How Animals Communicate: A Guide to Visual, Auditory, Chemical, and Tactile Signals

Animal communication is the transfer of information between individuals through signals that can be perceived by one or more sensory systems. The four main types are visual, auditory, chemical, and tactile signals, and most species use combinations of these modalities depending on context, distance, and the relationship between signaler and receiver. This article explains each communication type with examples from diverse species, compares their functions and limitations, and provides practical guidance for observing and recording animal communication in research, farming, and wildlife management settings.

At a Glance: Comparison of Animal Communication Modalities

The table below summarizes the four primary communication types, their typical purposes, example species, and key limitations. Use it as a quick reference when designing observation protocols or interpreting behavioral records.

The Scope of Animal Communication Studies

Animal communication research examines how signals are produced, transmitted, received, and interpreted. A signal is any behavior or physical feature that has evolved to carry information from a signaler to a receiver. The receiver may respond in ways that benefit itself, the signaler, or both. Communication is distinct from mere information transfer because signals are shaped by natural selection to influence receiver behavior.

The study of animal communication spans multiple disciplines including ethology, neurobiology, evolutionary biology, and conservation science. Researchers observe animals in natural and captive settings, record signal production and receiver responses, and analyze the physical properties of signals. For farmers and wildlife managers, understanding communication systems helps in detecting distress, managing breeding programs, reducing aggression, and assessing animal welfare.

Communication systems vary widely across taxa. Some species rely heavily on one modality while others integrate several. The Tasmanian devil, for example, uses visual, chemical, tactile, and auditory signals in its female-male interaction system, demonstrating that even species with relatively simple social structures employ multimodal communication [3]. The vocalizations of marsupials are generally low in intensity and easily overlooked by investigators, yet they exhibit complexity equivalent to the calls of many Eutherian mammals [3].

Visual Communication

Visual signals are among the most immediately observable forms of animal communication. They include body posture, movement patterns, color changes, facial expressions, and specialized structures such as crests, manes, or bright plumage. Visual signals are typically used over short to medium distances where the receiver can see the signaler clearly.

Posture and Movement

Body posture conveys information about aggression, submission, readiness to mate, and vigilance. A dominant animal may stand tall, enlarge its apparent size, or make direct eye contact. A submissive animal may crouch, avert its gaze, or make itself appear smaller. These signals reduce the need for physical combat by allowing animals to assess each other's condition and intent before engaging.

Movement-based signals include ritualized displays such as courtship dances, threat lunges, and play invitations. The speed, amplitude, and repetition of movements can carry additional meaning. For example, a slow deliberate approach may signal confidence while a rapid approach may signal alarm or aggression.

Color and Appearance

Color changes can signal reproductive readiness, emotional state, or species identity. Some fish and reptiles change color rapidly in response to social context. Permanent color differences between sexes or age classes help animals identify potential mates or rivals. Bright coloration may also serve as a warning signal to predators, indicating toxicity or unpalatability.

Visual Signals in Specific Species

The Tasmanian devil uses visual signals as part of its interaction system, particularly in the context of female-male encounters [3]. These signals include postural changes and movements that convey information about reproductive status and social intent. In captive settings, observing these visual signals helps researchers and zookeepers manage breeding pairs and reduce aggression.

Chimpanzees modify the modality of their gestures according to the strength of social bonds and personal network size [13]. Individuals with stronger social bonds use more tactile and visual gestures, while those with weaker bonds rely more on visual signals that maintain distance. This finding shows that visual communication is not fixed but is adjusted based on social context and relationship quality.

Limitations of Visual Signals

Visual signals require a clear line of sight. They are ineffective in darkness, dense vegetation, or murky water. Many nocturnal or forest-dwelling species therefore rely more heavily on auditory or chemical signals. Visual signals also fade quickly, meaning they cannot convey information over time the way scent marks can. For observers, visual signals are relatively easy to record but require good lighting and unobstructed views.

Auditory Communication

Auditory communication involves the production of sound through vocal cords, stridulation, drumming, or other mechanisms. Sound travels well through air and water, works in darkness, and can carry over long distances. Auditory signals are used for alarm calls, mate attraction, territorial defense, parent-offspring recognition, and coordination of group movement.

