What’s Up With the Male Dolphins of Shark Bay Who Don’t Use Sponges?

As discussed in detail in a recent AnimalWise post, a group of female bottlenose dolphins in Shark Bay, Western Australia has enjoyed quite a bit of attention of late for creatively using marine basket sponges as tools to assist them in rooting out bottom-dwelling fish. While the spotlight has been on the females, not much has been said about the males who (despite growing up fin-to-fin with sisters who learn how to use sponges) generally do not become spongers. The researchers studying the sponging behavior have not explored the lack of male sponging in depth, but have hypothesized that the males may be too focused on establishing and maintaining “alliances” to be able to devote the time and effort necessary to become specialized sponge-using foragers.1

I can't figure out that sponging thing either... (photo credit: Shark Bay Dolphin Project)

So what are these male alliances all about, and why are they so important?

Fortunately, a study published in the August 23, 2011 issue of Biology Letters2 provides some new detail and insight into male bottlenose dolphin alliances in Shark Bay.

In a surprise to most likely no one, the alliances are all about sex – maximizing a male’s chances of being able to mate. What is surprising, however, is the level of complexity of these male relationships.

Only humans and Shark Bay bottlenose dolphins are known to have multiple-level male alliances within a social network.

The researchers already knew that the Shark Bay males formed two distinct levels of alliance: first-order groupings of three (or, less frequently, two) males who cooperate to establish and maintain “consortships” with females, and second-order alliances comprised of two or more primary groups who band together to take females from other alliances and/or to defend against such “theft” attempts.

In itself, this degree of cooperation is notable, as alliances and coalitions within social groups are considered to be a hallmark of social complexity (for a posting on female elephant social networks, see here, and for hyena social dynamics, see here). The researchers put it succinctly: “Only humans and Shark Bay bottlenose dolphins are known to have multiple-level male alliances within a social network.” (AnimalWise aside: why are males less apt to have multi-tier social networks than females? Ok, perhaps I don’t need to ask….)

Are you in my second-order alliance? This is all so complicated! (photo credit: Shark Bay Dolphin Project)

In this most recent study, the research team describes a Shark Bay male dolphin society that is even more complex than previously reported – one that actually has three levels of nested alliances among males.

The researchers spent over five years observing 121 frequently-seen males in over 500 consortships, concluding that amicable low-level associations (i.e., third-order alliances) were regularly occurring between specific second-order alliances and trios or other second-order alliances. The researchers further noted that fights involving multiple groups of males suggested that the third-order alliances, like the second-order ones, are employed in conflicts over females, as higher-order alliances could be useful if second-order partners were not around when rivals appeared.

A few other interesting research findings include:

  • There was a nearly continuous range in the size of second-order alliances, which had between six and 14 members.
  • There did not appear to be a relationship between the size of the second-order alliances and how stable (long-lasting) their component first-order trios were.
  • Most of the males participated in second-order alliances, but a subset of five trios did not. Of these five trios, four were comprised of older males whose prior second-order alliance partners had disappeared over time. The researchers surmised that these particular dolphins may have participated in third-order alliances because they were particularly in need of assistance in protecting and obtaining females.
  • Most of the first-order trios associated with only one second-order alliance, but a small subset (around 3%) associated with more than one second-order alliance.

So, to sum up, while (a subset of about 1/11 of) the female bottlenoses of Shark Bay are engaging in specialized tool use with marine sponges, the males are absorbed in complex Machiavellian political relationships and sexual maneuvering. Hmm, sounds a bit familiar.

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1Mann, J., Sargeant, B., Watson-Capps, J., Gibson, Q., Heithaus, M., Connor, R., & Patterson, E. (2008). Why Do Dolphins Carry Sponges? PLoS ONE, 3 (12) DOI: 10.1371/journal.pone.0003868.

2Connor, R., Watson-Capps, J., Sherwin, W., & Krutzen, M. (2010). A new level of complexity in the male alliance networks of Indian Ocean bottlenose dolphins (Tursiops sp.) Biology Letters, 7 (4), 623-626 DOI: 10.1098/rsbl.2010.0852.

Cornered Rat Waves Poisoned Tool, Attacker Flees in Terror!

Screams the tabloid headline…

Is this the plotline for a sequel to The Planet of the Apes in which mistreated lab rats rebel against cruel animal experimenters?

No, it’s actually an accurate (ok, a bit sensationalized) description of the way in which a small African rat has opportunistically found a way to deploy a poison tool (yes, tool, see below) to defend itself from predators.

For years, observers had suspected something was up with the African crested rat (Lophiomys imhausi): it moves sluggishly, acts fearlessly – practically inviting predators to attack it – and twists around to display boldly-patterned black and white bands along its flanks when it’s excited or threated. Some have speculated that these displays could be designed to mimic the appearance of the spiny porcupine or skunk-like zorilla, and over the years there have been reports suggesting that the crested rat may be poisonous, based in part on anecdotes about dogs retreating in fear from the small rodents or showing signs of having been poisoned after crested rat run-ins.

The mystery of the crested rat was cleared up last week, when a team of researchers led by Jonathan Kingdon of the University of Oxford’s Department of Zoology, published their findings about the rat’s unique set of defenses online in the Proceedings of the Royal Society B1.

