agricultural machinery) can function as substitute habitats for grassland species; that reed strips along drainage ditches (common in the rice paddy landscapes of the Camargue) can function as substitute habitats to wetland species; and that hedgerows can function as substitute habitats to species that favour woodland edges. They did so by measuring how the local abundances of 14 bird species (nine typical of forest edges, 3 of grasslands, and two of reedbeds) respond to increasing coverage of either the three types of field margins or of the three types of semi-natural habitat. 

This is an elegant study design, yet – as is often the case with real field data – results are not as simple as expected. Indeed, for most species (11 out of 14) local abundances did not increase significantly with the area of their supposed primary habitat, undermining the assumption that they are strongly associated with (or dependent on) those habitats. Among the three species that did respond positively to the area of their primary habitat, one (a forest edge species) responded positively but not significantly to the area of field margins (hedgerows), providing weak evidence to the habitat compensation hypothesis. For the other two (grassland and a wetland species), abundance responded even more strongly to the area of field margins (grass and reed strips, respectively) than to the primary habitat, suggesting that the field margins are not so much a substitute but valuable habitats in their own right. 

It would have been good conservation news if field margins were found to be suitable habitat substitutes to semi-natural habitats, or at least reasonable approximations, to most species. Given that these margins have functional roles in agricultural landscapes (marking boundaries, access areas, water drainage), they could constitute good win-win solutions for reconciling biodiversity conservation with agricultural production. Alas, the results are more complicated than that, with wide variation in species responses that could not have been predicted from presumed habitat affinities. These results illustrate the challenges of conservation practice in complex landscapes formed by mosaics of variable land use types. With species not necessarily falling neatly into habitat guilds, it becomes even more challenging to plan strategically how to manage landscapes to optimize their conservation. The results presented here suggest that species’ abundances may be responding to landscape variables not taken into account in the analyses, such as connectivity between habitat patches, or maybe positive and negative edge effects between land use types. That such uncertainties remain even in a well-studied region as the Camargue, and for such a well-studied taxon such as birds, only demonstrates the continued importance of rigorous field studies testing explicit hypotheses such as this one by Mallet and colleagues. 

References

IUCN, & UN-WCMC (2023). Protected Planet. Protected Planet. https://www.protectedplanet.net/en 

Mallet, P., Béchet, A., Sirami, C., Mesléard, F., Blanchon, T., Calatayud, F., Dagonet, T., Gaget, E., Leray, C., & Galewski, T. (2023). Field margins as substitute habitat for the conservation of birds in agricultural wetlands. bioRxiv, 2022.05.05.490780, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.05.05.490780 

Rigal, S., Dakos, V., Alonso, H., Auniņš, A., Benkő, Z., Brotons, L., Chodkiewicz, T., Chylarecki, P., de Carli, E., del Moral, J. C. et al. (2023). Farmland practices are driving bird population decline across Europe. Proceedings of the National Academy of Sciences, 120, e2216573120. https://doi.org/10.1073/pnas.2216573120 

Sabatini, F. M., Burrascano, S., Keeton, W. S., Levers, C., Lindner, M., Pötzschner, F., Verkerk, P. J., Bauhus, J., Buchwald, E., Chaskovsky, O., Debaive, N. et al. (2018). Where are Europe’s last primary forests? Diversity and Distributions, 24, 1426–1439. https://doi.org/10.1111/ddi.12778 

Venter, O., Sanderson, E. W., Magrach, A., Allan, J. R., Beher, J., Jones, K. R., Possingham, H. P., Laurance, W. F., Wood, P., Fekete, B. M., Levy, M. A., & Watson, J. E. M. (2016). Sixteen years of change in the global terrestrial human footprint and implications for biodiversity conservation. Nature Communications, 7, 12558. https://doi.org/10.1038/ncomms12558 

The spatial distribution of a species is driven by both biotic and abiotic factors that may change over time (Soberón & Nakamura, 2009; Paquette & Hargreaves, 2021).  Therefore, species ranges are dynamic, especially in humanized landscapes where changes occur at high speeds (Sirén & Morelli, 2020). The distribution of many species is being reduced because of human impacts; however, some species are expanding their distributions, even over their niche (Lustenhouwer & Parker, 2022). One of the factors that may lead to a geographic niche expansion is behavioral flexibility (Mikhalevich et al., 2017), but the mechanisms determining range expansion through behavioral changes are not fully understood. 