Vocalizations and Their Functions

Vocalizations vary in frequency, amplitude, duration, and temporal pattern. These acoustic properties determine how far the signal travels and what information it conveys. Low-frequency sounds travel farther than high-frequency sounds and are often used for long-distance communication. High-frequency sounds are more easily localized and are often used for close-range communication.

The Tasmanian devil produces vocalizations that have been classified and described with respect to their probable function [3]. Four basic syllable types are defined, and these are related to at least four different functions. The vocalizations reflect adaptations for conveying information in several widely differing contexts [3]. This finding demonstrates that even species with relatively simple vocal repertoires use sound flexibly across social situations.

Prosody and Emotional Information

Across a wide range of animal taxa, prosodic modulation of the voice can express emotional information and is used to coordinate vocal interactions between multiple individuals [11]. Prosody refers to the rhythm, stress, and intonation of vocalizations. Emotional prosody allows receivers to assess the signaler's state, such as fear, aggression, or contentment, which influences their response.

Interactional prosody coordinates turn-taking and timing in vocal exchanges. Many species engage in antiphonal calling, where individuals alternate vocalizations in a structured pattern. This coordination is important for maintaining social bonds and synchronizing group behavior.

Tempo and Receiver Biology

A widespread animal communication tempo may resonate with the receiver's brain [7]. Field observations in Thailand found nearly identical tempos of co-located flashing fireflies and chirping crickets. A survey of published data shows that an abundance of evolutionarily distinct species communicate isochronously at approximately 0.5 to 4 Hz, suggesting this might be a tempo hotspot [7]. The hypothesis is that this timescale has a universal basis in the biophysics of the receiver's neurons, with small receiver circuits constructed from typical neurons being most responsive in this tempo range [7].

This finding has practical implications for playback experiments and acoustic monitoring. When designing auditory stimuli for research or management, matching the species-typical tempo may improve receiver response rates.

Auditory Communication in Farming Contexts

Farmers and livestock managers can use auditory signals to assess animal welfare. Changes in vocalization rate, amplitude, or frequency may indicate pain, distress, hunger, or illness. Establishing baseline vocalization patterns for healthy animals allows managers to detect deviations early. Recording equipment can monitor vocalizations continuously, providing data that supports early intervention.

Limitations of Auditory Signals

Auditory signals can attract predators, which creates a tradeoff between the benefits of communication and the risks of detection. Background noise from wind, water, or human activity can mask signals and reduce their effective range. Sound also degrades with distance, and different frequencies attenuate at different rates. For observers, acoustic recording requires appropriate equipment and careful placement to capture signals without distortion.

Chemical Communication

Chemical communication involves the production and detection of chemical signals, including pheromones and scent marks. These signals are detected through olfaction or taste and can convey information about species identity, individual identity, reproductive status, territory ownership, and emotional state. Chemical signals have the advantage of persisting in the environment after the signaler leaves, allowing information to be transmitted over time.

Pheromones and Scent Marking

Pheromones are chemical substances released by an individual that trigger a specific response in a member of the same species. They are involved in mate attraction, alarm signaling, trail following, and aggregation. Scent marking involves depositing chemicals on objects or surfaces through urine, feces, gland secretions, or rubbing. Marks convey information about the signaler's presence, identity, and condition.

The Tasmanian devil uses chemical signals as part of its interaction system [3]. Chemical communication is particularly important for solitary or nocturnal species that cannot rely on visual signals. Scent marks allow individuals to assess each other without direct contact, reducing the risk of aggressive encounters.

Individual Recognition

Chemical signals can encode information about individual identity, allowing animals to recognize mates, offspring, rivals, and group members. This recognition is important for maintaining social structure and avoiding inbreeding. The chemical profile of an individual may reflect its genetic makeup, diet, health status, and reproductive condition.

Chemical Communication in Management

Wildlife managers and farmers can use chemical signals to influence animal behavior. Synthetic pheromones are used in pest control to disrupt mating, lure insects into traps, or repel unwanted species. Understanding the chemical communication system of a target species allows managers to design more effective control strategies.

For livestock, chemical signals influence reproductive behavior, maternal bonding, and stress responses. Observing scent-marking behavior and investigating changes in chemical signaling can provide clues about animal health and social dynamics.

Limitations of Chemical Signals

Chemical signals transmit slowly compared to visual or auditory signals. The signaler cannot control the range or duration of the signal once it is released. Chemical signals may persist for hours or days, which can be advantageous for territorial marking but problematic if the signal becomes outdated. Environmental conditions such as rain, temperature, and wind affect signal persistence and detection.

