Laura Cunha Silva1,2*
Constanza Fellenberg3
Jerónimo Freudenthal4
Harish Kumar Tiwari5,6,7†
Salome Dürr1†- 1Vetsuisse Faculty, Veterinary Public Health Institute, University of Bern, Bern, Switzerland
- 2Graduate School for Cellular and Biomedical Sciences, University of Bern, Bern, Switzerland
- 3The University of Queensland, Brisbane, QLD, Australia
- 4Faculdade de Medicina Veterinária, Universidade de Lisboa, Lisbon, Portugal
- 5Indian Institute of Technology Guwahati, Guwahati, India
- 6Sydney Medical School, Faculty of Medicine and Health, University of Sydney, Sydney, NSW, Australia
- 7DBT Wellcome Trust India Alliance, Hyderabad, India
Understanding free-roaming dog (FRD) demographics and movement patterns is essential for effective rabies control interventions, such as mass dog vaccinations (MDV). This review assesses published studies on FRD movement and enumeration to assess existing knowledge. A scoping review was conducted following PRISMA guidelines. Three databases, namely, Embase, Scopus, and Web of Science databases, were searched for publications between 2012 and 2024. A total of 2,167 articles were screened through successive filtration process to select a final corpus of 52 publications. The studies were predominantly from India (n = 8), Brazil (n = 6), Indonesia (n = 5), Guatemala (n = 5) and Chad (n = 5) and mostly investigated FRD population size. Several techniques were used for FRD enumeration, with photographic mark capture-recapture being the most common. Most FRD movement studies focused on home ranges, influenced by the technique and population size. In many studies, advantages and disadvantages of the techniques employed remained unreported, leaving a scope for misleading conclusions when comparing the methods used. The review highlights significant research gaps in FRD movement and population studies in rabies-endemic regions, which are often overlooked in rabies control strategies. Addressing these gaps through targeted research is essential for developing more effective, evidence-based interventions.
1 Introduction
Canine mediated rabies is a neglected disease, and its elimination is hampered by the lack of comprehensive data, particularly in resource-limited, rabies-endemic countries of Africa and Asia (1). Reliable epidemiological data are crucial to understand the disease burden, to implement and evaluate control measures, and to guide policy decisions (2). However, rabies-endemic countries often lack robust surveillance systems and face administrative barriers (2), resulting in obscuring the true impact of rabies leading to misallocation of resources and accentuating the neglect surrounding the disease (3). In addition to the limited availability of data, inadequate diagnostic capacity and a lack of political commitment and allocated financial resources make implementation of intervention measures such as mass dog vaccination (MDV) and human post-exposure prophylaxis (PEP) difficult (4, 5).
The dog populations that are generally uncontrolled in rabies endemic countries are referred to as free-roaming dogs (FRD) (1). Usually abundant around human settlements (6), FRD home ranges are reported to include sites such as schools, temples, shopping centers, community markets, and carcass disposal sites (7, 8). However, the movement patterns and population densities vary greatly within and between countries (9–12). Various factors, such as culture, beliefs, education, and urbanization, influence these characteristics of dog populations (13–15).
The assessment of existing knowledge of FRD abundance and their movement patterns can help to strategize rabies control interventions, such as vaccination coverage, and effectively manage their populations (9, 11, 16–21). Unfortunately, most interventions in endemic countries rarely consider the targeted dog population estimates (2, 22–24). For example, in India, animal birth control (ABC) programs rarely considered the FRD demographic composition (such as sex ratios and age structure), an oversight which results in little reduction in the FRD populations (25). Similarly, a study in Malawi and a population dynamics model have demonstrated that MDV campaigns against rabies frequently fail to achieve the recommended 70% coverage, partly due to a lack of understanding of the roaming behaviors and home ranges of FRD subpopulations (8, 26). These findings emphasize the need for more studies on FRD enumeration and movement patterns to inform critical preintervention strategies, such as defining target populations, identifying vaccination areas, and understanding FRD behaviors to enhance the effectiveness of rabies control efforts.
An array of techniques originally developed for assessing wildlife abundance are applied to estimate FRD population sizes and behavior. A systematic review from 2015 identified techniques used to estimate FRD abundance such as direct and indirect counts, capture-recapture methods, and radio telemetry studies (27). World Organization for Animal Health (WOAH) identifies two methods, direct observation and mark-resight, for determining FRD population size and lists the potential downfalls of each (28). Both techniques rely on assumptions of equal visibility of marked and unmarked dogs, and no change in FRD population in the survey area, which may not always apply (28). A 2013 published systematic review of methods for estimating the size of restricted domiciliary dog populations found these methods for FRD to be generally questionable due to measurement bias and biases associated with length of sampling time, selection bias and non-response bias (29).
In contrast to enumeration studies, there are no reviews or guidelines outlining methods for investigating FRD movement. However, this does not imply that methods for analyzing FRD movement are absent. Similar to enumeration studies, movement methods rely on techniques already widely used in ecology research (30–33). The increased accessibility of GPS techniques in recent years has led to a rise in movement studies and published literature on FRD, with many articles focusing on home range analysis (34–38) and contact network analysis (7, 39–42).
In many settings where funding for rabies control is limited, relying on existing data and proven tools, supported by adaptable, evidence-based guidelines, can be a more feasible and equitable path toward effective implementation. The objective of this scoping review is to provide a comprehensive overview of the various methods used to estimate FRD population sizes and movements. Despite the wide range of techniques available, there is still a lack of critical evaluation by the authors of the published articles regarding the methods they employed, including a clear understanding of the advantages and limitations of these techniques within their studies. This review seeks to address that gap by exploring the methodological approaches used in existing research and examining where the studies were conducted, reducing the need for each country to generate local evidence through resource-intensive research. Specifically, we analyze studies conducted between 2012 and 2024 to assess current knowledge in FRD enumeration and dog movement methodologies.
2 Materials and methods
This scoping review followed the PRISMA extension for scoping reviews (PRISMA-ScR) guidelines (43).
2.1 Information sources and search
A search for scholarly literature on the subject was performed through three electronic web-based literature databases: Embase, Scopus and Web of Science using the search string “[(Free-roaming OR Free-ranging OR stray) OR (Free AND (roaming OR ranging))] AND (dog* OR canine*) AND [behavior OR behavior OR movement OR (population AND (enumeration OR size OR estimat*))]”.
2.2 Eligibility criteria
To be included in the review, studies needed to focus on population estimation or movement of FRDs. Peer-reviewed journal articles were included if they were published from, and including, 2012 up to and including the end of 2024, written in English, French, Portuguese, or Spanish, and constituted original research involving observational studies on FRDs. Origins of the articles were systematically analyzed and only those from studies conducted in rabies endemic countries with sporadic dog-mediated human rabies, and dog-mediated human rabies as per the WHO – The Global Health Observatory (44) and Regional Plan for the elimination of canine rabies (45).
Studies were excluded if full text was not available, or if they concerned specific FRD groups (e.g., pregnant bitches, neutered dogs etc.). Reviews, commentaries, pre-prints, conference papers and opinion pieces were excluded from the review.