Poisonous Defense

The researchers found that the crested rat gnaws and chews the roots and bark of local Acokanthera schimperi trees, which contain a substance called ouabain that is used in a traditional African arrow poison known to be capable of killing elephants by amplifying heart contractions. In chewing on the bark and roots, the rat creates a thick gel-like mixture of saliva and plant toxins, which it proceeds to slather onto the distinctively colored fur along its flanks. Here’s a video of the crested rat in which it briefly displays some grooming behavior:

As it turns out, the hairs of the fur in crested rat’s flank-area are highly specialized and extremely well-suited to deliver this self-applied poisonous mixture. These hairs are essentially perforated cylinders containing fiber-like strands that act as wicks, rapidly absorbing the slobbery, poisonous gel and drawing it up by capillary action. When the researchers chemically analyzed the hairs by infrared spectroscopy, they found strong evidence that that they were indeed absorbing and wicking up ouabain from the saliva mixture. Here’s another video of the hairs doing showing off their wicking abilities (that’s red dye in the video, not blood!):

Properly armed with this potent poison and benefited by some additional physical adaptations (an armored skull, enlarged vertebrae, and dense and thick skin), the crested rat enjoys a suite of defenses that allow it to stare down many a predator. The research paper describes the crested rat’s behavior when threatened:

Flaring of the fur is triggered by external interference or attack on the animal, whereupon white and black banding of the longer hairs on either side of the lateral line effects outlines of the tract in a bold white and black ‘target’ design. An aggravated rat pulls its head back into its shoulders and turns its flared tract towards its adversary as if actively soliciting an attack. This display may or may not be accompanied by vocalizations.

No, you don’t want to mess with Lophiomys imhausi.

The researchers characterize the crested rat’s poisonous defense as “toxicity by acquisition” never before reported for a placental mammal, noting that the closest mammalian analogy may be European hedgehogs, who are known to slather their spines with a mixture of toad venom and saliva, presumably to increase the pain and discomfort that their spines can inflict. By contrast, they point out that there’s no evidence that the crested rat needs to create any kind of a wound; rather, the would-be predator is poisoned when it bites – or even just mouths – the crested rat.

So, is the crested rat just a fascinatingly well-adapted defender, or is it a full-fledged tool user?

Tool Use

Tool user! (We at AnimalWise are never shy about making pronouncements … or speaking about ourselves in the “royal we.”)

Even poisonous rats like carrots! (photo credit: Susan Rouse)

Although not mentioned in the research report, the crested rat’s deployment of the plant toxins does indeed qualify as “tool use” as defined in Benjamin Beck’s Animal Tool Behavior, the most complete catalog of tool use in animals. The original 1980 version contained what remains one of the most widely-accepted scientific definitions of the term:

[T]he external employment of an unattached environmental object to alter more efficiently the form, position, or condition of another object, another organism, or the user itself, when the user holds or carries the tool during or just prior to use and is responsible for the proper and effective orientation of the tool.2

In 2011, this treatise was substantially revised and updated, and now contains the following definition:

The external employment of an unattached or manipulable attached environmental object to alter more efficiently the form, position, or condition of another object, another organism, or the user itself, when the user holds and directly manipulates the tool during or prior to use and is responsible for the proper and effective orientation of the tool.3

While it’s not all that much fun wading through the definitions (would they read better in verse?), the authors themselves make it clear that they would consider the crested rat’s “self-anointment” behavior to be tool use: the bark/roots are “unattached environmental objects,” the crested rat uses them to provide itself with a more efficient defensive position, it holds (carries) and manipulates the tool, and is responsible for properly and effectively orienting it.

In fact, the authors have come up with what they call modes of tool use, including several – Affix (attaching an object to the body with adhesive), Apply (attaching a fluid or object to the body without adhesive) and Drape (placing an object on the body temporarily) – which are directly applicable here.4

Moreover, considering only rodents (there are other examples elsewhere in the animal kingdom), the authors specifically call out a number of additional examples of “Affix, Apply, Drape” tool use by self-anointers: rice-field rats that apply the anal gland secretions of the weasel, one of their predators, presumably for concealment purposes; and California ground squirrels, rock squirrels, and Siberian chipmunks that anoint themselves with the scent of rattlesnakes by chewing shed snakeskin, applying dirt (substrate) the snake has been contacted with, and/or anointing themselves with snake urine, all most likely for “olfactory camouflage” purposes.5

So, there you have it. The crested rat is bold, it’s brave, it’s poison, and it’s a tool user!

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1Kingdon, J., Agwanda, B., Kinnaird, M., O’Brien, T., Holland, C., Gheysens, T., Boulet-Audet, M., & Vollrath, F. (2011). A poisonous surprise under the coat of the African crested rat Proceedings of the Royal Society B: Biological Sciences DOI: 10.1098/rspb.2011.1169.

2Beck, B.B. 1980. Animal tool behavior. New York: Garland (as quoted in Shumaker, Robert W.; Walkup, Kristina R.; Beck, Benjamin B.; Burghardt, Gordon M. (2011-04-28). Animal Tool Behavior: The Use and Manufacture of Tools by Animals (Kindle Locations 299-301). JHUP. Kindle Edition).

3Shumaker, Walkup; Beck & Burghardt 2011 (Kindle Locations 372-375).

4Id. (Kindle Location 601).

5Id. (Kindle Locations 1934-1943).

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.

Archerfish: Shooting Dinner from the Sky

Today’s featured guest is the fantastic archerfish, who merits this honor for several good reasons:

  • It is totally cool.
  • It performs highly complex cognitive tasks with tremendous efficiency.
  • It raises interesting questions about how we define tool use.

Archerfish in Action

First, the totally cool.

The archerfish is a small fish that earns a living by shooting prey – insects, spiders and even small lizards – out of the sky, knocking them off twigs and leaves and into the water with an incredibly accurate jet of water launched from its mouth. Here’s a brief video that shows off the archerfish’s hunting skills:

Pretty incredible, huh?

Complex Calculations

And this brings us to the topic of the archerfish’s specialized and complex cognitive abilities.

As the video notes, because the archerfish hunts from beneath the water’s surface, it must be able to take into account both the bending of light at the surface of the water and the curvature of the water stream it shoots toward a target perched as much as two meters away. I’m not aware of any studies on how adept humans are at shooting water pistols at above-ground targets while snorkeling, but in an accuracy contest, I’m betting on the archerfish.

Additionally, in a report published in Current Biology1 in 2006, researchers from Erlangen-Nürnberg University in Germany showed that the archerfish not only aim accurately, but are able to save energy by estimating the size of their prey and modifying the amount of water they shoot. Using high speed photography, the researchers “discovered that archerfish transfer systematically larger maximum forces to larger targets … for any given size of prey, the fish apply about ten times the forces the adhesive organs of prey of that size could maximally sustain.”