For researchers, chemical communication is more difficult to observe than visual or auditory communication. Detecting chemical signals often requires specialized equipment such as gas chromatography or behavioral bioassays. Field observations of scent marking and sniffing behavior provide indirect evidence of chemical communication.

Tactile Communication

Tactile communication involves physical contact between individuals. It includes grooming, touching, nudging, mounting, and fighting. Tactile signals require close proximity and are therefore used primarily in social interactions between individuals that already have a relationship. Tactile communication reinforces social bonds, provides reassurance, coordinates cooperative behavior, and conveys aggression or submission.

Grooming and Social Bonding

Grooming is a common form of tactile communication in many mammals and birds. It serves hygienic functions but also reinforces social bonds and reduces tension. Grooming can be reciprocal, with individuals taking turns, or directed, with dominant individuals receiving more grooming than subordinates. The distribution of grooming within a group reflects social structure and relationship quality.

Chimpanzees modify the modality of their gestures according to the strength of social bonds [13]. Individuals with stronger social bonds use more tactile gestures, indicating that touch is associated with closer relationships. This finding supports the view that tactile communication is important for maintaining and signaling social bonds.

Tactile Signals in Aggression and Mating

Tactile signals are also used in aggressive encounters and mating behavior. Physical contact can escalate or de-escalate conflicts, with gentle touches signaling submission and forceful contact signaling aggression. During mating, tactile signals coordinate positioning and timing between partners.

The Tasmanian devil uses tactile signals as part of its female-male interaction system [3]. Tactile signals in this context likely coordinate mating behavior and convey information about receptivity and readiness.

Vibration and Substrate-Borne Signals

Biotremology is the study of communication through substrate-borne vibrations [14]. Many insects, spiders, and other arthropods produce vibrations by drumming, stridulating, or tapping on surfaces. These vibrations travel through soil, plant stems, leaves, and other substrates. Substrate-borne signals are used for mate attraction, alarm, and territorial defense.

Vibrational communication is particularly important for small animals that cannot produce loud airborne sounds. It also works in darkness and dense vegetation where visual signals are ineffective. For researchers, detecting substrate-borne vibrations requires specialized sensors and careful experimental design.

Tactile Communication in Livestock Management

Farmers can use tactile communication to assess animal temperament and build trust. Gentle handling and regular physical contact can reduce stress responses in livestock. Observing how animals respond to touch provides information about their emotional state and health. Animals that avoid touch may be in pain or fearful, while animals that seek contact may be comfortable and content.

Limitations of Tactile Signals

Tactile signals require physical proximity, which limits their use to close-range interactions. They cannot transmit over distance or persist over time. Tactile signals also carry a risk of injury if contact becomes aggressive. For observers, tactile communication is difficult to record without close observation, and the meaning of specific touches may depend heavily on context and relationship history.

Multimodal Communication and Signal Integration

Most animals use multiple communication modalities, often simultaneously. Multimodal communication allows signalers to convey redundant information, which increases the likelihood that the receiver will detect and correctly interpret the signal. It also allows signalers to convey different types of information through different channels.

The Tasmanian devil uses visual, chemical, tactile, and auditory signals in its interaction system [3]. This multimodal approach allows the species to communicate effectively across different contexts and distances. The four basic vocalization types are related to at least four different functions, and they reflect adaptations for conveying information in several widely differing contexts [3].

Redundancy and Backup Signals

Redundant signaling occurs when the same information is transmitted through multiple modalities. This redundancy provides a backup if one channel is blocked or degraded. For example, an animal may combine a vocal alarm call with a visual display, ensuring that nearby receivers detect the alarm even if they cannot hear the call.

Complementary Signals

Complementary signaling occurs when different modalities convey different types of information. For example, a visual signal may indicate the signaler's identity while an auditory signal indicates its emotional state. Combining complementary signals allows for richer and more precise communication than any single modality could achieve.

Context-Dependent Modality Use

Animals adjust their use of communication modalities based on context. Factors such as distance, lighting, background noise, social relationship, and the presence of predators influence which modality is used. Chimpanzees modify the modality of their gestures according to the strength of social bonds and personal network size [13], demonstrating that social context shapes communication choices.

