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Crane pair coming home
Another clamor awoke me in the early dawn, around 4:00 a.m. There had been a heavy overnight frost and the two cranes were standing on the ice in the middle of the bog. Both Millie and Roy tossed their heads up in a quick motion, their bills opening wide during each call. We heard what sounded like a hammer hitting a metal bell or drum, as he opened his bill once to make each call and she chimed in at the same time but called and opened her bill twice to make two short similar cries of a higher pitch. It was a composite call made by both together; a duet. A third, distant crane responded. The distant calls, and the pair’s duet, were repeated back and forth, often and loudly.
The vigor of the pair’s unison calling was still palpable, even when the first morning light lit the sky and silhouetted the black spruces, and when I thought about the enormous effort they had invested to get here, I realized what was at stake: home ownership. The pair’s loud clanging calls attracted no others flying in from the distance; instead, the calls are a vocal “no trespassing” sign, one leaving no doubt that any potential challenger would be facing not just one bird, but a united, cooperating pair.
I watched the pair for another hour. She by then occasionally fluffed herself out and, as she had done the day before, continued periodically to squat where she had pulled at or dropped sedge and other potential nesting material.
The pair continued their slow, deliberate steps that morning while meandering from one end of the bog to the other, as though inspecting every square centimeter of it, and at 9:00 p.m. we saw Roy jump high and with outstretched wings dance by himself out on the ice. Millie dashed by him with fluttering wings, to round out a mutual performance. A little later we heard a purring call as she spread her wings to the sides and stood still. Roy, with outstretched neck and elevated bill, jumped onto her back and, balancing himself with a few wing beats, mated. He dismounted after a couple of seconds. Both then bowed to each other and continued their walk. They were home, intended to nest, and had now sealed the deal.
Still, the lone bird that had tried to intrude on their turf did not easily abandon its intended claim. But why would this crane want a claim without a mate to nest with? Was it Oblio, their grown colt from last summer? Was he, now that Millie and Roy were re-nesting, finally being “thrown out of the nest” by his parents, who no longer tolerated company of any sort? I suspect this was the case. The offspring of most birds have a strong attachment to home. This emotion is a biologically relevant drive, because home is where reproduction has proven to be successful. But the young have a lifetime ahead of them, and if for some reason the parents don’t make it back home, the offspring could inherit their territory. If the parents do come back to reclaim their home, at least nearby territory would be more like the old home than the far-off unknown. Even if a bird is without a mate, finding a suitable territory is often a prerequisite to getting one.
Shortly after we had breakfast the next morning, the lone crane flew over once again, and the pair immediately launched their synchronized duet as a vocal challenge. This time, although the lone crane did not land, the pair jumped into the air and chased it until all three were out of our sight and hearing. But the pair returned soon and then again performed several nest-building probes. They again mated in what would become a routine for the next several days: at least once in the morning and once in late afternoon or at night.
When I first saw the cranes the next dawn, each was standing on one leg with its head tucked into its back feathers. Ice had again solidly covered their pond, after having partially melted along the edges the day before. Both Millie and Roy seemed to be asleep, although he, balancing himself on one leg, occasionally reached up with the other to scratch his chin and head with the toes of the foot. But whenever a crane called from the distance, both their heads shot up instantly, and they renewed their spirited in-unison call, she making the two short notes and her mate making one, at a lower pitch.
We had so far not seen them feed. Indeed, it was hard to imagine that there was food available in any case. If lucky, they might by now catch a vole or find a few of last year’s cranberries, but this year that would probably not be likely until days later when the snow would melt. A crane’s large body size requires much more food than a small bird’s, but that same large size is an advantage in tiding them over during lean periods, and thus to return to their homes before the anticipated flush of food becomes available.
As it got lighter on this dawn, the cranes soon had company. A pair of swans and then a small flock of five Canada geese flew by. A robin sang. By about 7:00 a.m., the crane pair became animated as well, walking over the bog while picking here and there, and again mating.
We did not see the pair most of that afternoon. The lone crane, seemingly to take advantage of their absence, again flew in and this time landed, looked around, and repeatedly made a trilling call. But in about fifteen seconds the pair flew in as if out of nowhere, and one of them sped over the ice as if to attack the interloper, which immediately flew off. The pair gave chase, and all three disappeared from sight. In several minutes the single bird returned. Again the pair came and caught up to the lone crane before it had a chance to take off, and this time they attacked it viciously, in a flurry of flailing wings.
The pair had by now, after the third day, won the major part of the battle. Barring accidents, they would within days lay their two eggs and go on to raise their colt. Normally both eggs hatch, but as in some eagles and vultures, usually only one chick survives, probably because one gets fed less and then weakens and eventually starves. Presumably through evolutionary history, for the fast growth required to reach full development and readiness to migrate by August, there has not been enough food to raise two colts at once. One might suppose the cranes could simply lay only one egg, but sometimes an egg does not hatch, and the second is insurance.
