Visual Signals in Animal Communication: Displays, Colors, and Body Language

Animals use visual signals to convey information about identity, status, intent, and environmental conditions. These signals include color displays, body postures, facial expressions, and bioluminescent emissions. Visual communication operates when a sender produces a signal that alters the behavior of a receiver, and effective signals are conspicuous, redundant, and stereotypic to reduce ambiguity (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). This article explains the main categories of visual signaling, how animals produce and perceive these signals, and how observers can identify and interpret them in research, clinical, and field settings.

Visual signals differ from acoustic and olfactory signals in important ways. Visual signals can be adjusted rapidly for response during interactions between individuals at close or medium range, while olfactory signals persist in the environment and acoustic signals do not (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). This temporal flexibility makes vision a primary channel for moment-to-moment social negotiation in many species.

At a Glance: Categories of Visual Signals

The table below organizes visual signals by functional category, mechanism, and representative examples. This classification follows the functional framework proposed for interspecific visual signaling, which divides signals into categories such as anti-predator, food acquisition, reproductive, and agonistic signaling (Interspecific visual signalling in animals and plants: a functional classification, Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2017).

The Functional Framework of Visual Signaling

Visual signaling serves multiple ecological functions, and the same signal mechanism can serve different purposes depending on context. The functional classification of interspecific visual signaling identifies six major categories: anti-predator, food acquisition, anti-parasite, host acquisition, reproductive, and agonistic signaling (Interspecific visual signalling in animals and plants: a functional classification, Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2017). Each category is enabled by several distinct mechanisms, and understanding this structure helps observers predict what a signal means in a given context.

Anti-Predator Visual Signals

Anti-predator visual signals include camouflage, startle displays, and warning coloration. Cuttlefish and other cephalopods can change their skin color and texture rapidly to blend with their surroundings or to produce startling patterns that deter predators. These signals are adjusted in real time based on visual feedback from the environment.

Startle displays often combine color change with body posture to create a sudden, conspicuous appearance that interrupts a predator's attack sequence. Warning coloration, in contrast, is a persistent signal that advertises toxicity or unpalatability. The receiver learns to associate the visual pattern with a negative experience, reducing future attacks.

Food Acquisition Signals

Some animals use visual signals to attract prey. Deep-sea stomiid fish possess photophores, specialized light-emitting organs, that function as lures to attract prey toward their mouths (The Skin Photophores of Chauliodus sloani Bloch & Schneider, 1801 (Pisces: Stomiidae), Animals, 2025). These lures exploit the prey's attraction to light in an otherwise dark environment.

Anglerfish use a similar strategy with a modified dorsal fin spine that bears a luminous lure. The signal works because the prey mistakes the light for a small organism or other food source. This mechanism demonstrates how visual signals can manipulate receiver behavior for the sender's nutritional benefit.

Reproductive and Agonistic Signals

Reproductive visual signals advertise species identity, sex, and individual quality. The peacock's tail feathers are a classic example of a color ornament that females evaluate during mate choice. These displays are often costly to produce and maintain, which makes them honest indicators of condition.

Agonistic signals communicate dominance, submission, or threat during competitive interactions. Dogs use visual displays of relative dominance or submission to reduce the likelihood of physical conflict (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). A dog that rolls onto its back exposes its belly, signaling submission, while a dog that stands tall with erect ears and tail signals confidence or threat.

Color as a Visual Signal

Color vision relies on parallel inputs tuned to spectral content, and it is important for detection of objects such as mates, flowers, and oviposition sites (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021). Color signals can be static, such as feather or fur pigmentation, or dynamic, such as the rapid color changes seen in cephalopods and some fish.

Structural and Pigmentary Color

Animal colors arise from two main mechanisms: pigments that absorb specific wavelengths and structural features that reflect or scatter light. Structural color can produce iridescent effects that change with viewing angle, as seen in many butterfly wings and bird feathers. These structural colors often require precise nanoscale architecture, and their production is sensitive to developmental conditions.

Pigmentary color is produced by molecules such as melanins, carotenoids, and pterins. Carotenoids cannot be synthesized by animals and must be obtained from the diet, which makes carotenoid-based coloration an honest signal of foraging ability and health.