Animal Communication and the Question of Language

A central question in animal communication research is why no animal communication system has evolved into a simple language. Individuals of some animal species have been taught simple versions of human language despite their natural communication systems failing to rise to the level of a simple language [8]. This raises the question of why animals do not deploy this capacity in their own communication systems.

Design Features of Language

Researchers use key design features to evaluate the language-like properties of natural animal communication systems [8]. These features include productivity, displacement, arbitrariness, and cultural transmission. Bird song has several of these design features or their precursors, including social learning and cultural transmission of vocal signals [8]. However, although bird song communication is nuanced and complex, and has the acoustic potential for productivity, it is not productive, meaning it cannot be used to say many different things [8].

The Role of Cooperative Interests

The debate over whether animal communication should be viewed as a cooperative information transmission process or as a competitive process where signaler and receiver vie for control points to a necessary condition for the evolution of a simple language [8]. The degree to which the interests of the signaler and receiver align is critical. While strong cognitive and signal production mechanisms are necessary pre-adaptations for a simple language, they are not sufficient [8]. Also necessary is the existence of identical or near-identical interests of signaler and receiver and a socio-ecology that requires high-level cooperation across a range of contexts [8].

Commentary on this argument agrees that many animal species have a level of cognitive sophistication that should predispose them to have at least rudimentary languages, but such species do not meet the criterion of sufficient alignment of interest [9]. The emphasis on alignment of interest over cognitive capacity is a valuable contribution to the language evolution literature [9].

Phonemes and Animal Communication

The question of whether animal communication systems have phonemes remains open [12]. Phonemes are the smallest units of sound that distinguish meaning in human language. While some animal vocalizations show categorical perception and combinatorial structure, no animal system has been shown to have a phoneme inventory comparable to human language.

Implications for Communication Research

Understanding why animal communication systems are not languages helps researchers appreciate the unique features of human language and the evolutionary conditions that enabled its emergence. It also informs the design of communication-based management tools, such as playback experiments and acoustic monitoring, by clarifying what animal signals can and cannot convey.

Practical Observation and Recording Methods

Observing and recording animal communication requires systematic methods and careful attention to context. The following steps provide a framework for conducting communication observations in research, farming, or wildlife management settings.

Step 1: Define the Research Question

Clearly state what you want to learn about the communication system. Examples include identifying the signals used during courtship, determining how alarm calls vary with predator type, or assessing whether vocalization rate changes with pain level. A focused question guides data collection and analysis.

Step 2: Select Observation Methods

Choose between focal sampling, where one individual is observed for a set period, and scan sampling, where multiple individuals are observed at regular intervals. Focal sampling provides detailed data on individual behavior while scan sampling provides an overview of group activity. Consider using video and audio recording to supplement direct observation.

Step 3: Record Signal Production and Receiver Response

For each signal observed, record the signaler's identity, the signal type, the context, and the receiver's response. Note the distance between signaler and receiver, the time of day, and environmental conditions. This contextual information is essential for interpreting signal function.

Step 4: Establish Baseline Patterns

Collect data over multiple sessions to establish baseline patterns of communication. Baselines allow you to detect deviations that may indicate changes in welfare, health, or social dynamics. For example, an increase in alarm calls may indicate the presence of a predator or a source of chronic stress.

Step 5: Analyze and Interpret Data

Analyze the data to identify patterns in signal use and receiver response. Compare communication across contexts, individuals, and time periods. Interpret the findings in light of the species' ecology and social system, and consider alternative explanations for observed patterns.

Records and Measurements for Communication Monitoring

Maintaining systematic records of animal communication supports welfare assessment, breeding management, and research. The following measurements are useful for monitoring communication in captive and farmed animals.

Vocalization Rate and Amplitude

Record the number of vocalizations per unit time and their amplitude. Changes in vocalization rate may indicate hunger, pain, distress, or social disruption. Amplitude changes may indicate the signaler's distance from the receiver or its motivational state.

Signal Diversity

Record the number of distinct signal types observed and the contexts in which they occur. A reduction in signal diversity may indicate depression, illness, or social isolation. An increase may indicate heightened arousal or social instability.

Receiver Response Latency

Measure the time between signal production and receiver response. Short latencies indicate efficient communication and strong signal salience. Long latencies may indicate signal degradation, receiver inattention, or poor signal design.

