# Behavioural adaptability of wild geladas (Theropithecus gelada) in human-impacted areas revealed by encounters with free-roaming dogs and potential threats

Luca Pedruzzi, Alice Galotti, Martina Francesconi, Alberto Quartesan, Sheleme Abiyou Gamessa, Bezawork Afework Bogale, Giulio Petroni, Valentina Serra, Alban Lemasson, Elisabetta Palagi

► To cite this version:

Luca Pedruzzi, Alice Galotti, Martina Francesconi, Alberto Quartesan, Sheleme Abiyou Gamessa, et al.. Behavioural adaptability of wild geladas (Theropithecus gelada) in human-impacted areas revealed by encounters with free-roaming dogs and potential threats. Global Ecology and Conservation, 2025, 62, pp.e03738. 10.1016/j.gecco.2025.e03738. hal-05165530

HAL Id: hal-05165530

https://hal.science/hal-05165530v1

Submitted on 16 Jul 2025

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
<figure><img src="image_2.png" /></figure>

ELSEVIER

Contents lists available at [ScienceDirect](https://www.sciencedirect.com)

Global Ecology and Conservation

journal homepage: [www.elsevier.com/locate/gecco](https://www.elsevier.com/locate/gecco)

<figure><img src="image_3.png" /></figure>

Short communication

# Behavioural adaptability of wild geladas (*Theropithecus gelada*) in human-impacted areas revealed by encounters with free-roaming dogs and potential threats

Luca Pedruzzi<sup>a,b,*</sup>, Alice Galotti<sup>b</sup>, Martina Francesconi<sup>b</sup>, Alberto Quartesan<sup>b</sup>, Sheleme Abiyou Gamessa<sup>c</sup>, Bezawork Afework Bogale<sup>c</sup>, Giulio Petroni<sup>b</sup>, Valentina Serra<sup>b</sup>, Alban Lemasson<sup>a,d,1</sup>, Elisabetta Palagi<sup>b,e,1</sup>

<sup>a</sup> EthoS (Ethologie Animale et Humaine) - U.M.R 6552, Université de Rennes, Université de Normandie, CNRS, Rennes 35000, France

<sup>b</sup> Department of Biology, University of Pisa, Pisa, Italy

<sup>c</sup> Department of Zoological Sciences, College of Natural and Computational Sciences, Addis Ababa University, Addis Ababa 1000, Ethiopia

<sup>d</sup> Institut Universitaire de France, 1 Rue Descartes, 75231 Paris Cedex 05, France

<sup>e</sup> Natural History Museum, University of Pisa, Calci, Pisa, Italy

## ARTICLE INFO

**Keywords:**

Biodiversity conservation

Endemic species

Ethiopia

Human-wildlife conflicts

Interspecies interactions

Primates

Dogs

## ABSTRACT

Studying interactions between domestic species and wild primates is crucial for understanding the impacts of human expansion on wildlife. While free-roaming domestic dogs are one of the most common human-introduced carnivores and pose potential threats to primates, their direct influence on primate behaviour remains largely overlooked. Geladas (*Theropithecus gelada*), a primate species endemic to Ethiopia’s Afro-alpine grasslands with a specialized diet, are facing challenges from habitat loss and interactions with domestic animals throughout their geographical range. Here, we followed 17 gelada group units in an unprotected area of Ethiopia (6-month period, 598 h of observation) and explored their behavioural responses to different natural and human-related threats. We observed strong vigilance in response to the presence of humans chasing nearby groups, as well as in response to dogs and eagles. Other raptors and birds of prey (including vultures and kites) produced no vigilance response, or only a modest one (e.g., augur buzzard). We also report encounters and interactions (n = 16) between dogs and geladas. Although we observed some behaviours associated with agonism and anxiety from both sides, the presence of dogs at close proximity was mainly tolerated. This contrasts with data from populations inhabiting protected areas where geladas typically exhibit flight responses to encounters with dogs. These differences between populations hint at gelada behavioural plasticity to adapt and live in human-impacted environments with limited alternative feeding sites. Increased awareness and documentation of dog-primate interactive events are essential to better grasp the broader implications of human-wildlife coexistence, one of the most important challenges of the Anthropocene.

<footnote>* Corresponding author at: EthoS (Ethologie Animale et Humaine) - U.M.R 6552, Université de Rennes, Université de Normandie, CNRS, Rennes 35000, France.</footnote>

<footnote>E-mail addresses: luca.pedruzzi@phd.unipi.it, luca.pedruzzi@etudiant.univ-rennes.fr (L. Pedruzzi).</footnote>

<footnote n="1">¹ shared senior authorship</footnote>
# 1. Introduction

Habitat degradation and other anthropic pressures strongly impact not only the distribution and survival of wild animals but also their behaviour, social dynamics and genetic viability (Lande, 1998; Giuntini and Pedruzzi, 2023; Wong and Candolin, 2015). Currently, most primate species are threatened by the impacts of habitat anthropization, mainly due to large-scale habitat loss from agriculture, ranching, resource extraction, and infrastructure expansion (Estrada et al., 2017). Studying how species cope with these constraints is important not only to preserve animals but also to understand their underlying cognitive plasticity and ability to react in a resilient way. From this perspective, focusing on species that are not generalists and live in very specific habitats can be even more informative about how they navigate human-disturbed environments (Chapman et al., 2016; Nautiyal et al., 2024; Neves et al., 2024; Nowak and Lee, 2013).

The gelada (Theropithecus gelada) is a cercopithecine primate endemic to Afro-alpine grasslands of Ethiopia at elevations from 2000 to 4400 m (Chiou et al., 2022). Among the several evolutionary derived traits of the species, the graminivorous diet represents the most important peculiarity with major conservation issues (Palagi and Bergman, 2021). Despite habitat and diet specialization, geladas evolved a complex social system offering them the opportunity for adaptability. The genus Theropithecus is mono-specific with the possible (taxonomically debated) presence of three subspecies: *T. g. gelada*, in the northern part of the country, *T. g. obscurus* inhabiting the central highlands, and the southern isolated population of Arsi geladas, tentatively named *T. g. arsi* (Caldon et al., 2024; Chiou et al., 2022; Shotake et al., 2016; Trede et al., 2020; Zinner et al., 2018).

Geladas live in multi-level societies (Grueter et al., 2020) and display a complex sociobiology (Le Roux and Bergman, 2012; Snyder-Mackler et al., 2012; Tinsley Johnson et al., 2014), including variable repertoires of visual and vocal communicative strategies (Gustison et al., 2019; Mancini et al., 2013; Pedruzzi et al., 2024; Zanoli et al., 2022). Despite the global gelada population being in decline, due to limited data on their geographic range and demographic trends, they are still classified by IUCN as 'Least Concern' (Gippoliti et al., 2018). Moreover, the species is expected to be vulnerable to climate change due to the loss of suitable habitats (Ahmed et al., 2023; Dunbar, 1998) and, out of protected habitats, most populations inhabit anthropogenically-modified landscapes undergoing several challenges, including habitat loss and persecution due to crop foraging behaviour (Kifle, 2023). Importantly, most of the available data on gelada behaviour come from populations living in areas with limited human impact and thus much information is missing about unprotected areas. Interestingly, some anectdotal data suggest differences in gelada anti-predator behaviours from population to population, hinting at possible inter-area (or inter-subspecies) differences in behavioural responses to natural and human-induced threats (Iwamoto et al., 1996).

