Tag: animal cognition

  • Corvid Intelligence: Tool Use, Planning, and Why Crows Hold Funerals

    A New Caledonian crow named Betty, in a 2002 experiment at Oxford, bent a straight piece of wire into a hook to retrieve food from a tube. She had never seen wire before. She wasn’t trained to bend it. She looked at the problem — food at the bottom of a vertical tube, a straight wire that couldn’t reach it — and manufactured a tool from a novel material, on the spot, to solve a problem she’d never encountered. That single observation kicked off two decades of research into corvid cognition that has systematically demolished the assumption that complex intelligence requires a primate brain, a mammalian cortex, or 300 million years of shared evolutionary history with humans.

    Corvids — the family that includes crows, ravens, jays, magpies, and jackdaws — have brains the size of a human thumb. They have no neocortex, the structure that in mammals is responsible for the cognitive functions we associate with intelligence: planning, reasoning, abstract thought, self-awareness. They produce comparable cognitive outputs using entirely different neural architecture, which means either intelligence is less dependent on specific brain structures than neuroscience assumed, or corvids evolved their way to the same destination through a route nobody predicted.

    https://open.spotify.com/show/7kYYn1bsA68hcfeT1YYehh?si=ae8656690bc643fb

    Tool use: not the party trick it looks like

    New Caledonian crows are the corvid species with the most sophisticated tool use, and the research on them has gone well beyond “crow uses stick to get food.” In wild populations on the Pacific island of New Caledonia, these crows manufacture tools from pandanus leaves by tearing them into specific shapes — stepped, tapered, or wide — to probe insect larvae from tree bark. The tool shapes are consistent within populations and vary between populations, which means the techniques are culturally transmitted rather than genetically encoded. A young crow learns to make tools by watching older crows. If the older crows die before transmitting the technique, the knowledge disappears. This is culture — the same mechanism that transmits human skills across generations — operating in a bird with a brain that weighs 14 grams.

    In laboratory settings, the cognitive demands of corvid tool use have been tested with increasing rigor. Gruber and colleagues, in experiments published in Current Biology, presented New Caledonian crows with metatool problems: multi-step tasks where one tool must be used to obtain another tool, which is then used to reach food, with each stage of the problem out of sight of the others. The crows had to mentally represent the location and identity of tools and apparatuses they couldn’t see while planning and executing a sequence of tool behaviors. They succeeded — maintaining working representations of objects across spatial separation and planning one to two steps ahead. The researchers concluded that New Caledonian crows can use mental representations to solve sequential problems, a capacity previously attributed only to humans and great apes.

    A 2020 study in Proceedings of the Royal Society B pushed further. Crows learned a temporal sequence: they were shown a baited apparatus, given a choice of five objects five minutes later, and given access to the apparatus ten minutes after that. At test, the crows selected the correct tool for the specific apparatus they’d been shown — choosing the right tool for the right future task while ignoring previously useful tools and a low-value food item. The study’s conclusion: New Caledonian crows plan for specific future tool use. This capacity — selecting a tool now for a task that will occur later, based on a mental representation of what that future task requires — was previously considered a defining feature of human intelligence. Corvids and humans shared a common ancestor over 300 million years ago. Whatever cognitive machinery the crows are using, they evolved it independently.

    Funerals: danger assessment, not grief

    When a crow dies, other crows gather. They emit alarm calls — loud, repetitive scolding vocalizations — that attract additional crows to the scene. Dozens of birds may congregate around the body, observing it from nearby perches, sometimes flying down to inspect it, sometimes sitting in silence. To a human observer, it looks like mourning. The scientific explanation is more interesting than mourning.

    Kaeli Swift, a behavioral ecologist at the University of Washington working under corvid cognition researcher John Marzluff, conducted a two-year experiment across over a hundred sites in Washington State. She established feeding stations to attract local crows, then introduced a dead crow (a taxidermied specimen) while a masked human volunteer stood nearby. The crows responded to the dead crow with alarm calls and gathering behavior. More importantly, they subsequently avoided feeding at that location — and they associated the masked person with danger, responding with alarm calls when that person appeared again, even without the dead crow present. The crows learned from the death scene. They identified a potential threat (the person near the dead crow), memorized the threat’s face, and modified their behavior to avoid the area and the individual. Weeks and months later, they still recognized and responded to the mask.