Color Perception and Visual Modeling

Understanding what an animal sees requires knowledge of its visual system. Different species have different numbers and types of photoreceptors, which determine their color discrimination abilities. Butterflies, stomatopods, and jumping spiders have particularly well-developed color vision (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021).

Researchers use spectral measurements and visual modeling to estimate how colors appear to a given species. This approach involves measuring the reflectance spectrum of the signal and modeling how the receiver's photoreceptors would respond. The method is described in the zoological literature as a way to study animal coloration objectively (Using spectra and visual modeling to study animal coloration, Zoological Research, 2013).

Dynamic Color Change

Some animals can change their color rapidly in response to social or environmental cues. Cuttlefish and chameleons control specialized pigment cells called chromatophores to produce patterns that serve camouflage, courtship, or threat functions. These changes are under neural control and can occur in fractions of a second.

Dynamic color change requires a sophisticated integration of visual input and motor output. The animal must perceive its background or the social context and then produce an appropriate color pattern. This ability is particularly developed in cephalopods, which have large brains relative to body size.

Body Posture and Movement Signals

Body posture and movement provide a rich channel for visual communication, especially among social mammals. These signals can convey information about dominance, submission, play intent, and emotional state.

Canine Postural Signals

Dogs use a range of visual signals in dog-dog and dog-human communication. Communication is fundamental to maintaining affiliations, reducing competition, and identifying individuals in dogs (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). Veterinarians act as receivers for communication signals when greeting dogs as patients, and the information obtained from these signals informs clinical handling decisions (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997).

A dog's tail position, ear orientation, body weight distribution, and facial expression combine to signal its internal state. A relaxed dog holds its tail in a neutral position with soft eyes and a loose body. A fearful dog may tuck its tail, flatten its ears, and lower its body. An aggressive dog may stiffen its body, raise its hackles, and stare directly at the receiver.

Human perception of canine facial signals varies with breed morphology. Eye-tracking research shows that brachycephalic dogs elicit more visual attention from human viewers than normocephalic dogs, with the strongest effects for the ears, followed by the eyes, forehead, and snout (Analysis of human-oriented facial signals of the domestic dog using eye tracker technology, Frontiers in Veterinary Science, 2026). This finding has practical implications for reading dog behavior across breeds with different facial structures.

Primate Gestures and Multimodal Signals

Chimpanzees produce vocalizations in conjunction with other communicative signals. Data from 101 captive chimpanzees show that approximately 50% of vocal events were produced with another communicative modality, and directed vocalizations were more likely to include a signal from another modality than undirected vocalizations (Multimodal communication in chimpanzees, American journal of primatology, 2015). This multimodal pairing suggests that chimpanzees use their communicative signals strategically to meet specific socio-communicative ends.

Primate gestures include manual signals such as reaching, pointing, and begging, as well as facial expressions and body postures. These gestures can be combined with vocalizations to reinforce the message or to direct it to a specific individual.

Visual Signals in Bats

Bats rely heavily on sound and olfaction due to their nocturnal lifestyles, but visual signals still play a role in their communication (Social communication in bats, Biological reviews of the Cambridge Philosophical Society, 2018). Some bat species use visual cues in combination with sound, particularly species that emit constant frequency echolocation calls, to recognize conspecifics from heterospecifics.

The study of bat communication demonstrates that signal modalities are often used in combination instead of in isolation. A bat may use olfactory signals for species recognition and acoustic signals for individual identity, with visual signals providing supplementary information when light is available.

Bioluminescence as a Visual Signal

Bioluminescence, the emission of light catalyzed by luciferases, has evolved in many taxa from bacteria to vertebrates and is predominant in the marine environment (Bioluminescence and Photoreception in Unicellular Organisms: Light-Signalling in a Bio-Communication Perspective, International journal of molecular sciences, 2021). In animals with nervous systems capable of integrating light stimuli, bioluminescence triggers behavioral responses and plays a role in intra- or interspecific visual communication.

