Show Me the Honey! Honeyguides and Humans Team Up at Dinnertime

Humans have a long history of spotting superior abilities in other animals, and then training those animals to use those abilities to advance our own interests. Everyone’s familiar with how we’ve trained pigs to sniff out truffles for us with their sensitive snouts and how we’ve domesticated dogs to herd our livestock, alert us to intruders, guide us when our vision fails, and perform other services. Similar but less well-known examples include our training bees to detect the odor of explosives, cormorants to catch fish for us, and llamas to guard our sheep from coyotes and other predators.

It’s far less common, though, to find relationships where non-humans participate on a more equal footing, where they appear to train us at least as much as we train them. (People who are owned by cats should feel free to rebut this statement in the comment section below.)

Today’s post features one such relationship, the partnership between humans and the greater honeyguide (Indicator indicator), a bird that lives in the trees of sub-Saharan Africa.

Honeyguides and humans have very complementary appetites. Honeyguides get most of their food from beehives, feasting on larvae and wax that they extract from honeycombs (yes, they actually eat and can digest the wax!). Of course humans, too, seek out beehives, although our interest lies more in the bees’ sweet honey, and we’re generally more than happy to leave the wax and grubs for others to enjoy.

The bee-related skills of humans and honeyguides are relatively complementary as well. Honeyguides can fly swiftly across large areas and are expert at locating bee colonies, but have difficulty in extricating the combs on their own. Humans move more slowly along the ground and aren’t so adept at finding colonies, but once we have one in our sights, we’re able to overcome bee defenses and dig the combs out, even when the bees have nested deep within rock crevices and other hard-to-reach locations.

Out of this opportunity for mutualistic benefit, honeyguides and humans have worked out an elaborate interspecies communication system that allows them to work in tandem with certain signals understood by both parties.  This partnership has been formally documented in a three year field study conducted in the dry bush country of northern Kenya, focusing on the interactions between honeyguides and the nomadic Boran people who populate the area.

I’m the *real* Greater Honeyguide – don’t let imposters lead you astray! Visit me at http://safari-ecology.blogspot.com/ and see first comment below.

Each partner knows how to get the other’s attention. To attract the birds, the Borans call them with a penetrating whistle (known in the Boran language as Fuulido) that can be heard over a distances of greater than a kilometer and that is made by blowing air into clasped fists, modified snail shells, or hollowed-out palm nuts. Comparably, hungry honeyguides flag down humans by flying up close, moving restlessly from perch to perch, and emitting a double-noted, persistent “tirr-tirr-tirr-tirr” call. (Side note: I’ve been practicing this at home, and it doesn’t seem to attract much other than odd stares and raised eyebrows.)

The joint food expedition commences when the honeyguide flies briefly out of sight and then returns to a nearby, conspicuously visible perch. When the human companion approaches this perch, the honeyguide takes off, displaying its white outer tail feathers, and flies to a new resting place a short distance away, calling loudly when it lands. The Boran partner then approaches the new perch and the bird flies off again, repeating the pattern. As the Borans work with the bird, they whistle and shout to keep the bird interested in guiding. (Again, this doesn’t seem to work too well at home.)

The researchers found that the honeyguides signal the path and distance to the bee colony in a variety of ways. First, they indicate the correct direction through their flight paths, traveling consistently in the direction of the nest and increasing their precision as they near the target. It appears that the know in advance where the nests are located, as the researchers observed the honeyguides briefly visiting nests before dawn, peering into the entrances while it was still dark and the bees were docile.

Also, the honeyguides vary their behavior depending on distance to the hive. For example, when the hive is relatively distant, the birds begin the process with a relatively long disappearance during their first flight; conversely, their first disappearance is briefer when the hive is relatively nearby. Further, the honeyguides stop more frequently and the legs between perches become shorter as they and their human followers approach the nest, especially during the last 200 meters. Finally, the honeyguides select increasingly lower perches as they close in on the colony.

Upon arrival at the destination, the honeyguides perch close to the nest and emits an “indication call.” The researchers describe the scene as follows:

This call differs from the previous guiding call in that it has a softer tone, with longer intervals between successive notes. There is also a diminished response, if any at all, to whistling and shouting by humans. After a few indication calls, the bird remains silent. When approached by the searching gatherer, it flies to another perch close by, sometimes after circling around the nest. The resulting flight path finally reveals the location of the colony to the gatherer. If the honey collector does not (or pretends not to) detect the nest, the bird gives up after a while. It may then leave the area either silently or start a guiding session to another colony. In the latter case, it switches from the indication call to the guiding call and resumes a fairly direct flight pattern. Once the human team members find the nest, it becomes their turn to go to work and hold up their part of the bargain. After using smoky fires to reduce the bees’ aggression, the Boran honey gatherers use tools or their hands to remove the honey comb, and then break off pieces to be shared with their honeyguide partners.

To sum things up, here’s a great BBC video (featuring David Attenborough!) that describes the bird-human partnership and shows the honeyguides in action:

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ResearchBlogging.org
Isack, H., & Reyer, H. (1989). Honeyguides and Honey Gatherers: Interspecific Communication in a Symbiotic Relationship Science, 243 (4896), 1343-1346 DOI: 10.1126/science.243.4896.1343.

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Contagious Yawning Spreads to Birds

Apparently, parrots aren’t just smart, they’re competitive too. A couple of months ago, we covered recent research findings on contagious yawning in animals, reporting on the rarity of the phenomenon and its potential role as a form of social mimicry or even an indication of empathy. While certain primates clearly do yawn contagiously and dogs may yawn contagiously, the behavior hadn’t been reported in other animals and had been expressly ruled out in red-footed tortoises (although the tortoises may have had the last laugh, as they won the celebrated Ig Nobel Prize for their non-yawns).

Word of our mammal-centric coverage seems to have reached the small, oval ears of the always-influential parrot lobby, though, as just last week the journal Behavioural Processes published a study describing social yawning in budgerigars (Melopsittacus undulatus), the small Australian parrot often referred to as the parakeet. This study provides the first support for contagious yawning in a non-mammal, and even ups the ante by documenting what may be the first instance of contagious stretching, another stereotyped behavior that may play a social role for certain animals. Some may say that the paper’s timing is an utter coincidence and that only someone with delusions of grandeur would believe that it was even remotely linked to the AnimalWise post. We, speaking in our usual royal manner, prefer to think otherwise.