2.3 Selection of sources of evidence
A literature search following the criteria in section 2.1 and 2.2 identified 2,167 articles. After removing duplicates using Zotero,1 1,326 articles remained (Figure 1). The articles were divided equally between authors LCS and CF, who independently screened titles and abstracts. Non-conforming articles were excluded based on inclusion and exclusion criteria, and disagreements were resolved through discussion or by consulting a third party (SD, HKT). Rayyan software2 was used for this process, resulting in 93 articles meeting the inclusion criteria.
Figure 1. Prisma flowchart detailing the flow of information through the different phases of this scoping review. This chart showcases information on the number of records identified through literature search (Identification), number of records screened based on titles and abstract, and full text (Screening), and number of articles included in the scoping review (Included) and specific reasons for excluding articles after each selection process step.
Full-text screening was conducted by LCS, to exclude articles unrelated to dog population enumeration and movement techniques, thus narrowing the selection to 62 articles (Figure 1; Supplementary Table S1). During the data extraction step (section 2.4), and after consulting a third party (SD), 10 additional articles were again excluded (Supplementary Table S1), resulting in a final corpus of 52 articles.
2.4 Data charting and data items
A data-charting form was developed and discussed by the review team to identify key variables for extraction. Two researchers (LCS, JF) independently entered study characteristics, demographics, and other relevant data into an Excel spreadsheet, including details on study methodologies and the advantages and limitations of the methods as mentioned by the included articles’ authors. A list with data extracted is presented in Supplementary Figure S1. Preliminary extraction was tested with 20 items to ensure consistency. In cases of disagreement, a third researcher (SD or HKT) was consulted.
All data extraction was conducted using Microsoft Excel (Microsoft Corporation, Redmond, WA).
2.5 Synthesis of results
A descriptive analysis of the extracted data was conducted, including a narrative summary of key findings and article characteristics. Methodological differences, geographic variation, and sample sizes were considered when summarizing data. The advantages and limitations of methods used in enumeration and dog movement studies as stated by each included article authors were summarized in a table.
This study did not aim to conduct a meta-analysis; however, among movement studies, home range studies demonstrated consistent data collection processes, enabling comparisons of their applied methodologies. Median home range sizes presented in the articles were converted to hectares and illustrated with a bubble plot, showing variation by technique and study population size. Home range size variability across techniques was compared using the coefficient of variation (CoV) (46). Analyses were performed using R Statistical Software (v4.3.2; R Core Team 2023).
3 Results
3.1 Temporal and geographic distribution of the articles
A total of 2,167 articles meeting our search criteria were identified, with 518 from Embase, 637 from Scopus, and 1,012 from Web of Science. Following the full screening process, 52 articles were ultimately included in this review (Figure 1). The majority of included articles were from 2019 (n = 10), closely followed by 2021 (n = 9), while the fewest articles were recorded in 2012, 2013 and 2014 (n = 1 each) (Figure 2). No article from 2017 was included. Most of the articles focused on FRD enumeration (n = 39), whereas 14 articles addressed FRD movement investigations. One study (47) utilized enumeration and movement techniques simultaneously (Figure 2). Articles specifically centered on dog movement were reported mostly after 2018.
Figure 2. Barplot representing the number of articles per year and their primary focus of research. Articles were identified during a scoping review on dog population enumeration and movement in rabies endemic countries published between 2012 and 2024.
The 52 articles included in this analysis originate from 27 distinct countries. A significant portion of the articles were conducted in India (n = 8), Brazil (n = 6), Indonesia (n = 5), Guatemala (n = 5) and Chad (n = 5) (Figure 3). Notably, most countries (17 out of 27) reported only a single study conducted within their respective locations.
Figure 3. Choropleth Map depicting the number of articles published per country in the world. Articles were identified during a scoping review on dog population enumeration and movement in rabies endemic countries published between 2012 and 2024.
Twenty-two (56%) of the articles investigating FRD enumeration were conducted in urban sites, 13 (33%) in both rural and urban settings, and four (10%) exclusively in rural areas. Most (n = 6) movement articles occurred in both rural and urban areas (43%), whereas four movement articles (29%) were conducted in exclusively urban or rural areas.
Majority of included articles focused on both unowned and owned FRD (n = 35, 67%), whereas 14 (27%) focused specifically on owned FRD and three (6%) on unowned FRD.
3.2 Datasets and techniques used for FRD enumeration and movement studies
The datasets used for the FRD enumeration articles include transect and household surveys (n = 33), human population census (n = 6), dog census (n = 2), and photos from manual cameras (n = 16), unmanned aerial vehicles (such as drones, n = 1) and camera traps (n = 1). All articles on dog movements utilized GPS data, whereas direct observations and photographs were used in one study each on top of the GPS data.
All articles used between one and three different techniques within a single study (mean = 1.5 techniques). Photographic mark capture-recapture emerged as the most frequently used technique in FRD population estimates (n = 19) as well as dog:human ratio (n = 19), followed by simple transect count (n = 9) and mark-capture-recapture technique (n = 8). For FRD movement articles, the primary technique employed was the Minimum Convex Polygon (MCP) method (n = 8).
The datasets and techniques used in the included articles are presented in Table 1 for the enumeration studies and Table 2 for the movement studies. Supplementary Tables S2, S3 provide a detailed summary of each included article, covering its purpose, statistical methods, datasets, advantages, and limitations. Supplementary Table S4 offers a brief overview of the techniques used in the articles. Advantages and limitations to each technique have been reported by the authors of the included articles (Tables 1, 2). Most articles did not report any advantages (n = 12, 16%), limitations (n = 9, 12%), or both (n = 25, 32%) to their deployed techniques (Supplementary Tables S2, S3).
Table 1. Overview of the techniques for FRD population’s enumeration and their required dataset, advantages and limitations used in 39 articles identified during a scoping review between 2012 and 2024.
Table 2. Overview of the techniques to investigate FRD movements and their purposes, required dataset, advantages and limitations used in 14 articles identified during a scoping review between 2012 and 2024.
3.3 Summary of home range sizes
Home range estimation was conducted in 14 articles, with 11 providing analyzable results. These articles used minimum convex polygon (MCP) (n = 6), biased random bridges (BRB) (n = 5), kernel (n = 2), and time-localized convex hull (T-LoCoH) (n = 1) techniques. Five articles (Study ID 5,12, 38,48,52) applied the same method across different sites, while one article (Study ID 33) used both BRB and MCP at the same site. Median core home ranges ranged from 0.0027 to 228 ha (mean = 11.6 ha), and extended home ranges from 1.66 to 2,400 ha (mean = 258.6 ha). Variation is high, both for core and extended HR values. The BRB showed the most consistent core home range results (CoV = 0.22), while MCP had the high variability for both core (CoV = 1.35) and extended home range estimates (CoV = 1.01), and kernel showcased the highest variability for core home range estimates (CoV = 1.70) (Figures 4, 5).