Functions of Bioluminescent Signals

Bioluminescent signals serve multiple functions in marine environments. Photophores in fish can serve defense against predators, prey attraction, intraspecific communication and recognition, and counterillumination (The Skin Photophores of Chauliodus sloani Bloch & Schneider, 1801 (Pisces: Stomiidae), Animals, 2025). Counterillumination involves matching the downwelling light from the surface to conceal the animal's silhouette from predators below.

Fireflies use bioluminescent flashes for mate attraction, with each species having a characteristic flash pattern. The timing, duration, and intensity of flashes provide information about species identity and individual quality.

Bioluminescence in Unicellular Organisms

The function of light emission in unicellular organisms is less clear than in animals. It is currently thought that bioluminescence in these organisms evolved in an ecological framework to be perceived by visual animals (Bioluminescence and Photoreception in Unicellular Organisms: Light-Signalling in a Bio-Communication Perspective, International journal of molecular sciences, 2021). For example, bioluminescence may allow bacteria to be ingested by zooplankton or fish, providing favorable conditions for growth and dispersal, while luminous flashes emitted by dinoflagellates may have evolved as an anti-predation system against copepods.

Recent findings show that bacteria and protists possess photoreceptors capable of perceiving and integrating light stimuli of different wavelengths, and single-cell light perception produces responses ranging from phototaxis to more complex behaviors (Bioluminescence and Photoreception in Unicellular Organisms: Light-Signalling in a Bio-Communication Perspective, International journal of molecular sciences, 2021). This suggests that the interplay between light emission and reception could play multiple roles in unicellular organisms, similar to visual animals.

Symbiotic Bioluminescence

Some marine animals obtain their bioluminescent capabilities through symbiosis with luminous bacteria. The bacterium Vibrio fischeri synchronizes bioluminescence in marine animals through quorum sensing mechanisms (Vibrio fischeri Symbiotically Synchronizes Bioluminescence in Marine Animals via Quorum Sensing Mechanism, 2018). The bacteria produce light only when their population density reaches a threshold, which ensures that the light output is bright enough to be visible.

This symbiotic relationship requires careful host management of bacterial populations. The host animal provides nutrients to the bacteria and maintains them in specialized light organs, while the bacteria produce light that serves the host's communicative or defensive needs.

Polarized Light Signals

Some animals use polarized light for communication. Mantis shrimps produce and detect circularly polarized light, which serves as a communication signal in these species (Circularly Polarized Light as a Communication Signal in Mantis Shrimps, Current Biology, 2015). The visual systems of many arthropods are specialized for the detection of polarized light, which in addition to communication with conspecifics can be used for orientation and navigation (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021).

Polarized light signals are invisible to animals that lack the appropriate photoreceptors, which makes them a private communication channel. This privacy can be advantageous for signaling that needs to avoid detection by predators or competitors.

Visual Signal Production and Perception

The production and perception of visual signals are constrained by the physical limitations of vision. To achieve high spatial resolution, fine sampling in different directions is necessary, as demonstrated by the large eyes of dragonflies (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021). However, a comparatively tiny robber fly has similarly high visual resolution in the frontal visual field despite having eyes a fraction of the size of those of dragonflies.

Trade-Offs in Visual System Design

Visual systems face inherent trade-offs between resolution, sensitivity, and field of view. Optical superposition compound eyes perform particularly well for vision in low light, and other modifications to maximize photon capture involve large lenses, stout photoreceptors, and neural pooling of information (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021). Extreme adaptations even allow insects to see colors at very low light levels or to navigate using the Milky Way.

These trade-offs mean that a signal that is conspicuous to one species may be invisible to another. Understanding the receiver's visual system is essential for interpreting whether a particular color or pattern functions as a signal.

Signal Design Principles

Effective visual signals are conspicuous, redundant, and stereotypic (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). Conspicuous signals stand out from the background and are easy to detect. Redundant signals provide the same information through multiple features, which reduces the chance of miscommunication. Stereotypic signals are produced in a consistent, predictable manner that receivers can recognize.

These design principles apply across signal modalities. A dog's aggressive display combines multiple features such as raised hackles, bared teeth, and a stiff posture, all of which convey the same message. The redundancy ensures that the receiver perceives the threat even if one feature is obscured.

Observing and Recording Visual Signals

Systematic observation of visual signals requires a structured approach. The following steps provide a framework for recording visual communication in field or clinical settings.