Fascinating, simply fascinating...

Michael Miller, Andrew Gallup and other researchers from the University of the Binghamton conducted an observational study of yawning and stretching in a group of approximately 20 adult male and female budgerigars living together in an aviary as an established flock. Over a period of about a year and a half, the research team video recorded the flock on 23 separate occasions. The recording sessions, each of which lasted 90 minutes, were conducted at varying times of the day, and the researchers took a number of precautions (such as ignoring the first 15 minutes of each tape) to ensure that the flock’s behavior was as natural and undisturbed as possible. Trained reviewers then systematically reviewed all of the tapes, recording the time and occurrence of each yawn and stretch, and categorizing each stretch by whether the bird extended one or both legs.

The researchers’ hypothesis was that, if yawning and stretching were spreading contagiously among the birds, the behaviors would occur in nonrandom “clumps” – that is, rather than being evenly dispersed throughout the recording sessions, multiple yawns (or multiple stretches of the same type) would take place in closely-spaced bouts and then be followed by a long interval until a new priming behavior triggered another bout. Further, they predicted that, although there might be might be overall tendencies tied to particular times of the day (for example, the budgerigars might, on average, yawn more frequently during evening sessions), if the yawning and stretching really were being triggered contagiously, then specific clumping patterns would not repeat themselves when multiple same-time-of-day sessions were compared.

To test their hypotheses, the researchers performed detailed, session-by-session analyses of each type of behavior. For example, they tallied how frequently each behavior occurred, measured the time between adjacent stretches and yawns, and sorted the adjacent pairs into different “bins” depending on the length of the interval. They also analyzed each session for clumping by breaking it down into a large number of short (20 to 30 second) intervals, which allowed them to identify “runs” of consecutive intervals that either did, or did not, contain the behavior in question. Finally, they statistically analyzed their data in a variety of ways to identify patterns and associations.

And the results?

Both yawning and stretching behaviors were indeed clustered within trials, and the period between adjacent yawns and stretches was “strongly biased toward very short (< 20 sec) and very long (> 300 sec) intervals,” especially for the yawns. Also, as hypothesized, despite the clustering for both behaviors, “neither behavior routinely occurred at specific times from the start of a session across multiple recordings at the same time of day. This suggests that the clumping of these behaviors was due to social influences, and not to underlying physiological effects as a result of similar circadian patterns.”

The research team summarized its findings and suggested directions for future investigation as follows:

The observational results presented here suggest that yawning and stretching are at least mildly contagious in budgerigars under semi-natural flock-living conditions. In line with each behavior’s presumed physiological function, contagious yawning and stretching may ultimately coordinate mental state and a group’s collective movements, but future research needs to test these predictions.

So, kudos to the budgerigars! Parrots everywhere can take pride in these findings, which point to previously-unknown areas of avian social signaling and coordination, and which may open up new avenues for studying collective behavior.

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ResearchBlogging.orgMiller, M., Gallup, A., Vogel, A., Vicario, S., & Clark, A. (2011). Evidence for contagious behaviors in budgerigars (Melopsittacus undulatus): An observational study of yawning and stretching Behavioural Processes DOI: 10.1016/j.beproc.2011.12.012.

Zeroing In On Parrot Math Abilities

It may seem surprising, but the concept of “zero” is actually a relatively recent mathematical innovation. Indeed, the first rudimentary use of a zero-like notation didn’t appear until around 300 BC, when the Babylonians began using a special placeholder symbol to designate the absence of another value in their base-sixty number system. While revolutionary in its own right, the Babylonian null placeholder was still rather limited (for example, it couldn’t be used alone and never appeared at the end of a number), and another millennium passed before gifted Indian mathematicians and astronomers introduced a fully functional “true zero” as part of a formalized system of arithmetic operations. Some 1,500 years later, with this important mathematical foundation finally in place, Apple launched the iPhone on the AT&T wireless network.

Are there any parallels in the animal world, any similarly gifted nonhuman mathematicians that have innovated with the concept of zero?

The answer seems to be yes: Alex, the male African Grey Parrot of book and movie fame (Alex & Me), may go down in history as the parrot equivalent of Albert Einstein, revolutionizing parrot mathematics with his insight into concepts of nothingness.

How many crackers do I see? None! (image: The Alex Foundation)

It was in late 2003, early 2004 that Alex appears to have had his great breakthrough regarding the mathematical usefulness of zero-like concepts. At that time, Irene Pepperberg and Jesse Gordon of Brandeis University, who had been working with Alex over an extended period on a variety of cognitive and communicative studies, decided to conduct some experiments to explore the extent of his numerical competence.

Alex already was adept at tests requiring him to identify numbers of objects – he knew the English words for one through six, and could provide accurate verbal responses to questions about, for instance, how many green blocks were included in a mixed array of blue, red and green blocks and balls. Pepperberg and Gordon now wanted to see whether Alex really understood the numbers he was providing and could grasp the interchangeability of numerical questions.

To do so, they flipped things around: rather than asking Alex to provide the number of objects in particular groupings as he had in prior experiments, they went in the other direction by asking him to indicate which object groups were associated with a particular number. That is, they presented Alex with a tray of objects of various materials, colors and shapes (for example, six green plastic spoons, four yellow tops and three orange wooden sticks), and asked him questions such as “What color six?” and “What toy four?” Alex’s task was to look at the objects on the tray and then respond correctly (in this case, with “green” based on the six green spoons and “top” based on the four yellow tops).

(I know, this all sounds a bit like Jeopardy: “Please be sure to phrase your answer in the form of a question…”)

Perhaps not surprisingly, Alex aced the test, responding correctly to this new battery of questions over 80% of the time. More significant, though, is how Alex – apparently bored with the questioning – spontaneously extended the scope of the experiment:

On the 10th trial within the first dozen, Alex was asked “What color 3?” to a set of two, three, and six objects. He replied “five”; the questioner asked him twice more and each time he replied “five.” The questioner, not attending to the tray, finally said “OK, Alex, tell me, what color 5?” Alex immediately responded “none.”

