What Animals Use Echolocation? A Guide to Nature's Sonar Users

Echolocation is the biological process where an animal emits sound pulses and interprets the returning echoes to sense objects, prey, and obstacles in its environment. This active sensory system is used by a diverse range of species, including bats, toothed whales and dolphins, shrews, some birds, and even certain rodents. The most well-known echolocators are bats and dolphins, which emit bursts of sound and listen to the echoes that bounce back to detect objects in their surroundings. This guide provides a practical reference for students, researchers, and life-science professionals seeking a clear list of echolocating animals, their habitats, and the specific methods they use.

At a Glance: Echolocating Animals and Their Methods

The table below summarizes the primary animal groups that use echolocation, their typical habitats, and the general method of sound production and processing. This quick-reference format is designed for field identification and comparative study.

Defining Echolocation and Its Core Principles

Echolocation is the use of reflected sound to sense features of the environment. Animals that echolocate actively emit pulses of ultrasound and perceive the acoustic reflections to determine the position of objects in dark, often three-dimensional spaces. The specificity of echolocation depends on the properties of the calls emitted, such as frequency, duration, and call shape, and how this ultrasound is modulated, reflected, received, and interpreted.

A global macroevolutionary analysis identified 1,329 echolocating mammal and bird species with a predominantly pan-tropical distribution, plus an additional 117 unconfirmed echolocators. This analysis characterized calls based on 5,146 acoustic records from 2,838 terrestrial and aquatic study sites across 140 countries. The study established that echolocation calls were linked to call frequency and variation parameters associated with orientation function, and to call type and duration that mediated prey acquisition. Morphological and phylogenetic constraints substantially influenced echolocation call designs at the global scale.

The fundamental principle is that sound travels through a medium, reflects off surfaces, and returns to the emitter. The time delay between emission and return indicates distance, while changes in frequency and amplitude provide information about the size, shape, and movement of objects. Echolocating animals have evolved specialized auditory and vocal systems to produce and process these signals with remarkable speed and precision.

Bats: The Most Diverse Echolocating Mammals

Bats are the most well-studied echolocating animals, with approximately 700 species in the sub-order Microchiroptera, representing an estimated 25% of living mammals. Many echolocating bats are nocturnal predators that have evolved a biological sonar system to orient and forage in three-dimensional space. Acoustic signal processing and vocal-motor control are tightly coupled, and successful echolocation depends on the coordination between auditory and motor systems.

Laryngeal Echolocation and Its Evolution

Laryngeal echolocation refers to calls produced in the larynx. Recent molecular phylogenies suggest that laryngeal echolocation probably evolved in the ancestor of all extant bats. Echolocation might have subsequently been lost in Old World fruit bats, only to evolve secondarily by tongue clicking in this family. Remarkable acoustic features such as Doppler shift compensation, whispering echolocation, and nasal emission of sound each show multiple convergent origins in bats.

The extensive adaptive radiation in echolocation call design is shaped largely by ecology. Perceptual challenges imposed by the environment can often override phylogenetic constraints. For example, bats that forage in open spaces tend to use lower frequency, longer duration calls that travel farther, while bats that forage in cluttered environments use higher frequency, shorter calls that provide more detailed information about nearby objects.

The Stylohyal Bone Connection

A key anatomical feature of laryngeally echolocating bats is the stylohyal bone that connects the larynx to the auditory bulla. This bone is used for heterodyne detection of Doppler-shifted echoes, allowing very precise frequency resolution and phase-sensitive analysis of the returning echoes for determining the velocity of echolocated objects like insects. The proximal portion of the stylohyal bone fuses with the tympanic bone in laryngeally echolocating bats, a form previously only seen in this group.

Call Structure and Energy Expenditure

Echolocation is metabolically demanding. Studying the energy costs of echolocation helps researchers understand animal energy allocation and provides insights into the evolutionary constraints of acoustic signals. Research on the constant-frequency bat Rhinolophus nippon used a miniature electrocardiogram system and a custom servomotor that moved prey toward stationary bats. During the search phase, bats emitted isolated echolocation pulses characterized by long pulse durations and inter-pulse intervals, together with higher root mean square amplitude, pulse energy, and peak amplitude.

In the approach phase, call rate increased significantly by 3.15-fold, and bats predominantly produced sonar sound groups. Pulse duration, inter-pulse intervals, root mean square amplitude, and pulse energy decreased to 65.23%, 25.82%, 78.50%, and 86.32% of the corresponding search-phase values, whereas peak amplitude increased to 110.99%. This indicates that bats can flexibly adjust the structure of their echolocation calls. Despite the increased call rate, neither heart rate nor metabolic rate differed between phases, providing direct physiological evidence for understanding energy expenditure in bat echolocation.