    Crows respond far more strongly to dead crows than to dead birds of other species. They largely ignore dead pigeons, robins, or other non-corvid birds placed in their territories but react intensely to dead members of their own species. Some studies suggest they respond more strongly to familiar individuals than to unfamiliar crows, indicating they may recognize specific community members even in death.

    The “funeral” is not a ceremony. It’s a threat assessment protocol. The crows are investigating the scene to determine what killed the dead crow, whether that threat persists, and how to avoid it. The alarm calls broadcast the danger to the wider community. The subsequent avoidance behavior encodes the lesson into the population’s behavioral repertoire. Marzluff’s research demonstrated that crows can remember human faces that posed a threat for years — and they transmit this knowledge to crows that weren’t present for the original event. A crow that never saw the masked person holding a dead crow will nonetheless scold that person if other crows in the community do, because the social alarm response propagates through the group.

    The behavioral function is pragmatic: collective intelligence applied to mortality data. The emotional dimension — whether crows experience something analogous to grief — remains scientifically unresolvable. Swift’s position is candid: she believes crows have emotional intelligence, but testing that scientifically is impossible because there’s no way to access what’s happening at an emotional level inside an animal’s brain. What’s measurable is the behavioral output: crows process death, learn from it, remember the context, and share the information. Whether they feel anything while doing it is a question the methodology can’t answer.

    What corvid brains do differently

    The corvid brain lacks a neocortex. In mammals, the neocortex is the seat of higher cognitive function — the structure that expanded dramatically in primates and reached its maximum density in humans. Corvids achieve comparable cognitive outputs using a structure called the pallium, which is organized differently from the mammalian cortex but performs analogous functions. The neuron density in the corvid pallium is remarkably high relative to brain volume — corvid brains pack more neurons per gram than most mammalian brains.

    A 2025 paper in Animal Cognition by Veit and colleagues explored the “dimensions of corvid consciousness” — a research framework asking not whether corvids are conscious but what aspects of consciousness their neural architecture could support. The paper argues that corvid brains process sensory information, maintain working memory, and generate flexible behavioral responses through neural pathways that are structurally distinct from but functionally analogous to mammalian circuits. A German neurobiologist trained two crows — Glenn and Ozzy — to peck at “yes” or “no” targets to indicate whether they had detected a faint light, demonstrating analytical introspection: the crows reported on their own perceptual states, a capacity associated with subjective experience.

    The convergent evolution angle is what makes corvids matter for neuroscience rather than just for animal behavior. If complex cognition can evolve independently in a brain that is structurally unrelated to the primate brain, then intelligence is not a property of a specific neural architecture. It’s a property of certain computational principles — neuron density, connectivity patterns, feedback loops — that can be instantiated in multiple biological substrates. The corvid brain is evidence that there is more than one way to build a mind, and that the way mammals did it is not the only way it can be done.

    Corvids sit alongside octopuses as the strongest natural evidence that intelligence is convergent rather than unique. We cover corvid cognition alongside cuttlefish camouflage, electroreception, and the full landscape of how animal brains solve problems humans assumed required human brains across our Neurozoology course — including why a 14-gram brain that last shared an ancestor with yours 300 million years ago can plan for the future, manufacture tools from materials it’s never seen, and hold a funeral that’s more operationally useful than most of ours.

  • Elephant Mourning Rituals: What We Know About Animal Grief

    In 2024, researchers from the Indian Institute of Science Education and Research documented five cases of Asian elephants burying their dead calves. The elephants positioned the calves into muddy trenches, covered them with earth—leaving only the legs protruding—and then stood over the burial sites for extended periods. Footprints around the carcasses confirmed that adult elephants had spent considerable time at the locations. In one case, the adults trumpeted for nearly 60 minutes—sustained, unbroken vocalizations of the kind elephants don’t produce during routine social interaction. The study, published in the Journal of Threatened Taxa, provided the first systematic documentation of intentional burial behavior in Asian elephants. The calves were not abandoned. They were interred.

