Macaque Monkeys: Behavior, Habitat, and Conservation

Macaques are a genus of Old World monkeys found across Asia and North Africa, characterized by complex social systems, high cognitive flexibility, and remarkable adaptability to human-modified landscapes. This article provides a species-level overview of macaque behavior, habitat use, and conservation status, with emphasis on the practical implications of human-macaque interactions for farmers, land managers, and wildlife professionals. The genus Macaca includes more than 20 species ranging from the well-studied rhesus macaque (Macaca mulatta) to the endangered moor macaque (Macaca maura) of Sulawesi. Understanding macaque social organization, cognitive abilities, and habitat preferences is essential for developing effective coexistence strategies and conservation interventions.

At a Glance: Macaque Species and Conservation Context

The table below summarizes key macaque species, their general habitat associations, and the primary conservation and management considerations relevant to human-dominated landscapes.

Social Structure and Organization

Macaques are group-living primates that face constant challenges from complex social demands. Research on subcortical mechanisms of social behavior indicates that behavioral variables important for decision-making, including stimulus, action, and outcome, are associated with agent information in regions such as the amygdala, striatum, lateral hypothalamus, and dopaminergic midbrain nuclei. These self-relevant and other-relevant associative signals integrate into a social utility signal that allows decision makers to organize optimal behavior according to social demands. This neural architecture supports the sophisticated social navigation observed across macaque species.

Dominance Hierarchies and Social Tolerance

Macaque species vary considerably in their social tolerance and dominance style. Rhesus macaques exhibit a despotic dominance hierarchy with a well-documented facial display repertoire. In contrast, crested macaques demonstrate higher social tolerance and more complex facial behavior. A 2023 study testing the social complexity hypothesis on three macaque species found that the relative entropy of facial behavior was higher for the more tolerant crested macaques compared to the less tolerant Barbary and rhesus macaques across all social contexts. This indicates that crested macaques more frequently use a higher diversity of facial behavior. The context specificity of facial behavior was higher in rhesus macaques, demonstrating that Barbary and crested macaques used facial behavior more flexibly across different social contexts.

Individual Differences in Social Functioning

Within species, pronounced individual differences in social functioning are evident. The macaque Social Responsiveness Scale-Revised (mSRS-R) provides a quantitative measure of social functioning in natural social settings. Factor analysis of the mSRS-R in a large sample of male rhesus macaques yielded three factors: Poor Social Motivation, Poor Social Attractiveness, and Inappropriate Behavior. Animals with higher Poor Social Motivation were more likely to be observed alone and less likely to be observed in contact and grooming with conspecifics. Animals with higher Poor Social Attractiveness were less likely to be observed playing but more likely to be observed grooming with conspecifics. These factors mapped closely to social behavior and personality dimensions, providing support for the instrument's convergent and discriminant validity.

Personality, Dominance, and Health

Individual differences in personality, behavior, and social status are associated with health outcomes in macaques. A study of 85 socially housed rhesus macaques at two National Primate Research Centers used veterinary records to determine the number of injuries and illnesses for each macaque. The study measured personality using 12 items from a well-established primate personality questionnaire, performed focal observations of behaviors, and calculated dominance status from directional supplant data. Results indicated that more confident and more anxious macaques experienced fewer injuries. These findings demonstrate that personality dimensions, including Dominance, Confidence, Openness, Anxiety, and Friendliness, are independently associated with health outcomes.

Cognitive Abilities and Problem Solving

Macaques possess sophisticated cognitive abilities that enable them to solve complex problems and adapt to changing environments. A 2024 study introduced the Language of Problem Solving (LoPS) model as a quantitative framework to investigate the structure of problem-solving behavior through a language model. The model was applied to an adapted classic Pac-Man game as a cross-species behavioral paradigm to test both humans and macaque monkeys. The LoPS model extracted the latent structure embedded in the agents' gameplay, revealing the non-Markovian temporal dependency structure of their problem-solving behavior and the hierarchical structures of problem solving in both species. Both species evolved their LoPS grammars during learning, progressing from simpler to more complex ones, suggesting that the structure of problem solving evolves to support more sophisticated and efficient problem solving.