The PCI Ecology study by Summers et al. (2023) uses a very large database on the current and historic distribution of two species of grackles that have shown different trends in their distribution. The great-tailed grackle has largely expanded its range over the 20th century, while the range of the boat-tailed grackle has remained very similar. They take advantage of this differential response in the distribution of the two species and run several analyses to test whether it was a change in habitat availability, in the realized niche, in habitat connectivity or in in the other traits or conditions that previously limited the species range, what is driving the observed distribution of the species. The study finds a change in the niche of great-tailed grackle, consistent with the high behavioral flexibility of the species.

The two reviewers and I have seen a lot of value in this study because 1) it addresses a very timely question, especially in the current changing world; 2) it is a first step to better understanding if behavioral attributes may affect species’ ability to change their niche; 3) it contrasts the results using several complementary statistical analyses, reinforcing their conclusions; 4) it is based on the preregistration Logan et al (2021), and any deviations from it are carefully explained and justified in the text and 5) the limitations of the study have been carefully discussed. It remains to know if the boat-tailed grackle has more limited behavioral flexibility than the great-tailed grackle, further confirming the results of this study.
 
References

Logan CJ, McCune KB, Chen N, Lukas D (2021) Implementing a rapid geographic range expansion - the role of behavior and habitat changes. http://corinalogan.com/Preregistrations/gxpopbehaviorhabitat.html

Lustenhouwer N, Parker IM (2022) Beyond tracking climate: Niche shifts during native range expansion and their implications for novel invasions. Journal of Biogeography, 49, 1481–1493. https://doi.org/10.1111/jbi.14395

Mikhalevich I, Powell R, Logan C (2017) Is behavioural flexibility evidence of cognitive complexity? How evolution can inform comparative cognition. Interface Focus, 7, 20160121. https://doi.org/10.1098/rsfs.2016.0121

Paquette A, Hargreaves AL (2021) Biotic interactions are more often important at species’ warm versus cool range edges. Ecology Letters, 24, 2427–2438. https://doi.org/10.1111/ele.13864

Sirén APK, Morelli TL (2020) Interactive range-limit theory (iRLT): An extension for predicting range shifts. Journal of Animal Ecology, 89, 940–954. https://doi.org/10.1111/1365-2656.13150

Soberón J, Nakamura M (2009) Niches and distributional areas: Concepts, methods, and assumptions. Proceedings of the National Academy of Sciences, 106, 19644–19650. https://doi.org/10.1073/pnas.0901637106

Summers JT, Lukas D, Logan CJ, Chen N (2022) The role of climate change and niche shifts in divergent range dynamics of a sister-species pair. EcoEvoRxiv, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.32942/osf.io/879pe

​​​​​The persistence of populations facing the destruction of their habitat is a multifaceted question that has mobilized theoreticians and empiricists alike for decades. As an ecological question, persistence has been studied as the spatial rescue of populations via dispersal into remaining suitable habitats. The spatial aggregation of habitat destruction has been a key component of these studies, and it has been applied to the problem of coexistence by integrating competition-colonization tradeoffs. There is a rich ecological literature on this topic, both from theoretical and field studies (Fahrig 2003). The relationship between life-history strategies of species and their resilience to spatially structured habitat fragmentation is also an important component of conservation strategies through the management of land use, networks of protected areas, and the creation of corridors. In the context of environmental change, the ability of species to adapt to changes in landscape configuration and availability can be treated as an eco-evolutionary process by considering the possibility of evolutionary rescue (Heino and Hanski 2001; Bell 2017). However, eco-evolutionary dynamics considering spatially structured changes in landscapes and life-history tradeoffs remains an outstanding question. Finand et al. (2023) formulate the problem of persistence in fragmented landscapes over evolutionary time scales by studying models for the evolution of dispersal in relation to habitat fragmentation and spatial aggregation. Their simulations were conducted on a spatial grid where individuals can colonize suitable patch as a function of their competitive rank that decreases as a function of their (ii) dispersal distance trait. Simulations were run under fixed habitat fragmentation (proportion of unsuitable habitat) and aggregation, and with an explicit rate of habitat destruction to study evolutionary rescue.