Now, Alex had previously been trained to use the word “none” in a different context – comparing objects for similarity or difference (for example, to respond to a question about which of two identically-sized objects was bigger) – but he had never been taught to use “none” to describe a quantity that was not present. Fascinated, Pepperberg and Gordon randomly interspersed six more “none trials” into the ongoing experiment. It turned out that Alex’s response was no fluke – he gave the correct “none” response in five out of six of these trials, an accuracy rate of 83.3%.

Here’s a brief video in which Pepperberg describes the experiment and Alex’s unexpected use of the “none” concept:

Thus, it appears that Alex spontaneously used “none” in a zero-like manner to label a null set and designate an absence of objects. As the researchers summarized it, “the notion of none, even if already associated with absence of similarity and difference (and lack of size difference), is abstract and relies on violation of an expectation of presence; that Alex transferred the notion from other domains to quantity, without training or prompting by humans, was unexpected.”

While Alex’s use of “none” may not be as full and robust as the true zero concept that we use today, it nonetheless (no pun intended) is quite impressive. Moreover, Alex’s insight may prove to be quite practical, with the parrot concept of “none” providing helpful guidance as we attempt to answer some of the more pressing questions of our time, including:

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ResearchBlogging.orgPepperberg, I., & Gordon, J. (2005). Number Comprehension by a Grey Parrot (Psittacus erithacus), Including a Zero-Like Concept. Journal of Comparative Psychology, 119 (2), 197-209 DOI: 10.1037/0735-7036.119.2.197.

Back to the Future – Mental Time Travel in Tropical Birds?

I may not have a nuclear-powered DeLorean parked in my driveway, but I can travel in my own personal time machine anytime I want, and so can you.

Through what’s known as mental time travel, or MTT, you can move backwards and forwards through time – visiting the past when you remember a specific event you’ve already experienced, and then zipping forward to the future as you use this memory to predict, plan for and shape events that are yet to come.

Mental time travel is no mean feat: it implicates sophisticated cognitive processes and is thought to form the foundation for advanced forms of consciousness such as self-awareness and the ability to attribute independent thought, desires and intentions to others (an ability sometimes referred to as “theory of mind”).

So, can other animals engage in mental time travel? Perhaps not surprisingly, this has been a controversial topic, and many have argued that we humans are the only ones able to mentally flit about the fourth dimension, leaving all other animals stuck in the here and now. Although this may be partly attributable to our anthropocentric world view1, the language barrier between humans and other animals also poses a real problem, as it’s difficult to design MTT experiments that don’t involve interviews, since the best time travel evidence may consist of the voyager’s personal and subjective reports of the experience. Accordingly, solid evidence for MTT in other animals has been limited, and much of the evidence that does exist consists of anecdotal accounts and a small number of experiments involving great apes and western scrub-jays.

In a paper2 published in the October 14, 2011, issue of Behavioral Ecology, though, a research team led by Corina Logan of the Department of Experimental Psychology at the University of Cambridge proposed an intriguing new avenue for further research, one that might significantly expand the number of species that may be tested for MTT abilities. More specifically, Logan and her colleagues identified a specialized strategy among birds – army ant bivouac checking – that may provide conditions in the wild that could favor the development of mental time travel in a variety of species.

If I go back in time and shoot my grandmother, does that mean I’ll never be born?? (White Whiskered Puffbird, credit Glenn M. Duggan FZS)

While many tropical rain forest birds earn an opportunistic living by gobbling up insects and other small invertebrates flushed out of hiding by army ants on food raids, a subset go a step further – after raids by a specific army ant species (Eciton burchellii), these birds follow the ants back to their temporary nests (known as a bivouacs) in the evening, and then return to check on the bivouacs the next morning before the ants raid again. To date, twenty one different “bivouac-checking” bird species have been identified.

Tracking army ant bivouacs is more complicated than one might think. As the ants raise their young, they cycle through two distinct phases: one (approximately 20-day) phase during which they remain in a set location, conducting most of their raids at the beginning and end of the phase and relatively few during the middle two weeks; and a second (approximately 14-day) nomadic phase, during which they move their bivouac on a daily basis and conduct raids almost every day.

Aren’t you hungry? We haven’t eaten since tomorrow. (Ocellated Antbirds, credit PhilSlosberg)

From the perspective of a bivouac-checking bird seeking a reliable food source, these varying phases are significant. When the ant colony is stationary, it may be relatively easy to find, but its raids will be sporadic; when the colony is nomadic, it may be more difficult to find, but its raids will be quite regular. Clearly, a bird will do better if it can keep track of multiple stationary colonies that conduct raids only sporadically, if it can quickly find nomadic colonies based on their prior locations and previous movements, and if it can remember whether particular colonies are in phases in which they’re likely to conduct raids.

The researchers identified these conditions as providing a potential testing ground for MTT. While the birds obviously can’t be interviewed, their environment may elicit behavior that shows that they have an “episodic-like memory” (that is, they can recall the what-where-when aspects of past events) and that they can take action in anticipation of future motivational states independent of their current needs (that is, they can plan flexibly for the future).

After noting that the birds appear to form specific memories about locations (since they return in the morning after evening bivouac checks), the researchers hypothesize that the birds may remember which colonies are in which locations and what phase the colony is in, “and that they may be using episodic-like memory if they prefer to check those bivouacs from army ant colonies in the nomadic phase.” Moreover, they continue:

Time travel jokes never get old! (Spotted Antbird, credit Mike Danzenbaker)

We suspect that future planning could be involved in bivouac-checking bird behavior because birds check bivouacs when sated (conferring no immediate benefit), a behavior that does not make sense until the next morning on return to the bivouac when the bird finds the ants raiding again and encounters its next meal (a delayed benefit). Because bivouac checking occurs after foraging at a raid, there is no immediate benefit to conducting this behavior in terms of acquiring a meal in the next few minutes. Instead, the benefit occurs the next morning when the ants begin raiding again; bivouac-checking birds return and are the first to begin foraging at the raid. This could indicate a dissociation between their current state (sated) and a future need (will need to eat tomorrow), which suggests anticipation of future events. (Citations omitted.)