Step 1: Define the Context

Record the social and environmental context in which the signal occurs. Note the species, sex, age, and social status of the sender and receiver. Describe the physical environment, including lighting conditions, distance between individuals, and presence of other animals.

Step 2: Describe the Signal

Document the specific features of the visual signal. For color signals, record the hue, saturation, brightness, and pattern. For postural signals, describe the position of the head, ears, tail, limbs, and body axis. For bioluminescent signals, note the color, intensity, duration, and timing of the emission.

Step 3: Record the Response

Observe and record the receiver's behavioral response. Did the receiver change its behavior after perceiving the signal? Was the response immediate or delayed? Did the receiver produce a signal in return?

Step 4: Repeat and Compare

Collect multiple observations of the same signal type across different contexts and individuals. Compare the form and function of the signal across these observations to identify consistent patterns and contextual variations.

Step 5: Use Appropriate Technology

For detailed analysis, use video recording, photography, or spectral measurement. Eye-tracking technology can reveal which features of a signal attract visual attention, as demonstrated in studies of human perception of dog faces (Analysis of human-oriented facial signals of the domestic dog using eye tracker technology, Frontiers in Veterinary Science, 2026). Spectral measurements and visual modeling provide objective data on color signals (Using spectra and visual modeling to study animal coloration, Zoological Research, 2013).

Records and Measurements

Maintaining systematic records of visual signal observations supports both research and practical management. The following measurements are commonly used in the study of visual communication.

Behavioral Metrics

Behavioral metrics include the frequency, duration, and latency of specific signals. Frequency measures how often a signal occurs within a defined time period. Duration measures how long a single signal lasts. Latency measures the time between a stimulus and the signal response.

Visual Attention Metrics

Eye-tracking studies measure fixation duration, number of visits, and saccades across facial regions or other signal features (Analysis of human-oriented facial signals of the domestic dog using eye tracker technology, Frontiers in Veterinary Science, 2026). These metrics reveal which features of a signal receive the most visual attention and how attention is distributed across the signal.

Spectral Measurements

Spectral measurements quantify the reflectance or emission spectrum of a color signal. These measurements provide objective data that can be modeled to predict how the signal appears to a receiver with a particular visual system.

Temporal Measurements

For dynamic signals such as bioluminescent flashes or rapid color changes, temporal measurements record the timing and sequence of signal elements. These measurements are essential for understanding signals that convey information through temporal patterns.

Common Failure Patterns in Signal Interpretation

Misinterpreting visual signals can lead to incorrect conclusions in research and problematic outcomes in animal management. The following failure patterns are common.

Ignoring Receiver Visual Systems

A signal that is conspicuous to a human observer may be invisible to the intended receiver, or vice versa. Humans have trichromatic color vision, while many other species have different numbers and types of photoreceptors. Interpreting signals without knowledge of the receiver's visual system can lead to false conclusions about signal function.

Confusing Signal with Non-Signal

Not all conspicuous features are signals. A feature must alter the behavior of a receiver to qualify as a communication signal (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). Bright coloration may serve camouflage or thermoregulation instead of communication. Observers should verify that a feature functions as a signal before classifying it as one.

Overlooking Multimodal Integration

Many animals combine visual signals with acoustic, olfactory, or tactile signals. Chimpanzees produce vocalizations with other communicative modalities in approximately 50% of vocal events (Multimodal communication in chimpanzees, American journal of primatology, 2015). Focusing only on the visual channel can miss important information conveyed through other modalities.

Misreading Breed or Species Differences

Signal form can vary within a species due to breed morphology or individual variation. Brachycephalic dog breeds elicit different visual attention patterns than normocephalic breeds, with humans showing more attention to the ears, eyes, forehead, and snout of brachycephalic dogs (Analysis of human-oriented facial signals of the domestic dog using eye tracker technology, Frontiers in Veterinary Science, 2026). Observers should account for these differences when interpreting signals.

Welfare and Safety Context

Understanding visual signals has direct implications for animal welfare and human safety. Misreading a threat signal can lead to injury, while misreading a submission signal can cause unnecessary stress.