Their results reveal a balance between the selection for high dispersal under increasing habitat fragmentation and selection for lower dispersal in response to habitat aggregation. This balance leads to the coexistence of polymorphic dispersal strategies in highly aggregated landscapes with low fragmentation where high dispersers inhabit aggregated habitats while low dispersers are found in isolated habitats. The authors then integrate the spatial rescue mechanism to the problem of evolutionary rescue in response to temporally increasing fragmentation. There they show how rapid evolution allows for evolutionary rescue through the evolution of high dispersal. They also show the limits to this evolutionary rescue to cases where both aggregation and fragmentation are not too high. Interestingly, habitat aggregation prevents evolutionary rescue by directly affecting the evolutionary potential of dispersal. The study is based on simple scenarios that ignore the complexity of relationships between dispersal, landscape properties, and species interactions. This simplicity is the strength of the study, revealing basic mechanisms that can now be tested against other life-history tradeoffs and species interactions. Finand et al. (2023) provide a novel foundation for the study of eco-evolutionary dynamics in metacommunities exposed to spatially structured habitat destruction. They point to important assumptions that must be made along the way, including the relationships between dispersal distance and fecundity (they assume a positive relationship), and the nature of life-history tradeoffs between dispersal rate and local competitive abilities. 

References

Bell, G. 2017. Evolutionary Rescue. Annual Review of Ecology, Evolution, and Systematics 48:605–627. https://doi.org/10.1146/annurev-ecolsys-110316-023011 
Fahrig, L. 2003. Effects of Habitat Fragmentation on Biodiversity. Annual Review of Ecology, Evolution, and Systematics 34:487–515. https://doi.org/10.2307/30033784 
Finand, B., T. Monnin, and N. Loeuille. 2023. Evolution of dispersal and the maintenance of fragmented metapopulations. bioRxiv, 2022.06.08.495260, ver. 3 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.06.08.495260 
Heino, M., and I. Hanski. 2001. Evolution of Migration Rate in a Spatially Realistic Metapopulation Model. The American Naturalist 157:495–511. https://doi.org/10.1086/319927

Temperature-size rule (TSR) is a phenomenon of plastic changes in body size in response to temperature, originally observed in more than 80% of ectothermic organisms representing various groups (Atkinson 1994). In particular, ectotherms were observed to grow faster and reach smaller size at higher temperature and grow slower and achieve larger size at lower temperature. This response has fired the imagination of researchers since its invention, due to its counterintuitive pattern from an evolutionary perspective (Berrigan and Charnov 1994). The main question to be resolved is: why do organisms grow fast and achieve smaller sizes under more favourable conditions (= relatively higher temperature), while they grow longer and achieve larger sizes under less favourable conditions (relatively lower temperature), if larger size means higher fitness, while longer development may be risky? 

This evolutionary conundrum still awaits an ultimate explanation (Angilletta Jr et al. 2004; Angilletta and Dunham 2003; Verberk et al. 2021). Although theoretical modelling has shown that such a growth pattern can be achieved as a response to temperature alone, with a specific combination of energetic parameters and external mortality (Kozłowski et al. 2004), it has been suggested that other temperature-dependent environmental variables may be the actual drivers of this pattern. One of the most frequently invoked variable is the relative oxygen availability in the environment (e.g., Atkinson et al. 2006; Audzijonyte et al. 2019; Verberk et al. 2021; Woods 1999), which decreases with temperature increase. Importantly, this effect is more pronounced in aquatic systems (Forster et al. 2012). However, other temperature-dependent parameters are also being examined in the context of their possible effect on TSR induction and strength.

Food availability is among the interfering factors in this regard. In aquatic systems, nutritional conditions are generally better at higher temperature, while a range of relatively mild thermal conditions is considered. However, there are no conclusive results so far on how nutritional conditions affect the plastic body size response to acute temperature changes. A study by Bazin et al. (2023) examined this particular issue, the effects of food and temperature on TSR, in medaka fish. An important value of the study was to relate the patterns found to fitness. This is a rare and highly desirable approach since evolutionary significance of any results cannot be reliably interpreted unless shown as expressed in light of fitness. 

The authors compared the body size of fish kept at 20°C and 30°C under conditions of food abundance or food restriction. The results showed that the TSR (smaller body size at 30°C compared to 20°C) was observed in both food treatments, but the effect was delayed during fish development under food restriction. Regarding the relevance to fitness, increased temperature resulted in more eggs laid but higher mortality, while food restriction increased survival but decreased the number of eggs laid in both thermal treatments. Overall, food restriction seemed to have a more severe effect on development at 20°C than at 30°C, contrary to the authors’ expectations. 

I found this result particularly interesting. One possible interpretation, also suggested by the authors, is that the relative oxygen availability, which was not controlled for in this study, could have affected this pattern. According to theoretical predictions confirmed in quite many empirical studies so far, oxygen restriction is more severe at higher temperatures. Perhaps for these particular two thermal treatments and in the case of the particular species studied, this restriction was more severe for organismal performance than the food restriction. This result is an example that all three variables, temperature, food and oxygen, should be taken into account in future studies if the interrelationship between them is to be understood in the context of TSR. It also shows that the reasons for growing smaller in warm may be different from those for growing larger in cold, as suggested, directly or indirectly, in some previous studies (Hessen et al. 2010; Leiva et al. 2019). 