    This is the kind of evidence that makes the question of animal grief impossible to dismiss and difficult to answer. The elephants’ behavior meets every behavioral criterion a scientist could reasonably apply: they altered their routine in response to death, they attended the body, they performed sustained and unusual vocalizations, and they engaged in deliberate physical manipulation of the remains that serves no obvious survival function. If a human community performed the same sequence—gathering around a body, vocalizing, burying the dead, standing vigil—no one would hesitate to call it mourning. The question is whether using that word for elephants is scientific description or anthropomorphic projection.

    What elephants do around their dead

    The behavioral record is extensive enough to establish patterns rather than anecdotes.

    Elephants investigate the bones of dead elephants—touching them with their trunks, lifting them, carrying them, sometimes moving them to new locations. They do this with the remains of relatives and non-relatives alike, and they do it with old bones as well as fresh carcasses. Critically, they don’t do it with the bones of other species. Whatever is happening when an elephant examines elephant remains, it’s species-specific. The trunks that can detect vibrations through the ground, identify individual elephants by scent at distances of miles, and manipulate objects with the dexterity of a human hand are deployed over bones in patterns that researchers consistently describe as careful, deliberate, and sustained.

    When an elephant dies within a social group, the surviving members frequently refuse to leave the body. They stand over it for hours or days. They touch the carcass with their trunks repeatedly—the face, the ears, the mouth. They sometimes attempt to lift the dead animal or push it to its feet. Some researchers have observed elephants placing grass, leaves, and branches over the bodies, partially covering them. Others have documented elephants guarding carcasses from predators.

    The behavioral changes extend beyond the immediate vicinity of the body. After a death, herd members have been observed eating less, moving more slowly, showing reduced social interaction, and producing vocalizations described by researchers as unusually quiet and subdued—grumbling, low-frequency sounds distinct from normal communication calls. Marc Bekoff, an animal behavior expert, described observing a herd whose matriarch had died: “Their heads were down, ears drooping, tails hanging listlessly, and they were just walking here and there, moping around, apparently brokenhearted.” The behavioral shift persisted for days.

    Elephants also produce temporal gland secretions during encounters with dead elephants—fluid that streams from glands on the sides of the head, associated with states of heightened emotional arousal including stress, excitement, and what researchers cautiously describe as distress. Some observers have reported what appears to be tear production, though whether this represents emotional crying or a stress-related physiological response is unresolved.

    The Lawrence Anthony episode

    When conservationist Lawrence Anthony—known as “The Elephant Whisperer”—died suddenly in March 2012, the two herds of once-aggressive rogue African elephants he had rehabilitated at his Thula Thula reserve in South Africa traveled roughly 12 hours through the Zululand bush to arrive at his home. They hadn’t visited in over a year. They appeared on the day of his death and remained for what observers described as a two-day vigil. The timing and distance traveled are difficult to reconcile with coincidence, though how the elephants could have known of Anthony’s death—he died indoors, miles from where the herds were ranging—remains unexplained.

    The Anthony story is frequently cited in popular accounts of elephant grief and is worth noting precisely because it illustrates both the power and the limit of the evidence. The elephants’ arrival was real and documented. The interpretation—that they somehow learned of his death and traveled to mourn him—requires a mechanism that no one has identified. Elephants have extraordinary sensory capabilities, including infrasound communication over distances of miles and the ability to detect seismic vibrations through their feet. Whether any of these could account for detecting a human death at the reported distance is unknown. The episode is compelling enough to report and uncertain enough to resist a clean conclusion, which is where most of the honest evidence for animal grief sits.