Clinical Applications

Veterinarians and animal handlers act as receivers for communication signals when greeting animal patients (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). The information obtained from these signals informs handling decisions and can prevent bites or other injuries. A dog that shows signs of fear or aggression should be handled with additional precautions.

Breed-Specific Considerations

Breed morphology affects how humans perceive animal signals. Brachycephalic dogs elicit more visual attention from humans, which may affect how owners and handlers interpret their behavior (Analysis of human-oriented facial signals of the domestic dog using eye tracker technology, Frontiers in Veterinary Science, 2026). Understanding these differences can improve communication between humans and animals.

Environmental Enrichment

Providing opportunities for species-appropriate visual communication can improve welfare in captive animals. This may include providing visual barriers to reduce stressful visual contact, or providing mirrors or visual stimuli that encourage natural signaling behavior.

Limitations of Visual Signal Research

Research on visual communication faces several limitations that observers should recognize.

Taxonomic Bias

Research on visual communication is unevenly distributed across taxa. Some groups, such as primates and birds, are well studied, while others receive less attention. Bats, for example, have communication systems that are less studied than other taxa despite their diversity (Social communication in bats, Biological reviews of the Cambridge Philosophical Society, 2018).

Methodological Constraints

Studying visual signals in natural environments is challenging. Many signals are brief, context-dependent, or occur in habitats where observation is difficult. Deep-sea bioluminescence, for example, requires specialized equipment to observe.

Interpretive Challenges

Determining the function of a visual signal requires demonstrating that the signal alters receiver behavior. This demonstration is often difficult in field settings where many variables cannot be controlled. The functional classification of interspecific visual signaling provides a framework for organizing observations, but assigning signals to categories requires careful evidence (Interspecific visual signalling in animals and plants: a functional classification, Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2017).

Professional Escalation Criteria

Observers should seek professional consultation when they encounter situations that exceed their expertise.

When to Consult a Veterinarian

Consult a veterinarian when an animal shows persistent or unusual visual signals that may indicate pain, illness, or neurological dysfunction. Changes in posture, facial expression, or eye appearance can signal underlying health problems. A veterinarian can distinguish behavioral signals from clinical signs of disease.

When to Consult a Behavior Specialist

Consult a certified animal behavior specialist when visual signals are associated with aggression, fear, or other behavioral problems that pose a safety risk. A behavior specialist can develop a treatment plan that addresses the underlying causes of problematic signaling.

When to Consult a Research Methodologist

Consult a research methodologist when designing studies of visual communication, particularly studies involving spectral measurement, visual modeling, or eye-tracking technology. These methods require specialized expertise to implement correctly.

Frequently Asked Questions

What is the difference between a visual signal and a visual cue?

A visual signal is a feature that evolved specifically to convey information and that alters the behavior of a receiver. A visual cue is a feature that provides information to an observer but did not evolve for that purpose. Communication occurs when one individual produces a signal that alters the behavior of another individual (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). For example, a dog's tail wagging can be a signal if it evolved to communicate emotional state, while the dog's size is a cue that provides information about its physical capabilities.

How do animals produce color signals?

Animals produce color signals through pigments that absorb specific wavelengths and structural features that reflect or scatter light. Pigmentary color comes from molecules such as melanins and carotenoids, while structural color arises from nanoscale architecture that interacts with light. Some animals can change color rapidly by controlling pigment cells called chromatophores.

Why do some animals use bioluminescence for communication?

Bioluminescence is useful for communication in environments where ambient light is limited, such as the deep sea or at night. It allows animals to produce conspicuous signals without relying on external light sources. Bioluminescent signals serve functions including prey attraction, predator defense, and intraspecific communication (The Skin Photophores of Chauliodus sloani Bloch & Schneider, 1801 (Pisces: Stomiidae), Animals, 2025).

How can I tell if a visual feature is a communication signal?

A visual feature is a communication signal if it alters the behavior of a receiver. To test this, observe whether receivers respond differently when the feature is present versus absent. Effective signals are conspicuous, redundant, and stereotypic (Canine communication, The Veterinary clinics of North America. Small animal practice, 1997). If a feature does not change receiver behavior, it may be a cue instead of a signal.

Do animals combine visual signals with other communication channels?