Figure 4. Bubble plot illustrating the estimated median core home range size (in hectares and in a logarithmic scale) in eight articles included in the scoping review, visually separated by the technique used. The bubble size represents the sample size of dogs included in the respective article.
Figure 5. Bubble plot illustrating the estimated median extended home range size (in hectares and in a logarithmic scale) in eight articles included in the scoping review, visually separated by the technique used. The bubble size represents the sample size of dogs included in the respective article.
4 Discussion
The first milestone of the rabies elimination roadmap, defined by organization United Against Rabies, is building evidence related to various fields concerning dog rabies elimination, including the abundance and behavior of FRD (48). Yet, it is evident from our review that rabies endemic countries in Africa and Asia largely lack compiled information in the scientific literature on dog populations. Among the 81 countries worldwide that are considered endemic for dog-mediated rabies, i.e., countries with present or sporadic dog-mediated human rabies (44, 45), we have identified only 27 (33%) that have conducted studies on either FRD population estimates or movement. Among these 27 countries, 17 (63%) have only one scientific article published within their borders. This underscores a notable gap in local knowledge and a lack of understanding of the diversity in FRD populations and movement patterns between and within countries. Most action plans in rabies endemic countries exclude the need for understanding FRD movement, behavior and demography for effective intervention. In addition, the approach to apply Oral Rabies Vaccination (ORV) campaigns to poorly accessible FRD, as discussed by the WHO (49), is promoted in guidelines, but often disregarded. This can both be an effect, or a reason for the scarcity of dog population studies conducted in rabies endemic countries.
Within the two continents mostly affected by rabies, our review included fewer studies from African countries compared to Asian countries. The included 11 Asian countries represent 44% of the 25 Asian countries endemic for dog-mediated human rabies, whereas in Africa only 21% (9 out of 42) of rabies endemic countries were represented in the review (44). This finding may stem from each continent’s policies regarding rabies and FRD or dogs in general, which are largely lacking or if present, are not effectively implemented (50–53). When compared with the number of studies identified from outside Africa and Asia, we found that studies originated from 11 (79%) out of the 14 rabies endemic countries. This indicates that these countries, located mainly in Latin America, invested more into FRD research, and thus rabies control (45). Within Asia, India and Indonesia collectively produce 13 out of the 26 total articles (Figure 3). The heterogeneous distribution observed may be attributable to several factors, including a higher actual or perceived burden of FRD and rabies in these regions, larger human population sizes, or the presence of active research groups in these countries. Indeed, a single research group is responsible for at least half of the included publications from India, while another research group contributed the majority of included articles from Indonesia.
Estimating FRD population size is challenging and prone to bias. This is mainly due to the heterogeneity in the dog population, i.e., the presence of both owned and ownerless dogs, leading to variability in detection probabilities (27, 54, 55). Therefore, trade-offs between complexity in study design and data analysis, and simplification with potentially higher risk of bias need to be considered. For example, the simplest technique deployed, the dog:human ratio calculations, was used 19 times. However, it was typically presented as secondary results rather than being the primary focus of an article on dog population estimates. Ratio estimations are influenced by variations in human population density, making it difficult to make such findings universally usable (27). Also, simple methods like transect counts and distance sampling are effective and cost-efficient. However, both techniques do not consider heterogeneous probabilities of animal detection (27, 56). More specifically, simple transect counts only provide indicators of canine abundance rather than precise population estimates (57), while distance sampling’s random line placement on long roads can lead to overestimations (58).
Most studies on FRD enumeration (27 out of the total) have employed capture-recapture methods. Despite their common use in research, capture-recapture techniques assume a closed population, which was reported as a limitation in the here assessed articles (13, 47, 58–62). A closed population is only met if studies are conducted over short periods with negligible immigration and emigration of dogs, no loss of marks, no misclassification between marked and unmarked dogs, and homogeneous capture probabilities (27). This may be realistic for some, but not all studies conducted. The marking of the dogs can be done individually (e.g., by photos taken), or overall (e.g., by marking them with collars or paint without differentiating between individuals). The main advantage of using photography for marking FRD is that it accounts for individual heterogeneity, providing more accurate estimates compared to simple mark capture-recapture (55). Additionally, digital photography reduces costs associated with artificial marking, avoids handling dogs, and eliminates count variations due to trap-shy or trap-eager behavioral response. However, photographic capture-recapture has limitations, as recognizing individual FRD can be challenging and potentially limited to populations with many indistinguishable individuals (54, 60, 62, 63). Investment into research on more resource-friendly approaches to match individual dogs in photographs is thus demanded. Simple mark-capture recapture is faster, and it does not require the laborious task of reviewing photographs and identifying individual dogs, making it less prone to human error and observer fatigue. Another technique reported are spatial models, which has been used in two studies (64). These complex models require preparing spatial data before analysis which is challenging and time-consuming, hence limiting its use, but holds promise due to their versatility and ability to handle small sample sizes.
Data collection for dog enumeration studies is diverse, drawing from multiple sources. Dog counts during transect surveys is one data collection method, which, despite their straightforward application, must account for factors like topography, climate conditions, and lighting, as these elements impact FRD detection, photographic capture-recapture, and distance measurements (Supplementary Tables S2, S3). Household or school-based surveys (including aiming for full censuses) are also common methods, but they are labor-intensive, often underestimating, and time-consuming (27).
Overall, it can be said that enumeration methods are becoming more diverse and complex, moving beyond simpler techniques. This complexity, coupled with a lack of recognition of the importance of FRD population studies in the development of National Action Plans (NAPs), may contribute to the limited number of studies on this topic (2). Additionally, the lack of a gold standard methodology for estimating free-roaming dog populations increases uncertainty and limits the comparability between study findings. In the absence of a gold standard, population estimation methods from wildlife have gained acceptance (54). Nonetheless, selecting the most suitable technique is challenging and largely influenced by the resources and conditions available at the study site, and by the limitations of the methods, which has been presented in this review (27).
Despite the importance of FRD movement patterns for rabies control and disease spread (9, 65), no consensus on a gold standard for FRD movement studies has been met. Also, so far, this is the first attempt to provide an overview on techniques used to investigate FRD movement behavior alongside the included authors’ stated advantages and limitations of deployed techniques. Most articles on this topic are published after 2019, and all included movement articles reviewed here rely on GPS-collected data. They emerged after a study published from Australia in FRD in Aboriginal communities, using GPS collars and comparing methods for home range calculations (34). The GPS technology allows researchers to study animal movements without human interference and in a non-intrusive manner (66, 67). Advances in this technology over recent years have made GPS units increasingly lightweight and small to carry (68). The decreasing cost and greater market availability of GPS technology, along with the enhanced computational power to process extensive GPS datasets, have made conducting movement studies more feasible (66). These advancements have minimized the need for labor-intensive observations by researchers, likely leading to an increase in such studies in recent years.
