Spatiotemporal Population Growth Patterns and Interactions Among Sympatric Central European Mesocarnivores

Round 1

Reviewer 1 Report

Comments and Suggestions for Authors

Dear Authors,

The manuscript certainly touches on an interesting topic. Interrelationships of sympatric species of animals are always of interest. But the authors were not able to properly explain the selection of three objects. The spectrum of nutrition of these carnivorous mammals in Hungary should be given to assess their role and possible influence on each other. In this form, the manuscript cannot be published. An addition must be added to the title of the manuscript. The text of the manuscript is mixed up in different chapters and should be moved to the appropriate chapters. The comparative part of the discussion needs expansion and citation of additional sources of literature on other types of medium-sized carnivorous mammals. You should focus on writing the conclusion of the manuscript on the basis of the conducted hypothesis. The authors obtained interesting results, presented them, and based on the proposed hypothesis, they should be disclosed in the conclusions of the manuscript. After eliminating all noted manuscripts, the manuscript can be reviewed again.

Comments for author File: Comments.pdf

Author Response

Please see the attachment.

Author Response File: Author Response.docx

Reviewer 2 Report

Comments and Suggestions for Authors

This manuscript represents an impressive and substantial contribution to the field of mesocarnivore ecology and management in Europe. The authors have undertaken a formidable task by analyzing a long-term, national-scale dataset to address timely and ecologically significant questions regarding the interactions between expanding golden jackal populations and native mesocarnivores. The study is well-motivated, the analytical framework is sophisticated, and the conclusions are generally well-supported by the data. The use of Generalized Additive Models (GAMs) to tease apart spatiotemporal trends and species interactions is commendable. The writing is clear, and the figures effectively support the narrative. This work is of high relevance for both theoretical ecology and applied wildlife management. The following comments are offered in the spirit of constructive critique to help further strengthen an already excellent manuscript before publication.

Lines 30-31: I think that you should add these two important and recent references as examples to support your sentence: ”Species coexistence has been an important topic in community ecology, as these studies are essential for understanding the diversity and structure of ecological communities.”. I would like to suggest:

Bosso, L., Fichera, G., Mucedda, M., Pidinchedda, E., Veith, M., Smeraldo, S., ... & Ancillotto, L. (2025). Island Life and Interspecific Dynamics Influence Body Size, Distribution and Ecological Niche of Long‐Eared Bats. Journal of Biogeography, e70071.

Søndergaard, S. A., Fløjgaard, C., Ejrnæs, R., & Svenning, J. C. (2025). Shifting baselines and the forgotten giants: integrating megafauna into plant community ecology. Oikos, 2025(5), e11134.

Lines 35-36 I think that you should add these two important and recent references as examples to support your sentence: ” This partitioning of resources can be along the temporal, trophic, and spatial dimension.”. I would like to suggest:

Salinas-Ramos, V. B., Ancillotto, L., Cistrone, L., Nastasi, C., Bosso, L., Smeraldo, S., ... & Russo, D. (2021). Artificial illumination influences niche segregation in bats. Environmental Pollution, 284, 117187.

Říha, M., Vejřík, L., Rabaneda-Bueno, R., Jarić, I., Prchalová, M., Vejříková, I., ... & Peterka, J. (2025). Ecosystem, spatial and trophic dimensions of niche partitioning among freshwater fish predators. Movement Ecology, 13(1), 36.

Lines 42-46: The final sentence of the introduction is cut off ("lowering foraging efficiency, causing direct mortality from killing, and through inverse density-dependency [8], thereby impacting overall intraguild population dynamics."). This requires completion for logical flow.

Lines 54-58: The statement "Therefore, it is expected that the arrival of the jackal has likely made a profound impact..." is somewhat vague and could be sharpened. It would be beneficial to specify the hypothesized mechanisms of impact (e.g., competitive suppression, spatial exclusion, trophic shifts) based on the literature cited (e.g., Lanszki et al., 2006; Scheinin et al., 2006), rather than a general "profound impact."