    The scientific problem with grief

    Anthropologist Barbara J. King proposed a definition that has become the field’s working standard: to qualify as grief, surviving individuals who knew the deceased must alter their behavioral routine—eating or sleeping less, acting listless or agitated, attending the body. By this behavioral definition, elephants grieve. So do chimpanzees (who become subdued and eat less after a death in the group), dolphins (who carry dead calves for days or weeks), orcas (who push dead newborns for hours, refusing to let them sink), gorillas (Koko the gorilla became “very somber” with “her lip quivering” when told of Robin Williams’s death), wolves (whose surviving pack members show measurable behavioral depression after losing a companion), and corvids (crows gather around their dead in what researchers have called “funerals,” though the function appears to be threat assessment rather than mourning).

    King’s definition is useful because it’s measurable. It’s also deliberately agnostic about subjective experience—it describes what the animal does, not what the animal feels. This distinction is the central methodological problem. Grief, in humans, is an internal experience—a subjective state of emotional pain, longing, and loss. We can’t access the subjective experience of another species. We can only observe behavior and infer. The inference is strong when the behavior is complex, sustained, species-specific, and functionally unnecessary—which is why elephant bone investigation, calf burial, and extended vigils are so compelling. There’s no obvious survival benefit to standing over a dead body for two days or carrying bones from one location to another. The behavior suggests something beyond curiosity or confusion, but “beyond curiosity” is not the same as “grief in the way humans experience it.”

    Elephants have von Economo neurons—specialized brain cells previously documented only in humans, great apes, and cetaceans, associated with empathy, social awareness, and self-recognition. Their brains are the largest of any land animal, roughly three times the mass of a human brain, with a highly developed hippocampus (the structure associated with memory and emotion). They recognize individual elephants after years of separation. They form lifelong social bonds. They have the neurological infrastructure that, in every other species where it appears, is associated with complex emotional processing.

    What we’re actually arguing about

    The debate over animal grief is not about whether the behaviors exist—they’re documented, filmed, published, and reproducible. The debate is about whether the word “grief” applies to what’s happening inside the animal’s mind, and that debate is ultimately about consciousness: whether elephants (and apes, and cetaceans, and corvids) have subjective emotional experiences that are analogous to ours, or whether they have sophisticated behavioral responses to social disruption that look like grief from the outside but feel like nothing from the inside.

    The emerging scientific consensus, as surveyed by Emory University, is moving toward the former. Most researchers who study animal cognition now accept that many species possess emotional experiences with subjective qualities. The question has shifted from “do animals have emotions?” to “how do animal emotions compare to human experiences?” The answer is probably: similar in kind, different in degree, and impossible to access directly because we can’t be an elephant any more than we can be a bat.

    What the evidence supports is this: elephants respond to death with behaviors that are sustained, deliberate, species-specific, neurologically supported, and functionally unnecessary for survival. They bury their calves. They stand vigil over bodies. They return to bones years later and touch them with the organ most sensitive to individual identity they possess. They alter their behavior for days or weeks after a loss. Whether this constitutes grief depends on whether you require the subjective experience to use the word, and that requirement is a philosophical choice, not a scientific one. The elephants’ behavior doesn’t change based on which choice you make.

    We cover elephant mourning alongside orangutan self-medication, baboon politics, and the full landscape of animal cognition across our Animal Culture & Knowledge course—including why the hardest question in the study of animal minds isn’t what they do. It’s what they feel.

  • Orangutan Self-Medication: How Great Apes Choose Plants to Treat Their Own Wounds and Infections

    On June 22, 2022, researchers at the Suaq Balimbing research station in Sumatra’s Gunung Leuser National Park heard a series of long calls from the canopy—the vocalizations male orangutans produce during dominance confrontations. The next day, they noticed that a flanged male orangutan named Rakus had a fresh wound on his right cheek, just below the eye, probably from a fight with a neighboring male. Three days later, they watched him do something no wild animal had ever been documented doing: he selected a specific plant—a climbing vine called Fibraurea tinctoria, known locally as Akar Kuning—ripped off its leaves, chewed them for 13 minutes, and then spent seven minutes applying the resulting juice directly to his wound with his fingers. He didn’t swallow the leaves during the application phase. When flies began landing on the wound, he covered it entirely with the chewed plant material, creating a poultice. The next day, he returned to the same plant and ate more leaves. Within five days, the wound closed. By July 19—roughly a month after the injury—only a faint scar remained. No infection developed.

