Blue tits surviving in an ever-changing world
Identifying drivers of spatio-temporal variation in survival in four blue tit populations
How long individuals live has a large influence on a number of biological processes, both for the individuals themselves as well as for the populations they live in. For a given species, survival is often summarized in curves showing the probability to survive from one age to the next. However, these curves often hide a large amount of variation in survival. Variation can occur from chance, or if individuals have different genotypes or phenotypes that can influence how long they might live, or if environmental conditions are not the same across time or space. Such spatiotemporal variations in the conditions that individuals experience can lead to complex patterns of evolution (Kokko et al. 2017) but because of the difficulties to obtain the relevant data they have not been studied much in natural populations.
In this manuscript, Bastianelli and colleagues (2021) identify which environmental and population conditions are associated with variation in annual survival of blue tits. The analyses are based on an impressive dataset, tracking a total of almost 5500 adults in four populations studied for at least 19 years. The authors describe two core results. First, average annual survival is lower in deciduous forests compared to evergreen forests. The differences in average annual survival between the forest types link with previously described differences, with individuals having larger clutches (Charmantier et al. 2016) and higher aggression (Dubuc-Messier et al. 2017) in the populations where adult survival is lower. Second, there are huge fluctuations from one year to the next in the percentage of individuals surviving which occur similarly in all populations. Even though survival covaried across the four populations, this variation was not associated with any of the local or global climate indices the authors investigated.
Studies like these are fundamental to our understanding of population change. They are important from an applied side as they can reveal the sustainability of populations and inform potential management options. On a basic research side, they reveal how evolution operates in populations. Theoretical studies predict that individuals are often not adapted to average conditions they experience, but either selected to balance the extremes they encounter or to make the best during harsh conditions when it really matters (Lewontin & Cohen 1969).
This study also opens the door to new research, highlighting that demographic studies should pay attention to variation in survival and other life history traits. For blue tits specifically, the study shows that in order to understand the demography of populations we need a better mechanistic understanding of the environmental and physiological pressures influencing whether individuals die or not to make predictions whether and how climate or other ecological effects shape variation in survival.
Bastianelli O, Robert A, Doutrelant C, Franceschi C de, Giovannini P, Charmantier A (2021) Identifying drivers of spatio-temporal variation in survival in four blue tit populations. bioRxiv, 2021.01.28.428563, ver. 4 peer-reviewed and recommended by Peer community in Ecology. https://doi.org/10.1101/2021.01.28.428563
Charmantier A, Doutrelant C, Dubuc-Messier G, Fargevieille A, Szulkin M (2016) Mediterranean blue tits as a case study of local adaptation. Evolutionary Applications, 9, 135–152. https://doi.org/10.1111/eva.12282
Dubuc-Messier G, Réale D, Perret P, Charmantier A (2017) Environmental heterogeneity and population differences in blue tits personality traits. Behavioral Ecology, 28, 448–459. https://doi.org/10.1093/beheco/arw148
Kokko H, Chaturvedi A, Croll D, Fischer MC, Guillaume F, Karrenberg S, Kerr B, Rolshausen G, Stapley J (2017) Can Evolution Supply What Ecology Demands? Trends in Ecology & Evolution, 32, 187–197. https://doi.org/10.1016/j.tree.2016.12.005
Lewontin RC, Cohen D (1969) On Population Growth in a Randomly Varying Environment. Proceedings of the National Academy of Sciences, 62, 1056–1060. https://doi.org/10.1073/pnas.62.4.1056
Dieter Lukas (2021) Blue tits surviving in an ever-changing world. Peer Community in Ecology, 100085. 10.24072/pci.ecology.100085
Revision round #218 May 2021
Decision round #2
Thank you for addressing all the comments of the reviewers and me in such detail. The manuscript now contains additional details helpful to fully appreciate the inferences your study provides. I only have a few, very minor suggestions for changes that should be easy to address so I can recommend this study.
Comparison with previous study by Grosbois and colleagues:
In the introduction, there is a bit of a jump from the pace-of-life arguments (paragraph 105-132) to the study by Grosbois (lines 133-143) back to your framing around the pace-of-life (paragraph 144-150). I think it would help to see more clearly how the study by Grosbois et al fits within the frame of your study. Did Grosbois and colleagues also look at factors related to the pace-of-life arguments (e.g. habitat type) or is this framing one of the aspects where your study advances beyond those previous analyses, but not find any differences? If they only focused on climatic variables, I think you could move the description of this study to the earlier part of the introduction where you discuss climate (integrate lines 133-143 with/after the text in the paragraph starting in line 88).
In the final paragraph of the introduction, you state that the main aim of this study is to determine "a difference in adult survival between habitat types". Accordingly, I think it would be helpful to already mention the relevant result in the first paragraph of the discussion. When you discuss these results (line 660ff), you could also link to the broader literature again to show how your set of findings (including your earlier studies) provide insights beyond these populations (see e.g. Galipaud, M., & Kokko, H. (2020). Adaptation and plasticity in life-history theory: How to derive predictions. Evolution and Human Behavior, 41(6), 493-501.)
There are some minor typos you might want to fix for the final version. I focused on the new and revised parts of the manuscript, so you might want go through the whole manuscript for another check:
Line 103: first letter not capitalised: if large scale climatic factors
Line 114: missing word: such [as] pair fidelity
Line 117: missing reference: Nord & Nilsson 2016 Bio Let.
Table 4: final R2 value still has additional values after the decimal point
Line 380: I don't think you need the reference for U-CARE here in the results because you are already provide it in the methods
Line 431: missing space: quadratic effect
Line 518: change phrase: stronger dataset instead of strongest dataset
Line 641: change word: I think you mean provisioning rather than provisional
Revision round #101 Mar 2021
Decision round #1
In particular, I agree with both reviewers that more detail should be provided in the manuscript to explain the potential links between the factors you investigate and survival. I think this will make the manuscript clearer and help readers to see how your findings might transfer to other populations or other species. From my perspective, as someone who does not know a lot about your study species of blue tit, I think it could also help to have a paragraph in the introduction that briefly explains some key parameters of the species as they relate to survival (e.g. seasonal breeder, non-migratory but swarm forming in the winter?, when do individuals of which sex disperse how far?, is the highest mortality in the winter, during incubation in the nest, during feeding from predation?, what is the maximum longevity?). Having this information might help to explain better in the introduction why you chose these various factors and how they could potentially shape survival in the different populations. In turn, I think it would help if the introduction is focused only on those factors that are central to your study: for example, the abstract mentions boldness which has no direct relation to your study setup.
I have an additional comment about the presentation of the approach and results in the tables and figures. There is a lot of information, but it is sometimes difficult to understand what information there is to extract for someone not as familiar with the setup as you.
Table 1: Is this supposed to be a supplement? It currently appears at the end of the manuscript, and you later focus on the more specific models rather than presenting the full breadth of results from all the models.
Table 3: There are some formatting issues in the pdf. I also wonder whether you want to set the number of digits after the decimal points (e.g. have four digits) to make comparisons easier.
Table 5: I think it would be interesting to have your results here as well for comparison. You could also add a column that explains whether each site is deciduous or evergreen.
Figure 1: Could you maybe include photographs that illustrate the forest conditions at the four sites?
Figure 2: The label for the fourth population E-Muro in red is missing. Maybe you could combine this figure with the additional illustrations of year-by-year variation shown in Figure 3 and Figure 4a.
Figure 3/4a: As mentioned above, the year-by-year trends on their own are not particularly informative. I think they gain their information from the comparison with the other trends, to compare the extent of variation in each and whether yearly fluctuations might align.