Lines 62-66: The justification for the study is well-stated, but it could be strengthened by more directly contrasting the limitations of presence-only camera data (which often informs co-occurrence models) with the potential and limitations of hunting bag data (which provides an index of abundance over a much broader scale). This would better frame the unique niche this study fills.

Lines 68-73: The hypotheses are clear but could be more explicitly linked to underlying ecological theory. For instance, the expected negative fox-jackal correlation is rooted in interference competition, while the expected positive fox-badger correlation may stem from commensalism/facilitation or shared responses to environmental drivers. Making these theoretical links explicit would deepen the introduction.

Lines 77-84: A more detailed justification is needed for why hunting bag data is a reliable index of population growth rates for these species in Hungary, beyond the citation of national framework reducing bias. Specifically, discuss potential confounding factors: Are hunting quotas or effort directly linked to population size? Could cultural or economic shifts in hunting practices over 27 years create non-biological trends? A brief discussion on the validation of these data, perhaps referencing previous national studies that have successfully used this database (e.g., Csányi et al., 2010; Bijl et al., 2024), is crucial.

Lines 85-92: The calculation of λ and *r* per grid cell is standard. However, a potential issue arises with cells where N_t = 0. How were these handled? Were they excluded, set to a minimum value, or did the calculation of *r* become undefined? This must be clarified, as it affects the spatial coverage of the correlation and GAM analyses. The rationale for also calculating the regression slope of hunting bags is not entirely clear, as it seems to represent a similar long-term trend to the average *r*. The authors should justify why both metrics are necessary and how they complement each other, or consider focusing on one to avoid redundancy.

Lines 93-102: The choice of a 5-year "short-term" period is arbitrary. A more principled approach might be to use a moving window analysis or to select periods based on biological/management milestones (e.g., pre- and post-jackal saturation in a region). The justification "because then the European badger and golden jackal occupied the majority of the country" is circular if occupancy is an outcome. Consider refining this justification.

Lines 103-124: The population trend GAM is well-specified. However, the choice of the Tweedie distribution power parameter (p=1.453) should be justified. Was this selected via model selection (e.g., using gam.check or AIC)? Stating how this parameter was determined is important for reproducibility. For the species-interaction GAMs, using the annual growth rate *r* as the response variable is interesting. However, this approach assumes that the growth rate in year *t* is influenced by the growth rates of competitors in the same year *t*. Ecologically, competitive effects might be lagged (e.g., competition affecting reproduction or survival manifests in the next year's growth). The authors should discuss this assumption and consider if models with a 1-year lag might be more appropriate or should be tested as an alternative. The models control for spatiotemporal autocorrelation via smooths, which is good. However, they do not explicitly account for potential confounding environmental covariates (e.g., habitat type, prey availability, climate) that could drive synchronous growth rates among species, leading to spurious "positive associations." While the broad scale makes including such covariates challenging, this is a significant limitation that must be explicitly acknowledged in the discussion.Lines 125-133: The results from the population trend GAM are clearly reported. However, for spatial smooths, reporting only the edf and p-value is insufficient. A qualitative description of the key spatial patterns (e.g., "higher predicted bags in the southwest for jackal") would help the reader interpret the model before viewing the figures.

Lines 145-152 & Figures 4-6: The description of the spatial patterns in the text is somewhat difficult to follow without constant reference to the figures. The text should provide a more concise verbal summary of the main spatial findings (e.g., "Jackal growth was strongest in the southwest, coinciding with areas where its hunting bag now exceeds that of the red fox (Fig. 4), while fox and badger trends were more geographically uniform").

Lines 160-179: The consistent report of weak positive correlations (r < 0.3) is clear. However, the meaning of "significant" with very large sample sizes (thousands of grid-cell years) needs caution. With N this large, even trivial correlations become statistically significant. The authors should emphasize the ecological insignificance of these very weak correlations in the text, not just report the p-values. Effect size is key here. The exception (weak negative jackal-fox correlation based on slopes) is interesting but not discussed in the text. Why might long-term trends show a slight negative relationship while annual growth rates show a weak positive one? This discrepancy warrants a sentence of interpretation.