Audiovocal Control and Vocal Learning

Fine audiovocal control is a hallmark of human speech production and depends on precisely coordinated muscle activity guided by sensory feedback. Research on the echolocating bat Hipposideros armiger applied state feedback control theory to the analysis of call frequency adjustments. Individual bats exhibited highly distinct patterns of vocal compensation to spectrally altered calls, mirroring typical observations of speech control in humans listening to spectrally altered speech. The same computational principles of state feedback control apply to bat echolocation and human speech.

Vocal production learning, a core ability underlying human speech, is rare among mammals. Apart from humans, two of the four identified vocal production learning mammalian lineages, cetaceans and bats, are echolocators. Research on Pratt's roundleaf bats (Hipposideros pratti) used a psychophysical reinforcement learning paradigm to train bats to detect pure tones and virtual echoes. Bats learned to adapt their call amplitude to auditory tasks across various time scales, ranging from instantaneous to months. Bats modified the noise-induced call amplitude adaptation, a reflexive audio-vocal behavior known as the Lombard effect, in a task-specific manner. Perceptual demands for echolocation efficiency over longer time scales may have facilitated the evolution of this rare vocal production learning ability in echolocating mammals.

Individual Signatures in Bat Calls

Deep learning methods have revealed persistent individual signatures within bat echolocation calls. Recordings from 34 individuals of the greater leaf-nosed bat (Hipposideros armiger) under controlled laboratory conditions showed that a convolutional neural network achieved an average identification accuracy of 84% for single calls and 91% for call sequences. In contrast, traditional Discriminant Functional Analysis achieved accuracies of only 39% and 47%. Temporal patterning enhances individual classification accuracy, suggesting it contributes to the encoding of individual-specific information. Echolocation calls can contain stable individual identity that was previously undetectable.

Velocity and Information Sampling

Moving animals must gather information at sufficient rates, detail, and range relative to their velocity while filtering this information to that essential for a given task. Echolocators are exceptional models for investigating how animals filter and adjust information flow to motor patterns. Research on three sympatric bats of the same foraging guild, edge-space hawkers, but different niches, found that the trawler Myotis daubentonii and the hawker Pipistrellus pygmaeus exhibit similar flight and echolocation behavior, whereas the gleaner Myotis nattereri flies slower and produces calls of lower duration and intensity, greater bandwidth and call interval, but similar beam breadth.

These differences in echolocation behavior converge when accounting for flight speed. These species move equivalent distances between call emissions and travel through their respective sonar ranges in the same time interval. Each echolocation call's duration is related to the two-way travel time of its sonar range, and thus velocity, the same way across species. This similarity suggests general mechanisms of information processing and conserved traits underlying auditory attention in vespertilionid bats and perhaps other echolocators.

Toothed Whales and Dolphins: Aquatic Echolocators

Toothed whales and dolphins, collectively known as odontocetes, are the primary aquatic echolocators. They produce high-frequency clicks through nasal passages, which are focused by a fatty structure called the melon in the forehead. These clicks travel through water and return as echoes that provide detailed information about the underwater environment.

Odontocetes use echolocation for navigation, prey detection, and social communication in the often dark or turbid waters they inhabit. The evolution of echolocation in this group is distinct from bats, reflecting the different physical properties of sound transmission in water compared to air. Water is denser than air, allowing sound to travel faster and farther, which influences the design and use of echolocation signals.

Sperm whales produce some of the most powerful echolocation clicks in the animal kingdom, used to detect prey at depths where light does not penetrate. Dolphins are known for their ability to discriminate between objects of different sizes, shapes, and materials using echolocation alone. The auditory processing capabilities of odontocetes are highly specialized, with large auditory brain regions dedicated to interpreting returning echoes.

Shrews and Rodents: Terrestrial Echolocators

Several terrestrial mammals use echolocation, including shrews and some rodents. The soft-furred tree mice (Typhlomys) are a newly identified lineage of echolocating mammals. Behavioral experiments show that these mice can locate and avoid obstacles in darkness using hearing and ultrasonic pulses. The proximal portion of their stylohyal bone fuses with the tympanic bone, a form previously only seen in laryngeally echolocating bats.

Further, researchers found convergence of hearing-related genes across the genome and of the echolocation-related gene prestin between soft-furred tree mice and echolocating mammals. These findings suggest that soft-furred tree mice are capable of echolocation and represent a new lineage of echolocating mammals.