Since medaka fish represent predatory vertebrates, the results of the study contribute to the issue of global warming effect on food webs, as the authors rightly point out. This is an important issue because the general decrease in the size or organisms in the aquatic environment with global warming is a fact (e.g., Daufresne et al. 2009), while the question of how this might affect entire communities is not trivial to resolve (Ohlberger 2013). 

REFERENCES

Angilletta Jr, M. J., T. D. Steury & M. W. Sears, 2004. Temperature, growth rate, and body size in ectotherms: fitting pieces of a life–history puzzle. Integrative and Comparative Biology 44:498-509. https://doi.org/10.1093/icb/44.6.498

Angilletta, M. J. & A. E. Dunham, 2003. The temperature-size rule in ectotherms: Simple evolutionary explanations may not be general. American Naturalist 162(3):332-342. https://doi.org/10.1086/377187

Atkinson, D., 1994. Temperature and organism size – a biological law for ectotherms. Advances in Ecological Research 25:1-58. https://doi.org/10.1016/S0065-2504(08)60212-3

Atkinson, D., S. A. Morley & R. N. Hughes, 2006. From cells to colonies: at what levels of body organization does the 'temperature-size rule' apply? Evolution & Development 8(2):202-214 https://doi.org/10.1111/j.1525-142X.2006.00090.x

Audzijonyte, A., D. R. Barneche, A. R. Baudron, J. Belmaker, T. D. Clark, C. T. Marshall, J. R. Morrongiello & I. van Rijn, 2019. Is oxygen limitation in warming waters a valid mechanism to explain decreased body sizes in aquatic ectotherms? Global Ecology and Biogeography 28(2):64-77 https://doi.org/10.1111/geb.12847

Bazin, S., Hemmer-Brepson, C., Logez, M., Sentis, A. & Daufresne, M. 2023. Distinct impacts of food restriction and warming on life history traits affect population fitness in vertebrate ectotherms. HAL, ver.3  peer-reviewed and recommended by PCI Ecology. https://hal.inrae.fr/hal-03738584v3

Berrigan, D. & E. L. Charnov, 1994. Reaction norms for age and size at maturity in response to temperature – a puzzle for life historians. Oikos 70:474-478. https://doi.org/10.2307/3545787

Daufresne, M., K. Lengfellner & U. Sommer, 2009. Global warming benefits the small in aquatic ecosystems. Proceedings of the National Academy of Sciences USA 106(31):12788-93 https://doi.org/10.1073/pnas.0902080106

Forster, J., A. G. Hirst & D. Atkinson, 2012. Warming-induced reductions in body size are greater in aquatic than terrestrial species. Proceedings of the National Academy of Sciences of the United States of America 109(47):19310-19314. https://doi.org/10.1073/pnas.1210460109

Hessen, D. O., P. D. Jeyasingh, M. Neiman & L. J. Weider, 2010. Genome streamlining and the elemental costs of growth. Trends in Ecology & Evolution 25(2):75-80. https://doi.org/10.1016/j.tree.2009.08.004

Kozłowski, J., M. Czarnoleski & M. Dańko, 2004. Can optimal resource allocation models explain why ectotherms grow larger in cold? Integrative and Comparative Biology 44(6):480-493. https://doi.org/10.1093/icb/44.6.480

Leiva, F. P., P. Calosi & W. C. E. P. Verberk, 2019. Scaling of thermal tolerance with body mass and genome size in ectotherms: a comparison between water- and air-breathers. Philosophical Transactions of the Royal Society B 374:20190035. https://doi.org/10.1098/rstb.2019.0035

Ohlberger, J., 2013. Climate warming and ectotherm body szie - from individual physiology to community ecology. Functional Ecology 27:991-1001. https://doi.org/10.1111/1365-2435.12098

Verberk, W. C. E. P., D. Atkinson, K. N. Hoefnagel, A. G. Hirst, C. R. Horne & H. Siepel, 2021. Shrinking body sizes in response to warming: explanations for the temperature-size rule with special emphasis on the role of oxygen. Biological Reviews 96:247-268. https://doi.org/10.1111/brv.12653

Woods, H. A., 1999. Egg-mass size and cell size: effects of temperature on oxygen distribution. American Zoologist 39:244-252. https://doi.org/10.1093/icb/39.2.244