Logan and her colleagues call for additional field research and, if their hypotheses are supported, laboratory experiments that will enable experimenters to vary bivouac locations and colony phases under controlled circumstances, and to determine whether the birds use specific memories and flexible future planning or whether they engage in automatic behavior using vision, smell, circadian rhythm or other cues in checking on bivouacs.

At this point, the researchers’ hypotheses need additional experimental support, but it’s already clear that they’ve made some keen observations about specialized behavior in the wild and have opened the door to substantially expanded testing for mental time travel in animals. As more researchers come up with similarly elegant ways of investigating abilities previously thought to be unique to humans, I think we will see additional barriers fall.

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1As regular readers know, this is a recurrent AnimalWise theme. Time and again we humans have claimed that we are the only ones to have a particular skill, only to discover later on that, in fact, many animals may share the same or comparable abilities with us. Prior posts have discussed this phenomenon in areas including tool use (many different fish, crested rats, ants, dolphins, chimpanzees, crows), language (bees and prairie dogs), analogic reasoning (capuchins and baboons), grief and mourning in various animals, and self-recognition (dogs, magpies).

ResearchBlogging.org2Logan, C., O’Donnell, S., & Clayton, N. (2011). A case of mental time travel in ant-following birds? Behavioral Ecology DOI: 10.1093/beheco/arr104.

Analogical Reasoning in Animals

In today’s post, I’d like to explore some surprising recent findings about the abilities of animals in the area of analogical reasoning.

Reasoning by analogy is central to the way we think, enabling us to use familiar concepts to solve new problems. When a catastrophic event strikes Wall Street, economists inevitably point to analogous historical disruptions in their attempts to predict whether we’re facing long-term troubles or a quick recovery. When lawyers advocate on behalf of clients in new realms such as digital media, they often ground their arguments in principles that evolved centuries ago to protect real property interests. When scientists explain the motion of molecules and other phenomena that we cannot directly perceive, they frequently turn to concrete examples such as colliding billiard balls or streams of water.

On a more mundane level, analogical thinking underlies many of our idioms and permeates our everyday language. Think how lost you’d be if you were suddenly unable to understand phrases that explicitly or implicitly apply concepts from one context to events or actions in another. Conversations at work would confuse you (more than usual). Your boss’ suggestion that you take some time off to recharge your batteries would leave you scratching your head rather than looking for deals on tropical island vacations. You wouldn’t be able to follow political discussions (oh no!). You’d be the only one asking “oh my god, was it with guns or knives?” after hearing that one candidate outdueled another in a debate. You’d be the only one worrying about cannibalism after learning that the people were hungry for new leadership. You’d find sports to be newly upsetting, as you’d literally go into mourning after learning of your favorite team’s fatal missteps. (Ok, I take that back – nothing has changed here, especially for Boston Red Sox fans.)

Relational Matching Tests

One of the most common tests used to assess an individual’s ability to solve analogy problems is known as relational matching-to-sample or RMTS. In its classic form, RMTS involves first showing the subject a sample set consisting of two or more objects that are either identical (for example, two circles) or nonidentical (for example, a square and a circle). Sets containing identical objects are sometimes referred to as reflecting the “identity relation” and those containing nonidentical objects are said to reflect the “nonidentity relation.” Next the subject is shown two comparison sets containing novel objects, one embodying the identity relation (e.g., two triangles) and the other the nonidentity relation (e.g., a rectangle and a triangle). To succeed, the subject must choose the comparison set that matches the relationship demonstrated by sample set. For instance, the correct choice for a subject shown two circles in the sample would be the comparison set containing the two triangles, whereas the correct choice for the subject initially shown the square and the circle would be the comparison set containing the rectangle and the triangle.

RMTS is particularly well suited for testing the abilities of non-human animals, as it poses an analogy problem in a strictly visual manner, not relying in any way on linguistic skills. In essence, success requires the subject to not only make a “first order comparison” between same and different, but also to make a “second order comparison” by applying this underlying distinction to a novel environment. Many researchers consider this ability to lie at heart of analogical reasoning.

A “Profound Disparity”?

Until recently, studies have suggested that humans and a select few great apes stand far apart from all other animals in terms of analogical reasoning abilities. While many animals can successfully distinguish between same and different shapes or colors, they tend to struggle when it comes to making second order comparisons of the sort required by RMTS tasks. Since only humans and some chimpanzees, gorillas and orangutans have performed well at RMTS testing, researchers have proposed that a “profound disparity” exists between the analogical reasoning capacity of hominids and other animals.

For example, several studies have shown that some baboons and pigeons can learn to pass RMTS tests if they involve large-sample and comparison sets (e.g., comparisons involving 4 x 4 grids of 16 all identical and 16 all different objects), but that their performance rapidly deteriorates as the size of the grid decreases as well as when the distance between the objects in the grid increases. According to researchers, one reason why animals do better with larger sample sets may be that there’s a greater amount of variation or “entropy” between non-analogous grids in larger sample and comparison sets, which makes the task of distinguishing between potential answers easier.

Notwithstanding these prior findings, however, two studies published in the last few months now pose a challenge to the “profound disparity” concept, suggesting that a suitable testing environment can showcase robust analogical reasoning skills in non-apes.

Clever Capuchins

In the first study, which was published in PLoS ONE in August 2011, researchers led by Valentina Truppa and Elisabetta Visalberghi of the National Research Council in Rome, Italy, found that New World tufted capuchin monkeys (Cebus apella) were capable of solving RMTS tasks involving sample and comparison sets involving sets of as few as two objects.

What are all of those freaking squiggles in that diagram above my head? (image credit: Charlesjsharp)

The research team studied five capuchin monkeys, testing them over and over again on RMTS tasks involving varying numbers of icons. While the specific tests varied, the general approach was to start by giving the monkeys trials involving a relatively small pool of different icons and, only if and when a monkey achieved proficiency (as measured by percentages of correct answers) over the course of thousands of trials, to introduce novel icons for comparison. Also, in one of the experiments, if the monkey did not ultimately reach the proficiency threshold on a two-icon comparison test, “entropy” was increased and the monkey was given an easier four-icon test.