Scent Marking Frequency

Record the frequency and location of scent-marking behavior. Changes in marking frequency may indicate territorial pressure, reproductive cycling, or stress. Marking location provides information about how the animal structures its environment.

Grooming and Contact Rates

Record the frequency and duration of grooming and other tactile contact. Changes in contact rates may indicate changes in social bonding, dominance relationships, or individual health.

Common Failure Patterns in Communication Observation

Several common errors can compromise the validity of communication observations. Being aware of these failure patterns helps observers design better studies and interpret data more accurately.

Observer Effects

The presence of an observer can alter animal behavior. Animals may reduce or increase communication in response to human presence. Mitigate observer effects by using remote cameras, one-way glass, or habituation periods before formal data collection begins.

Context Neglect

Interpreting signals without recording context leads to errors. The same signal may have different meanings in different contexts. Always record the social, environmental, and temporal context of each signal.

Modality Bias

Observers often focus on the most conspicuous modality, such as loud vocalizations, and miss quieter or less obvious signals. The vocalizations of marsupials are generally low in intensity and easily overlooked by an investigator [3]. Systematic observation protocols should include attention to all modalities.

Sampling Bias

Observing only certain individuals, times, or locations can bias results. Ensure that sampling covers the full range of individuals, social contexts, and environmental conditions relevant to the research question.

Equipment Limitations

Recording equipment may not capture the full range of signals. Microphones may miss low-amplitude or low-frequency sounds, and cameras may miss rapid movements or color changes. Test equipment before data collection and use multiple recording methods when possible.

Welfare and Safety Context

Understanding animal communication has direct applications for animal welfare assessment and management. Communication signals provide a window into the internal state of animals, allowing caregivers to detect pain, fear, stress, and social disruption.

Communication in Euthanasia Decision-Making

Euthanasia in companion animal practice represents one of the most emotionally and ethically challenging decisions in veterinary medicine, requiring clinical judgment, effective communication, and sensitivity toward both animal welfare and pet owner well-being [4]. Most pet owners assess their animal's condition through personal observation or veterinary advice, whereas structured quality-of-life tools are rarely used [4]. Understanding animal communication helps owners and veterinarians interpret behavioral signals that indicate suffering or quality of life.

Pet owners who explicitly reported receiving emotional support from their veterinarian experienced significantly lower emotional burden and greater satisfaction with communication than those who explicitly reported no emotional support [4]. This finding highlights the central role of communication in improving companion animal euthanasia decision-making [4].

Communication in Wildlife Rescue and Rehabilitation

Wildlife rescue is a morally motivated practice that seeks to alleviate suffering in free-ranging, predominantly native, wild animals [6]. However, actions to help individual animals often have far-reaching and sometimes unintended consequences for animal welfare and raise wider concerns regarding animal ethics and ecological integrity [6]. Understanding the communication signals of wild animals helps rescuers assess their condition and make informed decisions about intervention.

Decision-making competencies and communication can be promoted through further training in ethical reasoning and the adoption of an ethical review process [6]. Wildlife rehabilitators should be familiar with the normal communication patterns of the species they care for so they can detect deviations that indicate distress or illness.

Communication in Animal Disease Control Policy

Policy briefs are concise documents designed to communicate information to policymakers to inform the policymaking process [5]. The importance of achieving a balance between conflicting demands in content and formatting was identified as a key aspect of policy brief preparation [5]. Policy briefs were identified as an important component of effective science-policy communication within the context of a trusting relationship between scientists and policymakers [5].

For animal health researchers, communicating findings about animal communication and welfare to policymakers requires clear, accessible language and a focus on actionable recommendations. Understanding the communication preferences of decision-makers improves the impact of research findings.

Limitations and Professional Escalation Criteria

Animal communication research has inherent limitations that should be acknowledged when interpreting findings and making management decisions.

Limitations of Signal Interpretation

The function of a signal cannot be determined from its form alone. The same signal may serve different functions in different contexts, and the receiver's response is influenced by its own state and experience. Interpretations of signal function should be based on systematic observation and, where possible, experimental manipulation.

Limitations of Cross-Species Generalization

Communication systems are species-specific, and findings from one species may not apply to another. Even closely related species can have very different communication systems. Management decisions should be based on species-specific evidence instead of general assumptions.

Limitations of Captive Studies

Observations of communication in captive settings may not reflect natural behavior. Captive environments differ from natural habitats in space, social composition, and environmental complexity. The Tasmanian devil interaction system was analyzed in captivity [3], and while this provided valuable data, captive observations should be complemented with field studies where possible.