Shrews use echolocation primarily for close-range orientation in their underground tunnels and dense ground vegetation. Their echolocation calls are typically low-intensity ultrasonic pulses that provide information about nearby obstacles and potential prey. Unlike bats, which use echolocation for long-range navigation and prey capture, shrews use it for short-range spatial awareness in confined environments.

Birds That Echolocate

Two groups of birds are known to use echolocation: oilbirds and swiftlets. These birds inhabit caves and use echolocation for obstacle avoidance in complete darkness. Unlike bats, which produce ultrasonic calls, these birds use audible clicks or short calls that are within the human hearing range.

Oilbirds (Steatornis caripensis) are nocturnal fruit-eating birds that roost and nest in caves. They produce rapid clicks that allow them to navigate through cave passages in darkness. Swiftlets, which are small insectivorous birds found in Southeast Asia and the Pacific, produce similar echolocation calls for navigating their cave roosts.

The echolocation method in these birds is less sophisticated than in bats or dolphins, providing primarily coarse information about the presence and location of large obstacles instead of detailed information about prey. This reflects the different ecological demands on these species, which use echolocation mainly for spatial orientation instead of prey capture.

Human Echolocation: A Learned Skill

Some blind people have learned to echolocate, making mouth clicks and using the returning echoes from those clicks to sense obstacles and objects of interest in their surroundings. Blind echolocation experts can sense small differences in the location of objects, differentiate between objects of various sizes and shapes, and even between objects made of different materials, just by listening to the reflected echoes from mouth clicks.

Neuroimaging has shown that the processing of echoes activates brain regions in blind echolocators that would normally support vision in the sighted brain. The patterns of these activations are modulated by the information carried by the echoes. Echolocation may enable some blind people to do things that are otherwise thought to be impossible without vision, potentially providing them with a high degree of independence in their daily lives. This demonstrates that echolocation can serve as an effective mobility strategy in the blind and sheds new light on the plasticity of the human brain.

Practical Assessment Steps for Identifying Echolocating Animals

For researchers and life-science professionals conducting field assessments, the following steps provide a structured approach to identifying and studying echolocating animals.

Step 1: Determine the Environment

Identify whether the study site is terrestrial, aquatic, or aerial. This immediately narrows the potential echolocators. Bats dominate aerial and cave environments, odontocetes dominate marine and some freshwater environments, and shrews and rodents are found in terrestrial habitats. Oilbirds and swiftlets are restricted to cave environments in specific geographic regions.

Step 2: Use Appropriate Detection Equipment

Ultrasonic detectors are essential for detecting bat and rodent echolocation calls, which are typically above the human hearing range. Aquatic hydrophones are required for recording dolphin and whale clicks. For birds and human echolocators, standard audio recording equipment may suffice since their calls are often audible.

Step 3: Record Call Characteristics

Document the frequency, duration, intensity, and pattern of echolocation calls. These characteristics vary by species and provide diagnostic information. Note whether calls are constant frequency, frequency modulated, or a combination. Record the inter-pulse interval and whether calls are emitted singly or in groups.

Step 4: Observe Behavioral Context

Note the behavior of the animal during echolocation. Is it searching for prey, navigating, or avoiding obstacles? Bats increase their call rate significantly during prey approach, as observed in Rhinolophus nippon where call rate increased 3.15-fold from search to approach phase. Behavioral context influences call structure and provides clues about the function of echolocation in different situations.

Step 5: Consider Morphological Features

Examine anatomical features that support echolocation. The fusion of the stylohyal bone with the tympanic bone is a key indicator of laryngeal echolocation in bats and has also been found in soft-furred tree mice. In odontocetes, the presence of a melon and specialized nasal passages indicates echolocation capability.

Records and Measurements for Echolocation Studies

Maintaining systematic records is critical for echolocation research. The following measurements are standard in the field and provide comparable data across studies.

When recording echolocation calls, standardize the distance between the animal and the recording device, as amplitude measurements are distance dependent. Record environmental conditions including temperature, humidity, and ambient noise levels, as these affect sound transmission and echo characteristics. For aquatic recordings, note water temperature, salinity, and depth.

Common Failure Patterns in Echolocation Observation

Several common errors can compromise echolocation studies. Recognizing these patterns helps researchers avoid invalid conclusions.

Misidentifying Non-Echolocation Sounds

Many animals produce sounds that are not echolocation. Social calls, alarm calls, and territorial vocalizations can be confused with echolocation signals. Echolocation calls are typically stereotyped, repetitive, and tightly coupled to movement or orientation behavior. Confirm that the sound is used for spatial sensing before classifying it as echolocation.