From the articles included in this review, we found that authors of movement studies primarily focus their efforts on estimating FRDs’ home ranges. We found that home range sizes are conditional to the type of technique and study population size used, with less complex techniques (such as the MCP) producing more inconsistent results (Figures 4, 5). The widespread adoption of simpler techniques in movement studies often overlooks either the spatial, temporal, or both complexities inherent in animal movement (69, 70), making it difficult to compare home range studies across different countries and regions. The BRB method was discussed to deliver the most reliable home range estimates, likely due to its highly effective method for addressing serial autocorrelation in movement data, frequent in animal tracking studies (71). Additionally, BRB’s ability to decompose spatial usage into frequency and repetition components allows gathering information on an animal’s number of visits to particular locations and the average time spent there (71). Such detailed spatial information is often lacking in simpler techniques like MCP and conventional kernel methods. This pattern is particularly evident in the results obtained using the MCP. Although MCP estimates exhibited substantial variability, they were among the lowest home range values reported, even across populations of varying sizes. This outcome is somewhat unexpected, given that MCP is known to be highly sensitive to outliers and typically tends to overestimate home range size due to the influence of a few wide-ranging individual fixes (69, 72). A plausible explanation for these findings is the lack of standardization in sampling protocols and inherent differences between dog populations (21). Regardless of population size, when sampling regimes (e.g., number of location fixes, tracking duration, spatial coverage) are not standardized, MCP estimates are prone to remain highly variable (72). In addition, values may vary according to the method specifications used to define core and extended home ranges, as these are determined by varying percentage thresholds applied to the sampled data (e.g., excluding the top 5% of outliers to calculate an extended home range encompassing 95% of all recorded fixes).
Research on habitat selection and contact networks of FRD is limited in number, revealing a significant knowledge gap in rabies-endemic countries. Despite some research on the impact of dog movement on rabies outbreaks from rabies endemic areas (8, 9, 37, 73), most such has been carried out in non-endemic regions like Australia (42, 74–78). Investing in habitat selection and social network analysis research of FRD in rabies endemic regions are thus needed to better guide rabies control interventions, such as where to deposit ORV and which dogs primarily to be targeted for vaccination in case resources are limited.
The number of articles identified that investigated FRDs’ movement in rabies endemic countries is limited. Additionally, many enumeration studies are often commissioned by government entities and remain unpublished, meaning that such information is not available in the scientific literature. There is, however, a growing trend among authors to consider the complexities of animal movement in time and space by selecting more sophisticated techniques and acknowledging the limitations of simpler techniques. In contrast, authors using these more advanced techniques rarely discuss their limitations.
We used a scoping review methodology rather than a systematic review due to the exploratory nature of the research question and the diversity of study designs and outcomes in the identified articles (79). We acknowledge that our search string and eligibility criteria may be restrictive, potentially excluding studies that investigated dog enumeration and movement but may have not been captured. However, the objective of this review was to provide a comprehensive overview of the methods currently employed in the scientific literature to study FRD movement and enumeration. By focusing on rabies endemic regions, this review enhances relevance of the research in this field and identified knowledge gaps in areas most affected by the disease. Furthermore, several included studies did not report the advantages and limitations of their techniques, restricting our ability to fully assess the authors’ understanding of the strengths and weaknesses of the methods they employed in their research.
We here presented a large range of studies on FRD populations and movement in rabies-endemic regions using diverse technologies. At the same time, it became pertinent that research is limited to selected countries, hindering the development of locally adapted rabies control strategies, as these require detailed understanding of the local dog populations (9, 80, 81). Moreover, the high resource demands of the techniques used, and the absence of standardized methods, complicates the design of future studies and the comparisons across studies. Nevertheless, it may not be essential for each country and region to conduct their own research on dog populations; instead, they can draw on existing studies for valuable insights. This approach can be enhanced by developing comprehensive guidelines that countries can adopt for their own context and implement effectively.
Author contributions
LC: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Software, Validation, Visualization, Writing – original draft, Writing – review & editing. CF: Data curation, Validation, Writing – review & editing. JF: Data curation, Validation, Writing – review & editing. HT: Conceptualization, Investigation, Methodology, Supervision, Writing – review & editing, Writing – original draft. SD: Conceptualization, Data curation, Investigation, Methodology, Project administration, Supervision, Writing – review & editing, Writing – original draft.
Funding
The author(s) declare that financial support was received for the research and/or publication of this article. Open access funding by University of Bern.
Conflict of interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.
Generative AI statement
The authors declare that no Gen AI was used in the creation of this manuscript.
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Supplementary material
The Supplementary material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fvets.2025.1567807/full#supplementary-material
Footnotes
References
2. Fahrion, AS, Taylor, LH, Torres, G, Müller, T, Dürr, S, Knopf, L, et al. The road to dog rabies control and elimination-what keeps us from moving faster? Front Public Health. (2017) 5:103. doi: 10.3389/fpubh.2017.00103
3. Swedberg, C, Bote, K, Gamble, L, Fénelon, N, King, A, and Wallace, RM. Eliminating invisible deaths: the woeful state of global rabies data and its impact on progress towards 2030 sustainable development goals for neglected tropical diseases. Front Trop Dis. (2024) 5:1303359. doi: 10.3389/fitd.2024.1303359
4. Changalucha, J, Hampson, K, Jaswant, G, Lankester, F, and Yoder, J. Human rabies: prospects for elimination. CAB Rev. (2021) 16:39. doi: 10.1079/pavsnnr202116039
5. Kanda, K, Jayasinghe, A, Jayasinghe, C, and Yoshida, T. A regional analysis of the Progress of current dog-mediated rabies control and prevention. Pathogens. (2022) 11:1130. doi: 10.3390/pathogens11101130
6. Cunha Silva, L, Friker, B, Warembourg, C, Kanankege, K, Wera, E, Berger-González, M, et al. Habitat selection by free-roaming domestic dogs in rabies endemic countries in rural and urban settings. Sci Rep. (2022) 12:1–10. doi: 10.1038/s41598-022-25038-z
7. Warembourg, C, Fournié, G, Abakar, MF, Alvarez, D, Berger-González, M, Odoch, T, et al. Predictors of free-roaming domestic dogs’ contact network centrality and their relevance for rabies control. Sci Rep. (2021) 11:1–13. doi: 10.1038/s41598-021-92308-7
8. De la Puente-Arévalo, M, Motta, P, Dürr, S, Warembourg, C, Nikola, C, Burdon-Bailey, J, et al. Ranging patterns and factors associated with movement in free-roaming domestic dogs in urban Malawi. Ecol Evol. (2022) 12:8498. doi: 10.1002/ece3.8498
9. Raynor, B, de la Puente-León, M, Johnson, A, Díaz, EW, Levy, MZ, Recuenco, SE, et al. Movement patterns of free-roaming dogs on heterogeneous urban landscapes: implications for rabies control. Prev Vet Med. (2020) 178:104978. doi: 10.1016/j.prevetmed.2020.104978
10. Villatoro, FJ, Sepúlveda, MA, Stowhas, P, and Silva-Rodríguez, EA. Urban dogs in rural areas: human-mediated movement defines dog populations in southern Chile. Prev Vet Med. (2016) 135:59–66. doi: 10.1016/j.prevetmed.2016.11.004
11. Fèvre, EM, Bronsvoort, BMDC, Hamilton, KA, and Cleaveland, S. Animal movements and the spread of infectious diseases. Trends Microbiol. (2006) 14:125–31. doi: 10.1016/j.tim.2006.01.004
12. Jackman, J, and Rowan, AN. Free-roaming dogs in developing countries: the benefits of capture, neuter, and return programs In: DJ Salem and AN Rowan, editors. The state of the animals 2007. Washington, DC: Humane Society Press (2007). 55–78.