Bodily Self-Recognition

Macaques demonstrate a hierarchical bodily self that develops in stages. Research on mirror self-recognition found that macaque monkeys trained for mirror self-recognition could spontaneously recognize themselves when viewing video images of their back from a third-person perspective, but this generalization required a few minutes of exploration. Through visual-tactile synchronization training similar to that used in human out-of-body studies, naive monkeys could recognize themselves from third-person perspective back images and demonstrated mirror self-recognition within seconds. In human infants, third-person perspective back self-recognition developed spontaneously several months after mirror self-recognition. These results demonstrate the existence of hierarchical bodily self in macaques and similar stepwise development of bodily self in human infants.

Facial Musculature and Communication Ontogeny

The development of facial musculature in rhesus macaques provides insight into the ontogeny of social communication. Dissection of facial masks from 13 infant rhesus macaque cadavers revealed that, unlike adults, infant rhesus macaques have extrinsic external ear muscles and muscles associated with the lower lip that are very gracile and poorly developed. Musculature associated with the upper lip and nares are well developed, individually distinct, and robust, like those of adults. However, researchers were unable to locate the zygomaticus major muscle in any fetal or infant sample. These results may reflect variation in mimetic muscle ontogeny, differences related to facial allometry, or a behaviorally meaningful adaptation to different life stages of macaque ontogeny.

Habitat Use and Ecological Flexibility

Macaques demonstrate considerable ecological flexibility in habitat use. A study of wildlife distribution in Cambodia's Prey Lang Wildlife Sanctuary, a lowland evergreen forest ecosystem, recorded seven mammals and two birds, with the long-tailed macaque among the detected species. Wildlife richness and abundance were higher in evergreen-dominated habitats than in mixed deciduous-evergreen forests. Certain K-selected species, including pileated gibbon, Indochinese silvered langur, and great hornbill, were highly specialized and preferred intact forests, whereas generalist species such as northern red muntjac, long-tailed macaque, and wild pig showed ecological flexibility in habitat use.

Forest Fringe and Human Settlement Interfaces

Macaque habitat use frequently extends into human-modified landscapes. Research on crop-raiding rhesus macaques in the Eastern Ghats, India, evaluated the influence of habitat context on the performance of five crop protection strategies over three consecutive cropping seasons. Field experiments were conducted across two contrasting habitat types: forest fringe and forest fringe associated with human settlements. Forest fringe sites associated with human settlements supported larger, more frequently encountered macaque troops and experienced greater crop damage than forest-edge sites. Crop damage was strongly associated with macaque visitation in both habitat types.

Roadside Habitat Use and Provisioning

Human activities such as provisioning and road construction can alter macaque habitat use patterns. A study of human-moor macaque coexistence in Sulawesi, Indonesia, compared data from 2016-2017 to 2023-2025 and examined changes in provisioning patterns, macaque roadside use, and people's perceptions of macaques. Results showed that although provisioning frequency remained stable, hand-feeding became increasingly common and macaques increased their use of roadside habitat. People's perceptions shifted from excitement and novelty to fear and normalization. Decreasing tolerance, coupled with increased risks associated with roadside behavior, highlights the system's potential to transition to a state incompatible with coexistence.

Human-Macaque Conflict and Management

Human-macaque conflict represents a major conservation and livelihood challenge in human-modified landscapes. The long-term effectiveness of mitigation strategies remains poorly understood because behavioral habituation may reduce deterrent efficacy over time.

Crop Protection Strategies

A field study in the Eastern Ghats, India, compared low-cost solar fencing, nylon netting, bear scare devices, agri cannons, and conventional manual watching against crop-raiding rhesus macaques over three consecutive cropping seasons. Crop damage differed significantly among treatments, habitats, and seasons, with deterrent-based methods showing progressive declines in effectiveness across seasons, consistent with behavioral habituation. In contrast, low-cost solar fencing provided consistently effective protection throughout the study. Comparative economic performance mirrored ecological effectiveness, with solar fencing providing the most favorable returns under commercial farming conditions.

Nonlethal Management in Plantation Systems

Research on bonnet macaque (Macaca radiata) infestation in cardamom plantations in the Cardamom Hill Reserves, Kerala, India, evaluated nine non-lethal techniques for managing monkeys. The percentage of incidence and intensity of attack was lower in cardamom plants applied with BoRep, followed by the hanging of rubber snakes, placing sawdust with kerosene, glitter paper fencing, application of red chilli extract, and Neem Seed Kernel Extract. The olfactory repellent BoRep was reported for the first time as a repellent for M. radiata and primates. Because of the primates' high level of intelligence, the study recommended using multiple effective strategies instead of sticking to one strategy.