Ignoring Environmental Effects on Sound Transmission

Temperature, humidity, and vegetation density affect sound propagation and echo characteristics. Calls recorded under different environmental conditions may not be directly comparable. Standardize recording conditions or account for environmental variables in analysis.

Overlooking Individual Variation

Echolocation calls vary between individuals. Deep learning analysis of Hipposideros armiger revealed individual signatures within echolocation calls that traditional methods could not detect. Studies that pool calls across individuals without accounting for individual variation may miss important patterns or draw incorrect conclusions about species-level call characteristics.

Assuming Static Call Structure

Echolocation calls are highly flexible and change with behavioral context. Bats adjust call amplitude, duration, and rate based on task demands and environmental conditions. Studies that assume a fixed call structure for a species will fail to capture the adaptive nature of echolocation.

Limitations of Echolocation Research

Echolocation research faces several inherent limitations that should be acknowledged in any study design.

Technical Constraints

Recording ultrasonic calls requires specialized equipment that may not be available in all field settings. The directionality of echolocation calls means that recordings depend heavily on the position of the microphone relative to the animal's head. Calls emitted away from the microphone may be severely attenuated or missed entirely.

Species Coverage Gaps

While a global macroevolutionary analysis identified 1,329 echolocating mammal and bird species, an additional 117 species remain unconfirmed echolocators. Many species have not been studied in sufficient detail to determine whether they echolocate. The absence of evidence for echolocation in a species does not confirm the absence of echolocation.

Captivity Effects

Laboratory studies of echolocation may not reflect natural behavior. Bats recorded in controlled laboratory conditions may produce different call structures than those in the wild. The study of Hipposideros armiger individual signatures was conducted under controlled laboratory conditions, and the authors noted the need for future studies to monitor animals in more dynamic environments.

Ethical Considerations

Recording animal vocalizations, particularly from UAVs or other intrusive methods, raises ethical considerations. Researchers must balance the value of acoustic data against potential disturbance to animals. The first documented successful recordings of animals in their natural habitat using UAVs demonstrated that these tools can be important for bioacoustic monitoring, but the ethical implications of such monitoring require careful consideration.

Welfare and Safety Context

Studying echolocating animals requires attention to both animal welfare and researcher safety.

Animal Welfare

Minimize disturbance to echolocating animals during observation and recording. Bats are particularly sensitive to disturbance at roost sites, and repeated disturbance can cause roost abandonment. Use passive acoustic monitoring where possible to reduce direct interaction with animals. When capture or handling is necessary, follow institutional animal care protocols and minimize handling time.

Researcher Safety

Field studies of echolocating animals may involve working in caves, at night, or in aquatic environments. Cave work requires appropriate safety equipment and training. Night work in terrestrial habitats requires awareness of local hazards. Aquatic work requires appropriate boating safety measures and awareness of the behavior of large odontocetes.

Regulatory Compliance

Research involving echolocating animals may require permits under wildlife protection laws. Many bat species are protected, and disturbing roosts may be illegal without appropriate authorization. Marine mammal research is regulated in many jurisdictions. Check local regulations before beginning any study involving echolocating animals.

Professional Escalation Criteria

Researchers should seek expert consultation when encountering situations beyond their expertise. The following criteria indicate when escalation to a specialist is appropriate.

Unusual Call Patterns

If recorded echolocation calls do not match known species descriptions, consult an expert in bioacoustics. Novel call patterns may indicate undocumented species, unusual behavioral states, or recording artifacts.

Morphological Anomalies

If anatomical examination reveals structures inconsistent with known echolocation adaptations, such as unusual stylohyal bone morphology, consult a comparative anatomist. Such findings may represent new evolutionary insights or pathological conditions.

Conservation Concerns

If observations suggest population declines, habitat threats, or disturbance impacts on echolocating species, escalate to conservation authorities. The integration of animal culture into conservation planning is an emerging consideration, with culturally transmitted migration routes found to be 20% more resilient to habitat disruptions than genetically inherited migration routes.

Equipment Limitations

If standard recording equipment cannot capture the signals of interest, consult with specialists who have access to advanced acoustic technology. UAV-based recording, deep learning analysis, and other emerging methods may be necessary for certain research questions.

Frequently Asked Questions

What is the difference between echolocation and passive listening?