Dinner is served! (photo credit: Peter Arnold)

In a later study published in Science2, the same research group elaborated on the efficiency and speed with which archerfish are able to swim to the precise spot where their prey will land after being hit by a water jet. (Because archerfish hunt in groups and are surrounded by other surface-feeders, they have to be able to swim to fallen prey extremely quickly or they will lose it to another hungry mouth.)

The researchers found that archerfish are able react to the motion of falling prey and start swimming to the correct spot at the correct speed within as little as 40 milliseconds, 1/20 of a second. Moreover, the archerfish accomplish this complex task (which requires them to process a host of variables, including the initial height of the prey, the speed of the fall, and the direction in which it is falling) using relatively few neurons and without reference to a priori information regarding the trajectory of the water jet that hit the prey. As the researchers summarized it, “our data show that even complex decisions can be made rapidly and accurately by a relatively small number of neurons.”

So, as we consider the meaning of the archerfish’s impressive skills, we should bear in mind that sophisticated cognitive behavior can evolve to address the particular tasks and challenges facing a species, and that even an animal with a small, non-mammalian brain can accomplish “super-human” cognitive feats if those feats help the animal to successfully adapt to its ecological niche.

Tool Use?

One final question is whether the archerfish is engaging in tool use when it shoots down its dinner with jets of water. We touched on this in the earlier post regarding the fearsome clam-smashing tuskfish, noting Jane Goodall’s definition of tool use as “the use of an external object as a functional extension of mouth or hand in the attainment of an immediate goal.” While there will undoubtedly continue to be debate and disagreement over the definition of tool use, some points to ponder for now include:

  • Does the water used by the archerfish constitute an “external object” within the meaning of the Goodall definition? On the one hand, the water was external to the archerfish until it decided to use it to shoot down prey; on the other, the water obviously is not external right at the moment it is launched.
  • Does the “external object” need to be solid, as alluded to in the earlier tuskfish post? Why should the consistency of the object matter?
  • Can one argue that the archerfish is transforming the nature of the water (from a surrounding environmental medium into a targeted projectile)? If yes, does this imply that the archerfish’s use is more sophisticated than, say, simply picking up a rock lying outside on the ground (or a monkey wrench hanging on an Ace Hardware rack) and using it as is?
  • Some researchers have described behavior that meets some, but not all, of the requirements of a strict tool use definition as “proto” or “borderline” tool use. Is that what we are talking about here?
  • Should the behavior speak for itself without attempting to attach a label to it? Why does it matter whether or not we categorize the behavior as tool use? Is there something anthropomorphic about the “tool use” label in the first place?
These are all interesting questions, at least for those of us who are not preoccupied with shooting our dinner out of the sky.

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1Schlegel, T., Schmid, C., & Schuster, S. (2006). Archerfish shots are evolutionarily matched to prey adhesion Current Biology, 16 (19) DOI: 10.1016/j.cub.2006.08.082.

2Schlegel, T., & Schuster, S. (2008). Small Circuits for Large Tasks: High-Speed Decision-Making in Archerfish Science, 319 (5859), 104-106 DOI: 10.1126/science.1149265.

On the Branch of a Tree, Not at the Top of a Ladder

Every so often, it’s good to see something clearly illustrating that it’s not all about us, that evolution doesn’t simply progress its way up a ladder, climbing ever higher until it reaches humans on the top rung.

Genetic comparisons offer one such clear illustration.

For example, now that we’ve fully sequenced the human and chimpanzee genomes, we can take a close look at the different paths our genes have travelled during the six or seven million years since we parted ways from a common ancestor. In that time, humans and chimpanzees have plainly diverged quite a bit — on the one hand, humans have learned to walk on two legs, experienced dramatic growth in brain size, and now excel at speech, language and a whole host of cognitive functions; on the other hand, although chimpanzees are clearly intelligent primates, they still retain many of the physical and behavioral characteristics that they had millions of years ago.

Obviously, then, our genes have undergone the greater process of Darwinian natural selection … right?

Wrong.

The Human-Chimpanzee Genome Comparison

Do you think those humans are ever going to evolve like us? (photo credit: Delphine Bruyere)

A team of researchers led by Margaret Bakewell and Jianzhi Zhang of the University of Michigan1 decided to systematically compare the human and chimpanzee genomes to find out which species’ lineage has undergone more positive Darwinian selection over time. In essence, they lined up the two genomes to identify where they differed, and then used the DNA of the rhesus macaque, which shares an older ancestor with each of us, to figure out whether differences were due to changes in the human or in the chimpanzee DNA.

Moreover, since some DNA changes have no impact on protein production, the team was able to use statistical methods to look at the changes that do impact protein production and identify which of these were positive in the sense that they conferred a survival or reproductive advantage. (Without getting into the mathematical details, genes where a disproportionate number of the DNA changes do impact protein production are the ones where positive selection is taking place.)

In all, the researchers scanned nearly 14,000 genes (greater than 50% of the genes in the primate genome), and carefully controlled for relatively quality differences in the available genomic sequences. Using their most conservative data, they identified 154 genes that were under positive selection in the human lineage and 233 in the chimpanzee lineage. In other words, chimpanzees have 51% more positively-selected genes than humans have.

The research team summed up these findings:

[I]n sharp contrast to common belief, there were more adaptive genetic changes during chimp evolution than during human evolution. Without doubt, we tend to notice and study human-specific phenotypes more than chimp-specific phenotypes, which may have resulted in the prevailing anthropocentric view on human origins.

Interestingly, the types of genes undergoing positive change are not particularly correlated to the areas where we have seen the greatest physical divergence, such as brain size. Rather, as the below charts indicate, the areas of positive selection are widely distributed through biological processes, molecular functions and tissue groups (in the charts, PSG stands for “positively selected gene”) :

What Explains the Greater Positive Selection in Chimps?