13. Smith, LM, Quinnell, R, Munteanu, A, Hartmann, S, Villa, PD, and Collins, L. Attitudes towards free-roaming dogs and dog ownership practices in Bulgaria, Italy, and Ukraine. PLoS One. (2022) 17:368. doi: 10.1371/journal.pone.0252368
14. Smith, LM, Quinnell, RJ, Goold, C, Munteanu, AM, Hartmann, S, and Collins, LM. Assessing the impact of free-roaming dog population management through systems modelling. Sci Rep. (2022) 12:5049. doi: 10.1038/s41598-022-15049-1
15. De Melo, SN, da Silva, ES, Barbosa, DS, Teixeira-Neto, RG, Lacorte, GA, Horta, MAP, et al. Effects of gender, sterilization, and environment on the spatial distribution of free-roaming dogs: an intervention study in an urban setting. Front Vet Sci. (2020) 7:289. doi: 10.3389/fvets.2020.00289
16. Conan, A, Akerele, O, Simpson, G, Reininghaus, B, Van Rooyen, J, and Knobel, D. Population dynamics of owned, free-roaming dogs: implications for rabies control. PLoS Negl Trop Dis. (2015) 9:4177. doi: 10.1371/journal.pntd.0004177
17. Hampson, K, Dushoff, J, Cleaveland, S, Haydon, DT, Kaare, M, Packer, C, et al. Transmission dynamics and prospects for the elimination of canine rabies. PLoS Biol. (2009) 7:53. doi: 10.1371/journal.pbio.1000053
18. Morters, MK, Mckinley, TJ, Restif, O, Conlan, AJK, Cleaveland, S, Hampson, K, et al. The demography of free-roaming dog populations and applications to disease and population control. J Appl Ecol. (2014) 51:1096–106. doi: 10.1111/1365-2664.12279
19. Tiwari, HK, Bruce, M, O’dea, M, and Robertson, ID. Utilising group-size and home-range characteristics of free-roaming dogs (FRD) to guide mass vaccination campaigns against rabies in India. Vaccines (Basel). (2019) 7:136. doi: 10.3390/vaccines7040136
20. Tiwari, HK, Gogoi-Tiwari, J, and Robertson, ID. Eliminating dog-mediated rabies: challenges and strategies. Anim Dis. (2021) 1:1–13. doi: 10.1186/s44149-021-00023-7
21. Warembourg, C, Wera, E, Odoch, T, Malo Bulu, P, Berger-González, M, Alvarez, D, et al. Comparative study of free-roaming domestic dog management and roaming behavior across four countries: Chad, Guatemala, Indonesia, and Uganda. Front Vet Sci. (2021) 8:7900. doi: 10.3389/fvets.2021.617900
22. Mbilo, C, Kabongo, JB, Pyana, PP, Nlonda, L, Nzita, RW, Luntadila, B, et al. Dog ecology, bite incidence, and disease awareness: A cross-sectional survey among a rabies-affected community in the democratic republic of the congo. Vaccines (Basel). (2019) 7:98. doi: 10.3390/vaccines7030098
23. Nel, LH. Factors impacting the control of rabies In: RM Atlas and S Maloy, editors. One health: People, animals, and the environment. Washington, DC: American Society for Microbiology (2014)
24. Ogun, AA, Okonko, IO, Udeze, AO, Shittu, I, Garba, KN, Fowotade, A, et al. Feasibility and factors affecting global elimination and possible eradication of rabies in the world. J Gen Mol Virol. (2010) 2:1–27. doi: 10.1128/9781555818432.ch7
25. Tiwari, HK, Robertson, ID, O’Dea, M, and Vanak, AT. Demographic characteristics of free-roaming dogs (FRD) in rural and urban India following a photographic sight-resight survey. Sci Rep. (2019) 9:992. doi: 10.1038/s41598-019-52992-y
26. Belsare, A, and Vanak, AT. Modelling the challenges of managing free-ranging dog populations. Sci Rep. (2020) 10:18874. doi: 10.1038/s41598-020-75828-6
27. Belo, VS, Werneck, GL, Da Silva, ES, Barbosa, DS, and Struchiner, CJ. Population estimation methods for free-ranging dogs: a systematic review. PLoS One. (2015) 10:e0144830. doi: 10.1371/journal.pone.0144830
29. Downes, MJ, Dean, RS, Stavisky, JH, Adams, VJ, Grindlay, DJC, and Brennan, ML. Methods used to estimate the size of the owned cat and dog population: a systematic review. BMC Vet Res. (2013) 9:121–12. doi: 10.1186/1746-6148-9-121
30. Lichti, NI, and Swihart, RK. Estimating utilization distributions with kernel versus local convex hull methods. J Wildl Manag. (2011) 75:413–22. doi: 10.1002/jwmg.48
31. Cross, SL, Tomlinson, S, Craig, MD, and Bateman, PW. The time local convex hull method as a tool for assessing responses of fauna to habitat restoration: a case study using the perentie (Varanus giganteus: Reptilia: Varanidae). Aust J Zool. (2020) 67:27–37. doi: 10.1071/ZO19040
32. Worton, BJ. Kernel methods for estimating the utilization distribution in home- range studies. Ecology. (1989) 70:164–8. doi: 10.2307/1938423
33. Kie, JG, Matthiopoulos, J, Fieberg, J, Powell, RA, Cagnacci, F, Mitchell, MS, et al. The home-range concept: are traditional estimators still relevant with modern telemetry technology? Philos Trans R Soc B Biol Sci. (2010) 365:2221–31. doi: 10.1098/rstb.2010.0093
34. Dürr, S, and Ward, MP. Roaming behaviour and home range estimation of domestic dogs in aboriginal and Torres Strait islander communities in northern Australia using four different methods. Prev Vet Med. (2014) 117:340–57. doi: 10.1016/j.prevetmed.2014.07.008
35. Dürr, S, Dhand, NK, Bombara, C, Molloy, S, and Ward, MP. What influences the home range size of free-roaming domestic dogs? Epidemiol Infect. (2017) 145:1339–50. doi: 10.1017/S095026881700022X
36. Sousa, FM, Warembourg, C, Abakar, MF, Alvarez, D, Berger-Gonzalez, M, Odoch, T, et al. Investigation of optimized observation periods for estimating a representative home range of free-roaming domestic dogs. Sci Rep. (2023) 13:22750. doi: 10.1038/s41598-023-49851-2
37. Muinde, P, Bettridge, JM, Sousa, FM, Dürr, S, Dohoo, IR, Berezowski, J, et al. Who let the dogs out? Exploring the spatial ecology of free-roaming domestic dogs in western Kenya. Ecol Evol. (2021) 11:4218–31. doi: 10.1002/ece3.7317
38. Pérez, GE, Conte, A, Garde, EJ, Messori, S, Vanderstichel, R, and Serpell, J. Movement and home range of owned free-roaming male dogs in Puerto Natales, Chile. Appl Anim Behav Sci. (2018) 205:74–82. doi: 10.1016/j.applanim.2018.05.022
39. Sparkes, J, Körtner, G, Ballard, G, Fleming, PJS, and Brown, WY. Effects of sex and reproductive state on interactions between free-roaming domestic dogs. PLoS One. (2014) 9:e116053. doi: 10.1371/journal.pone.0116053