Technology-Based Deterrent Systems

Technology-based solutions for macaque management are being developed and tested. One proposed system, called Smart Repeller, uses ultrasonic sound waves and Calcium Carbide Cannon, along with computer vision technology and artificial intelligence to detect the presence of monkeys and activate repelling mechanisms automatically. The proposed device eliminates the need for human intervention, making it efficient and cost-effective. Another AI-driven solution developed for rural Nepal uses AI-powered cameras to scan and monitor target areas and identify rhesus macaques in real-time using the YOLO algorithm, trained using a custom dataset of monkey images. Upon detection, the system activates an ultrasonic deterrent subsystem. The YOLO-based detection model achieved a precision of up to 1.00, an F1-score of 0.97, and a recall of 0.99.

Macaques in Research and Captive Management

Macaque species, specifically rhesus macaques, are the most common nonhuman primates used in biomedical research due to their suitability as models of high priority diseases, cost-effective breeding and housing compared to most other nonhuman primates, and close evolutionary relationship to humans. With this close evolutionary relationship, however, is a shared adaptation for a socially stimulating environment, without which both their welfare and suitability as a research model are compromised.

Social Housing Requirements

Outdoor social group housing provides the best approximation of a social environment that matches macaque behavioral biology in the wild. However, this is not always possible at all facilities, where animals may be housed indoors in small groups, in pairs, or alone. Animals may experience many housing changes in their lifetime depending on project needs, changes in social status, management needs, or health concerns. Research has documented the physiological and health effects of social housing changes and the potential impacts on research outcomes for studies using macaques. Because most nonhuman primates are adapted for sociality, a social context is likely important for improving repeatability, reproducibility, and external validity of research findings.

Effects of Individual Housing

The duration of individual housing affects social behavior in macaques. A study on adult male bonnet macaques examined the effect of duration of individual housing on social behavior. The title and publication metadata indicate that housing conditions influence subsequent social interactions, though specific findings were not available in the source record. Similarly, research on captive Indochinese and insular long-tailed macaques following transfer to a new facility examined social behavior changes associated with facility relocation.

Social Affiliation and Huddling Behavior

Social affiliation in macaques can be measured through huddling behavior. Research on Taihangshan macaques used huddling behavior as an indicator of social affiliation patterns. Huddling serves as a thermoregulatory and social bonding behavior that reflects the strength of social relationships within groups.

Neural Basis of Social Behavior

The neural mechanisms underlying macaque social behavior provide insight into both normal social functioning and potential disruptions. Research on the impact of early amygdala damage on juvenile rhesus macaque social behavior found that amygdala-lesioned animals generated fewer aggressive and affiliative signals and spent less time in social interactions with familiar peers. When observed alone or with an unfamiliar peer, amygdala-lesioned animals spent more time being inactive and physically explored the environment less. Although overall frequencies of many observed behaviors did not differ between groups, the general patterning of social behavior distinguished the amygdala-lesioned animals in a discriminant function analysis.

Subcortical Contributions to Social Decision-Making

Emerging evidence suggests that subcortical regions mediate the collection and processing of information from other agents. Behavioral variables important for decision-making, including stimulus, action, and outcome, are associated with agent information in subcortical regions such as the amygdala, striatum, lateral hypothalamus, and dopaminergic midbrain nuclei. Such self-relevant and other-relevant associative signals are then integrated into a social utility signal, presumably at the level of midbrain dopamine neurons. This social utility signal allows decision makers to organize their optimal behavior in accordance with social demands.

Visual Processing and Neural Representation

Macaques serve as important models for understanding visual processing pathways. Deep spiking neural networks have been used to model the visual cortex of macaques and mice. Similarity analyses revealed that depths of the layers with the highest similarity scores exhibit little differences across mouse cortical regions but vary significantly across macaque regions, suggesting that the visual processing structure of mice is more regionally homogeneous than that of macaques. The different performance in fitting macaque neural representations under different stimuli exhibits the functional specialization of information processing in macaques.

Conservation Status and Threats

Macaque conservation status varies considerably across species. While some species such as rhesus and long-tailed macaques are widespread and adaptable, others face significant threats from habitat loss, hunting, and human-wildlife conflict.