The researchers believe that the principal explanation for the findings is that, for most of the time that humans and chimpanzees have evolved separately, the average population size of chimps was 3-5 times as large as that of humans. This is significant, as population genetic theories predict that positive selection is less effective in smaller populations (i.e., in a small group there are simply fewer opportunities for the occurrence of beneficial mutations with survival and/or reproductive advantages).

Now, there are a few caveats to this story. For one, this type of comparison does not necessarily capture recent or ongoing changes that are not yet “fixed” in the genome, so recent positive changes to the human genome may have gone undetected. Also, while this analysis adds up the relative number of genes undergoing positive change, it does not take into account the fact that some changes may be more important than others, as a change to a single gene can sometimes have a dramatic impact. Also, the study focuses on changes to genes that impact the proteins that they produce, but not the way in which those genes are expressed (e.g., whether and when the genes are turned on or off), and gene expression can account for very significant differences between species.

Nevertheless, even with these caveats, this study is an eye-opener.

From a human perspective, we naturally see our distinguishing characteristics as critically important, and often assume that they reflect something special from an evolutionary standpoint. When we look closely, though, we sometimes find that our views are far more subjective than objective. From a broader perspective, we have been just a single species with (for most of our history) a relatively small population, and have accordingly undergone a correspondingly slower rate of natural selection.

Makes one wonder whether the chimps, with all of those positive genetic changes, could have evolved a way for handling debt ceilings and political consensus. Now that would be an adaptation that would come in handy these days!

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1Bakewell, M., Shi, P., & Zhang, J. (2007). More genes underwent positive selection in chimpanzee evolution than in human evolution Proceedings of the National Academy of Sciences, 104 (18), 7489-7494 DOI: 10.1073/pnas.0701705104

Asian Elephant Social Networkers

In a terrific new study in this month’s BMC Ecology1, a team of researchers led by Shermin de Silva of the University of Pennsylvania Biology Department has published the results of extensive, multi-year research regarding the social dynamics of a population of Asian elephants (Elephas maximus) at Uda Walawe National Park in Sri Lanka. The researchers studied 286 adult female elephants from September 2006 to December 2008, observing the social relationships they formed on a one-to-one basis, in small groups, and at the overall population level.

While group social behavior in African savannah elephants (Loxodonta africana) has been studied extensively, this new research is the first detailed, quantitative study of a wild Asian elephant population over such a lengthy time period … and what the researchers found was quite surprising.

You spend all your time social networking! First do your homework, then you can go on Facebook (photo credit: HelpElephants.com)

Prior less comprehensive studies had suggested that Asian elephants form less complex social networks than do African savannah elephants, with Asian elephants forming smaller and looser social groups based primarily on mother/daughter bonds, and rarely if ever involving relationships between unrelated females. In this in-depth longitudinal study, though, a different, more nuanced, portrait of Asian elephant society emerged.

Although, on any given day, the researchers would see only small groups of elephants that didn’t appear to interact extensively, over time, individual elephants formed larger social units that could be remarkably stable across years, even while associations among such units varied quite a bit across seasons.

One-to-One Relationships (Dyads)

The researchers started out by measuring how much time pairs of adult females spent together and found that, at a high level, the frequency of their associations was highly correlated across all five seasons in the National Park (Sri Lanka has a highly seasonal environment, with two separate monsoon seasons, two dry seasons, and a transitional season) – that is, pairs who associated in one season tended to associate in all seasons, and those who did not associate in a given season weren’t likely to associate at all.

Yeah, let's just hang and make nice for now, then we'll hit the rice paddies when nobody's looking! (photo credit: EleAid.com)

In studying one-to-one relationships, the researchers turned their attention to 51 “core” elephants who they thought would provide particularly good data, since these elephants were observed frequently and during all seasons of the year. These elephants formed a total of 478 pair relationships, which the researchers divided out statistically as follows:

  • A total of six (1.3%) of the pairings were “strong” and stable relationships, as measured by the relative percentage of time these pairs spent together during all seasons. Nine of the elephants (17.6%) participated in relationships in this category.
  • A total of 433 (90.6%) of the pairings were “temporary,” with the association peaking during a single season (most of the peaks were in either the transitional or dry seasons). All 51 of the elephants had at least one relationship that fell into this category.
  • A total of 39 (8.2%) of the pairings were “cyclical,” with the associations peaking in frequency during the two dry seasons (interesting, the researchers did not find relationships where the peaks were during the two wet seasons). Thirty two (62.7%) of the elephants had relationships that were cyclical.

Next, the researchers analyzed whether the identities of an elephants’ preferred companions changed over time. Overall, they found that the elephants spent slightly more than 20% of their time with their long-term companions (the top five companions over five seasons) and slightly more than 30% of their short-term companions (the top five for the current season). On an individual level, there was quite a bit of variation: eight (15.7%) of the elephants maintained 4 to 5 of their top five companions for all five seasons, while 16 (31.4%) completely changed their top-five companions during the study.

The researchers cite the example of two elephants, Kamala and Kanthi, who spent nearly all their time together – they were part of the “K” unit (Kamala, Kanthi, Karin, Kavitha and Kalyani, but no Kardashians) that was particularly close – and contrasted this kloseness to an individual named “471” that had few stable companions. (I wonder if this was due to distress over only receiving a number for a name.)

Additionally, the researchers noticed that the elephants who had the most relationships tended to form weaker bonds with each individual partner, whereas those with relatively few pairings tended to spend a relatively large amount of time with each of their companions.

Hmm, these elephants are beginning to sound quite a bit like people…

Small Group Associations (Ego Networks)

At the next level up, the researchers studied so-called “ego networks,” social networks consisting of an elephant and all of the other individual elephants with whom she associated at least once. The researchers focused on 88 of the adult females who they observed in every season, and calculated five measurements for each: (1) the number of her direct companions, (2) the number of ties between the direct companions, (3) the total number of potential ties between each of these direct companions, (4) the ratio of actual to potential ties, and (5) the number of individuals within two degrees of separation of the subject (number of friends plus number of “friends of friends”).