40. Bombara, CB, Dürr, S, Machovsky-Capuska, GE, Jones, PW, and Ward, MP. A preliminary study to estimate contact rates between free-roaming domestic dogs using novel miniature cameras. PLoS One. (2017) 12:e0181859. doi: 10.1371/journal.pone.0181859
41. Laager, M, Mbilo, C, Madaye, EA, Naminou, A, Léchenne, M, Tschopp, A, et al. The importance of dog population contact network structures in rabies transmission. PLoS Negl Trop Dis. (2018) 12:e0006680. doi: 10.1371/journal.pntd.0006680
42. Brookes, VJ, VanderWaal, K, and Ward, MP. The social networks of free-roaming domestic dogs in island communities in the Torres Strait, Australia. Prev Vet Med. (2020) 181:104534. doi: 10.1016/j.prevetmed.2018.09.008
43. Tricco, AC, Lillie, E, Zarin, W, O’Brien, KK, Colquhoun, H, Levac, D, et al. PRISMA extension for scoping reviews (PRISMA-ScR): checklist and explanation. Ann Intern Med. (2018) 169:467–73. doi: 10.7326/M18-0850
44. WHO. The Global Health Observatory–Presence of dog-transmitted human rabies: 2021. Geneva: WHO. (2023). Available online at: https://www.who.int/data/gho/data/themes/topics/rabies.
46. Shechtman, O. The coefficient of variation as an index of measurement reliability In: O Shechtman, editor. Methods of clinical epidemiology. Berlin, Heidelberg: Springer (2013). 39–49.
47. Dias, RA, Guilloux, AGA, Borba, MR, Guarnieri, MC d L, Prist, R, Ferreira, F, et al. Size and spatial distribution of stray dog population in the University of São Paulo campus, Brazil. Prev Vet Med. (2013) 110:263–73. doi: 10.1016/j.prevetmed.2012.12.002
48. United Against Rabies. The Rabies Roadmap United Against Rabies Forum. Available online at: https://www.unitedagainstrabies.org/the-rabies-roadmap/ (Accessed Dec 29, 2023).
49. WHO Rabies Modelling Consortium. Zero human deaths from dog-mediated rabies by 2030: perspectives from quantitative and mathematical modelling. Gates Open Res. (2020) 3:1564. doi: 10.12688/gatesopenres.13074.2
50. Gongal, G, and Wright, AE. Human rabies in the WHO Southeast Asia region: forward steps for elimination. Adv Prev Med. (2011) 2011:1–5. doi: 10.4061/2011/383870
51. Haselbeck, AH, Rietmann, S, Tadesse, BT, Kling, K, Kaschubat-dieudonné, ME, Marks, F, et al. Challenges to the fight against rabies—the landscape of policy and prevention strategies in Africa. Int J Environ Res Public Health. (2021) 18:1–15. doi: 10.3390/ijerph18041736
52. Miranda, MEG, and Miranda, NLJ. Rabies prevention in Asia: institutionalizing implementation capacities In: MEG Miranda, editor. Rabies and Rabies Vaccines. Berlin: Springer International Publishing (2020). 103–16.
53. Rupprecht, CE, Abela-Ridder, B, Abila, R, Amparo, AC, Banyard, A, Blanton, J, et al. Towards rabies elimination in the Asia-Pacific region: from theory to practice. Biologicals. (2020) 64:83–95. doi: 10.1016/j.biologicals.2020.01.008
54. Tiwari, HK, Robertson, ID, O’Dea, M, Gogoi-Tiwari, J, Panvalkar, P, Bajwa, RS, et al. Validation of application superduplicates (AS) enumeration tool for free-roaming dogs (FRD) in urban settings of Panchkula municipal corporation in North India. Front Vet Sci. (2019) 6:458134. doi: 10.3389/fvets.2019.00173
55. Tiwari, HK, Vanak, AT, O’Dea, M, Gogoi-Tiwari, J, and Robertson, ID. A comparative study of enumeration techniques for free-roaming dogs in rural Baramati, district Pune, India. Front Vet Sci. (2018) 5:368159. doi: 10.3389/fvets.2018.00104
56. Iván Peña, G, Florangel Vidal, F, and Aliesky, HR. Stray dog population of the municipality of Camagüey, Cuba. Rev Investig Vet Peru. (2016) 27:840–4. doi: 10.15381/rivep.v27i4.12570
57. Flores, V, Viozzi, G, Rauque, C, Mujica, G, Herrero, E, Ballari, SA, et al. A cross-sectional study of free-roaming dogs in a Patagonian city: their distribution and intestinal helminths in relation to socioeconomic aspects of neighborhoods. Vet Parasitol Reg Stud Rep. (2022) 33:100747. doi: 10.1016/j.vprsr.2022.100747
58. Meunier, NV, Gibson, AD, Corfmat, J, Mazeri, S, Handel, IG, Gamble, L, et al. A comparison of population estimation techniques for individually unidentifiable free-roaming dogs. BMC Vet Res. (2019) 15:1–10. doi: 10.1186/s12917-019-1938-1
59. Gill, GS, Singh, BB, Dhand, NK, Aulakh, RS, Ward, MP, and Brookes, VJ. Stray dogs and public health: population estimation in Punjab, India. Vet Sci. (2022) 9:75. doi: 10.3390/vetsci9020075
60. Shamsaddini, S, Ahmadi Gohari, M, Kamyabi, H, Nasibi, S, Derakhshani, A, Mohammadi, MA, et al. Dynamic modeling of female neutering interventions for free-roaming dog population management in an urban setting of southeastern Iran. Sci Rep. (2022) 12:4781. doi: 10.1038/s41598-022-08697-w
61. Cárdenas, M, Grijalva, CJ, and de la Torre, S. Free-roaming dog surveys in Quito, Ecuador: experiences, lessons learned, and future work. Front Vet Sci. (2021) 8:8. doi: 10.3389/fvets.2021.766348
62. Smith, LM, Goold, C, Quinnell, RJ, Munteanu, AM, Hartmann, S, Villa, PD, et al. Population dynamics of free-roaming dogs in two European regions and implications for population control. PLoS One. (2022) 17:636. doi: 10.1371/journal.pone.0266636
63. Cleaton, JM, Blanton, JD, Dilius, P, Ludder, F, Crowdis, K, Medley, A, et al. Use of photography to identify free-roaming dogs during sight-resight surveys: impacts on estimates of population size and vaccination coverage, Haiti 2016. Vaccine X. (2019) 2:100025. doi: 10.1016/j.jvacx.2019.100025
64. Thanapongtharm, W, Kasemsuwan, S, Wongphruksasoong, V, Boonyo, K, Pinyopummintr, T, Wiratsudakul, A, et al. Spatial distribution and population estimation of dogs in Thailand: implications for rabies prevention and control. Front Vet Sci. (2021) 8:790701. doi: 10.3389/fvets.2021.790701
65. Dürr, S, Wera, E, Brookes, VJ, Warembourg, C, Griss, S, and Fahrion, AS. The role of dog ecology in canine rabies prevention and control in Asia: lessons from Indonesia and the oceanic region In: S Dürr, editor. One health for dog-mediated rabies elimination in Asia: A collection of local experiences. Wallingford: CABI International (2023). 142–59.