    The paper, published in Scientific Reports in May 2024 by Isabelle Laumer and Caroline Schuppli of the Max Planck Institute of Animal Behavior, called it “the first known case of active wound treatment in a wild animal with a medical plant.” The emphasis on “active” is deliberate. Animals have been observed swallowing plants with medicinal properties before—chimpanzees chew bitter pith, gorillas and bonobos swallow rough leaves whole to mechanically dislodge intestinal parasites, Bornean orangutans rub chewed plants on their limbs. But those behaviors involve ingestion or generalized application. What Rakus did was topical, targeted, and sequential: he applied the plant’s juice specifically to the wound, on no other body part, repeated the application multiple times, and then covered the wound with plant material. He treated his own injury the way a human would treat a cut—clean it, apply medicine, bandage it.

    Why Akar Kuning matters

    Fibraurea tinctoria is not a random plant. It’s a climbing liana found across Southeast Asia—Indonesia, Malaysia, Thailand, Vietnam—and it’s used extensively in traditional medicine to treat dysentery, diabetes, malaria, and infections. Chemical analysis of the plant has identified furanoditerpenoids and protoberberine alkaloids with documented antibacterial, anti-inflammatory, antifungal, antioxidant, and analgesic properties. The plant also contains jatrorrhizine, which has antimicrobial and anticancer properties, and palmatine, which has anti-inflammatory and antiviral effects. This isn’t a plant that happens to have healing properties. It’s a plant whose healing properties are well-characterized enough that humans have been using it medicinally for centuries.

    Rakus’s population at Suaq Balimbing rarely eats it. In 21 years and roughly 390,000 feeding observations at the site, Fibraurea tinctoria appeared in only 0.3 percent of feeding scans. This wasn’t a plant the orangutan was already eating when he happened to touch his wound. He selected it specifically, used it in a way that doesn’t correspond to normal feeding behavior, and applied it exclusively to the injury. The researchers were careful to note that in 21 years and 28,000 observation hours, they had never previously seen an orangutan use leaves to treat a wound.

    How deliberate was it?

    This is the question the paper addresses directly and honestly. The behavior appeared intentional: Rakus selectively treated only his facial wound, not other body parts. He repeated the application multiple times. He used both the juice (liquid application) and the solid plant material (poultice). The entire process—feeding on the plant, applying the juice, covering the wound—took a considerable amount of time and was sustained across two consecutive days. The sequence is difficult to explain as accidental.

    But the researchers offer two possible origin stories, and they’re transparent about not being able to distinguish between them. The first is “accidental individual innovation”—Rakus may have been feeding on the plant, accidentally touched his wound while chewing, felt immediate pain relief from the plant’s analgesic effects, and then repeated the behavior because it worked. Under this model, the behavior was discovered by accident and reinforced by its consequences, which is how a lot of animal tool use and self-medication originates. The second possibility is social learning—Rakus wasn’t born at Suaq Balimbing. Male orangutans disperse from their natal area during or after puberty, sometimes traveling long distances. Rakus may have observed the behavior in his birth population, carried the knowledge across dispersal, and applied it when the situation required. If so, the behavior represents a cultural tradition transmitted between individuals, not an individual invention.

    The researchers can’t determine which explanation is correct because they don’t know where Rakus was born or what behaviors are practiced in that unknown population. This ambiguity is frustrating but honest—and it’s the central challenge of studying animal self-medication in the wild. You’re observing rare behaviors in long-lived animals across vast landscapes with limited coverage, and the most interesting questions (was it invented or learned?) require data from populations you may never have access to.

    The broader landscape of animal self-medication

    Rakus’s wound treatment is the most dramatic documented case, but self-medication in animals—zoopharmacognosy—is a recognized field with decades of evidence across multiple species and continents.