(Note that, assuming at least one of the researchers is within five degrees of separation of Kevin Bacon, this would mean that the entire ego network would be within seven degrees of separation of Kevin Bacon.)

Without getting into the full statistical analysis, the researchers’ principal conclusion was that:

[W]hile a subject’s direct companions do change over time, she has a few that are almost always present; even those that are not present continuously may have been companions in previous seasons. Thus, individuals maintain long-term relationships with others even though they may be apart for one or several seasons and [the amount of time spent together is small].

In other words, the elephants remember their friends and reestablish their relationships even after having been apart for long periods.

Population Level

Finally, the researchers looked at the social structure of the entire population. They found that the elephants in the overall popular had an extensive and well-connected social network, and that the distinct social units within the population were two to three times larger than had previously been seen in the field. Moreover, they observed that many of the social units maintained their integrity across seasons, even as individuals switched units and the connections between the units changed.

For those of you who like to look at dot patterns, below is a colorful series of diagrams depicting the connections between elephants, measured at different societal levels and during different seasons (T1 is the transitional season, D1 and D2 are the dry seasons, and W1 and W2 are, you guessed it, the wet seasons):

Figure 5 from Research Paper

Recap

While the strength of the associations among these Asian elephants (as measured by percentage of time that individuals spent together) is generally a good bit lower than that of the associations among African savannah elephants, most of these elephants had a few strong ties as well as consistent ties that they maintained over several seasons. Further, the Asian elephants were hardly asocial – while their mix of companions did fluctuate over time, they often returned to a subset of preferred companions.

Moreover, through their years of observation and statistical analysis of the elephants at the population level, the researchers found that the elephants’ social units were much larger than had been observed in prior studies, and that these social units were more stable across the years than were the companions of individual elephants.

The researchers speculated that one reason for the surprising findings is that the elephants stay in touch in ways that are hard for humans to detect, allowing the elephants to maintain bonds and relationships that we fail to observe. For example, elephants can communicate acoustically over great distances, and often use scent to follow one another’s paths at night (and, for that matter, even when the other elephants would be in plain sight, at least from the human perspective).

Finally, the researchers are planning to perform a detailed genetic study of the population in order to analyze the degree to which relatedness impacts the social organization of Asian elephant society. We’ll be waiting!

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1de Silva, S., Ranjeewa, A., & Kryazhimskiy, S. (2011). The dynamics of social networks among female Asian elephants BMC Ecology, 11 (1) DOI: 10.1186/1472-6785-11-17.

Sheep: Barnyard Brainiacs

It turns out that sheep are far more intelligent than their reputation for barnyard slowness would lead one to believe. In recent research published in PLoS ONE1, Professor Jenny Morton of the Department of Pharmacology at the University of Cambridge and her colleague Laura Avanzo reported that domestic sheep can perform extremely well on tests of designed to measure cognitive abilities, possibly as well as any animal other than primates.

Professor Morton, who had been studying Huntington’s disease, wanted to find out whether transgenic sheep with a specific genetic defect might be useful in preclinical research regarding potential treatments for this neurodegenerative disease. Because Huntington’s is characterized by cognitive deterioration, Morton was particularly interested in seeing how well sheep would perform cognitively, since suitable research subjects for neurologic disorders like Huntington’s inevitably must undergo systematic cognitive testing relevant to the disease.

Accordingly, Morton and Avanzo devised a series of tests that they gave to seven female Welsh Mountain sheep, six of whom completed the whole study. No word on why all of the ungulate volunteers were female, although my guess is that the males were off rollicking around with male bottlenose dolphins who were avoiding sponge fishing duty.

Welsh Mountain ewe: wool-giver and five-time Jeopardy champion (photo credit: Vertigogen)

The Tests

The tests were designed to measure the ability of the sheep to perform in three areas (discrimination learning, reversal learning and “attentional set-shifting”), which are relevant to what the researchers refer to as executive function – that is, the “ability to learn associations between stimuli, actions and outcomes, and to then adapt ongoing behavior to changes in the environment.” While the sheep took a large number of very specific tests, the tests fell into the following general categories:

  1. Simple discrimination tests. Sheep must choose between two feed buckets that are identical except one is blue the other is yellow. One color contains a food reward; the other is empty. Later “retention tests” repeat the original tests after time has passed to see how well the sheep remember.
  2. Simple discrimination reversal tests. Sheep must relearn the correct answer after sneaky researchers reverse the color of the bucket containing the food reward. (Note: we encountered this type of testing in the earlier AnimalWise post about the clever Anole lizards). Again, later “retention tests are given.
  3. Compound discrimination tests. The rewarded color is the same as in 1 above, but the relevantly-colored objects are now “perforated sports cones” rather than buckets. Additional buckets of irrelevant colors (one black, one green) are placed next to the sports cones, with the food reward in whichever bucket happens to be next to the correctly-colored sports cone.
  4. Intradimensional shift tests. Now, the sheep are presented with new shapes (rhomboids and cones) and new colors (purple and green). The sheep must still make a correct choice based on color, but need to learn the new color to apply.
  5. Intradimensional shift reversal tests. Same as 4, but sheep must relearn correct answer after the researchers change the rewarded color.
  6. Extradimensional shift tests. Again, the sheep are presented purple or green cones or rhomboids, but this time they must figure out now that the reward is based on choosing the correct shape, rather than a particular color.
  7. Extradimensional shift reversal test. Same as 6, but sheep must relearn after researchers swap which shape is rewarded.

Of the above tests, 1 & 3 measure “discrimination learning”; 2, 5 & 7 measure “reversal learning”; and 4 & 6 measure “attentional set-shifting.”

The Results

In a nutshell, the sheep did amazingly well.

They very quickly learned to pass the initial simple discrimination test (within seven sets of eight discriminations). When presented with the first reversal test, their performance initially dropped off, but they learned the new correct answer within three days of testing (11 sets of discriminations). For the compound discrimination testing, their performance again dropped slightly at the outset, but within two days they had this new puzzle figured out as well. Moreover, the retention tests showed that the sheep were able to remember the correct answer after time had passed (six weeks in the case of the simple discrimination test; two weeks for the simple reversal test).