Lines 180-194 & 195-210: The low explanatory power (adj. R² = 0.04 to 0.16) of these models is a critical result that needs more emphasis. It should be stated upfront in the results narrative that while associations are statistically significant, they explain very little of the variance in growth rates. This strongly supports the main conclusion of "no substantial competitive interference." The comparison of effect sizes (β) between national and regional models is insightful. However, the text states "the effect size became weaker compared to the national-scale model." This should be quantified for the key relationships (e.g., β for fox on badger changed from 0.51 to 0.59, which is actually stronger. Clarify which specific coefficients became weaker).

Lines 214-225: The discussion appropriately highlights spatial heterogeneity and temporal partitioning as mechanisms for coexistence. However, the argument would be stronger if it more directly engaged with the "niche variation hypothesis" and cited foundational works (e.g., Schoener's dimensions of niche partitioning). The sentence on "periodic behaviours" is vague; clarify if this refers to diel or seasonal activity shifts.

Lines 226-240: The paragraph on context-dependence is excellent, citing relevant case studies. To improve synthesis, the authors could explicitly frame these within a conceptual model where the outcome of interactions (facilitation, coexistence, competition) depends on an axis of, for example, resource abundance/predictability and population density.

Lines 241-250: This is a crucial point. The discussion on synchronous management as a driver of positive correlations is perhaps the most parsimonious explanation for the observed patterns and should be elevated. Consider moving this paragraph earlier in the discussion. It directly addresses the major alternative hypothesis to the "coexistence via niche partitioning" narrative.

Lines 246-247: I think that you should add these two important and recent references as examples to support your sentence: ”Overall, these interactions suggest that coexistence and competition are context-dependent, ranging from facilitation to competition depending on local conditions and behavioural flexibility.”. I would like to suggest:

Fraissinet, M., Ancillotto, L., Migliozzi, A., Capasso, S., Bosso, L., Chamberlain, D. E., & Russo, D. (2023). Responses of avian assemblages to spatiotemporal landscape dynamics in urban ecosystems. Landscape Ecology, 38(1), 293-305.

Bonte, D., Keith, S., & Fronhofer, E. A. (2024). Species interactions and eco-evolutionary dynamics of dispersal: the diversity dependence of dispersal. Philosophical Transactions B, 379(1907), 20230125.

Lines 251-260: The conclusions are sound. The call for integrating fine-scale behavioral data is well-placed. To make it more actionable, suggest specific integrative frameworks, such as "Integrated Population Models" that could combine hunting bag data with telemetry-derived survival estimates or camera-trap occupancy data.

Author Response

Please see the attachment.

Author Response File: Author Response.docx

Reviewer 3 Report

Comments and Suggestions for Authors

The authors present an informative analysis of long-term population trends and broad-scale coexistence patterns among European badgers, red foxes, and golden jackals in Hungary. They used nearly three decades of hunting bag data as proxies for occurrence and abundance. The central finding—that all three mesocarnivores show increasing population trends and largely positive, albeit weak to moderate, correlations in annual growth rates. They conclude that there is no broad-scale evidence of competitive suppression of badgers or foxes by the expanding golden jackal, and that these species can coexist at national and regional scales.

The abstract, introduction, and main text frame the study well within the broader literature on carnivore guilds, niche partitioning, and the rapid expansion of golden jackals in Europe. The statistical methodology is appropriate. ​

The discussion of limitations around hunting bags is reasonable but could be made more explicit. The authors acknowledge that hunting bags are affected by variation in effort, and they argue that Hungary’s mandatory, systematic reporting framework reduces bias; also noting that accounting for the number of hunters did not substantially change the results. However, the potential influence of “hunter affinities” (preferences for or against particular species) and shifting management priorities (e.g., predator control focus, legal status changes) deserves more direct treatment, because these factors can decouple harvest numbers from underlying population dynamics and may confound inferences about interspecific relationships. At present, these issues are mentioned but not fully integrated into the interpretation of growth rates and correlations.