Human-derived threats for wildlife include domestic species that can indirectly and directly compete with animal populations, as it occurs between grazing livestock and geladas (Abebe et al., 2022; Abie and Bekele, 2016; Kifle and Bekele, 2020). Human impact can also include conflicts via introduced predatory species like free-roaming domestic dogs. Dogs are the most widespread human-introduced carnivores, exhibiting varying degrees of free-ranging behaviour and frequently interacting with wildlife (Hughes and Macdonald, 2013; Sepúlveda et al., 2014). Dogs pose significant threats to wildlife conservation by increasing the risk of disease transmission (Lyngdoh et al., 2014). For primates, one of the most common dangers is direct killing, particularly when they travel on the ground (rhesus macaques, *Macaca mulatta*, Chakraborty et al., 2024; vervet monkeys, *Chlorocebus pygerythrus*, Chapman et al., 2016; black howler monkeys, *Alouatta pigra*, Pozo-Montuy et al., 2013). Even indirect cues, such as the dog barking, can negatively affect primate behavioural dynamics; for instance, dog vocalizations have been shown to elevate physiological stress and vigilance behaviours in howler monkeys (*Alouatta palliata*) (Rangel-Negrín et al., 2023). The presence of dogs can also induce behavioural shifts in primates; some species can adapt by avoiding humans and dogs to protect themselves and their feeding activities (blue monkeys, *Cercopithecus mitis*, Cords, 2014), for instance allocating more time to feeding and traveling and less to social behaviours (Himalayan langur, *Semnopithecus schistaceus*, Nautiyal et al., 2024). Other species may face disruptions in their behavioural repertoire due to dog harassment (long-tailed macaques, *Macaca fascicularis*, Gumert et al., 2013).

Despite having become rather ubiquitous, interspecies encounters between dogs and primates remain poorly studied, likely due to the challenges involved in observing these interactions and to the bias against publishing anecdotal data (Ramsay and Teichroeb, 2019; Waters et al., 2023). In many cultivated unprotected areas in Ethiopia, where high competition arises between geladas raiding the crops and local human populations (Kifle, 2023), dogs are often used, with variable efficacy, to chase and eventually kill the geladas (Abie and Bekele, 2016; Abu et al., 2018; Gippoliti et al., 2018; Kifle and Bekele, 2020; Yihune et al., 2009). Geladas have been reported to generally show immediate flight responses to dogs in the Guassa Community Conservation Area (Venkataraman et al., 2015) with no tolerance for co-presence with dogs.

Here, we followed several gelada groups inhabiting an unprotected area of the central highlands of Ethiopia (Debre Libanos). First, we analysed the behavioural vigilance of the study groups to different natural (i.e., different species of birds of prey) vs anthropogenic (i.e., human chasing groups on the opposite side of the cliff, free-roaming dogs) heterospecific encounters to investigate the possible variability in the perception of threats associated to human direct or indirect presence. Then, we report and describe a series of video-recorded encounters between the monkeys and free-roaming dogs to assess their tolerance level, here indicated by the co-presence of both species in the grazing area. Through this report, we aim to highlight the behavioural adaptability and resilience of geladas while emphasizing the differences that seem to emerge from different populations inhabiting protected or more pristine habitats.
## 2. Methods

### 2.1. Study site and scan-sampling

The study was carried out between February-April and October-December 2024 in the unprotected area of Debre Libanos, in the central highlands of Ethiopia (Northwest Shewa zone, Oromia Region) (Abie and Bekele, 2017). Specifically, 18 group units (One-Male Units, OMUs) familiar to the presence of researchers (Galotti et al., 2025) were followed from 8AM to 4PM during the study period, as they daily visited two grazing areas in Set Deber (Site 1, February-April, 11 OMUs) and Godo (Site 2, October-December, 7 OMUs), two sites inhabited by gelada groups (Fig. 1a). The area has undergone a significant increase in human population and activities such as livestock grazing and agriculture (Abie and Bekele, 2016). We used scan sampling at 15-min intervals recording the number and identity of the OMUs present on the site, the time of day, the study site, and the presence of other species (humans, livestock, dogs, other primates) within an area of about 200 m from the geladas. The 200-meter radius was chosen as it represented the maximum distance within which we could reliably detect and identify humans, livestock, dogs, and other species in the field, given the local topography. This choice ensured consistency in data collection across scans and reflects a methodological threshold aimed at maximizing observational accuracy.

### 2.2. Reactions to potential threats

We coded the behavioural responses of geladas to the following species: i) birds of prey/raptors flying over the herd of monkeys, ii) humans chasing non-study gelada groups on nearby cliffs visible and audible by study groups, iii) dogs entering the grazing area (either feral, living around human settlements or belonging to someone but left loose). According with literature on the avifauna in the Debre Libanos area (Desalegn and Negussie, 2022), we recorded the presence of yellow-billed kite (Milvus aegyptius), hooded (Necrosyrtes monachus), white-backed (Gyps africanus) and bearded (Gypaetus barbatus) vultures, augur buzzard (Buteo augur), Verreaux (Aquila verreauxii), black-chested snake (Circaetus pectoralis), and tawny eagle (Aquila rapax). We noted the time/day, the weather (sunny (S), partially clouded (PC), clouded (C)), the number of OMUs present in the herd, and at least 10 % of the subjects in the groups showed vigilance, which can be defined as the presence of at least one of the following behaviours (Allan and Hill, 2018): looking towards the potential threat, alarm calling, moving away, standing on hindlimbs while looking, often occurring with geladas lifting their head towards the stimulus and pausing the typical hand movements associated with gelada grazing behaviour (Pedruzzi et al., 2025). The 10 % threshold was selected as a conservative criterion to distinguish group-level responses from individual or incidental behaviours, allowing us to focus on events likely perceived as meaningful threats by a significant portion of the group. As geladas were generally grazing during the observations and as their feeding implies a seated position with their head facing the ground (Dunbar, 1977), this allowed us to easily observe a change in the gaze direction of the animals towards the potential threats.

### 2.3. Data analyses

We ran a Generalized Linear Mixed Model (GLMM) to investigate which factors affected the likelihood of vigilance (presence/

<figure><img src="image_4.png" /><figcaption>**Fig. 1.** a) Map of the two study sites in Debre Libanos (Ethiopia, 9.711944°N, 38.8475°E). We generated the red and blue areas by connecting the GPS points recorded during scan-sampling. b) Bar plot indicating the percentage of observation time (obtained from 15-min scan sampling), with humans or livestock present (red part) on the grazing areas used by gelada groups followed.</figcaption></figure>
absence distribution = binomial). The fixed factors included were the i) *Type of potential threat* (YB Kites/Eagles/Vultures/Augur buzzards/Humans/Dogs), ii) *Number of OMUs*, iii) *Weather* (S/PC/C), and iv) *Time of the day* (rounded to the nearest full hour, continuous numeric variable). The *Day* was included as random factor. Multicollinearity was assessed via the check collinearity function from the performance package (version 0.4.4) using Variance Inflation Factors (VIFs: 1.42–1.55). The fixed factors in the models showed low correlation. Model significance was evaluated by comparing the full model to a null model, including only the random effect using the Likelihood Ratio Test (LRT) with the Chisq test argument (Dobson and Barnett, 2018). To determine the p-value of predictors, LRTs were performed between the full model and a model without the specific predictor, using the Anova function (Barr et al., 2013). For pairwise comparisons involving factors with more than two levels, the emmeans package was used to perform the Tukey test (Bretz et al., 2016; Lenth et al., 2021).