Habitat Loss and Fragmentation

Habitat loss and fragmentation represent primary threats to macaque populations. The study in Prey Lang Wildlife Sanctuary highlighted that protecting intact evergreen forests via REDD+ supports specialized species, while adaptive management in mosaic landscapes benefits generalists such as long-tailed macaques. Conservation strategies must account for the differential habitat preferences of macaque species, with some requiring intact forest and others thriving in modified landscapes.

Human Perceptions and Coexistence

Human perceptions of macaques significantly influence conservation outcomes. The Sulawesi study on moor macaques found that people's perceptions shifted from excitement and novelty to fear and normalization over time. Decreasing tolerance, coupled with increased risks associated with roadside behavior, highlights the system's potential to transition to a state incompatible with coexistence. Interventions that enable safer roadside crossing for the macaques and community outreach programs that make use of people's empathy for the macaques' welfare can promote resilient coexistence.

Genetic and Biobank Resources

Macaque genotype-phenotype resources are being developed to support both biomedical research and conservation. A macaque biobank is planned in three phases: the initial phase will establish standardized protocols and collect pilot data, followed by large-scale cohort expansion and multimodal data acquisition. The final phase will focus on the application of artificial intelligence for multimodal data fusion and analysis.

Practical Assessment and Management Steps

For farmers, land managers, and wildlife professionals dealing with macaque conflicts, the following assessment and management steps are recommended based on available evidence.

Step 1: Document the Problem

Maintain systematic records of macaque activity including visitation frequency, troop size, time of day, crop damage extent, and seasonal patterns. Record which crops are affected and the stage of crop development when damage occurs. This documentation provides the baseline data needed to evaluate management effectiveness.

Step 2: Assess Habitat Context

Determine whether the affected area is forest fringe or forest fringe associated with human settlements. Research indicates that sites associated with human settlements support larger, more frequently encountered macaque troops and experience greater crop damage. This assessment informs the selection of management strategies and expectations for conflict intensity.

Step 3: Select Deterrent Strategies

Choose deterrent methods based on evidence of effectiveness and awareness of habituation risks. Low-cost solar fencing provided consistently effective protection throughout a three-season study, while deterrent-based methods showed progressive declines in effectiveness across seasons consistent with behavioral habituation. For plantation systems, multiple nonlethal strategies used together are more likely to succeed than single approaches.

Step 4: Rotate and Combine Methods

Because macaques habituate to deterrents, rotate methods and combine multiple strategies. The recommendation from research on bonnet macaques in cardamom plantations is to use multiple effective strategies instead of sticking to one strategy. Technology-based solutions that automate detection and deterrent activation may reduce habituation by providing consistent and unpredictable responses.

Step 5: Monitor and Adjust

Evaluate management effectiveness regularly using the baseline data collected in Step 1. Track crop damage, macaque visitation, and economic performance of management strategies. Adjust approaches when deterrent effectiveness declines. Comparative economic performance should mirror ecological effectiveness, with the most effective strategies providing the most favorable returns under commercial farming conditions.

Step 6: Engage Community and Professional Support

Community engagement is essential for sustainable coexistence. Outreach programs that make use of people's empathy for macaque welfare can support conservation goals. When conflicts escalate or involve protected species, consult with wildlife authorities and conservation professionals for species-specific guidance and permitted management options.

Records and Measurements

Maintaining accurate records is essential for effective macaque management. The following measurements provide the data needed for evidence-based decisions.

Common Failure Patterns in Macaque Management

Several common failure patterns emerge from research on macaque management. Recognizing these patterns helps managers avoid ineffective approaches.

Habituation to Deterrents

Deterrent-based methods show progressive declines in effectiveness across seasons, consistent with behavioral habituation. Macaques learn to ignore or adapt to scare devices, agri cannons, and similar methods over time. Managers should anticipate habituation and plan for method rotation or investment in more durable solutions such as solar fencing.

Single-Strategy Dependence

Relying on a single management strategy is unlikely to succeed given macaque intelligence and adaptability. Research on bonnet macaques explicitly recommends using multiple effective strategies instead of sticking to one strategy. Integrated approaches that combine physical barriers, repellents, and technology-based detection systems are more likely to maintain effectiveness.

Ignoring Habitat Context

Management strategies that do not account for habitat context may fail. Sites associated with human settlements support larger, more frequently encountered macaque troops and experience greater crop damage. Strategies that work in forest-edge sites may be insufficient in settlement-associated sites.