Many animals combine visual signals with acoustic, olfactory, or tactile signals. Chimpanzees produce vocalizations with other communicative modalities in approximately 50% of vocal events, and directed vocalizations are more likely to include another modality than undirected vocalizations (Multimodal communication in chimpanzees, American journal of primatology, 2015). Bats also combine olfactory and acoustic signals for recognition (Social communication in bats, Biological reviews of the Cambridge Philosophical Society, 2018).

Why do different species perceive visual signals differently?

Different species have different visual systems with varying numbers and types of photoreceptors. These differences determine which wavelengths an animal can detect and how it discriminates between colors. Butterflies, stomatopods, and jumping spiders have particularly well-developed color vision (Stark trade-offs and elegant solutions in arthropod visual systems, The Journal of experimental biology, 2021). A signal that is conspicuous to one species may be invisible to another.

What should I do if I observe unusual visual signals in an animal?

If you observe persistent or unusual visual signals in an animal, particularly signals that may indicate pain, fear, or aggression, consult a veterinarian or animal behavior specialist. Changes in posture, facial expression, or eye appearance can signal underlying health or welfare problems. Professional assessment can determine whether the signals indicate a medical or behavioral issue.

How do researchers study animal coloration objectively?

Researchers use spectral measurements to quantify the reflectance or emission spectrum of color signals, then use visual modeling to predict how the signal appears to a receiver with a particular visual system (Using spectra and visual modeling to study animal coloration, Zoological Research, 2013). This approach provides objective data that can be compared across species and contexts.

Related Articles

• Competition in Biology: Definition, Types, and Examples
• Types Of Mutations
• Animal Biology
• Animal Research
• Types Of Sampling In Biostatistics

References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Interspecific visual signalling in animals and plants: a functional classification.. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2017.
• Canine communication.. The Veterinary clinics of North America. Small animal practice, 1997.
• Feline vocal communication.. Journal of veterinary science, 2020.
• Social communication in bats.. Biological reviews of the Cambridge Philosophical Society, 2018.
• Waisman Center.. International journal of developmental neuroscience : the official journal of the International Society for Developmental Neuroscience, 2002.
• Bioluminescence and Photoreception in Unicellular Organisms: Light-Signalling in a Bio-Communication Perspective.. International journal of molecular sciences, 2021.
• Multimodal communication in chimpanzees.. American journal of primatology, 2015.
• Stark trade-offs and elegant solutions in arthropod visual systems.. The Journal of experimental biology, 2021.
• A bibliometric analysis of alpha-synuclein in Parkinson's disease from 2015 to 2024.. 2026.
• The evolution of healthcare through the eye: From ancient superstition to the ophthalmoscope.. 2026.
• Virtual reality training for gait disorders in Parkinson’s disease: a bibliometric and visual analysis of hotspots and trends using CiteSpace and VOSviewer
• Analysis of human-oriented facial signals of the domestic dog using eye tracker technology.. 2026.
• Case Report: Preliminary evaluation of time-resolved imaging of contrast kinetics magnetic resonance imaging for assessing temporal enhancement patterns in small animal head and neck tumors.. 2026.
• Mitigating Blue-Light Risk in Display-Based Digital Therapeutics: A Practical Framework to Support Clinical Efficacy. 2026.
• Vibrio fischeri Symbiotically Synchronizes Bioluminescence in Marine Animals via Quorum Sensing Mechanism. 2018.
• Circularly Polarized Light as a Communication Signal in Mantis Shrimps.. Current Biology, 2015.
• The Skin Photophores of Chauliodus sloani Bloch & Schneider, 1801 (Pisces: Stomiidae): A Morphological, Ultrastructural and Immunohistochemical Study. Animals, 2025.
• Communication and language in animals. Bulletin De La Societe Royale Des Sciences De Liege, 2002.
• [Using spectra and visual modeling to study animal coloration].. Dong Wu Xue Yan Jiu Zoological Research Dong Wu Xue Yan Jiu Bian Ji Wei Yuan Hui Bian Ji, 2013.
• Visual communication in animals: Applying a Portmannian and Uexküllian biosemiotic approach. Visual Communication, 2014.

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