The pair seemed more animated after their last fight with the lone intruder, and by evening they again mated (for at least the third time that day). In most birds one mating is enough to fertilize the eggs. Perhaps several matings are insurance, but this seemed more than enough for insurance. Perhaps, like their dances, mating is additionally part of their bonding ritual.
A pair of mallards, and then a pair of pintails, arrived in the evening, and the ducks swam next to each other near the cranes at the edge of the pond, where some open water had reappeared during the day. The cranes ignored them and again walked in their stately manner back and forth across the ice of the pond, and now they pecked in the low vegetation being exposed along the edges. They were by now finding overwintered cranberries exposed by the melting snow.
On the evening before I would leave for my journey home, Christy and George hosted a potluck party. Shadows fell on the white frozen middle of the pingo as the western sky turned yellow and orange and the spruces became dark silhouettes. A pair of pintails again landed in the open water along the pond’s edge. The cranes were standing, each on one leg, their heads tucked into their back feathers. People crowded around the spotting scope in the living room, watching them occasionally shift position, lower a leg, poke a head out to look around. Suddenly the person then at the scope erupted with an exclamation: “They are mating!” She had seen the male approach the female with her spread wings, mount, flutter, and jump off. The pair had bowed to each other. Suddenly many people crowded around the scope to watch.
Why, I wondered, would anyone, or almost everyone, want to watch cranes mate? Why was nobody interested in watching the mating activity of the two ducks, or of the numerous redpolls? Could it be, I wondered, because we feel a closer kinship with cranes than with other birds?
Cranes are similar to us in many ways. Some are nearly as tall as a person. They walk on two long legs like us, albeit with a much more graceful and deliberate gait, so that they remind one of a caricature of a gentleman or an elegant woman on a leisurely stroll. The sandhill crane’s red bald pate and sharp yellow eye add to the caricature. Cranes form lifelong pairs and stay together as families, but they are also gregarious and join up into large groups. They form a strong attachment to their home. They not only make music with trumpeting calls that sound like bugles, but they also dance, and do so on various occasions.
All of the fourteen species of the world’s cranes dance. Crane dancing involves running, leaping into the air, flapping the wings, turning in circles, stiff-legged walking, bowing, stopping and starting, pirouetting, and even throwing sticks. Dancing is primarily done by pairs and presumably functions in cementing pair bonds and/or synchronizing reproduction. But it can also be induced at any time, and it stimulates other cranes to dance. Even the young colts perform some of the species’ dance. Possibly it serves as practice and could be motivated by the same basic emotions of joy that are an indicator of health important to mating.
Cranes’ dances often stimulate humans to dance as well and have been mimicked in many cultures all over the world where cranes live. Crane dances were performed by ancient Chinese, Japanese, southern African, and Siberian people. If not emulated, cranes are admired. In the Blackfoot tribe of Native Americans of northern Montana, the last name “Running Crane” is common.
Nerissa Russell, an anthropologist, and Kevin McGowan, an ornithologist from Cornell University, revealed that eighty-five hundred years ago at a Neolithic site in what is now Turkey, people probably performed crane dances using crane wings as props that were laced to the arms. Furthermore, someone of these people apparently hid a single crane wing in a narrow space in the wall of a mud-brick house along with other special objects (a cattle horn, goat horns, a dog head, and a stone mace head). Russell and McGowan also found evidence that vultures may have been hunted for their feathers for presumably a much different costume worn as well for a ceremonial purpose. The authors inferred that the cranes were linked with happiness, vitality, fertility, and renewal (since they arrived in the spring). While the crane dance was one of life and birth, and possibly marriage and rebirth, the vulture dance was associated with death and perhaps return to the afterlife.
Russell and McGowan believe that the crane wing interred in the wall of the house was never intended to be seen. It was a symbolic object related to marriage and construction of a new home and may have been coincident with a particular human marriage and home-making. The associations among dancing, pairing, and raising young and home would have been natural for people who saw cranes return to their home ground, just as I had seen Millie and Roy do. Seeing the close parallels in the biology of the birds with their own lives, and understanding the cranes’ dancing as helping to make or cause the good things that followed, Neolithic people would have been compelled to symbolically emulate the crane dance of homecoming and of new life.
BEELINING (#ulink_023ec351-cb60-54c0-8f4c-006a10f6452b)
Observation sets the problem; experiment solves it, always presuming that it can be solved.