At first, the sheep performed no better than chance on the more difficult intradimensional shift test, but they soon were performing at over 90% correct. They also experienced a large drop off in performance on the extradimensional shift test, but improved gradually until they reached 80% correct on the fourth day of testing. The sheep learned also were able to learn the reversals (within eight sets of discriminations for the intradimensional reversal and within 10 sets for the extradimensional reversal).

Morton and Avanzo summarized the results as follows:

We show that not only can normal can sheep perform discrimination reversal learning tasks, but they can also perform attentional set shifting tasks that test executive function. To our knowledge, this is the first time that these executive functions have been demonstrated in any large animal, apart from primates.

They were surprised by this success, conceding that they hadn’t been expecting the sheep to do well on the more difficult tests and indicating that they were “driven more by curiosity than expectation” in even giving the tests to them.

So, given these results, sheep seem to have gotten a bum rap for intelligence. There are relatively few studies on ovine intelligence, although research has shown that they can learn and remember how to navigate complex maze2 and that they are very good at remembering faces3.  And then there’s my favorite, that they’ve learned to roll their way across hoof-proof metal cattle grids in order to raid villagers’ valley gardens4!

One reason for the mistaken impression about sheep cognition may be that we have a bit of a blind spot when it comes to intelligence. We expect it in ourselves and a few other select animals, but even scientists can be quite surprised when it pops up elsewhere. Perhaps the main lesson here is that we should do our best to remain open to finding intelligence in unexpected places – if nothing else, this sort of a mental stretch will be a good test of our own cognitive abilities.

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1Morton, A., & Avanzo, L. (2011). Executive Decision-Making in the Domestic Sheep PLoS ONE, 6 (1) DOI: 10.1371/journal.pone.0015752.

2LEE, C., COLEGATE, S., & FISHER, A. (2006). Development of a maze test and its application to assess spatial learning and memory in Merino sheep Applied Animal Behaviour Science, 96 (1-2), 43-51 DOI: 10.1016/j.applanim.2005.06.001.

3Kendrick, K., da Costa, A., Leigh, A., Hinton, M., & Peirce, J. (2007). Sheep don’t forget a face Nature, 447 (7142), 346-346 DOI: 10.1038/nature05882.

4See, e.g., BBC News, “Crafty sheep conquer cattle grids,” July 30, 2004.

Female Dolphins Sponge Their Way to Success

After 27 years, scientists finally appear to have unraveled most of the mystery surrounding a very enterprising group of (primarily) female bottlenose dolphins (tursiops aduncus) who live in Shark Bay, off the coast of Western Australia.

Why are those dolphins looking at me like that? (photo credit: Eric Patterson, Shark Bay Dolphin Project)

The story opens in 1984, when observers first noticed that some of the Shark Bay dolphins were breaking off conical marine basket sponges and wearing them over their beaks (rostra). Because only a small percentage of the dolphins in the area engaged in this behavior and it was very difficult to see what they were doing with the sponges, especially when they were underwater, the first research on this behavior wasn’t published until over a decade later.

Preliminary Findings: Tool Use by a Few Females

In a 1997 article in Ethology1, a team of researchers led by Janet Mann of Georgetown University described their initial findings: five female dolphins were regularly seen with sponges, and four additional dolphins (only one of which was a male) were each seen carrying sponges on a single occasion. The regular sponge users were relatively solitary, tended to use the sponges in a deep water channel area, and did not participate in the group feeding and social aggregations to which other dolphins in the group were attracted.

The researchers weren’t sure what the dolphins were doing with the sponges, but they assumed that there had to be some sort of functional advantage, since the sponges were often quite large, covering a large portion of the dolphin’s face, interfering with normal use of the mouth, contributing to hydrodynamic drag, and potentially impacting the ability to engage in echolocation. They considered three possibilities: that the dolphins were playing with the sponges, that the sponges contained some medicinal or other useful compound, or that the dolphins were using the sponges as a tool to aid in foraging.

They concluded that it wasn’t likely that the sponges were being used as toys, as the spongers were relatively solitary, used the sponges methodically for hours at a time, year after year, and didn’t engage in typical play postures, splashing or vocalizations as they carried the sponges. Similarly, they determined that medicinal or similar uses were unlikely, since, among other things, the regular sponge users all seemed healthy and there were no indications that they were ingesting the sponges (although the researchers conceded that this could be difficult to observe).

Hi ho, hi ho, it's off to sponge I go! (photo credit: Eric Patterson, Shark Bay Dolphin Project)

On the other hand, it did seem likely that the dolphins were using the sponges to help them forage for prey: they were seen eating fish when engaging in sponging behavior; they invested an amount of time in carrying sponges similar to that invested by other foraging dolphins; and they made sounds and generally behaved in ways consistent with foraging. The researchers speculated that sponges might be used to protect the dolphin’s face, either from spines or stingers of prey animals or from the abrasive sea floor as they flushed out burrowing prey. In either case, they believed that this would constitute “tool use,” something that had been reported in captive dolphins but never before in the wild.

Finally, the researchers drew no conclusions on why males didn’t engage in sponging, except to note that perhaps it required a degree of solitary living that was at odds with their need to form and maintain cohesive and cooperative alliances.