Ultimately, after a total of 21,888 trials (yes, that’s correct!) one of the five capuchins, Roberta, proved to be a real overachiever. As the researchers put it:

The current study demonstrates the acquisition of abstract concepts based on second-order relations by one capuchin monkey, Roberta. She was first successful with four-item stimuli and then with two-item stimuli, the latter being the most difficult condition previously thought to be mastered only by apes. Since her performance was robust across different types of stimuli and well above that of the other subjects, we can argue that relational analogies are very difficult for capuchins, but under specific circumstances not impossible.

Way to go, Roberta!

Bright Baboons

In a second study, published on September 20, 2011, in Psychological Science, a research team headed by Joël Fagot of the Centre National de la Recherche Scientifique at the Université de Provence reported that guinea baboons (Papio papio) can learn to perform surprisingly well at RMTS tasks … and then retain this ability over a 12-month period. In this study, 29 baboons with no language training and little or no experience with relational matching tests participated in various RMTS experiments involving geometric shape comparisons.

All this RMTS stuff gives me a headache (image: Animal Globe)

The first experiment consisted of classic RMTS trials, each involving a sample set made up of pairs of identical or nonidentical geometric shapes, and two comparison sets with new geometric shapes, with only one of the comparison sets matching the relationship demonstrated by the sample set. At first, the testing pairs were selected randomly from among 10 geometric shapes, but once a baboon had achieved an accuracy level of 80% or better in three consecutive sessions of 100 trials, new geometric shapes were introduced up to a maximum of 90 shapes by the end of the experiment. Six of the 29 baboons were able to make it to the 80% threshold level, and five were ultimately able to proceed through testing until they reached all 90 shapes.

The second experiment included changes designed to make the challenge more difficult: the geometric shapes were moved further apart and, perhaps more significantly, in half of the tests the “incorrect” comparison pair, rather than containing all new geometric shapes, actually contained one of the shapes from the sample pair. In other words, even though this comparison pair was incorrect from the standpoint of analogy testing, it contained a shape that was directly linked to the sample set, potentially confusing the baboon if it was focused on the similarity of the shapes rather than the conceptual relationship between the shapes.

In spite of the enhanced degree of difficulty, all five of the baboons who participated – the same baboons who had been successful in the first experiment – performed at above chance levels throughout the second experiment (although, not surprisingly, their performance tailed off somewhat in the trials where the incorrect response shared a shape with the sample set).

Finally, the research team retested the five successful baboons in accordance with the first experiment methodology after a one-year lapse during which the baboons had no practice at RMTS tasks. All five baboons reached the 80% success level far more quickly than they had the first time around, providing strong evidence that they had been able to retain their relational matching skills over this one-year period.

As with the capuchin monkeys, the baboons were not naturals at these tests – they went through thousands upon thousands of trials and only gradually acquired their relational-matching skills. Once again, though, the research strongly suggests that there is not a bright line “profound disparity” between the capabilities of hominids and those of other animals, and that other animals can demonstrate the cognitive foundation necessary for abstract analogical reasoning.

So, as in other areas, the more we explore the abilities of animals, the more we find that we have been wrong about what we thought were cognitive barriers. As we become more adept at designing experiments that are patiently conducted and thoughtfully tailored to the skills and natural adaptations of the specific animals we are studying (rather than the skills and adaptations of college undergraduates), we should continue to see the breakdown of additional barriers.

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ResearchBlogging.orgTruppa, V., Piano Mortari, E., Garofoli, D., Privitera, S., & Visalberghi, E. (2011). Same/Different Concept Learning by Capuchin Monkeys in Matching-to-Sample Tasks PLoS ONE, 6 (8) DOI: 10.1371/journal.pone.0023809

Fagot, J., & Thompson, R. (2011). Generalized Relational Matching by Guinea Baboons (Papio papio) in Two-by-Two-Item Analogy Problems Psychological Science, 22 (10), 1304-1309 DOI: 10.1177/0956797611422916

They’ll Take Two in the Bush – Crows and Ravens Show Patience

We live in an “act now!” world that frequently tests us, luring us with temptations and encouraging us to indulge. We may clearly see the importance of living within our budget yet still be dazzled by the shiny appeal of that new sports car; we may strongly believe in the benefits of a healthy diet yet still be weakened with lust for that large slab of double chocolate cake.

Nevertheless, we do sometimes succeed in delaying immediate gratification for the sake of something better in the future, in remembering those clichés about “good things come to those who wait” that our parents and grandparents inflicted on us. Undoubtedly, this is something we’re able to achieve because we’re humans, because we can be goal-directed and can prevail over our impulses, because we are more than unthinking animals who are captives to their immediate needs. Right?

Not so fast.

It is true that many animals seem unable to defer gratification, with prior experiments showing that animals such as rats, pigeons and chickens will rarely choose a delayed food reward over an immediate one, even if the delayed reward is much more attractive and the delay is only a few seconds. (From an evolutionary standpoint, this sort of impatience may make a lot of sense when an animal faces competition and future opportunities for food are unknown. “Life is uncertain, have dessert first!”)

To date, the major exception has appeared to be in primates: chimpanzees, bonobos, rhesus macaques and capuchin monkeys have demonstrated that they can wait for up to five minutes or so if that enables them to obtain a desirable food reward – a level of performance comparable to that of humans. (Interestingly, tests have shown that we humans seem to be much better at deferring money rewards than food rewards. Perhaps this, too, has a basis in natural selection, as food has been obviously always been an imperative, whereas money has existed for only an evolutionary blink of the eye.)

Also, while all of this might lead one to conclude that the ability to delay gratification lies solely within the province of humans and our closest relatives, it now turns out that corvids, the famously smart bird family (see prior AnimalWise posts here and here and here and here) that includes ravens and crows, may be every bit as patient.