## 2.4. Video recording of encounters with dogs

In the event of a dog sighting, all dog and gelada behaviours and interactions were video recorded on an *ad libitum* basis (Altmann, 1974). The video recording went in parallel with scan sampling; each encounter began with the arrival of dog(s) in the grazing area of geladas and finished when the dog(s) left for at least 30 min. Data collected during these encounters included the time of day, the number of dogs and OMUs/geladas present, as well as estimates of the dogs' size, sex, and their proximity to the nearest gelada. Additionally, by a continuous group focal sampling, we also coded from video recordings the possible presence of chases or attempted physical interactions by the dogs, flight responses by geladas toward the nearest cliffs, and other interspecific interactions such as alarm-calling, scratching, yawning, and various visual and vocal signals exchanged between dogs and geladas.

# 3. Results

## 3.1. Scan-sampling

The study groups were observed for a total of 598 h (2389 scans, 1502 in February-April 2024, 887 in October-December 2024). Geladas and domestic dogs were co-present in the same area in 22 scans (5.5 h, 16 scans in Site 1, 6 scans in Site 2). To further testify the level of anthropogenic impact of the area, our scan sampling revealed that in about 70 % of observation time (1749/2469 scans) humans were present around the groups, while livestock was present in about 78 % of observation time (1902/2469 scans) (Fig. 1b).

## 3.2. Reaction to potential threats

The total encounters with potential threats were 214 (augur buzzard, n = 7, dogs, n = 12, eagles, n = 6, humans, n = 40, vultures,

<figure><img src="image_5.png" /></figure>

**Fig. 2.** a) Error plots representing the likelihood of vigilance behaviours (mean ± SE) shown by the focal gelada groups in response to different anthropic vs natural potential threats (see also Table 1); b) Mosaic plot representing the different possible vigilance responses elicited by the different anthropogenic vs natural potential threats, namely No reaction (in green), gazing behaviour (vigilance, in dark yellow), gazing behaviour together with alarm and moving (vigilance + alarm/moving, in red). The width of the bars (x-axis) is proportional to the number of observations for each type of potential threat.
n = 62, yellow-billed kite, n = 87). The full model significantly differed from the null one ($X_5^2$= 81.54, p < 0.0001). The Type of potential threat ($X_4^2$=17.88, p = 0.003) significantly affected the likelihood of vigilance (Fig. 2a, Table 1). Geladas were more likely to display vigilance behaviours in response to humans chasing other gelada groups (humans vs kites: estimate=5.182, p = 0.001; humans vs vultures: estimate=5.652, p = 0.012), dogs (dogs vs kites: estimate=5.643, p = 0.007; dogs vs vultures: estimate=6.113, p = 0.026) arriving in the grazing area or to eagles (eagles vs kites: estimate=5.941, p = 0.023; eagles vs vultures: estimate=6.412, p = 0.050) compared to yellow-billed kites or vultures flying over them (Fig. 2a). Fig. 2b represents the type of behavioural responses recorded (only looking vs looking and alarm-calling or moving further from potential threat) according to the species encountered.

### 3.3. Video recording of encounters with dogs

Table 2 summarizes the key information about the encounters. We could individually recognize 13 different dogs in 16 different interactions with geladas, occurring on 12 different days across the study period and lasting up to about 13 min (Table 2). As soon as dogs entered the grazing area, either alone (n = 13) or in small groups (n = 3), they approached geladas. In all cases dogs came closer than 20 m to geladas (Fig. 3). Interestingly, geladas never showed flight responses at the arrival of dogs and never left the grazing area despite attempts of chasing by the dogs. Although in two encounters the arrival of dogs led to geladas moving about 20 m, they did not escape to the cliff and eventually got closer to the dogs after few minutes. In few cases, the adult gelada male of the OMUs even chased back or threatened the dog (eyebrow raise, Lazow and Bergman, 2020). Dogs chasing the monkeys (n = 7 cases) were always males, and the monkeys generally remaining closer to the dogs were males, either subadults or adults. Although there were only two instances in which more than one adult dog approached the geladas, in both cases the dogs consistently chased the monkeys. However, the geladas' response did not appear to vary substantially between encounters with single dogs and those with groups (Table 2). Signals displayed by dogs during encounters included barking, growling, attempts to bite, and play bowing, whereas geladas typically responded with eyebrow raising (threats), jaw fencing/bared-teeth lip-flip when chasing back the dog, and self-scratching/yawning (Table 2).

## 4. Discussion

Studying encounters between domestic species and wild primates is crucial to understand side-effects of human expansion in rural areas overlapping with threatened or endemic species that, in turn, will need to behaviourally adapt to such new challenges (Waters et al., 2023). Despite being the most widespread human-introduced carnivore (Hughes and Macdonald, 2013), the direct influence of dogs on the behaviour of wild primates has not been extensively explored (Nautiyal et al., 2024).

The preliminary part of our report highlighted the anthropogenic impact on our project field site, as humans and livestock were present around the study groups of geladas (Theropithecus gelada) in ~70% and ~78% of the observation time respectively, with possible negative effects on geladas due to grazing competition (Abebe et al., 2022). We also report the behavioural responses of geladas to possible natural vs anthropogenic threats. We show that the strongest vigilance reactions (looking, alarm calling, moving) were in response to humans chasing nearby groups (other side of the cliff) and to dogs entering the grazing area compared to birds of prey flying over (i.e., kites, vultures, buzzards). Increased vigilance in response to dogs and humans may reduce time available for foraging, potentially affecting energy intake and fitness. Similar trade-offs have been documented in other primates (Teichroeb and Sicotte, 2012), where vigilance during feeding incurs measurable costs. Such behavioural shifts may also reflect underlying stress, especially in human-modified landscapes. Despite the low number of encounters, eagles elicited geladas' vigilance responses. This result fits with the data on the predatory role of eagles on other phylogenetically close monkeys such as hamadryas baboons (Zinner and Peláez, 1999). This result may also indicate that geladas recognize birds at the species level based on visual characteristics, such as

<caption>**Table 1**</caption>

Estimated parameters (Coeff), Standard Error (SE), and results of the Likelihood Ratio Tests ($\chi^2$) of the GLMM. Significant P values are bold; df= degree(s) of freedom; - = not applicable.