    Chimpanzees at multiple African field sites chew the bitter pith of Vernonia amygdalina, a plant with antiparasitic compounds, when they’re suffering from intestinal infections. The behavior is targeted: chimps eat it when sick and avoid it when healthy, suggesting they’re responding to internal cues rather than eating it as a regular food. Gorillas, chimpanzees, and bonobos swallow rough, hairy leaves from Aspilia and other plants whole and without chewing—the leaves pass through the digestive tract intact and physically dislodge intestinal parasites, which researchers have confirmed by examining fecal samples and finding parasites wrapped in leaf material. This is mechanical self-medication: the plant’s physical properties, not its chemistry, provide the therapeutic effect.

    Bornean orangutans have been observed rubbing chewed leaves of Dracaena cantleyi on their limbs, producing a lather that may have anti-inflammatory or antiparasitic properties. Capuchin monkeys rub citrus fruits and certain plants on their fur, potentially as insect repellent. Some moth species lay their eggs on alkaloid-rich plants when infected by parasitoid wasps, effectively medicating their offspring by ensuring the larvae consume antiparasitic compounds. Even fruit flies preferentially consume alcohol-containing food when infected by parasitoid wasps—the ethanol kills the wasp larvae developing inside them.

    The pattern across these examples is consistent: animals with no understanding of chemistry, pharmacology, or infection select specific substances with specific biological activity in response to specific health conditions. The behavior isn’t random foraging. It’s condition-dependent, substance-specific, and in many cases targeted to the affected body region. The question isn’t whether animals self-medicate. They do. The question is what cognitive mechanism enables it.

    What it means for the origins of medicine

    The earliest known human medical manuscript, from Mesopotamia around 2200 BCE, describes wound treatment with plant-based remedies. But if a Sumatran orangutan—separated from the human lineage by roughly 14 million years of evolution—independently applies a biologically active plant to a wound and covers it with a poultice, the implication is that the cognitive capacity for wound treatment predates the human lineage entirely. Laumer and Schuppli suggest that “medical wound treatment may have arisen in a common ancestor shared by humans and orangutans,” and that the behavior observed in Rakus may reflect deep evolutionary roots rather than a recent invention.

    The alternative—that Rakus and the Mesopotamian scribe independently arrived at the same solution—is possible but requires the same cognitive prerequisites: recognizing that a wound needs treatment, selecting a substance with appropriate properties, applying it specifically to the injury, and sustaining the behavior long enough for healing to occur. Whether the common ancestor had this capability or whether it evolved convergently in hominids and orangutans, the conclusion is the same: medicine didn’t start with humans. It started with primates who paid attention to what made them feel better and repeated it.

    Traditional healers in Indonesian Borneo have reportedly learned plant-based remedies by observing orangutan behavior—the knowledge transmission running from ape to human rather than the reverse. If Rakus learned his wound treatment from his natal population, and if human populations learned similar treatments from watching orangutans, then the same medicinal knowledge has been transmitted across species boundaries in both directions. The forest pharmacy has always been open. The question is who figured out the inventory first.

    We cover orangutan self-medication alongside baboon politics, ant collective intelligence, and the full landscape of animal cognition across our Animal Culture & Knowledge course—including why the first pharmacist may not have been a person. It may have been a primate with a cheek wound and the sense to reach for the right vine.

  • Mirror Neurons Across the Animal Kingdom: From Apes to Parrots to Dolphins

    In 1992, a neuroscientist at the University of Parma named Giacomo Rizzolatti was studying the premotor cortex of macaque monkeys—specifically, the neurons that fired when a monkey reached for a peanut. Standard motor mapping stuff. Electrode in the brain, monkey grabs food, neuron fires, graduate student logs it, everybody goes home. Except one afternoon, a researcher reached for his own lunch in front of the monkey, and the same neuron fired. The monkey wasn’t moving. It was watching someone else move. And the cell lit up like it couldn’t tell the difference.

    That’s the origin story of mirror neurons, and it’s one of those moments in neuroscience where a single observation cracks open a door that everyone then spends thirty years arguing about the size of. The finding was replicated, published in 1996, and promptly became one of the most overhyped discoveries in the history of brain science—V.S. Ramachandran called them “the driving force behind the great leap forward in human evolution,” which is the neuroscience equivalent of calling a rookie quarterback the next Tom Brady after one preseason game. The actual data, as usual, is more interesting than the hype, and considerably more complicated.