Additional Findings: A Cultural Tradition of Tool Use among a Related Group of Females

Next, in 2005, Mann’s researcher team expanded on its findings in a paper published in the Proceedings of the National Academy of Sciences2, with salient points of the research including the following:

  • Sponges Are Foraging Tools. By this time, the researchers had found 15 adults in the community who regularly used sponges, only one of whom was a male. Although not a focus of the paper, it appears that the researchers had concluded by this time that the dolphins were indeed using the sponges as tools to protect their rostra as they foraged for prey on the sea floor.
  • “Sponging Eve.” The researchers tested the mitochondrial DNA of the regular spongers and found that sponging had been passed on mainly along a single matriline (line of descent from mother to daughter) and that, due to the high degree of genetic relatedness, all spongers likely descended from one recent “Sponging Eve.”
  • Female Social Culture. After considering in detail whether the sponging behavior could have resulted from either a genetic propensity or some unique aspect of the deep-water channels where the most of the sponging occurred, the researchers found the evidence for these alternatives lacking and concluded that by far the best explanation was that the sponge use was being socially learned and transmitted from mother to daughter. The researchers weren’t overly surprised by this finding, given that studies had already shown that dolphins have uncommonly complex cognitive and imitative skills and the ability to excel at vocal and social learning.
  • Uncommon Cultural Diversity. It was particularly rare to see this sort of cultural phenomenon in a small subset of the overall population (a single maternal line comprising only about 10% of the females in the group). In other studies (for example, involving apes), this type of culturally learned behavior is seen across the entire population.
  • Can’t Explain Males. Once again, the researchers surmised that perhaps males didn’t engage in sponging because they had to associate at high levels with alliance partners, but they left this point open.

The Story Continues: Spongers Are Fit

The story continued to unfold in 2008, when Mann and her team published a paper in PLoS ONE3 that focused in more on whether sponging was an advantageous behavior, or whether the spongers were in some fashion subordinate or less competitive and were making the “best of a bad situation.”

I don't know what you mean, it's no more elaborate than the other hats at the Royal Wedding... (photo credit: Eric Patterson, Shark Bay Dolphin Project)

By this point, recurrent sponging had been seen in 41 of the dolphins and a few more of them were male (29 were females, 6 were males, and 6 were of unknown sex). This still represented a small percentage (about 11% of adult females were spongers) and, although it now appeared that more than one matriline was involved, the data continued to show that the behavior was consistently passed down from mother to daughter, and less frequently from mother to son: there were no instances observed where a calf adopted the behavior if its mother wasn’t a sponger, and of 19 offspring born to sponger females who could be observed and whose sex was known, 91% of the daughters (10 of 11) and 25% of the sons (2 of 8) adopted sponging.

Further, the researchers found that the spongers were highly specialized, not using other hunting techniques and spending approximately 96% of their foraging time using sponges. In fact, the researchers concluded that, due to their lifestyle and specialization, spongers actually used tools more than any non-human animal.

So, was the sponging advantageous or a way of coping for not particularly well-adapted dolphins? Well, the researchers did find that spongers were more solitary and spent more time foraging at deeper depths and on longer dives, but noted that they really didn’t seem to suffer from any kind of fitness cost, as their calving success was equivalent to that of other females in the population.

Since there was no evidence that any kind of competition for food was relegating the spongers to their strategy, the research concluded that sponging simply seemed to be an “all-or-none phenomenon,” that required a specialized approach and a commitment to a single foraging type, but that most likely opened up a particular hunting niche in a diverse environment. While other dolphins could theoretically adopt the strategy, the researchers noted that daughters in particular tend to adopt their mothers’ foraging strategy, and unless the mother was a sponger, a daughter might simply not have had sufficient exposure to develop this highly specialized technique while a calf.

Once again, the team hypothesized about the males, stating: “Male offspring are exposed to sponging as often as female offspring, but do not seem to adopt the behaviour early, if at all. … [M]ales likely range more widely post-weaning, focus on establishing long-term alliances, and cannot afford to adopt foraging tactics that both demand extensive effort and specialization and limit their range and access to females.”

The researchers offered no opinions about whether the male dolphins were simply slow on the uptake or whether they associated sponges with housework to be avoided.

The Latest Chapter: Explaining the Purpose of Sponging

While all of this research had answered many questions and shed light on a fascinating example of tool use in wild female dolphins, one fundamental question remained. Dolphins are great at using echolocation to detect prey (even prey that is buried), so why do the Shark Bay spongers probe the debris-covered sea floor with their noses, risking injury (even with the protection afforded by the sponges) instead of minimizing sea floor contact by simply echolocating for buried prey as they do in other locations (for example, the Bahamas)?

What a mess! This sea floor needs a good sponging! (photo credit: Eric Patterson, Shark Bay Dolphin Project)

This is the question is answered in the latest chapter, a research paper published last week in PLoS ONE4. Mann’s research team had fun with this one, grabbing poles and going sponging themselves. What they found, aside from the fact that dolphins are far more graceful than people, was that the nature of the prey turned up by sponging helps explain the dolphins’ behavior.

It turns out that most of the bottom-dwelling fish that hide in Shark Bay the sea bottom lack swim bladders, gas-filled chambers used by fish to control their buoyancy as they swim up and down. Because they lack the major characteristic that distinguishes their density from sea water, they generate relatively weak acoustic signals and are difficult to detect with echolocation. In addition, the debris (rock, shell and coral) on the sea floor in the area seemed likely to cause “interfering reverberation and echo clutter,” which would further reduce the effectiveness of echolocation.

Moreover, it’s worth it to go after these swim bladderless fish. They are attractive targets, as they are reliably present on the sea floor and exhibit consistent, predictable behavior when rousted out of their hiding places, allowing the dolphins to adopt a single efficient technique as they sponge. Further, bladderless fish tend to have a relatively high fat content, providing hungry dolphins with a particularly energy-rich meal.

So, the sponging female dolphins of Shark Bay really are quite remarkable. They have established a mother-daughter subculture of tool use in the wild, successfully devising a highly specialized way of exploiting an attractive niche in their diverse environment.

You go girl(s)!

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1Smolker, R., Richards, A., Connor, R., Mann, J., & Berggren, P. (2010). Sponge Carrying by Dolphins (Delphinidae, Tursiops sp.): A Foraging Specialization Involving Tool Use? Ethology, 103 (6), 454-465 DOI: 10.1111/j.1439-0310.1997.tb00160.x.

2Krutzen, M. (2005). Cultural transmission of tool use in bottlenose dolphins Proceedings of the National Academy of Sciences, 102 (25), 8939-8943 DOI: 10.1073/pnas.0500232102.