66. Hebblewhite, M, and Haydon, DT. Distinguishing technology from biology: a critical review of the use of GPS telemetry data in ecology. Philos Trans R Soc B Biol Sci. (2010) 365:2303–12. doi: 10.1098/rstb.2010.0087
67. Cochrane, MM, Brown, DJ, and Moen, RA. GPS Technology for Semi-Aquatic Turtle Research. Diversity. (2019) 11:34. doi: 10.3390/d11030034
68. McMahon, LA, Rachlow, JL, Shipley, LA, Forbey, JS, Johnson, TR, and Olsoy, PJ. Evaluation of micro-GPS receivers for tracking small-bodied mammals. PLoS One. (2017) 12:e0173185. doi: 10.1371/journal.pone.0173185
69. Burgman, MA, and Fox, JC. Bias in species range estimates from minimum convex polygons: implications for conservation and options for improved planning. Anim Conserv. (2003) 6:19–28. doi: 10.1017/S1367943003003044
70. Kernohan, BJ, Gitzen, RA, and Millspaugh, JJ. Analysis of animal space use and movements In: BJ Kernohan, editor. Radio tracking and animal populations. Cambridge, MA: Academic Press (2001). 125–66.
71. Benhamou, S. Dynamic approach to space and habitat use based on biased random bridges. PLoS One. (2011) 6:e14592. doi: 10.1371/journal.pone.0014592
72. Börger, L, Franconi, N, De Michele, G, Gantz, A, Meschi, F, Manica, A, et al. Effects of sampling regime on the mean and variance of home range size estimates. J Anim Ecol. (2006) 75:1393–405. doi: 10.1111/j.1365-2656.2006.01164.x
73. Colombi, D, Poletto, C, Nakouné, E, Bourhy, H, and Colizza, V. Long-range movements coupled with heterogeneous incubation period sustain dog rabies at the national scale in Africa. PLoS Negl Trop Dis. (2020) 14:e0008317. doi: 10.1371/journal.pntd.0008317
74. Ferguson, EA, Hampson, K, Cleaveland, S, Consunji, R, Deray, R, Friar, J, et al. Heterogeneity in the spread and control of infectious disease: consequences for the elimination of canine rabies. Sci Rep. (2015) 5:18232. doi: 10.1038/srep18232
75. Hudson, EG, Brookes, VJ, Dürr, S, and Ward, MP. Domestic dog roaming patterns in remote northern Australian indigenous communities and implications for disease modelling. Prev Vet Med. (2017) 146:52–60. doi: 10.1016/j.prevetmed.2017.07.010
76. Sparkes, J, Körtner, G, Ballard, G, and Fleming, PJS. Spatial and temporal activity patterns of owned, free-roaming dogs in coastal eastern Australia. Prev Vet Med. (2022) 204:105641. doi: 10.1016/j.prevetmed.2022.105641
77. Yoak, AJ, Reece, JF, Gehrt, SD, and Hamilton, IM. Optimizing free-roaming dog control programs using agent-based models. Ecol Model. (2016) 341:53–61. doi: 10.1016/j.ecolmodel.2016.09.018
78. Maher, EK, Ward, MP, and Brookes, VJ. Investigation of the temporal roaming behaviour of free-roaming domestic dogs in indigenous communities in northern Australia to inform rabies incursion preparedness. Sci Rep. (2019) 9:1–12. doi: 10.1038/s41598-019-51447-8
79. Munn, Z, Peters, MDJ, Stern, C, Tufanaru, C, McArthur, A, and Aromataris, E. Systematic review or scoping review? Guidance for authors when choosing between a systematic or scoping review approach. BMC Med Res Methodol. (2018) 18:143. doi: 10.1186/s12874-018-0611-x
80. Davlin, SL, and VonVille, HM. Canine rabies vaccination and domestic dog population characteristics in the developing world: a systematic review. Vaccine. (2012) 30:3492–502. doi: 10.1016/j.vaccine.2012.03.069
81. Franka, R, Smith, TG, Dyer, JL, Wu, X, Niezgoda, M, and Rupprecht, CE. Current and future tools for global canine rabies elimination. Antivir Res. (2013) 100:220–5. doi: 10.1016/j.antiviral.2013.07.004
82. Ochoa, AY, Falcón, PN, Zuazo, RJ, and Guevara, PB. Estimated population of stray dogs in the district of Los Olivos, Lima, Peru. (2014) 25.
83. Wu, X, Yu, VY, Huang, Z, Lu, J, Tang, W, Shen, S, et al. Estimation of the rural dog population within a mega-city: An example in Jiading District, Shanghai. Front Vet Sci. (2021) 8:630180. doi: 10.3389/fvets.2021.630180
84. de la Reta, M, Muratore, M, Perna, S, Polop, J, and Provensal, MC. Abundance of stray dogs and its relationship with environmental factors in Río Cuarto (Córdoba, Argentina). Revista Veterinaria. (2018) 29:113–8.
85. Carolina Chávez, V, Néstor Falcón, P, León, D, and Daniel, Sánchez R. Stray dogs inside and outside of official markets of villa El Salvador district in Lima, Peru. Revista de Investigaciones Veterinarias del Peru. (2016) 27:176–82.