When's dinner going to be ready? (from Wikipedia, photo credit: Cj005257)

As described in a paper published last week in Biology Letters, a team led by Valérie Dufour of the University of Strasbourg recently found that crows (Corvus corone) and ravens (Corvus corax) are able to tolerate delays of over five minutes in order to obtain a better reward, and that they may use the same sort of tactics to distract themselves while they wait as humans do.

In this study, six crows and six ravens were first trained to exchange tokens for food rewards, and then were given a series of “delayed exchange” tests. In each test, a bird would be handed an initial piece of food, which it could either eat immediately or, upon receipt of a signal after a designated waiting period, exchange for a more a desirable reward that it could see throughout the testing period. If the bird ate the initial reward or tried to exchange it too early, the test would end, but if it waited until the proper signal after the waiting period had elapsed – success, a better reward!

The researchers ran the tests with different types of reward (which they labeled as low-, medium- and high-quality) and with varying waiting periods (from 2 to 640 seconds).

Not surprisingly, the birds were generally more willing to exchange for the most highly preferred rewards and, as the following graphic illustrates, had a harder time as the delay period increased (with both crows and ravens maxing out at 320 seconds, or slightly over five minutes):

Interestingly, when the birds had to wait 20 seconds or longer before being able to exchange, they usually placed the “reward in the hand” on the ground and/or cached it in nearby crevices. The researchers believed this to be a distractive strategy, as “[t]hese behaviours probably alleviate the cost of waiting: not having to hold the food distracts the bird’s attention from it.”

As someone who routinely has to put snack food out of reach or even out of sight in order to prevent Homer Simpson-like devouring, this explanation makes a lot of sense to me. (For those of you who would prefer a more uplifting example of a strategy for avoiding temptation, I invite you to think about Ulysses having himself lashed to his ship’s mast so that he can safely listen to the songs of the Sirens.)

In any event, delaying gratification is significant because it involves, on some level, making a judgment about the future and the likelihood of achieving a prospective reward. While it’s not clear whether this entails a full “sense of self,” it is worth (re)noting that corvids are one of the few animals that have demonstrated the ability to recognize themselves in mirrors, a cognitive test that’s often used to measure whether an animal has at least rudimentary self-awareness.

Once again, corvids are no bird brains!

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ResearchBlogging.orgDufour V, Wascher CA, Braun A, Miller R, & Bugnyar T (2011). Corvids can decide if a future exchange is worth waiting for. Biology letters PMID: 21920957.

Rosati, A., Stevens, J., Hare, B., & Hauser, M. (2007). The Evolutionary Origins of Human Patience: Temporal Preferences in Chimpanzees, Bonobos, and Human Adults Current Biology, 17 (19), 1663-1668 DOI: 10.1016/j.cub.2007.08.033.

Heilbronner, S., & Platt, M. (2007). Animal Cognition: Time Flies When Chimps Are Having Fun Current Biology, 17 (23) DOI: 10.1016/j.cub.2007.10.012.

Reconciling and Reassuring Ravens

Welcome to the elaborate, conflict-laden world of raven (Corvus corax) social dynamics!

Expanding on prior research demonstrating that ravens sometimes console fellow ravens who’ve been victims of aggression, researchers have now found that ravens who’ve been in conflicts often reconcile with their former opponents, the first time this behavior has been seen in birds.

Reconciling Ravens

In a study published this year in PloS ONE1, University of Vienna biologists Orlaith Fraser and Thomas Bugnyar found that reconciliation behavior does indeed occur between ravens who’ve had conflicts, particularly when the participants share a valuable relationship. While this sort of post-conflict kiss-and-make-up behavior is believed to play an important role in reducing stress and repairing relationships in primates and certain other mammals, it hadn’t been found in prior studies of birds, leading researchers to hypothesize that perhaps birds use different strategies to maintain social harmony or that reconciliation isn’t so important for birds, as their most important relationships are their pair bonds with mates, where they may be able to avoid significant conflicts in the first place.

Will we fight again? Nevermore! (photo credit: Audubon Guides)

Fraser and Bugnyar studied seven captive sub-adult ravens (who were too young to have formed pair bonds) for 13 months, measuring their behavior after a total of 197 aggressive conflicts (defined as incidents involving hitting, chasing or forced retreat). They then documented “affiliative behavior” (friendly interactions involving contact sitting, preening, beak-to-beak or beak-to-body touching) after each conflict, and compared it to the behavior occurring during neutral periods when no aggression had taken place.

They found that reconciliation (friendly contact occurring within 10 minutes of the end of the conflict) occurred after 37 of 197 conflicts and, in a significant majority of the cases, friendly interactions took place more quickly after a conflict than during the matched control period. Moreover, birds who were related or in “high value relationships” (pairs who had previously been observed to preen or sit in contact with one another) were more likely to reconcile. Interestingly, neither the sex-combination of the opponents nor the intensity of the conflict (measured by whether the birds hit each other and how many times a bird was chased or forced to retreat) impacted the likelihood of reconciliation.

The researchers did note that the behavior of ravens in the wild might differ from those in captivity, and that additional study would be needed to determine whether other factors, such as a history of food sharing, might also impact reconciliation behavior.

This study is significant in that it suggests that, through a convergent process and despite very different evolutionary histories, ravens have developed conflict resolution strategies that are similar to those employed by primates, reconciling with each other to preserve valuable relationships and reduce the chance of further discord.

Reassuring Ravens

This 2011 reconciliation research follows closely on the heels of a comparably-structured study2 that Fraser and Bugnyar published in 2010, also in PLoS ONE, establishing that ravens may possess a sense of empathy (yet another trait once thought to belong to humans alone, at least before evidence of empathy began turning up in primates and other animals).

In the 2010 study, Fraser and Bugnyar attempted to measure whether “bystander” ravens (those who’d witnessed but not been involved in an aggressive conflict) would console the conflict victim through “affiliations” (the same sort of friendly behavior – contact sitting, preening, beak-to-beak or beak-to-body touching – as was measured in the more recent “reconciliation” study).