<table><thead><tr><th>Fixed Effects</th><th>Coeff</th><th>SE</th><th>χ<sup>2</sup></th><th>df</th><th>P</th></tr></thead><tbody><tr><td colspan="6"><strong>Model - Likelihood of vigilance responses to potential threats</strong></td></tr><tr><td>Intercept</td><td></td><td></td><td>-</td><td>-</td><td>-</td></tr><tr><td colspan="6"><strong>Tested variables</strong></td></tr><tr><td>Type of potential threat<sup>a</sup></td><td></td><td></td><td>12.15</td><td>5</td><td>0.033</td></tr><tr><td>Type of potential threat (Dogs)</td><td>4.748</td><td>2.494</td><td></td><td></td><td></td></tr><tr><td>Type of potential threat (Eagles)</td><td>4.571</td><td>2.679</td><td></td><td></td><td></td></tr><tr><td>Type of potential threat (Humans)</td><td>3.884</td><td>2.143</td><td></td><td></td><td></td></tr><tr><td>Type of potential threat (Vultures)</td><td>-21.296</td><td>16170.900</td><td></td><td></td><td></td></tr><tr><td>Type of potential threat (YB Kite)</td><td>-1.257</td><td>1.706</td><td></td><td></td><td></td></tr><tr><td>Number of OMUs</td><td>0.138</td><td>0.289</td><td>0.23</td><td>1</td><td>0.631</td></tr><tr><td>Weather<sup>a</sup></td><td></td><td></td><td>0.74</td><td>2</td><td>0.690</td></tr><tr><td>Weather (PC)</td><td>-1.668</td><td>2.474</td><td></td><td></td><td></td></tr><tr><td>Weather (S)</td><td>-2.034</td><td>2.461</td><td></td><td></td><td></td></tr><tr><td>Time of the day</td><td>-0.181</td><td>0.331</td><td>0.30</td><td>1</td><td>0.584</td></tr></tbody></table>

$N_{observations}$= 214, $N_{days}$= 37. Random factors: Day, Variance= 3.6, SD= 1.9.

<footnote>aThese predictors were dummy coded, with “Type of potential threat (Augur buzzard)” and “Weather (Clouded, C)” being the reference categories.</footnote>
<table><caption>**Table 2**
Summary of key characteristics of the 16 encounters between geladas and dogs at Debre Libanos, Ethiopia recorded in February-April and October-December 2024.</caption><thead><tr><th>Encounter</th><th>Date, hour</th><th>Site</th><th>Geladas present (groups, subjects)</th><th>N° of dogs</th><th>Size of dog(s) compared to a gelada adult male</th><th>Micro-habitat</th><th>Dog (s) sex</th><th>Dog chasing geladas/trying to physically interact</th><th>Flight response of geladas</th><th>Description of the interaction</th><th>Duration of interaction (mm:ss)</th><th>Visual signals</th><th>Vocalisations/alarm calls</th></tr></thead><tbody><tr><td>1</td><td>15/02/ 2024, 9:40 am</td><td>1</td><td>3, 44</td><td>1</td><td>Smaller</td><td>Livestock grazing area</td><td>?</td><td>No</td><td>No</td><td>A dog approaches geladas at &lt; 20 m while they continue feeding activity</td><td>03:48</td><td>0</td><td>0</td></tr><tr><td>2</td><td>17/02/ 2024, 1:02 pm</td><td>1</td><td>4, 69</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>F</td><td>No</td><td>No</td><td>A female dog approaches geladas at &lt; 10 m; geladas continue grazing; the adult male of the closest group yawns while two young subjects start self-scratching. Eventually the dog goes away.</td><td>12:58</td><td>0</td><td>0</td></tr><tr><td>3</td><td>17/02/ 2024, 2:30 pm</td><td>1</td><td>4, 77</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>No</td><td>No</td><td>A dog approaches the adult gelada male of a group at &lt; 5 m while he continues feeding activity</td><td>02:35</td><td>0</td><td>0</td></tr><tr><td>4</td><td>23/02/ 2024, 8:10 am</td><td>1</td><td>1, 22</td><td>4</td><td>1 small, 3 comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>Male dogs approaching on the group of geladas at &lt; 5 m and trying to chase without biting, the adult gelada male threaten and tries to chase them back; geladas move few meters away and restart grazing; the geladas closest to dogs yawn and self-scratch</td><td>08:52</td><td>Geladas: eyebrow raiser (threat)</td><td>0</td></tr><tr><td>5</td><td>5/3/2024, 2:47 am</td><td>1</td><td>3, 57</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>F</td><td>No</td><td>No</td><td>A female dog approaches geladas at &lt; 10 m; geladas continue grazing</td><td>04:36</td><td>0</td><td>0</td></tr><tr><td>6</td><td>11/03/ 2024, 11:03 am</td><td>1</td><td>2, 38</td><td>2</td><td>1 small, 1 comparable</td><td>Livestock grazing area</td><td>F, ?</td><td>No</td><td>No</td><td>A female dog and her cub approaches geladas at &lt; 20 m while they continue feeding activity</td><td>07:13</td><td>0</td><td>0</td></tr><tr><td>7</td><td>12/03/ 2024, 8:49 am</td><td>1</td><td>1, 22</td><td>3</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>Dogs approach geladas at &lt; 20 m and then try to chase the adult male of the group; geladas move few metres away</td><td>07:10</td><td>0</td><td>0</td></tr><tr><td>8</td><td>13/03/ 2024, 12:40 pm</td><td>1</td><td>3, 57</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>A male dog approaches a subadult gelada male at &lt; 5 m; the subadult moves away to his group and the dog chase them; the adult male of the group</td><td>8:48</td><td>Geladas: eyebrow raiser (threat)</td><td>0</td></tr></tbody></table>

(continued on next page)
<table><caption>Table 2 (continued)</caption><thead><tr><th>Encounter</th><th>Date, hour</th><th>Site</th><th>Geladas present (groups, subjects)</th><th>N° of dogs</th><th>Size of dog(s) compared to a gelada adult male</th><th>Micro-habitat</th><th>Dog (s) sex</th><th>Dog chasing geladas/trying to physically interact</th><th>Flight response of geladas</th><th>Description of the interaction</th><th>Duration of interaction (mm:ss)</th><th>Visual signals</th><th>Vocalisations/alarm calls</th></tr></thead><tbody><tr><td>9</td><td>17/03/ 2024, 1:42 pm</td><td>1</td><td>3, 48</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>No</td><td>No</td><td>A male dog approaches geladas at &lt; 20 m; geladas continue grazing</td><td>00:55</td><td>0</td><td>0</td></tr><tr><td>10</td><td>22/03/ 2024, 12:30 pm</td><td>1</td><td>1, 19</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>F</td><td>No</td><td>No</td><td>A female dog approaches geladas at &lt; 10 m; geladas continue grazing but the closest to dogs self-scratch</td><td>00:24</td><td>0</td><td>0</td></tr><tr><td>11</td><td>26/03/ 2024, 8:25 am</td><td>1</td><td>5, 74</td><td>1</td><td>Smaller</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>A male dog approaches geladas very closely (&lt;2 m) while they slowly move away and then chase them; two subadult try to chase back the dog; geladas closest to dogs self-scratch</td><td>00:48</td><td>0</td><td>Dog: barking</td></tr><tr><td>12</td><td>26/03/ 2024, 9:31 am</td><td>1</td><td>5, 74</td><td>1</td><td>Smaller</td><td>Livestock grazing area</td><td>M</td><td>No</td><td>No</td><td>A male dog approaches and barks very close (&lt;2 m) to a subadult gelada male; the gelada self-scratch but continues feeding (Fig. 3d)</td><td>03:29</td><td>0</td><td>Dog: barking</td></tr><tr><td>13</td><td>14/11/ 2024, 10:47 am</td><td>2</td><td>7, 111</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>A male dog chases the geladas who move slowly and then stay in proximity (&lt;10 m) for several minutes</td><td>00:55</td><td>0</td><td>0</td></tr><tr><td>14</td><td>25/11/ 2024, 9:37 am</td><td>2</td><td>5, M, d, b, c, r</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>A male dog repeatedly tries to physically interact with an adult gelada male from a very close distance (&gt;1 m, Fig. 3b) trying to bite him and play-bowling; the gelada yawns and then chases the dog away.</td><td>05:12</td><td>Gelada: jaw fencing, eyebrow raiser (threat); Dog: play bows</td><td>Dog: growling</td></tr><tr><td>15</td><td>26/11/ 2024, 8:53 am</td><td>2</td><td>4, 71</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>Yes</td><td>No</td><td>A male dog repeatedly tries to physically interact with an adult gelada male from a very close distance (&gt;1 m); the gelada yawns, then he chases the dog away (Fig. 3c)</td><td>03:14</td><td>Gelada: jaw fencing, eyebrow raiser (threat); Dog: play bow</td><td>0</td></tr><tr><td>16</td><td>26/11/ 2024, 9:18 am</td><td>2</td><td>5, 87</td><td>1</td><td>Comparable</td><td>Livestock grazing area</td><td>M</td><td>No</td><td>No</td><td>A male dog approaches geladas at &lt; 10 m; geladas continue grazing</td><td>01:23</td><td>0</td><td>0</td></tr></tbody></table>