86. Tavlian, S, Stevenson, MA, Webb, B, Sharma, K, Pearson, J, Britton, A, et al. Prediction of the size and spatial distribution of free-roaming dog populations in urban areas of Nepal. Spat Spatiotemporal Epidemiol. (2024) 49:100647. doi: 10.1016/j.sste.2024.100647
87. Rinzin, K, Tenzin, T, and Robertson, I. Size and demography pattern of the domestic dog population in Bhutan: Implications for dog population management and disease control. Prev Vet Med. (2016) 126:39–47. doi: 10.1016/j.prevetmed.2016.01.030
88. Tenzin, T, Ahmed, R, Debnath, NC, Ahmed, G, and Yamage, M. Free-roaming dog population estimation and status of the dog population management and rabies control program in Dhaka City, Bangladesh. PLoS Negl Trop Dis. (2015) 9:e0003784. doi: 10.1371/journal.pntd.0003784
89. Tenzin, T, McKenzie, JS, Vanderstichel, R, Rai, BD, Rinzin, K, Tshering, Y, et al. Comparison of mark-resight methods to estimate abundance and rabies vaccination coverage of free-roaming dogs in two urban areas of south Bhutan. Prev Vet Med. (2015) 118:436–48. doi: 10.1016/j.prevetmed.2015.01.008
90. Tenzin, T, Hikufe, EH, Hedimbi, N, Athingo, R, Shikongo, MB, Shuro, T, et al. Dog ecology and rabies knowledge, attitude and practice (KAP) in the Northern Communal Areas of Namibia. PLoS Negl Trop. (2024) 18:e0011631. doi: 10.1371/journal.pntd.0011631
91. Sambo, M, Hampson, K, Changalucha, J, Cleaveland, S, Lembo, T, Lushasi, K, et al. Estimating the size of dog populations in Tanzania to inform rabies control. Vet Sci. (2018) 5:77. doi: 10.3390/vetsci5030077
92. Warembourg, C, Berger-González, M, Alvarez, D, Sousa, FM, Hernández, AL, Roquel, P, et al. Estimation of free-roaming domestic dog population size: Investigation of three methods including an Unmanned Aerial Vehicle (UAV) based approach. PLoS One. (2020) 15:e0225022. doi: 10.1371/journal.pone.0225022
93. Paschoal, AMO, Massara, RL, Bailey, LL, Kendall, WL, Doherty, PF, Hirsch, A, et al. Use of Atlantic Forest protected areas by free-ranging dogs: Estimating abundance and persistence of use. Ecosphere. (2016) 7:e01480. doi: 10.1002/ecs2.1480
94. Punjabi, GA, Athreya, V, and Linnell, JDC. Using natural marks to estimate free-ranging dog canis familiaris abundance in a MARK-RESIGHT framework in suburban Mumbai, India. Trop Conserv Sci. (2012) 5:510–20. doi: 10.1177/194008291200500408
95. Özen, D, Böhning, D, and Gürcan, IS. Estimation of stray dog and cat populations in metropolitan Ankara, Turkey. Turk J Vet Anim Sci. (2016) 40:7–12. doi: 10.3906/vet-1505-70
96. Silva, JE, Oliveira Rodrigues, T, Silva, AJ, and Queiroz, LH. Evaluating the movement of free-roaming dogs using georeferencing and the photographic capture-recapture method. Acta Veterinaria Brasilica. (2019) 13:70–6. doi: 10.21708/avb.2019.13.2.7779
97. Mustiana, A, Toribio, JA, Abdurrahman, M, Suadnya, IW, Hernandez-Jover, M, Putra, AAG, et al. Owned and unowned dog population estimation, dog management and dog bites to inform rabies prevention and response on Lombok Island, Indonesia. PLoS One. (2015) 10:e0124092. doi: 10.1371/journal.pone.0124092
98. Kalthoum, S, Ben Salah, C, Rzeigui, H, Gharbi, R, Guesmi, K, Ben Salem, A, et al. Owned and free-roaming dogs in the North West of Tunisia: estimation, characteristics and application for the control of dog rabies. Heliyon. (2021) 7. doi: 10.1016/j.heliyon.2021.e08347
99. Bouaddi, K, Bitar, A, Ferssiwi, A, Bouslikhane, M, Fitani, A, Mshelbwala, PP, et al. Socioecology of the canine population in the Province of El Jadida, Morocco. Vet Med Int. (2018) 2018:4234791. doi: 10.1155/2018/4234791
100. Jagriti Bhalla, S, Kemmers, R, Vasques, A, and Tamim, Vanak A. “Stray appetites”: a socio-ecological analysis of free-ranging dogs living alongside human communities in Bangalore, India. Urban Ecosyst. (2021) 24:1245–1258. doi: 10.1007/s11252-021-01097-4
101. Emiliano, A, and Adrián, S. Free-roaming dogs in Ushuaia City, Tierra del Fuego, Argentina. How many and why. Urban Ecosyst. (2023) 26:559–74. doi: 10.1007/s11252-022-01320-w
102. Nasiry, Z, Mazlan, M, Noordin, MM, and Mohd Lila, MA. Evaluation of Dynamics, Demography and Estimation of Free-Roaming Dog Population in Herat City, Afghanistan. Animals. (2023) 13:1126. doi: 10.3390/ani13071126
103. de Melo, SN, da Silva, ES, Ribeiro, RAN, Soares, PHA, Cunha, AKR, de Souza Gonçalves, CM, et al. The Influence of Community Feeders and Commercial Food Outlets on the Spatial Distribution of Free-Roaming Dogs—A Photographic Capture and Recapture Study. Animals. (2023) 13:824. doi: 10.3390/ani13050824
104. de Santi, CE, Chiba de Castro, WA, Sibim, AC, Lopes, RD, Galvão, SR, Kurtz, GM, et al. Spatial distribution and population dynamics of free-roaming (stray and semi-domiciled) dogs in a major Brazilian city. Front Vet Sci. (2024) 11:1417458. doi: 10.3389/fvets.2024.1417458
105. Kwaghe, AV, Okomah, D, Okoli, I, Kachalla, MG, Aligana, M, Alabi, O, et al. Estimation of dog population in Nasarawa state Nigeria: A pilot study. Pan Afr Med J. (2019) 34. doi: 10.11604/pamj.2019.34.25.16755
106. Wilson-Aggarwal, JK, Goodwin, CED, Moundai, T, Sidouin, MK, Swan, GJF, Léchenne, M, et al. Spatial and temporal dynamics of space use by free-ranging domestic dogs Canis familiaris in rural Africa. Ecol Appl. (2021) 31:e02328. doi: 10.1002/eap.2328
107. Wilson-Aggarwal, JK, Ozella, L, Tizzoni, M, Cattuto, C, Swan, GJF, Moundai, T, et al. High-resolution contact networks of free-ranging domestic dogs Canis familiaris and implications for transmission of infection. PLoS Negl Trop Dis. (2019) 13:e0007565. doi: 10.1371/journal.pntd.0007565
Keywords: rabies endemic countries, enumeration, zero by 30, disease elimination, dog-mediated rabies
Citation: Cunha Silva L, Fellenberg C, Freudenthal J, Tiwari HK and Dürr S (2025) Free-roaming dog populations and movement methodologies for global rabies elimination: knowns and unknowns – a scoping review. Front. Vet. Sci. 12:1567807. doi: 10.3389/fvets.2025.1567807
Edited by:
Eyal Klement, Hebrew University of Jerusalem, IsraelReviewed by:
Julie Cleaton, Centers for Disease Control and Prevention (CDC), United StatesRicardo Castillo-Neyra, University of Pennsylvania, United States
Copyright © 2025 Cunha Silva, Fellenberg, Freudenthal, Tiwari and Dürr. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Laura Cunha Silva, bGF1cmEuZGFzaWx2YUB1bmliZS5jaA==
†These authors have contributed equally to this work