Don't worry, you're much better looking than he is... (photo credit: pdphoto.org)

This time, they studied 11 sub-adult and two adult ravens raised in captivity, reviewing behavior after a total of 152 conflicts and in matching control periods and finding that both spontaneous and solicited (that is, initiated by the victim) bystander affiliations were likely to occur after conflicts.

More specifically, they found that unsolicited bystander affiliations were more likely to occur after more intense conflicts as well as when the ravens were related or shared valuable relationships, factors which suggested to the researchers that the affiliations served a distress-alleviating, or consoling, function. Also, the bystanders generally had stronger ties to the victims than to the aggressors, leading the researchers to conclude that it was unlikely that the bystanders were either acting as proxies for the aggressor to try to repair relationship between the combatants or trying to protect themselves from redirected attacks from the victims.

Based on these findings, Fraser and Bugnyar concluded that the best explanation for the bystanders’ unsolicited friendly behavior was that they were acting to console and alleviate the distress of the victims. The summarized the significance of this as follows:

Consolation is a particularly interesting interaction because it implies a cognitively demanding degree of empathy, known in humans as ‘sympathetic concern’. In order for a bystander to console a victim, they must first recognize that the victim is distressed and then act appropriately to alleviate that distress, requiring a sensitivity to the emotional needs of others previously attributed only to humans.

While the researchers noted some caveats, including the fact that study didn’t attempt to record vocalizations and that research on ravens in the wild was still necessary, they concluded that “the findings of this study … suggest that ravens may be responsive to the emotional needs of others.”

So, before you leave, here’s a multiple choice test regarding the moral of this story:

  1. Ravens are super smart, just like crows, nutcrackers, magpies and other corvids.
  2. We keep finding more and more ways in which other animals are able to do “uniquely human” things.
  3. If you plan on having an argument with a raven, you should make sure you bring all your raven buddies with you for support.
  4. All of the above.

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1Fraser, O., & Bugnyar, T. (2011). Ravens Reconcile after Aggressive Conflicts with Valuable Partners PLoS ONE, 6 (3) DOI: 10.1371/journal.pone.0018118.

2Fraser, O., & Bugnyar, T. (2010). Do Ravens Show Consolation? Responses to Distressed Others PLoS ONE, 5 (5) DOI: 10.1371/journal.pone.0010605.

Birds Display Their Animal Magnetism

In today’s post, I’d like to talk about something that many animals can do, and that humans simply cannot. I’m not referring to flying, breathing through gills, spinning webs, or running at 70 miles an hour across the African Savanna. I don’t even mean echolocation or pheromone detection or electroreception or polarized light detection, although some of these may be topics for future posts.

No, today I’d like to focus on magnetoreception, or the ability to sense the Earth’s magnetic fields as a means of perceiving one’s direction or location.

Many animals are able to sense magnetic fields, including honeybees, fruit flies, sea turtles, newts, lobsters, salmon, sharks and even bacteria, but some of the more in-depth and interesting research has been centered around migratory birds’ ability to use the Earth’s magnetic fields as a navigational aid as they make their long journeys.

For some reason, I've always felt strangely drawn to the North... (photo from Wikipedia, credit: Thermos)

While there are many aspects of this phenomenon that are still a mystery to us, scientists currently believe that there are two primary biophysical pathways that may explain how birds are able to navigate using the Earth’s geomagnetic fields. These two pathways are currently believed to coexist and complement each other, even though they involve very different processes.1

No, it's simple: just take a left at Norway and then bear right at Finland ... you can't miss it. (photo from Wikipedia, credit: Ernst Vikne)

The first pathway involves the use of iron-based receptors (crystals of a mineral known as magnetite) in the upper beak that can receive and transmit signals of a magnetic field directly to the bird’s brain via a specific nerve (the trigeminal nerve); the second pathway involves the activation of proteins called cryptochromes in the bird’s retina that, after being exposed to blue light, are sensitive to magnetic fields, enabling retinal cells to convert magnetic signals into visual ones and to transmit these signals to a specific sensory processing region (known as Cluster N) within the area of the bird’s forebrain responsible for vision.2

This second pathway may actually enable the bird to create images based on the magnetism fields – in other words, to actually see the Earth’s magnetism and use it as a navigational aid during migration, even during the dark of night.

Aside from being this being pretty cool, what is one to make of this from the standpoint of comparative cognition? Do we chalk one up for the birds (and the honeybees, fruit flies, sea turtles, etc.) and concede that humans are rather lacking in an important cognitive area?

I can already hear the objections. How can you compare this to those characteristics that truly set humans apart from the rest of animals? This is simply a heightened sensory ability, kind of like good eyesight or a fine sense of smell, and not that significant from the standpoint of higher cognition.

This is a fair point and an understandable (if anthropocentric) way to look at it, but there are still a couple of things you may want to consider:

First, we shouldn’t be overly dismissive of the amount cognitive sophistication involved in the long distance navigational feats. Think about it: migratory birds are able to travel, day and night, in all kinds of weather conditions, over great distances – sometimes thousands of miles – with a level of precision beyond our reach until the advent of GPS systems (thank you, Garmin). This sort of navigation is no simple process, either, as sensory input from multiple sources must be assessed, weighted, properly prioritized, reconciled and synthesized, all on a dynamic basis and in an environment when cues are constantly changing.

Also, consider whether this is another area where we humans may be tempted to slant the playing field to our advantage. While it’s easy to imagine our downplaying the significance of magnetoreception from a cognitive standpoint (after all, I just did), remember that we have no trouble in justifying to ourselves why “non-cognitive” human features, such as opposable thumbs and bipedalism, ought to be considered as important factors in distinguishing our capabilities from those of other animals. The point is not to argue that opposable thumbs weren’t critical to our becoming highly sophisticated tool-makers and users (I assume they were), but rather to suggest that we should be as flexible in thinking about factors pertinent to animal cognitive abilities as we are to those pertinent to our own.

Anyhow, time to fly away for the evening – see you soon!

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1University of Oldenburg, Department of Biology and Environmental Sciences, Animal Navigation website, visited July 21, 2011.