Number of individually recognized dogs= 13
<figure><img src="image_6.png" /><figcaption>**Fig. 3.** Screenshots from video recordings of gelada and free-roaming dog encounters, showcasing the variability in the nature of these interactions. The images illustrate moments of close co-presence (a), instances of dog disturbance and aggressive behaviour (b), as well as chasing between geladas and dogs (c).</figcaption></figure>

flight patterns, silhouette, and bird size, with a subsequent association with risk perception. A more in-depth analysis of these aspects could provide interesting insights into the role of avian predation in the behavioural ecology of primates and on the evolution of their sensory modalities' sensibility.

The high vigilance reactions of geladas towards humans and dogs in the field site confirm that geladas perceive these two species as a real threat. Beyond our focal observations, a growing body of research highlights the diverse ways in which free-ranging dogs influence primate behaviour and ecology across species and contexts (Hughes and Macdonald, 2013; Waters et al., 2023). In various environments, dogs have been associated with direct predation events (Chakraborty et al., 2024; Chapman et al., 2016), as well as with subtler disruptions such as elevated stress levels and increased vigilance (Rangel-Negrín et al., 2023) and shifts in time budgets (Nautiyal et al., 2024), while others show signs of harassment and altered social dynamics (Gumert et al., 2013). Although these findings reflect species-specific and context-dependent responses, they underline a broader, underexplored conservation issue.

Then, we further examined the diversity of behaviours associated with encounters between geladas and free-roaming dogs. These interactions offered a rare yet valuable opportunity to gain deeper insights into gelada responses and adaptive strategies (Waters et al., 2023). Indeed, here we report and describe a series of video-recorded interactions between free-roaming dogs (n = 16) and wild geladas, representing the largest number of such encounters documented to date in an unprotected area. Interestingly, and in contrast
with previous reports on geladas in protected areas (Iwamoto et al., 1996; Venkataraman et al., 2015), although the encounters were mostly of slightly negative valence, the monkeys never showed flight responses to the arrival of dogs, even when they approached within 10 m and attempted to chase geladas. In Ethiopia, especially in agricultural landscapes, dogs are actively employed to deter geladas from crops (Abie and Bekele, 2016; Abu et al., 2018; Kifle and Bekele, 2020). Yet, behavioural responses seem to vary: despite the few possible comparisons, previous studies on geladas reported high rates of flight responses in populations inhabiting protected areas (Guassa, Venkataraman et al., 2015) or in more isolated sites from human settlements (northern populations, Iwamoto et al., 1996), hinting at possible ecological or genetic factors affecting the observed behavioural variability. Dogs displayed barking, growling, and play-bowing, while geladas responded with eyebrow raising, jaw fencing, and self-scratching, all behaviours that are proxies of negative emotional states (Lazow and Bergman, 2020; Maestripieri et al., 1992). Moreover, several adult gelada males occasionally threatened or chased back dogs. The behavioural responses to dogs indicate a mix of flight and fight reactions, likely influenced by the level of perceived danger. Nevertheless, co-presence of geladas and dogs never appeared as a major concern. Such surprising tolerance may stem from the limited availability of alternative feeding sites in the Debre Libanos area, which could heighten the geladas' motivation to stay despite the presence of dogs. Indeed, the extent to which prey respond to cues from potential predators depends on both their ecological (i.e., present-day) and evolutionary (i.e., historical) exposure to predation (Hettena et al., 2014), with the likelihood of losing such discrimination skills increasing when defensive responses impose significant energetic costs (Lahti et al., 2009). Whether geladas are simply habituated to the regular presence of dogs, or whether certain gelada populations adapted over time to specific dogs' activities remains to be explored. In particular, further research should investigate the factors influencing tolerance towards dogs such as dog role (the effectiveness of using dogs to hunt geladas is reported to vary), dog individual features (e. g., age, sex, size) and the type of human management (e.g., leashed, free-roaming, human-fed, or feral, Sepúlveda et al., 2014). Here, comparing the size and number of dogs involved in past studies could help assess whether these factors influenced the differences in their reactions. Additionally, it is possible that our gelada study population has not experienced recent successful predation events by dogs, reducing their perceived risk. The level of care and nutrition provided to dogs by local human populations could also be a relevant factor (Sepúlveda et al., 2014). Finally, another aspect worth exploring is whether geladas adapt not just to the general presence of dogs but even to specific individuals known to the groups frequenting a specific grazing area of the highland. Future studies could also expose the monkeys to unfamiliar dogs to distinguish between a general habituation to free-roaming dogs and the recognition of familiar individuals. A similar approach could also be applied to human presence, testing gelada reactions to strangers compared to individuals regularly present in their habitat.

While our findings provide valuable insights into gelada responses to anthropogenic stressors, some limitations must be acknowledged. First, the relatively small number of observed dog encounters limits the generalizability of our results. Second, the absence of longitudinal data prevents us from assessing potential long-term effects of these interactions on gelada behaviour, such as habituation or cumulative stress. Future studies with broader temporal coverage and experimental designs could help overcome these constraints.

Whatever the adaptive mechanism at play, the behavioural resilience shown by geladas might be one of the key strategies allowing the endemic species to persist despite the high anthropogenic changes their habitats are going through (Kifle, 2023). Reports on dog-primate interactions are crucial yet often overlooked to understand animal behavioural flexibility in the Anthropocene, as they offer a valuable model to test the coping strategies of primates to anthropogenic pressures (Waters et al., 2023). Increased awareness and documentation of these events are essential to better grasp the broader implications of human-wildlife coexistence.