Neglecting Human Dimensions

Management failures often stem from neglecting human perceptions and behaviors. Provisioning, including hand-feeding, increases macaque use of human-dominated areas and shifts human perceptions from excitement to fear. Effective management must address human behavior alongside macaque deterrence.

Limitations and Knowledge Gaps

Several limitations and knowledge gaps affect macaque management and conservation. The long-term effectiveness of mitigation strategies remains poorly understood because behavioral habituation may reduce deterrent efficacy over time. Most studies of crop protection strategies cover limited time frames, and multi-year data are needed to assess durability of management approaches.

Research on social behavior in captive settings provides insights into macaque social needs but may not fully reflect wild populations. Studies of individual housing effects on bonnet macaques and social behavior following facility transfer in long-tailed macaques contribute to understanding captive management but have limited direct application to wild conflict situations.

The neural basis of social behavior in macaques is increasingly well understood, but translating these findings into practical management applications requires further research. Understanding how social information processing is altered in disease conditions may inform welfare assessments in captive settings.

Welfare and Safety Context

Macaque management involves both animal welfare and human safety considerations. Macaques are highly social animals adapted for socially stimulating environments. Housing or management practices that compromise social needs can compromise both welfare and behavioral suitability. In research settings, social housing is important for improving repeatability, reproducibility, and external validity of studies.

Human safety is a significant concern in macaque conflict situations. Macaques that become habituated to humans may pose risks including property damage, food theft, and health risks. Roadside behavior associated with provisioning increases risks for both macaques and humans. Management strategies should aim to reduce these risks while maintaining humane treatment of macaques.

Nonlethal management techniques are preferred for macaque conflict resolution. Research on bonnet macaques evaluated nine non-lethal techniques, and technology-based solutions aim to deter macaques without causing harm. When selecting management strategies, prioritize methods that effectively reduce conflict while minimizing animal suffering.

Professional Escalation Criteria

Certain situations warrant escalation to wildlife professionals, conservation authorities, or veterinary experts. Consult professionals when any of the following conditions apply.

Protected Species Involvement

If conflict involves macaque species with protected status or endangered classification, consult with wildlife authorities before implementing management measures. Species-specific regulations may restrict available management options.

Escalating Conflict Intensity

When crop damage or macaque visitation increases despite management efforts, professional assessment may identify factors not apparent to local managers. Persistent conflict may require landscape-level interventions beyond individual farm management.

Public Health Concerns

Macaque interactions that pose disease transmission risks or other public health concerns require professional involvement. Health authorities and wildlife professionals should be consulted when macaques frequent human settlements or food storage areas.

Welfare Concerns

If management activities risk causing harm to macaques, or if macaques appear injured, sick, or distressed, consult veterinary professionals and wildlife authorities. Humane treatment should guide all management decisions.

Research and Monitoring Opportunities

Situations that offer opportunities for systematic data collection may benefit from collaboration with research institutions. Studies of macaque behavior, habitat use, and management effectiveness contribute to the evidence base for future conservation and conflict management.

Frequently Asked Questions

What is the social structure of macaque groups?

Macaques live in multi-male multi-female groups with complex dominance hierarchies. Social tolerance varies by species, with rhesus macaques exhibiting despotic hierarchies and crested macaques showing higher social tolerance. Individual differences in social functioning are evident within species, measurable through instruments like the macaque Social Responsiveness Scale-Revised.

How do macaques communicate with each other?

Macaques use facial displays, vocalizations, and body postures for communication. Facial behavior complexity correlates with social tolerance, with more tolerant species like crested macaques using a higher diversity of facial behavior. Rhesus macaques have a well-documented facial display repertoire associated with their despotic dominance hierarchy.

What cognitive abilities do macaques possess?

Macaques demonstrate sophisticated problem-solving abilities, hierarchical planning, and bodily self-recognition. They can learn complex tasks and evolve their problem-solving strategies from simpler to more complex structures during learning. Macaques trained for mirror self-recognition can generalize to third-person perspective self-recognition.

Why do macaques raid crops?

Crop raiding occurs when macaques access agricultural areas for food, particularly in forest fringe habitats associated with human settlements. Larger, more frequently encountered macaque troops are associated with greater crop damage. Habitat context influences conflict intensity, with settlement-associated sites experiencing more damage than forest-edge sites.