— Jean-Henri Fabre
CRANES FLY AN ENORMOUS DISTANCE TWICE ANNUALLY, BUT relative to their size, bees also fly huge distances — up to ten kilometers — and the foragers may perform such trips hourly. We can experiment with them to find out how they navigate. What we know about bee homing so far is nothing less than astounding, and it is built on a long history of research, primarily pioneered by the imaginative experiments dreamed up and performed by an Austrian named Karl von Frisch and his colleagues that date back over a half-century. Arguably, our knowledge dates back still further to early American frontiersmen trying to find bees’ treasure troves of honey.
In 1782, Hector St. John Crèvecoeur, a writer and farmer from Orange County in New York State, wrote:
After I have done sowing, by way of recreation, I prepare for a week’s jaunt in the woods, not to hunt either the deer or the bear, as my neighbors do, but to catch the more harmless bees … I proceed to such woods as one at a distance from any settlements. I carefully examine whether they abound in large trees, if so, I make a small fire on some flat stones, in a convenient place; on the fire I put some wax; close by this fire, on another stove, I drop honey in distinct drops, which I surround with small quantities of vermillion, laid on the stones; and I retire carefully to watch whether any bees appear. If there are any in the neighborhood, I rest assured that the smell of burnt wax will unavoidably attract them; they will find the honey, for they are fond of preying on that which is not their own; and in their approach they will necessarily tinge themselves with some particles of vermillion, which will adhere long to their bodies. I next fix my compass, to find out their course — and, by the assistance of my watch, I observe how long those are returning which are marked with vermillion. Thus possessed of the course, and, in some measure the distance, which I can easily guess at, I follow the first, and seldom fail of coming to the tree where those republics are lodged. I then mark it [presumably with his name to claim ownership].
James Fenimore Cooper, author of the Leatherstocking Tales of the American frontier, of which The Last of the Mohicans is probably best known, in 1848 published the novel The Oak-Openings; or, The Bee-Hunter. Here Cooper depicts a different, perhaps more reliable method than Crèvecoeur’s of the frontier activity that came to be called “beelining.” Cooper’s story takes place during July 1812, in the “unpeopled forest of Michigan,” where, due to the Native Americans’ lighting periodic fires to clear the ground, there were many flowers among the scattered oaks. This was ideal honeybee habitat, and here the bee hunter Benjamin Boden, nicknamed “Ben Buzz,” practices his art. Ben captures a bee from a flower by placing a glass tumbler over it and sliding his hand underneath. He then places the tumbler with the captured bee on a stump next to a piece of filled honeycomb. He puts his hat over the tumbler and the honeycomb so the bee will not be able to escape. He waits as the bee, stumbling around in the dark, eventually finds the honey. Once it is preoccupied with imbibing the honey, it quits buzzing, and the silence is the signal for Ben to remove the hat and then the glass, as the bee will stay to finish its feast and will fly up, circle the honeycomb, and depart directly toward its nest. He then follows the bee to the tree, chops it down, and is rewarded with just over one hundred kilograms of honey. Easier said than done.
American honey hunters eventually added refinements to their beelining techniques. The main improvement was the invention and use of a “bee box,” a small wooden box designed to catch a bee and get it “drunk” on a hunk of honeycomb. It was used in Maine when I was a kid (I still own mine). George Harold Edgell, a lifelong bee tree hunter from New Hampshire, wrote in 1949 in a pamphlet titled The Bee Hunter that “one’s first task is to catch a bee and paint its tail blue” and “this must be done gently [because] bees do not like to be painted. To paint a bee, it is best to wait until it is eagerly sucking up a thick sugar syrup and is too pre-occupied to notice.”
By 1901 Maurice Maeterlinck, the Belgian playwright and Nobel laureate in literature, described in The Life of the Bee his scientific experiments on bees that were individually identified with daubs of paint, from which he deduced that these insects could communicate their discoveries of food bonanzas to hive mates that would then navigate directly to the food. However, American woodsmen not only had used similar methods, but had also, through their beelining, already gleaned that same surprising insight into what the bees could do. Maeterlinck credited his American predecessors for their discoveries and wrote, “The possession of this faculty [to communicate food locations to hive mates that then can navigate to the food] is so well known to American bee hunters that they trade upon it when engaged in searching for nests.”
Although early American woodsmen, whose lives depended almost directly on the knowledge gained by close contact with nature, were beelining devotees who had deduced that honeybees recruit hive mates, it would remain for Karl von Frisch to unravel the marvelous story of how the bees communicate within the hive. He earned the Nobel Prize in Physiology or Medicine for this work. I feel lucky that a Maine neighbor, Floyd Adams, took me beelining when I was eleven years old, and that when I was a teenager, my father gave me an inspiring little book by von Frisch entitled Bees: Their Vision, Chemical Senses, and Language. It explained the experiments that he and colleagues had performed. They were mesmerizing because they connected the practical experience of beelining in the Maine woods with the imaginative power of a scientist who had penetrated into the core of the bees’ world, their hive, their home.