Professional Escalation Criteria

Seek professional advice when communication observations indicate potential welfare problems that exceed your expertise. Escalate to a veterinarian if you observe signs of pain, illness, or injury. Escalate to an animal behaviorist if you observe persistent aggression, stereotypic behavior, or social disruption. Escalate to a wildlife authority if you encounter injured or orphaned wild animals that require specialized care.

Frequently Asked Questions

What is animal communication?

Animal communication is the transfer of information between individuals through signals that can be perceived by one or more sensory systems. The four main types are visual, auditory, chemical, and tactile signals. Most species use combinations of these modalities depending on context, distance, and the relationship between signaler and receiver.

What are examples of animal communication systems?

Examples include the vocalizations of Tasmanian devils, which have been classified into four basic syllable types related to different functions [3]. Fireflies use flashing light signals while crickets use chirping sounds, and both communicate at similar tempos [7]. Chimpanzees use gestures and modify their modality according to social bond strength [13]. Many insects and spiders use substrate-borne vibrations for communication [14].

How do animals use visual communication?

Animals use visual signals such as posture, movement, color change, and facial expression to convey information about aggression, submission, reproductive status, and identity. Visual signals require line of sight and are most effective over short to medium distances. The Tasmanian devil uses visual signals as part of its female-male interaction system [3].

How does auditory communication work in animals?

Auditory communication involves the production of sound through vocal cords, stridulation, drumming, or other mechanisms. Sound travels well through air and water, works in darkness, and can carry over long distances. Prosodic modulation of the voice can express emotional information and coordinate vocal interactions [11]. Many species communicate at tempos of approximately 0.5 to 4 Hz [7].

What is chemical communication in animals?

Chemical communication involves the production and detection of chemical signals, including pheromones and scent marks. These signals convey information about species identity, individual identity, reproductive status, and territory ownership. Chemical signals persist in the environment after the signaler leaves, allowing information to be transmitted over time.

What is tactile communication in animals?

Tactile communication involves physical contact between individuals, including grooming, touching, nudging, and fighting. It reinforces social bonds, provides reassurance, and coordinates cooperative behavior. Chimpanzees with stronger social bonds use more tactile gestures [13]. Biotremology is the study of communication through substrate-borne vibrations [14].

Why are no animal communication systems simple languages?

Animal communication systems lack the productivity of human language, meaning they cannot be used to say many different things [8]. A necessary condition for the evolution of a simple language is a high degree of alignment of interests between signaler and receiver [8]. Many animal species have the cognitive capacity for language but lack the socio-ecology that requires high-level cooperation across a range of contexts [8].

How can I observe animal communication in the field?

Define a clear research question, select appropriate observation methods, record signal production and receiver response, establish baseline patterns, and analyze data systematically. Record contextual information such as distance, time, and environmental conditions. Be aware of observer effects, context neglect, modality bias, and equipment limitations.

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Communication in the tasmanian devil (Sarcophilus barrisii) and a survey of auditory communication in the marsupialia.. Zeitschrift fur Tierpsychologie, 1975.
• Factors Influencing Decision-Making in Companion Animal Euthanasia: A Mixed-Methods Study of Pet Owners and Veterinarians.. 2026.
• What policymakers want: policy briefs as tools to inform animal disease control strategies.. 2026.
• Easy rescues, difficult decisions: Advancing the ethical dialogue in wildlife triage, treatment and rehabilitation.. 2026.
• A widespread animal communication tempo may resonate with the receiver's brain.. 2026.
• Why Are No Animal Communication Systems Simple Languages?. Frontiers in Psychology, 2021.
• Commentary: Why Are no Animal Communication Systems Simple Languages?. Frontiers in Psychology, 2021.
• Commentary: Why Are No Animal Communication Systems Simple Languages?. Frontiers in Psychology, 2021.
• Emotional and Interactional Prosody across Animal Communication Systems: A Comparative Approach to the Emergence of Language. Frontiers in Psychology, 2016.
• Do Animal Communication Systems Have Phonemes?. Trends in Cognitive Sciences, 2015.
• Wild chimpanzees modify modality of gestures according to the strength of social bonds and personal network size. Scientific Reports, 2016.
• Biotremology. Current Biology, 2016.

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