Echolocation is an active sensory system where the animal emits sound and interprets the returning echoes. Passive listening involves receiving and interpreting sounds produced by other sources. Echolocation provides the animal with information about objects that do not produce their own sounds, while passive listening only reveals information about sound-producing objects.

Do all bats use echolocation?

No. While most bats use laryngeal echolocation, Old World fruit bats do not use laryngeal echolocation. Molecular phylogenies suggest that echolocation may have been lost in Old World fruit bats and then evolved secondarily by tongue clicking in this family. Some bats rely primarily on vision and smell for orientation and foraging.

Can humans learn to echolocate?

Yes. Some blind people have learned to echolocate by making mouth clicks and using the returning echoes to sense obstacles and objects. Blind echolocation experts can differentiate between objects of various sizes, shapes, and materials. Neuroimaging shows that echo processing activates brain regions in blind echolocators that normally support vision in sighted individuals.

How do echolocating bats avoid deafening themselves with their own calls?

Echolocating bats have specialized middle ear muscles that contract during call emission, reducing the transmission of the outgoing call to the inner ear. This allows the auditory system to remain sensitive to the quieter returning echoes. The coordination between vocal production and auditory processing is tightly coupled in echolocating bats.

What is the stylohyal bone and why is it important?

The stylohyal bone connects the larynx to the auditory bulla in laryngeally echolocating bats. It is proposed to be used for heterodyne detection of Doppler-shifted echoes, allowing precise frequency resolution and phase-sensitive analysis of returning echoes. The proximal portion of this bone fuses with the tympanic bone in laryngeally echolocating bats, and this same fusion was found in soft-furred tree mice.

How fast can echolocating animals process echoes?

The speed at which the brain and muscles can respond to information about prey location constrains visual and echolocating predators in similar ways. Echolocating bats adjust their call rate and structure based on the distance to targets, with call duration related to the two-way travel time of the sonar range. The neural computations must occur within the time between call emission and echo return.

Do echolocating animals use echolocation for communication?

Echolocation calls primarily serve for orientation and prey detection, but they can also carry individual identity information. Deep learning analysis revealed persistent individual signatures within echolocation calls of Hipposideros armiger, suggesting that echolocation calls can contain stable individual identity. However, social communication in echolocating animals typically involves separate vocalizations distinct from echolocation calls.

Which bird species use echolocation?

Oilbirds and swiftlets are the two groups of birds known to use echolocation. These birds inhabit caves and use audible clicks for obstacle avoidance in darkness. Their echolocation is less sophisticated than that of bats or dolphins, providing primarily coarse information about large obstacles instead of detailed prey information.

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Echolocation in humans: an overview.. Wiley interdisciplinary reviews. Cognitive science, 2016.
• Cynthia F. Moss.. Current biology : CB, 2021.
• The brain limit.. eLife, 2021.
• Echolocation in soft-furred tree mice.. Science (New York, N.Y.), 2021.
• The evolution of echolocation in bats.. Trends in ecology & evolution, 2006.
• Sensory error drives fine motor adjustment.. Proceedings of the National Academy of Sciences of the United States of America, 2022.
• Laryngeally echolocating bats.. Nature, 2010.
• Neurobiology of echolocation in bats.. Current opinion in neurobiology, 2003.
• Energy Expenditure Optimization in the Echolocation of &lt,i&gt,Rhinolophus nippon&lt,/i&gt,: Evidence from Heart Rate Stability.. 2026.
• Task learning drives adaptive vocal adjustments in echolocating bats.. 2026.
• Deep Learning Reveals Persistent Individual Signatures in Bat Echolocation Calls of the Greater Leaf-nosed Bat. 2026.
• Decoding the functional adaptation of animal echolocation: a global macroevolutionary analysis.. 2026.
• Integrating Animal Culture into Red List Assessments and Recovery Planning for Migratory Species to Enhance Conservation Strategies. Journal of Animal Environment, 2025.
• Recording animal vocalizations from a UAV: bat echolocation during roost re-entry. Scientific Reports, 2018.
• Velocity as an overlooked driver in the echolocation behavior of aerial hawking vespertilionid bats.. Current Biology, 2025.
• Introduction to special issue, "how nature shaped echolocation in animals". Frontiers in Physiology, 2013.
• Echolocation in bats, odontocetes, birds, and insectivores. Exploring Animal Behavior Through Sound Volume 1 Methods, 2022.
• Bats, Model Animals for Hearing and Echolocation Studies. Progress in Biochemistry and Biophysics, 2022.
• Different as night and day: wild bats modify echolocation in complex environments when visual cues are present. Animal Behaviour, 2020.

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