2See, e.g., Heyers, D., Zapka, M., Hoffmeister, M., Wild, J., & Mouritsen, H. (2010). Magnetic field changes activate the trigeminal brainstem complex in a migratory bird Proceedings of the National Academy of Sciences, 107 (20), 9394-9399 DOI: 10.1073/pnas.0907068107, and Heyers, D., Manns, M., Luksch, H., Güntürkün, O., & Mouritsen, H. (2007). A Visual Pathway Links Brain Structures Active during Magnetic Compass Orientation in Migratory Birds PLoS ONE, 2 (9) DOI: 10.1371/journal.pone.0000937.

Something to Crow About

Some of you may be aware that crows (who are corvids, like magpies and Clark’s Nutcrackers) are excellent problem solvers and that they are one of the few birds known to engage in tool use.

While there have been a variety of popular press articles describing tool use by New Caledonian crows, in this post I wanted to showcase a few videos that demonstrate visually just how impressive these crows are.

The first video features a New Caledonian crow creating a bent wire hook to fish out a food treat after realizing that a straight piece of wire won’t do the trick. Check it out; it’s pretty incredible:

In a second demonstration of cognitive abilities, the crow employs a sequence of three tools to obtain food reward – using a short stick to withdraw a medium-length stick, using the medium-length stick to obtain a long stick, and then using the long stick to reach the food. As the video notes, this is the first time a non-human animal with no explicit training has been observed using three different tools in the correct sequence to achieve a goal. Again, the video illustrates this feat quite nicely:

Finally, a recent Wired1 article, together with accompanying video, features a New Caledonian crow finding a novel use for a tool, poking a rubber spider. This sort of flexible tool use is quite rare, and crows are the first non-mammals who have demonstrated that they can use a single tool in multiple ways. Here’s the video:

I love how the crow gingerly pokes at the rubber spider and then jumps back – talk about a a familiar looking reaction!

For more information and videos relating to tool usage by New Caledonian crows, you can explore the tool use website2 of the Behavioural Ecology Research Group at the University of Oxford.

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1Wired, “Clever Crows Use Tools in New Way,” January 5, 2011.

2Visited on July 11, 2011.

The Rational Ant

In a recent post I described how pigeons are better than humans at solving the Monty Hall problem and might therefore prove to be formidable competitors on Let’s Make a Deal. In this post, I have some good news and some bad news for those of you readers who are human (I make no assumptions in this blog). The good news is that I have yet to see any research showing that pigeons can triumph over humans at Jeopardy. The bad news is that the top two winners on Let’s Make a Deal could well end up being a pigeon and an ant, leaving the human contestants to go home with nothing more than an electronic version of the game (and perhaps a goat or two).

An article in ScienceNOW1 provides the backdrop:

Ants enjoying a nectar lunch on a sunny day (photo: Wikipedia)

Consider the following scenario: You want to buy a house with a big kitchen and a big yard, but there are only two homes on the market–one with a big kitchen and a small yard and the other with a small kitchen and a big yard. Studies show you’d be about 50% likely to choose either house–and either one would be a rational choice. But now, a new home comes on the market, this one with a large kitchen and no yard. This time, studies show, you’ll make an irrational decision: Even though nothing has changed with the first two houses, you’ll now favor the house with the big kitchen and small yard over the one with the small kitchen and big yard. Overall, scientists have found, people and other animals will often change their original preferences when presented with a third choice.

Not so with ants. These insects also shop for homes but not quite in the way that humans do. Solitary worker ants spread out, looking for two main features: a small entrance and a dark cavity. If an ant finds an outstanding hole–such as the inside of an acorn or a rock crevice–it recruits another scout to check it out. As more scouts like the site, the number of workers in the new hole grows. Once the crowd reaches a critical mass, the ants race back to the old nest and start carrying the queen and larvae to move the entire colony.

The article goes on to describe some research on ant decision-making conducted by Stephen Pratt, an Arizona State University behavioral ecologist, and Susan Edwards, of the Department of Ecology and Evolutionary Biology at Princeton University. In this research, published in Proceedings of the Royal Society: Biological Sciences2, Pratt and Edwards designed a series of possible nests for 26 ant colonies:

The duo cut rectangular holes in balsa wood and covered them with glass microscope slides. The researchers then drilled holes of various sizes into the glass slides and slipped plastic light filters under the glass to vary the features ants care about most. At first, the colonies only had two options, A and B. A was dark but had a large opening, whereas B was bright with a small opening. As with humans, the ants preferred both options equally: The researchers found no difference between the number of colonies that picked A versus B.

Then the scientists added a third option, called a decoy, that was similar to either A or B in one characteristic but clearly worse than both in the other (a very bright nest with a small opening, for example). Unlike humans, the ants were not tricked by the decoy, the team reports online today in the Proceedings of the Royal Society B. Although a few colonies picked the third nest, the other colonies did not start favoring A or B and still split evenly between the two.

Ants can make better decisions because they take advantage of collective wisdom and do not “overthink” their options the way humans are prone to do. As Pratt noted in an article published in PhysOrg.com3, “Typically we think having many individual options, strategies and approaches are beneficial, but irrational errors are more likely to arise when individuals make direct comparisons among options.”

This research is particularly fascinating in that it poses a direct challenge to our core belief that we will always enjoy a large advantage over other animals when there is an intellectual way to solve a problem: sure, animals may have highly-evolved senses of smell, they may be fast, they may have impressive reflexes and their instincts may be powerful, but where we humans are able to harness our large brains, we will inevitably prevail.

In fact, though, we should hold off before patting ourselves on the back. As this (and other) research shows, we suffer from biases and flaws in the way we approach thought problems that can lead to irrational decisions and that can even put us at a disadvantage vis-à-vis other animals, including the birds and the ants.

Something to think about.

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1ScienceNOW, “Can’t Decide? Ask an Ant,” July 22, 2009.

2Edwards SC, Pratt SC. Rationality in collective decision-making by ant colonies. Proc Biol Sci. 2009 October 22; 276(1673): 3655–3661, published online 2009 July 22 (doi: 10.1098/rspb.2009.0981).

3PhysOrg.com, “Ants more rational than humans,” July 24, 2009.

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