3Mann, J., Sargeant, B., Watson-Capps, J., Gibson, Q., Heithaus, M., Connor, R., & Patterson, E. (2008). Why Do Dolphins Carry Sponges? PLoS ONE, 3 (12) DOI: 10.1371/journal.pone.0003868.

4Patterson, E., & Mann, J. (2011). The Ecological Conditions That Favor Tool Use and Innovation in Wild Bottlenose Dolphins (Tursiops sp.) PLoS ONE, 6 (7) DOI: 10.1371/journal.pone.0022243.

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.

Portia, Queen of Spiders

Do you think that spiders are mindless machines, driven by pure instinct to make their webs and then attack intruders? Well, it’s time you met Portia:

The formidable Portia! (photo credit: Akio Tanikawa)

In an emerald rainforest of northeastern Australia, a sunbeam pierces the canopy, touches broad green leaves on the way down, and beams onto a lichen-spotted rock surface. In the beam’s circle, the slow, careful motions of a brownish jumping spider are illuminated. The jumping spider belongs to the genus Portia and it is stalking its prey, a different species of spider sitting in its own web. Portia steps cautiously from the rock surface out onto the web and stops. Delicately, Portia begins to pluck the web with its palps and legs, making signals that mimic the struggles of a trapped insect. When the prey spider ignores Portia’s plucking, Portia varies the characteristics of the signals, generating a kaleidoscopic of what appears to be a random selection of signals. Eventually, in response to one of these signals, the prey spider swivels toward Portia. Immediately, Portia backtracks to that particular signal and repeats it again and again. There being no further response from the prey, Portia eventually reverts to broadcasting a kaleidoscope of signals. When the prey spider still moves no farther, Portia adopts another ploy.

Now Portia slowly and carefully stalks across the web toward the resident spider, intermit­tently making a variety of signals. From time to time, a soft breeze blows, ruffling the web. The ruffling creates background noise in the web, and Portia exploits these moments, during which the resident spider’s ability to detect an intruder is impaired, by stalking faster and farther during these periods than when the air is still. Nearing the resident spider, Portia makes a signal that elicits from the resident spider a sudden, rapid approach. However, the spider advances very aggressively, and Portia scrambles to the edge of the web, then turns around to look over the scene. Soon Portia moves away from the web and undertakes a lengthy detour, first going away from the prey and around a large projection on the rock surface, losing sight of the prey spider along the way.

About an hour later, Portia appears again, but now is positioned above the web on a small overhanging portion of the rock. After anchoring itself to the rock with a silk dragline, Portia next slowly lowers itself down through the air, not touching the web at all. Arriving level with the resident spider, Portia suddenly swings in, grabs hold of the unsuspecting spider, and sinks its poison-injecting fangs into the hapless victim.

I’m going to leave the light on tonight, too.

This rather dramatic account is from a chapter in The Cognitive Animal written by Stim Wilcox (Department of Biology, SUNY Binghamton) and Robert Jackson (Department of Zoology, University of Canterbury, New Zealand)1.

For those of you who like visuals, here’s a brief video of Portia:

Clearly, you don’t want to mess with Portia.

In their essay, Wilcox and Jackson note how tricky it is to discuss cognition in animals, with almost as many definitions of the term as people using them. Rather than trying to choose a single definition, they instead apply a framework designed to raise questions about six separate cognitive properties: reception (taking in information), attention (focusing on particular tasks), representation (maintaining a mental image or cognitive map), memory (retaining information), problem solving (deriving pathways to the achievement of goals), and communication language (influencing other individuals by manipulating symbols).

They then run through these properties point by point, in the process illustrating the cognitive abilities of Portia.

Reception: In this area, Wilcox and Jackson emphasize Portia’s amazing eyesight, which apparently is more acute in distinguishing spatial features than that of any known animal of comparable size and even rivals that of primates: “Portia can precisely locate and identify spiders from a distance of 30–40 body lengths away, monitor the spider’s orientation and behavior during the course of a predatory sequence, and in general quickly gain critical information for predatory decisions during complex interactions with a dangerous prey.” For an in-depth article on Portia eyesight, you can check out this piece2 by Jackson and Duane Harland (I love the title: “‘Eight-legged cats’ and how they see”).

Attention: Wilcox and Jackson highlight the extraordinary, prolonged attention paid by Portia as she hunts, especially as she singles out and zeros in on a prey spider in an extended bout of stalking that involves multiple tactics and focused flexibility over time. They note how she must be particularly attentive, given the extreme sensitivity of her prey’s own web to movement and weight, and describe how she manipulates the signals she sends across the target’s web (aggressive mimicry) and opportunistically takes advantage of wind-caused background noise and vibration to move more quickly across the web than she does when the air is still (smokescreen tactics).

Representation, Memory, Problem Solving: For these areas, Wilcox and Jackson point to her planned detours, which demonstrate problem-solving and suggest mental maps and prolonged memory. They observe that, by planning ahead and formulating a solution before executing her maneuver, Portia comes particularly close to what we might typically call “thinking.”

Communication: Finally, while acknowledging that Portia clearly does not have any sort of verbal language or use symbols with arbitrarily assigned meanings, Wilcox and Jackson describe the way in which Portia strings together series of signals as she engages in aggressive mimicry, noting how this involves a complex, flexible and dynamic sequence of interactions between her and her target. As they put it, “Studying Portia’s signal-making strategy from this perspective may bring us closer than we initially expected to something like the cognitive implications of verbal language.”

I don’t have much to add here.  Hail Portia!

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1Wilcox, S. and Jackson, R. (2002). Jumping Spider Tricksters: Deceit, Predation, and Cognition. In M. Bekoff, C. Allen & G. Burghardt (Eds.). The cognitive animal: Empirical and theoretical perspectives on animal cognition. pp. 27–33. Cambridge, MA: MIT Press.

2Harland, D. and Jackson, R. (2000). ‘Eight-legged cats’ and how they see – a review of recent research on jumping spiders (Araneae: Salticidae). Cimbebasia 16: 231–240.