Similarly, while the manuscript does discuss the absence of a dedicated limitations section, several key constraints emerge from the methods and discussion that should also be forwarded more systematically. First, reliance on hunting bag data as a proxy for abundance is a structural limitation, as these data are sensitive to effort, regulation, and management changes, not just population size. Second, the models include smooth terms for time and space but do not incorporate explicit environmental covariates such as land cover, prey availability, or climate variables, which likely account for a substantial portion of the unexplained variation in growth rates. Third, the use of annual data means that fine-scale temporal processes—such as seasonal or diel partitioning—cannot be detected, even though these mechanisms are mentioned in the discussion as plausible drivers of coexistence. ​

The interpretation that positive correlations in growth rates reflect apparent coexistence is plausible but remains correlative. The low explanatory power of interaction models, combined with the possibility of synchronous drivers (e.g., shared prey increases, changes in management, landscape changes) means that the positive associations could be driven by common external factors rather than direct facilitation or the absence of competition. The authors are careful on this point, but a more explicit statement about the correlative nature of the findings and potential confounders—such as coordinated predator control campaigns affecting all three species, will strengthen the argument and be more convincing to the reader.

The manuscript already points toward several important avenues for improvement and future work, and emphasizing these would help transform a broad-scale analysis into a more comprehensive framework for mesocarnivore coexistence. Integrating hunter effort or presenting side-by-side models with and without effort as a covariate would enhance confidence in the robustness of the inferences drawn from hunting bags. Incorporating environmental or resource covariates, and exploring lagged effects of competitor abundance (e.g., previous-year growth of jackals on current-year fox growth), could account for some of the unexplained deviance and reveal delayed or masked competitive effects. Methodologically, the authors might also consider multi-species hierarchical models that jointly estimate species’ responses while sharing spatial and temporal random effects.

 Finally, the discussion of coexistence mechanisms could be better anchored to specific ecological hypotheses. Existing work on diet overlap and spatial ecology of jackals and foxes in the region indicates shared reliance on small mammals and anthropogenic resources, but also potential partitioning in habitat and foraging strategies. Explicitly linking the observed weak positive associations in growth rates to these trophic and spatial patterns, while explaining how fine-scale avoidance or interference could be averaged out at regional scales, would provide a tighter connection between statistical results and ecological theory.

Overall, the study makes a valuable contribution by documenting broad-scale, long-term trends and apparent coexistence in a rapidly changing mesocarnivore guild, and with clearer articulation of limitations and more explicit integration of environmental and behavioral dimensions, it could serve as a strong template for similar analyses elsewhere.

 

Author Response

Please see the attachment.

Author Response File: Author Response.docx

Round 2

Reviewer 1 Report

Comments and Suggestions for Authors

Dear Authors,

The manuscript has been improved. This is undeniable. I hope the authors will avoid making mistakes in future studies and publications. I am somewhat concerned that the study results contain no evidence that jackals preyed on foxes or badgers. However, further research may uncover new facts. Overall, everything else is unquestionable. The review of the study results was chosen appropriately, as were the statistical methods used for its analysis. The article has taken into account the comments on the methodology. The analysis for each chapter is sufficient and does not raise objections. The results are taken into account with previous studies by other authors. I recommend this article for the journal Life.

Author Response

Thank you for taking the time to review our manuscript.

Reviewer 2 Report

Comments and Suggestions for Authors

Dear Authors,

I would like to thank you for submitting the revised version of your manuscript. I am pleased to see that you have carefully considered several of the points raised during the previous round of review, and that some important issues have been appropriately addressed.

However, after a thorough re-evaluation of the revised manuscript, I note that there are still several aspects that require further clarification, improvement, or correction before the work can be considered satisfactory. These remaining issues concern methodological and interpretative elements, as well as clarity and presentation.