## Ethics

Even though the study was completely non-invasive, formal approval for the observations and protocols used throughout the project was sought and obtained from the Bioethical Committee of the University of Pisa. Moreover, research procedures strictly adhered to the approved guidelines of the Ethiopian Wildlife Conservation Authority (EWCA).

## Funding

The research has been funded by the Leakey Foundation (Science for reconciliation: What an Ethiopian monkey tells about peace-making, grant n° S202310431) and by the following zoos and foundations (funders of BRIDGES project, UNIPI, AOO "BIO" - 0005878/ 2022), in alphabetic order: Dudley Zoo (UK), Fondazione ARCA (Italy), Giardino Zoologico di Pistoia (Italy), NaturZoo Rheine (Germany), Parc des Félins (France), Parco Natura Viva (Italy), Parco Zoo Falconara (Italy), Rotterdam Zoo (The Netherlands), Saint-Félicien (Canada), Wildlife Conservation Benefit (Italy), Wilhelma Zoo (Germany), Zoo de Cerza (France), Zoo de La Boissière du doré (France). Rennes Métropole and the Région Bretagne (France) covered the field expenses for the first author while the University of Pisa and University of Rennes funded student fellowships.

## CRediT authorship contribution statement

**Sheleme Abiyou Gamessa:** Investigation, Writing – review & editing, Methodology. **Bezawork Afework Bogale:** Funding acquisition, Writing – review & editing, Project administration. **Martina Francesconi:** Formal analysis, Investigation, Writing – review & editing, Methodology, Data curation. **Alberto Quartesan:** Methodology, Data curation, Writing – review & editing, Investigation. **Valentina Serra:** Writing – review & editing, Funding acquisition. **Alban Lemasson:** Writing – review & editing, Investigation, Supervision, Conceptualization, Formal analysis, Methodology. **Giulio Petroni:** Funding acquisition, Writing – review & editing. **Luca
**Pedruzzi:** Formal analysis, Visualization, Data curation, Writing – review & editing, Methodology, Writing – original draft, Investi-
gation, Conceptualization. **Alice Galotti:** Writing – review & editing, Investigation, Methodology, Conceptualization, Visualization,
Data curation. **Elisabetta Palagi:** Supervision, Formal analysis, Project administration, Data curation, Writing – review & editing,
Funding acquisition, Conceptualization.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to
influence the work reported in this paper.

Acknowledgments

We extend our gratitude to the Ethiopian Wildlife Conservation Authority (EWCA) and the Oromia Forest and Wildlife Service for
granting us permission to conduct our research, as well as to Debre Libanos Woreda and Monastery for their support during our stay in
the community. We are especially grateful to Achim Johann, coordinator of the Gelada EEP, for his invaluable assistance in fund-
raising. Our heartfelt thanks go to Hailu Tilahun for the support in the field, as well as to Birhanu and, in particular, Kabebew Asefa
Ylma for always looking after us during fieldwork. Lastly, we sincerely thank the community and the children of Set Deber and Godo
for their respect for our research and for the wonderful moments we shared—we will always cherish your kindness.

**Appendix A. Supporting information**

Supplementary data associated with this article can be found in the online version at doi:10.1016/j.gecco.2025.e03738.

Data availability

The raw data used for the study has been uploaded as Supplementary Material.

References

Abebe, B., Ashagrie, M., Moges, E., 2022. The activity and ranging patterns of gelada monkeys as behavioral responses to the effects of livestock grazing in the Simien Mountains National Park, Ethiopia. Int. J. Zool. 2022, 1–12. https://doi.org/10.1155/2022/8107527

Abie, K., Bekele, A., 2016. Threats to Gelada Baboon (Theropithecus gelada) around Debre Libanos, Northwest Shewa Zone, Ethiopia. Int. J. Biodivers. 2016, 1–7.
https://doi.org/10.1155/2016/3405717.

Abie, K., Bekele, A., 2017. Population estimate, group size and age structure of the Gelada Baboon (Theropithecus gelada) around Debre-Libanos, Northwest Shewa Zone, Ethiopia. Glob. J. Sci. Front. Res. 17 (1), 27–33.

Abu, K., Mekonnen, A., Bekele, A., Fashing, P.J., 2018. Diet and activity patterns of Arsi geladas in low-elevation disturbed habitat south of the Rift Valley at Indetu, Ethiopia. Primates 59 (2), 153–161. https://doi.org/10.1007/s10329-017-0640-9.

Ahmed, A.S., Chala, D., Kufa, C.A., Atickem, A., Bekele, A., Svenning, J.-C., Zinner, D., 2023. Potential changes in the extent of suitable habitats for geladas (Theropithecus gelada) in the Anthropocene. BMC Ecol. Evol. 23 (1), 65. https://doi.org/10.1186/s12862-023-02173-3.

Allan, A.T.L., Hill, R.A., 2018. What have we been looking at? A call for consistency in studies of primate vigilance. Am. J. Phys. Anthropol. 165 (S65), 4–22. https://doi.org/10.1002/ajpa.23381.

Altmann, J., 1974. Observational study of behavior: sampling methods. Behaviour 49 (3-4), 227-266. https://doi.org/10.1163/156853974X00534.

Barr, D.J., Levy, R., Scheepers, C., & Tily, H.J. (2013). *Random effects structure for confirmatory hypothesis testing: Keep it maximal*. https://doi.org/10.1016/j.jml.2012.11.001.

Bretz, F., Hothorn, T., Westfall, P., 2016. Multiple Comparisons Using R. CRC Press.

Caldon, M., Mutti, G., Mondanaro, A., Imai, H., Shotake, T., Oteo Garcia, G., Belay, G., Morata, J., Trotta, J.R., Montinaro, F., Gippoliti, S., Capelli, C., 2024. Gelada genomes highlight events of gene flow, hybridisation and local adaptation that track past climatic changes. Mol. Ecol. 6, 1–23. https://doi.org/10.1111/mec.17514.

Chakraborty, B., Pithva, K., Mohanty, S., McCowan, B., 2024. Lethal dog attacks on adult rhesus macaques (Macaca mulatta) in an anthropogenic landscape. Primates 65 (3), 151–157. https://doi.org/10.1007/s10329-024-01122-y.

Chapman, C.A., Twinomugisha, D., Teichroeb, J.A., Valenta, K., Sengupta, R., Sarkar, D., Rothman, J.M., 2016. How do primates survive among humans? Mechanisms employed by vervet monkeys at Lake Nabugabo, Uganda. In: Waller, M. (Ed.), Ethnoprimatology. Developments in Primatology: Progress and Prospects. Springer, Cham, pp. 77–94. https://doi.org/10.1007/978-3-319-30469-4-5.

Chiou, K.L., Janiak, M.C., Schneider-Crease, I.A., Sen, S., Ayele, F., Chuma, I.S., Knauf, S., Lemma, A., Signore, A.V., D'Ippolito, A.M., Abebe, B., Haile, A.A., Kebede, F., Fashing, P.J., Nguyen, N., McCann, C., Houck, M.L., Wall, J.D., Burrell, A.S., Snyder-Mackler, N., 2022. Genomic signatures of high-altitude adaptation and chromosomal polymorphism in geladas. Nat. Ecol. Evol. 6 (5), 630–643. https://doi.org/10.1038/s41559-022-01703-4.