    So what do mirror neurons actually do? The basic mechanism is straightforward: these are neurons in the premotor and parietal cortex that fire both when an animal performs an action and when it observes another individual performing the same action. Grab a peanut, the cell fires. Watch someone else grab a peanut, the same cell fires. The neuron doesn’t distinguish between doing and seeing—or more precisely, it encodes both, which is a meaningfully different claim than the pop-science version where your brain “simulates” everything it sees like some kind of empathy PlayStation.

    The pop-science version went roughly like this: mirror neurons are the biological basis of empathy, imitation, language, theory of mind, and possibly the entire foundation of human civilization. You can still find TED talks making this argument. The actual neuroscience community has, over the past two decades, walked most of that back—not because mirror neurons aren’t real or important, but because the leap from “this neuron fires during observation and execution” to “this neuron explains human culture” requires about fourteen intermediate steps that nobody has convincingly demonstrated.

    Here’s what we actually know, species by species.

    Macaques remain the best-studied case because you can do single-neuron recordings in them, which you generally cannot do in humans for obvious ethical reasons involving the part where you stick an electrode into someone’s brain. Rizzolatti’s lab and subsequent groups have mapped mirror neurons primarily in area F5 of the ventral premotor cortex and in the inferior parietal lobule. These neurons are action-specific—they respond to hand grasping, mouth actions, tool use—and they’re modulated by context. A macaque mirror neuron that fires when it watches another monkey grasp a peanut to eat it may not fire when the same monkey grasps the same peanut to place it in a container. The neuron isn’t just mirroring movement. It’s encoding the goal of the action, which is a much more interesting finding than the simple mirror story.

    The caveat—and this matters—is that macaques are actually terrible imitators. They don’t readily copy novel behaviors from observation. So if mirror neurons are supposedly the neural substrate of imitation, we have a problem, because the species in which they were discovered doesn’t really imitate. This is the kind of inconvenient fact that tends to get footnoted rather than headlined.

    Great apes are a different story. Chimpanzees, bonobos, gorillas, and orangutans all demonstrate genuine imitation—learning novel motor sequences by watching others perform them. The problem is that single-neuron recordings in great apes are extremely rare for ethical and practical reasons, so the direct electrophysiological evidence for mirror neurons in apes is thin. What we have instead is a lot of fMRI and behavioral data suggesting that homologous brain regions (the ape equivalents of F5 and the inferior parietal cortex) are active during action observation. The inference is reasonable—these are our closest relatives, the anatomy is conserved, the behavior is consistent—but it’s still an inference, not a measurement. We’re reading the box score, not watching the game.

    Humans are where the story gets both more exciting and more contentious. You can’t ethically do single-neuron recordings in healthy humans, but a handful of studies in epilepsy patients with implanted electrodes (who were being monitored for seizure localization, not mirror neuron research) have found neurons in the supplementary motor area and medial temporal lobe that respond to both observed and executed actions. Iacoboni’s UCLA group published some of this work in the 2010s. The broader human evidence comes from fMRI, EEG mu-suppression studies, and transcranial magnetic stimulation—all of which point to a “mirror neuron system” distributed across premotor cortex, inferior parietal lobule, and the superior temporal sulcus. The system is real. The question is what it actually does versus what we’d like it to do.

    The honest answer, as of 2026: mirror neurons in humans are probably involved in action understanding—recognizing what someone is doing and predicting what they’ll do next. There’s decent evidence they contribute to motor learning through observation. The link to empathy is much weaker than the popular narrative suggests, and the link to language is speculative at best. Gregory Hickok’s 2014 book The Myth of Mirror Neurons did a pretty thorough job of separating the signal from the noise here, and the field has been more careful since.

    Now, here’s where it gets genuinely weird. Because mirror neurons—or at least mirror-like neural systems—aren’t limited to primates.

