Abstract
The function of holding territories is primarily to have access to resources like food and mates. However, it is costly in terms of energy and time investment. Solitary-living, territorial species are known to reduce these costs by being more aggressive towards unfamiliar strangers and less aggressive towards neighbors. However, in social, territorial species, neighbors can impose a greater threat than strangers. We tested whether the highly social Asiatic wild dogs/dholes (Cuon alpinus) exhibit the “nasty neighbor” or the “dear enemy” phenomena in Tadoba Andhari Tiger Reserve (TATR), Maharashtra, India. We conducted scat translocation experiments where we presented fresh scats collected from unique donor groups to a resident dhole group and tested the type and the intensity of behavioral response (duration) to the stimulus. Dholes responded differentially to the two treatments suggesting they exhibit neighbor-stranger discrimination. Overall, strangers elicited a stronger response with longer duration and larger packs were less likely to respond to the stimulus than smaller packs. Differences found between categories of dhole scent marks establish the importance of olfactory communication, especially “counter-marking” in the species. Within recipient packs, individual status affected the response to trials wherein the alpha pair reacted more intensively to strangers than others. Our study provides experimental evidence to demonstrate that dholes exhibit the “dear enemy” phenomenon.
Significance statement
Animals defend territories from other members of their own species, but intrusions are commonplace in the wild. Different intruders may pose different levels of threats, and hence, intruders are treated differentially to minimize the energetic costs of territorial defense. In some animals, neighbors with well-established territories may become less aggressive towards each other. This is known as the dear enemy effect. By contrast, at times neighbors may represent a greater threat than strangers which is known as the “nasty neighbor” effect. We experimentally show that dholes exhibit the dear enemy phenomenon by responding more intensively to strangers than familiar neighbors. We show how response varied based on hierarchy in a pack as well as the pack sizes. Furthermore, we found that, both in core as well as buffer areas of their own territory, this relationship was consistent.






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All data generated or analyzed during this study are included in this published article [and its supplementary information files].
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The R Code has been attached as a supplementary file.
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Acknowledgements
We would like to thank Ms. Akanksha Saxena, Mr. Stephen Salazar, and Dr. Sahas Barve for their valuable inputs in statistical analysis and Mr. Nakul Markandey for his cartographic inputs. We are extremely grateful to all the reviewers and editors for recognizing the value of our study and patiently providing clear and elaborate inputs to improve the manuscript. We would also like to thank our field assistants, drivers, guides, naturalists, and tourists who helped us immensely in tracking dhole packs in the field.
Funding
This work was supported by Maharashtra Forest Department (MFD) (Permit No. D-22(8)/WL/Research/CT-722/(12–13)/2934/2013) and the National Tiger Conservation Authority (NTCA) (Letter No. 1–30/2003-PT(Part-I)).
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All authors (P. G., P. N., B. H.) contributed to the study conception and design. Material preparation, data collection, and analysis were performed by P. G. The first draft of the manuscript was written by P. G. and all authors (P. G., P. N., B. H.) commented on all versions of the manuscript. All authors read and approved the final manuscript.
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The authors obtained a Dhole capture permit (MFD-SPP-12/05.11.2016) for the radio-collared dholes in the study. All applicable international, national, and/or institutional guidelines for the use of animals were followed.
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The authors declare no competing interests.
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Communicated by T. Stankowich.
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Supplementary Information
Below is the link to the electronic supplementary material.
Supplementary 1
Excel sheet of the data used for analysis from experiment trials (XLS 44 kb)
Supplementary 2
R code for used for analysis (R 7 kb)
Supplementary 3
Map depicting the study area with dhole home ranges and locations of the stranger scats (Tadoba Andhari Tiger Reserve, Umred Karhandla Wildlife Sanctuary, Pench Tiger Reserve). (JPG 23157 kb)

Supplementary 4
Predicted intensity of response showing a gradual decrease in response as pack size increases from scat translocation experiments on dholes, Tadoba Andhari Tiger Reserve, India from model output of the top model (GLMM1). Line denotes mean predicted intensity values, while ribbon (shaded areas) represent 95% confidence intervals (PNG 67 kb)

Supplementary 5
Scaled beta co-efficient estimates (dots) with 95% confidence intervals (whiskers) of factors affecting the response of dholes to scat translocation experiments in Tadoba Andhari Tiger Reserve, Maharashtra, India. Confidence intervals overlapping 0 (dashed line) indicate no significant effect. (PNG 127 kb)
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Ghaskadbi, P., Nigam, P. & Habib, B. Stranger Danger: Differential response to strangers and neighbors by a social carnivore, the Asiatic wild dog (Cuon alpinus). Behav Ecol Sociobiol 76, 86 (2022). https://doi.org/10.1007/s00265-022-03188-4
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DOI: https://doi.org/10.1007/s00265-022-03188-4