Cords, M., 2014. Blue monkeys and bridges: Transformations in habituation, habitat and people. In: Strier, K.B. (Ed.), Primate Ethnographies. Pearson Education Inc, pp. 207–217.

Desalegn, T., Negussie, F., 2022. Diversity, species composition, and relative abundance of Avifauna at Debre Libanos Monastery, Ethiopia. Int. J. Zool. 2022, 1–9.
https://doi.org/10.1155/2022/8697842.

Dobson, A.J., Barnett, A.G., 2018. An Introduction to Generalized Linear Models, 4th ed.). CRC Press.

Dunbar, R.I.M., 1977. Feeding ecology of gelada baboons: a preliminary report. In: Clutton-Brock, T.H. (Ed.), Primate Ecology: Studies of Feeding and Ranging Behavior in Lemurs, Monkey and Apes. Academic Press, pp. 251–273. https://doi.org/10.1016/B9780-12-176850-8.50014-7.

Dunbar, R.I.M., 1998. Impact of global warming on the distribution and survival of the gelada baboon: a modelling approach. Glob. Change Biol. 4 (3), 293–304.
https://doi.org/10.1046/j.1365-2486.1998.00156.x.
Estrada, A., Garber, P.A., Rylands, A.B., Roos, C., Fernandez-Duque, E., Di Fiore, A., Nekaris, K.A.-I., Nijman, V., Heymann, E.W., Lambert, J.E., Rovero, F., Barelli, C., Setchell, J.M., Gillespie, T.R., Mittermeier, R.A., Arregoitia, L.V., de Guinea, M., Gouveia, S., Dobrovolski, R., Li, B., 2017. Impending extinction crisis of the world's primates: Why primates matter. Sci. Adv. 3 (1). https://doi.org/10.1126/sciadv.1600946.

Galotti, A., Francesconi, M., Pedruzzi, L., Abiyou Gamessa, S., Lemasson, A., Bogale, B.A., Palagi, E., 2025. Bridging species divides: affiliative interactions between theropithecus gelada and Papio anubis in an unprotected area in Ethiopia. Int. J. Primatol. 46 (1), 4-8. https://doi.org/10.1007/s10764-024-00450-6.

Gippoliti, S., Mekonnen, A., Burke, R., Nguyen, N., Fashing, P.J., 2018. Theropithecus gelada. IUCN Red. List Threat. Species. https://doi.org/10.2305/IUCN.UK.2019-3.RLTS.T21744A17941908en.

Giuntini, S., Pedruzzi, L., 2023. Sex and the patch: the influence of habitat fragmentation on terrestrial vertebrates' mating strategies. Ethol. Ecol. Evol. 35 (3), 269-298. https://doi.org/10.1080/039493702.2022.2059787.

Grueter, C.C., Qi, X., Zinner, D., Bergman, T., Li, M., Xiang, Z., Zhu, P., Migliano, A.B., Miller, A., Krützen, M., Fischer, J., Rubenstein, D.I., Vidya, T.N.C., Li, B., Cantor, M., Swedell, L., 2020. Multilevel organisation of animal sociality. Ecol. Evol. 35 (9), 834-847. https://doi.org/10.1016/j.tree.2020.05.003.

Gumert, M.D., Hamada, Y., Malaijvijntond, S., 2013. Human activity negatively affects stone tool-using Burmese long-tailed macaques Macaca fascicularis aurea in Laem Son National Park, Thailand. Oryx 47 (4), 535-543. https://doi.org/10.1017/S0030605312000130.

Gustison, M.L., Tinsley Johnson, E., Beehner, J.C., Bergman, T.J., 2019. The social functions of complex vocal sequences in wild geladas. Behav. Ecol. Sociobiol. 73 (1), 14. https://doi.org/10.1007/s00265-018-2612-5.

Hettena, A.M., Munoz, N., Blumstein, D.T., 2014. Prey responses to predator's sounds: a review and empirical study. Ethology 120 (5), 427-452. https://doi.org/10.1111/eth.12219.

Hughes, J., Macdonald, D.W., 2013. A review of the interactions between free-roaming domestic dogs and wildlife. Biol. Conserv. 157, 341-351. https://doi.org/10.1016/j.biocon.2012.07.005.

Iwamoto, T., Mori, A., Kawai, M., Bekele, A., 1996. Anti-predator behavior of gelada baboons. Primates 37 (4), 389-397. https://doi.org/10.1007/BF02381374.

Kifle, Z., 2023. The mystery of gelada (Theropithecus gelada) survival and adaptation in the highly anthropogenically-modified landscapes in the Ethiopian Highlands: a review. Glob. Ecol. Conserv. 47, e02669. https://doi.org/10.1016/j.gecco.2023.e02669.

Kifle, Z., Bekele, A., 2020. Human-gelada conflict and attitude of the local community toward the conservation of the Southern Gelada (Theropithecus gelada obscurus) around Borena Saynit National Park, Ethiopia. Environ. Manag. 65 (3), 399-409. https://doi.org/10.1007/s00267-019-01246-8.

Lahti, D.C., Johnson, N.A., Ajie, B.C., Otto, S.P., Hendry, A.P., Blumstein, D.T., Coss, R.G., Donohue, K., Foster, S.A., 2009. Relaxed selection in the wild. Trends Ecol. Evol. 24 (9), 487-496. https://doi.org/10.1016/j.tree.2009.03.010.

Lande, R., 1998. Anthropogenic, ecological and genetic factors in extinction and conservation. Popul. Ecol. 40 (3), 259-269. https://doi.org/10.1007/BF02763457.

Lazow, S.P., Bergman, T.J., 2020. The structural and motivational role of the unique lip-flip movement in the gelada (Theropithecus gelada) facial display repertoire. Am. J. Phys. Anthropol. 172 (2), 280-290. https://doi.org/10.1002/ajpa.24031.

Le Roux, A., Bergman, T.J., 2012. Indirect rival assessment in a social primate, Theropithecus gelada. Anim. Behav. 83 (1), 249-255. https://doi.org/10.1016/j.anbehav.2011.10.034.

Lenth, Russell, A., Buerkner, P., Herve, M., Love, J., Singmann, H., Lenth, M.R.V., 2021. Package 'emmeans' R topics documented: 34(1), 216-221. https://doi.org/10.1080/00031305.1980.10483031>License.

Lyndgohd, S., Gopi, G.V., Selvan, K.M., Habib, B., 2014. Effect of interactions among ethnic communities, livestock and wild dogs (Cuen alpinus) in Arunachal Pradesh, India. Eur. J. Wildl. Res. 60 (5), 771-780. https://doi.org/10.1007/s10344-014-0846-8.

Maestripieri, D., Schino, G., Aureli, F., Troisi, A., 1992. A modest proposal: displacement activities as an indicator of emotions in primates. Anim. Behav. 44 (5). https://doi.org/10.1016/S0003-3472(05)08592-5.