    Songbirds have what might be the most compelling mirror system outside of mammals. In zebra finches and other oscine songbirds, neurons in a region called the HVC (used to stand for “High Vocal Center” but now it’s just HVC because the original name was anatomically inaccurate, which is the neuroscience version of a company rebranding after a scandal) fire both when the bird sings a specific note sequence and when it hears the same sequence sung by another bird. These aren’t just auditory neurons responding to sound—they’re sensorimotor neurons that link production and perception of the same vocalization. The parallel to primate mirror neurons is striking, and it evolved completely independently, which tells you something about how useful this computational architecture must be.

    The songbird mirror system is deeply involved in vocal learning—young birds learn their species’ song by listening to a tutor and gradually matching their own output to the template, and the mirror-like neurons in HVC are a critical part of that error-correction loop. This is arguably a cleaner example of mirror neurons supporting imitation than anything in the primate literature, which is both fascinating and slightly embarrassing for the people who spent two decades claiming mirror neurons were a uniquely primate innovation.

    Parrots are the other avian case worth knowing. Alex the African Grey—Irene Pepperberg’s famous research subject—could label objects, understand concepts like “same” and “different,” and produce novel combinations of learned words. Parrots are vocal learners like songbirds, but they’re not closely related to them—vocal learning evolved independently in parrots, songbirds, and hummingbirds, which means the mirror-like neural circuitry that supports it likely evolved independently too. Parrot neuroscience is less developed than songbird work (partly because parrots are harder to work with and live approximately forever), but the behavioral evidence for action-perception coupling is strong. A parrot that watches you wave and then waves back is doing something that macaques—the species where we actually found mirror neurons—basically can’t do.

    Dolphins present maybe the most interesting case because they combine vocal learning, complex social cognition, and a brain that is anatomically very different from a primate brain. Dolphins can imitate novel motor behaviors on command (the “do this” paradigm developed by Louis Herman’s lab at the University of Hawaii in the 1990s), and they engage in vocal mimicry—copying signature whistles of other dolphins, which functions as something like calling someone by name. The neural basis is largely unknown because, to state the obvious, you cannot put a dolphin in an fMRI scanner with any meaningful cooperation, and single-neuron recordings in cetaceans are essentially nonexistent. What we have is behavioral evidence that strongly implies a mirror-like system, layered on top of a brain with a completely different cortical organization—dolphins have an insular cortex that may serve some of the functions that premotor cortex serves in primates, but honestly, cetacean neuroanatomy is still more question marks than answers.

    The pattern that emerges across all these species is that mirror-like neural mechanisms seem to pop up wherever you find sophisticated social learning—whether that’s vocal imitation in songbirds, motor imitation in apes, or behavioral mimicry in dolphins. And these systems evolved independently in lineages that diverged hundreds of millions of years ago, which suggests that coupling action perception to action production is such a useful computational trick that evolution keeps reinventing it. It’s convergent evolution at the neural architecture level, which is roughly as cool as neuroscience gets.

    What the pop-science narrative got wrong was the specificity of the claim. Mirror neurons aren’t the secret to human empathy or the origin of language or the biological basis of civilization. They’re a neural mechanism for linking what you see to what you do—one piece of a much larger puzzle that includes prefrontal cortex, temporal lobe social cognition networks, and a dozen other systems that we’re still mapping. But what the pop-science narrative got right, even if accidentally, was the intuition that something deep is happening when one brain watches another brain act and encodes that observation in the language of its own motor system. That’s not empathy, exactly. But it’s the scaffolding that makes empathy—and imitation, and social learning, and maybe culture—mechanistically possible.

    The fact that an octopus, which diverged from our lineage over 500 million years ago, can watch another octopus open a jar and then do it themselves raises the question of whether mirror-like computation might be even more widespread than we currently think. We genuinely don’t know. The electrophysiology hasn’t been done. But the behavioral signatures keep showing up in species we didn’t expect, and every time they do, the story gets bigger.

    We cover mirror neurons—and the broader neuroscience of social cognition across the animal kingdom—in depth across several lectures in our Neurozoology course, which traces the evolution of cognition from mycelial networks to primate brains across 48 lectures and 69 hours of audio. If the octopus jar thing made you want to know more, that’s a good sign.