What crop protection methods work against macaques?

Low-cost solar fencing provided consistently effective protection across multiple seasons in one study. Deterrent-based methods such as scare devices and agri cannons showed progressive declines in effectiveness due to habituation. Multiple nonlethal strategies used together are more effective than single approaches, and technology-based detection and deterrent systems are being developed.

How does provisioning affect macaque behavior?

Provisioning, including hand-feeding, increases macaque use of human-dominated areas and roadside habitat. Human perceptions shift from excitement to fear as provisioning continues. Decreasing tolerance and increased risks associated with roadside behavior can transition the system to a state incompatible with coexistence.

What is the conservation status of macaques?

Conservation status varies by species. Some species such as rhesus and long-tailed macaques are widespread and adaptable, while others such as moor macaques face threats from habitat loss and changing human perceptions. Habitat preferences differ, with some species requiring intact forest and others showing ecological flexibility in modified landscapes.

How should individual macaques be housed in captivity?

Macaques are adapted for sociality, and social housing is important for welfare and behavioral suitability. Outdoor social group housing best approximates wild behavioral biology. Pair-housing is the most common social housing configuration for macaques on research projects, though housing changes can have physiological and health effects.

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Subcortical encoding of agent-relevant associative signals for adaptive social behavior in the macaque.. Neuroscience and biobehavioral reviews, 2021.
• Laboratory rhesus macaque social housing and social changes: Implications for research.. American journal of primatology, 2017.
• The factor structure of the macaque social responsiveness scale-revised predicts social behavior and personality dimensions.. American journal of primatology, 2021.
• Higher social tolerance is associated with more complex facial behavior in macaques.. eLife, 2023.
• Macaques.. Current biology : CB, 2006.
• Rhesus macaque personality, dominance, behavior, and health.. American journal of primatology, 2018.
• The impact of early amygdala damage on juvenile rhesus macaque social behavior.. Journal of cognitive neuroscience, 2013.
• Development of Rhesus Macaque (Macaca mulatta) Facial Musculature: Implications for Macaque Social Behavior Ontogeny.. American journal of primatology, 2025.
• Behavioural habituation undermines deterrents in rhesus macaque crop raiding in Eastern Ghats, India. 2026.
• Habitat Preference of Key Wildlife Species in One of the Largest Lowland Evergreen Forests in Southeast Asia. 2026.
• Balancing risk and reward: Toward resilient human-primate coexistence in a rapidly changing environment in Sulawesi, Indonesia.. 2026.
• Camera-Trap Assessment of Terrestrial Mammals and Ground-Dwelling Birds in the Zhangjiajie Chinese Giant Salamander National Nature Reserve, China. 2026.
• Bodily Self in Macaque Monkeys and Human Infants. bioRxiv, 2025.
• Macaque's (Macaca radiata) Infestation and its Low-Cost Nonlethal Management on Cardamom in the Cardamom Hill Reserves, Kerala. Indian Journal of Entomology, 2025.
• A language model of problem solving in humans and macaque monkeys.. Current Biology, 2024.
• SMART REPELLER: A Smart system to prevent Rhesus Macaque Trespassing in Human Settlements and Agricultural Areas. EAI Endorsed Transactions on Internet of Things, 2024.
• Learning to See Through a Baby's Eyes: Early Visual Diets Enable Robust Visual Intelligence in Humans and Machines. arXiv.org, 2025.
• Deep Spiking Neural Networks with High Representation Similarity Model Visual Pathways of Macaque and Mouse. AAAI Conference on Artificial Intelligence, 2023.
• Design And Development of An AI-Driven Solution For Rhesus Macaque. Journal of Advanced College of Engineering and Management, 2026.
• Macaque genotype-phenotype resources: A prismatic portal into human biology and disease. Zoological Research, 2026.
• Social behavior of captive Indochinese and insular long-tailed macaques (Macaca fascicularis) following transfer to a new facility. International Journal of Primatology, 2002.
• The Effect of Duration of Individual Housing on Social Behavior of Adult Male Bonnet Macaques (Macaca radiata). Contemporary Topics in Laboratory Animal Science, 1998.
• Social affiliation of Taihangshan macaque based on huddling behavior. Acta Theriologica Sinica, 2013.

This article is educational and does not replace institutional policy, professional advice, or applicable safety and regulatory requirements.