Mancini, G., Ferrari, P.F., Palagi, E., 2013. In play we trust. Rapid facial mimicry predicts the duration of playful interactions in geladas. PLoS ONE 8 (6), e66481. https://doi.org/10.1371/journal.pone.0066481.

Nautiyal, H., Mathur, V., Gajare, K.H., Teichroeb, J., Sarkar, D., Diogo, R., 2024. Predatory dogs as drivers of social behavior changes in the central Himalayan langur (semnopithecus schistaceus) in agro-forest landscapes. Biology 13 (6), 410. https://doi.org/10.3390/biology13060410.

Neves, E., Cherkaooui, S.I., Amhaouch, Z., Duperron, C., Ménard, N., Le Gouar, P., 2024. Conflicts between humans and endangered barbary macaques (macaca sylvanus) at the edge of an agricultural landscape in Morocco. Int. J. Primatol. 45 (4), 860-881. https://doi.org/10.1007/s10764-024-00422-w.

Nowak, K., Lee, P.C., 2013. "Specialist" Primates Can Be Flexible in Response to Habitat Alteration. Primates in Fragments. Springer New York, pp. 199-211. https://doi.org/10.1007/978-1-4614-8839-2_14.

Palagi, E., Bergman, T.J., 2021. Bridging captive and wild studies: behavioral plasticity and social complexity in theropithecus gelada. Animals 11 (10), 3003. https://doi.org/10.3390/ani11103003.

Pedruzzi, L., Francesconi, M., Galotti, A., Bogale, B.A., Palagi, E., Lemasson, A., 2025. Wild gelada monkeys detect emotional and prosocial cues in vocal exchanges during aggression. PLoS One 20 (5), e0323295. https://doi.org/10.1371/journal.pone.0323295.

Pedruzzi, L., Francesconi, M., Palagi, E., Lemasson, A., 2024. The sound of yawns makes geladas yawn. Sci. Rep. 14 (1), 1-7. https://doi.org/10.1038/s41598-023-49797-5.

Pozo-Montuy, G., Serio-Silva, J.C., Chapman, C.A., Bonilla-Sánchez, Y.M., 2013. Resource use in a landscape matrix by an arboreal primate: evidence of supplementation in black howlers (Alouatta pigra). Int. J. Primatol. 34 (4), 714-731. https://doi.org/10.1007/s10764-013-9691-y.

Ramsay, M.S., Teichroeb, J.A., 2019. Anecdotes in primatology: temporal trends, anthropocentrism, and hierarchies of knowledge. Am. Anthropol. 121 (3), 680-693. https://doi.org/10.1111/aman.13295.

Rangel-Negrín, A., Gómez-Espinosa, E.E., Chavira-Ramírez, D.R., Dias, P.A.D., 2023. Dog barks influence the physiological stress and behavior of a wild primate. Sci. Total Environ. 882, 163585. https://doi.org/10.1016/j.scitotenv.2023.163585.

Sepúlveda, M.A., Singer, R.S., Silva-Rodríguez, E., Stowhas, P., Pelican, K., 2014. Domestic dogs in rural communities around protected areas: conservation problem or conflict solution? PLoS ONE 9 (1), e86152. https://doi.org/10.1371/journal.pone.0086152.

Shotake, T., Saijuntha, W., Agatsuma, T., Kawamoto, Y., 2016. Genetic diversity within and among gelada (Theropithecus gelada) populations based on mitochondrial DNA analysis. Anthropol. Sci. 124 (3), 157-167. https://doi.org/10.1537/ase.160717.

Snyder-Mackler, N., Beehner, J.C., Bergman, T.J., 2012. Defining higher levels in the multilevel societies of geladas (Theropithecus gelada). Int. J. Primatol. 33 (5), 1054-1068. https://doi.org/10.1007/s10764-012-9584-5.

Teichroeb, J.A., Sricotte, P., 2012. Cost-free vigilance during feeding in folivorous primates? Examining the effect of predation risk, scramble competition, and infanticide threat on vigilance in ursine colobus monkeys (Colobus vellerosus). Behav. Ecol. Sociobiol. 66 (3), 453-466. https://doi.org/10.1007/s00265-011-1292-1.

Tinsley Johnson, E., Snyder-Mackler, N., Beehner, J.C., Bergman, T.J., 2014. Kinship and dominance rank influence the strength of social bonds in female geladas (Theropithecus gelada). Int. J. Primatol. 35 (1), 288-304. https://doi.org/10.1007/s10764-013-9733-5.

Trede, F., Lemkul, A., Atickem, A., Beehner, J.C., Bergman, T.J., Burke, R., Fashing, P.J., Knauf, S., Mekonnen, A., Moges, A., Nguyen, N., Roos, C., Zinner, D., 2020. Geographic distribution of microsatellite alleles in geladas (Primates, Cercopithecidae): Evidence for three evolutionary units. Zool. Scr. 49 (6), 659-667. https://doi.org/10.1111/zsc.12451.

Venkataraman, V.V., Kerby, J.T., Nguyen, N., Ashenafi, Z.T., Fashing, P.J., 2015. Solitary Ethiopian wolves increase predation success on rodents when among grazing gelada monkey herds. J. Mammal. 96 (1), 129-137. https://doi.org/10.1093/jmammal-gyu013.

Waters, S., Watson, T., Farris, Z.J., Bornbusch, S., Valenta, K., Clarke, T.A., Chetry, D., Randriana, Z., Owen, J.R., El Harrad, A., Pal, A., & Home, C. (2023). *Dogs, Primates, and People: A Review* (pp. 61–81). https://doi.org/10.1007/978-3-031-11736-7_5.

Wong, B.B.M., Candolin, U., 2015. Behavioral responses to changing environments. Behav. Ecol. 26 (3), 665-673. https://doi.org/10.1093/beheco/aru183.

Yihune, M., Bekele, A., Tefera, Z., 2009. Human-gelada baboon conflict in and around the Simien Mountains National Park, Ethiopia. Afr. J. Ecol. 47 (3), 276-282. https://doi.org/10.1111/j.1365-2028.2007.00917.x.
Zanoli, A., Gamba, M., Lemasson, A., Norscia, I., Palagi, E., 2022. Inter-sexual multimodal communication during mating in wild geladas: the leading role of females. Curr. Zool. 65 (8), 570–580. https://doi.org/10.1093/cz/zoab093.

Zinner, D., Atickem, A., Beehner, J.C., Bekele, A., Bergman, T.J., Burke, R., Dolotovskaya, S., Fashing, P.J., Gippoliti, S., Knauf, S., Knauf, Y., Mekonnen, A., Moges, A., Nguyen, N., Chr. Stenseth, N., Roos, C., 2018. Phylogeography, mitochondrial DNA diversity, and demographic history of geladas (Theropithecus gelada). PLoS ONE 13 (8), 1–18. https://doi.org/10.1371/journal.pone.0202303.

Zinner, D., Peláez, F., 1999. Verreaux's eagles (Aquila verreauxi) as potential predators of hamadryas baboons (Papio hamadryas hamadryas) in Eritrea. Am. J. Primatol. 47 (1), 61–66. https://doi.org/10.1002/(SICI)1098-2345(1999)47:1<61::AID-AJP7>3.0.CO;2-K.