In addition, I must stress a very important point: the manuscript must be thoroughly revised by a native English speaker or a professional language editing service. The current version contains numerous grammatical errors, awkward phrasing, and syntactic inaccuracies that significantly affect readability. As a result, the text is often difficult to follow and lacks fluency, which undermines the overall quality and clarity of the work.

Please find below a detailed list of the points that still need to be addressed.

Lines 31-32: I think that you should add these two relevant and recent references as examples to support your sentence: ”Species coexistence has been an important topic in community ecology, as these studies are essential for understanding the diversity and structure of ecological communities.”. I would like to suggest:

Bosso, L., Fichera, G., Mucedda, M., Pidinchedda, E., Veith, M., Smeraldo, S., ... & Ancillotto, L. (2025). Island Life and Interspecific Dynamics Influence Body Size, Distribution and Ecological Niche of Long‐Eared Bats. Journal of Biogeography, e70071.

Wang, M., Pan, X., Yue, Z., Deng, R., Li, Z., & Wang, J. (2025). Seasonal variation drives species coexistence and community succession in microbial communities of stratified acidic pit lakes. Journal of Environmental Management, 381, 125177.

Lines 34-36 I think that you should add these two relevant and recent references as examples to support your sentence: ” Consequently, sympatric species can develop different strategies for using resources, resulting in niche differentiation and divergence in order to reduce interspecific competition”. I would like to suggest:

Salinas-Ramos, V. B., Ancillotto, L., Cistrone, L., Nastasi, C., Bosso, L., Smeraldo, S., ... & Russo, D. (2021). Artificial illumination influences niche segregation in bats. Environmental Pollution, 284, 117187.

Říha, M., Vejřík, L., Rabaneda-Bueno, R., Jarić, I., Prchalová, M., Vejříková, I., ... & Peterka, J. (2025). Ecosystem, spatial and trophic dimensions of niche partitioning among freshwater fish predators. Movement Ecology, 13(1), 36.

Lines 69-76: What are your predictions? Please add the hypothesis in bullet format.

Line 82: Why did you choose the cell grid? Please explain the reasons.

Lines 104 – 158: All the parameters added in the formula must be correctly and deeply explained in the main text.

Line 140: It is generalised additive model (GAM).

Line 157: All the R codes used in this study must be added in the supplementary materials (Well commented!).

Line 189: Please use colorblind (daltonic) colour friendly for the bars in this figure.

Line 226: Please add the north symbol and the scale in the map. Please move this figure in the supplementary materials. Please use colorblind (daltonic) colour friendly for the bars in this figure.

Line 230: Please change the colour scale for the fox and jackal. I don’t see nothing with this colour scale. To add the scale and the north symbol in the map. Please use colorblind (daltonic) colour friendly for the bars in this figure.

Line 233: To add the scale and the north symbol in the map.

Lines 290 – 400: Please add two paragraphs in your discussion: 1) limitation of your study; 2) management recommendation.

Line 419 – 421: I think that you should add this relevant and recent references as examples to support your sentence: ” Importantly, field studies should not only focus on cases of intraguild competition but also on its absence, especially since the coexistence of species within a guild is necessary for biodiversity to exist in the first place.”. I would like to suggest:

Fraissinet, M., Ancillotto, L., Migliozzi, A., Capasso, S., Bosso, L., Chamberlain, D. E., & Russo, D. (2023). Responses of avian assemblages to spatiotemporal landscape dynamics in urban ecosystems. Landscape Ecology, 38(1), 293-305.

Comments on the Quality of English Language

In addition, I must stress a very important point: the manuscript must be thoroughly revised by a native English speaker or a professional language editing service. The current version contains numerous grammatical errors, awkward phrasing, and syntactic inaccuracies that significantly affect readability. As a result, the text is often difficult to follow and lacks fluency, which undermines the overall quality and clarity of the work.

Author Response

Please see the attachment.

Author Response File: Author Response.docx

Reviewer 3 Report

Comments and Suggestions for Authors

I am satisfied with the revision.

Author Response

Thank you for taking the time to review our manuscript.