Floyd’s family’s home was the farm four hundred meters down our dirt road. It was populated by chickens, geese, cows, pigs, plus all the other usual and unusual wildlife that lives in a place with a tolerance for disorder. Along with Floyd, my companions were the four Adams boys, Butchy, Billy, Jimmy, and Robert, an in-law of theirs. Floyd, a dark-haired, mustachioed, wounded Marine Corps veteran recently returned from the Pacific, had a bad limp and a thirst for Black Label beer. Leona, his blond, petite wife, appreciated his fondness for honey but less so his taste for beer. He and the “boys,” after a hot day haying, sometimes went fishing on our nearby Pease Pond in the evening, but in August our big draw was always the beelining.
After we found a bee tree, we carved our initials into the bark to proclaim ownership (property lines were irrelevant with regard to bee trees; finders keepers was the rule), and at some convenient time we returned with crosscut saw, axes, wedges, a beehive, and pails and kettles for honey. Getting part of our living from the land was fun, and it meant understanding and using the bees’ homing behavior to find their hollow trees in the forest and resettling them into a new home, which we brought back to the farm and set up at a window in the attic of the house.
Fast-forward to a quarter-century later: My nephew Charlie Sewall and I are in a patch of goldenrod blooming in a pasture where each fall the wild honeybees gather nectar to top off their honey stores for the coming winter. We start by capturing a single bee in our bee box, a simple four-sided wooden box that has a ten-by-fifteen-centimeter piece of honeycomb with sugar syrup filling out the bottom. We dab the box with a drop of anise for scent and capture our bee by holding the box under her after she has landed on a flower and then slapping the box cover over her. At first the captive buzzes in the box trying to escape, but the buzzing stops when she stumbles onto the sugar syrup and starts to tank up, which will take her a minute or two. We then remove the cover and set the open box onto a pole that reaches to just above the tips of the goldenrod. We gently daub her with a spot of paint while she is absorbed in sucking up syrup, as I remembered Floyd doing. We then hunker down into the goldenrod and wait as she continues sucking up her newfound sweets that she will soon share with her hive mates. After about two minutes, her honey stomach is filled. She crawls out onto the edge of the box, stops to wipe her antennae with her front feet, lifts off, and flies back and forth downwind of the box. We duck lower to keep her silhouetted in sight against the sky as she starts flying loops, which become increasingly wider and oriented in one direction. Finally she straightens her flight path and takes off, making a “beeline” into the distance. Knowing that nobody in that direction keeps bees, it’s clear that she is on her way to a bee tree. She will soon be back with others, and we then consult our wristwatches to time her trip. A bee flies about four hundred meters a minute, and it may take her three to six minutes in the hive to regurgitate and unload her honey stomach’s contents into the mouths of begging, receiving bees.
We settle down and wait, and after perhaps ten minutes or less a bee suddenly appears and makes very rapid zigzagging flights just downwind of the box. The sound of her fight has a higher pitch than that of the bees foraging on the nearby goldenrod flowers. This means that she is more motivated and has a higher body temperature because of the rich food she is expecting. She settles into the box and starts imbibing the syrup. More bees will come soon, and when they get near our bee box, they will be guided in by the scent of the anise that marks the spot. After they tank up, we watch their flight directions.
If the food is in the immediate home vicinity, the bee does a “round dance” on the honeycomb when she returns to her home. She repeatedly runs in small circles while shaking her abdomen, and she regurgitates small samples of her find at intervals during her dance. If they become motivated after receiving information about the quality and scent of the food advertised, her hive mates leave the hive and search for the advertised food. If it is beyond a few hundred meters, the bee alters her dance to also contain information concerning location. The distance of the journey to the food is proportional to the duration of the waggle runs, and the angle of the straight runs with respect to the vertical direction informs the bees in what direction to fly when they leave the hive. If the straight run is in the up-direction on the honeycombs (which always hang vertically in the hive), the food source is in the direction toward the sun. If the food location is, for example, at an angle of ten degrees to the right of the vertical, the food direction is ten degrees to the right of the horizontal component of the sun direction when the bee would fly from the hive. Thus, her behavior is a symbolic representation in body movements of the flight to the food.
The first steps in the evolution of recruitment likely involved simple alerting signals in or at the nest entrance before takeoff. Other bees could have followed those signaling bees, probably by scent, for at least a short distance in flight. Through a few million years, the alerting likely became modified to take on an ever-greater leading function by bees flying in an ever more conspicuous manner in the direction of the food, so that followers could start off flying with ever greater accuracy in the right direction. These flights, later in the evolutionary progression, were eventually restricted to a buzz run directly on the top of the combs, but still in the food direction. We can infer this, because such “primitive” recruitment is still found in some tropical honeybee species that have their combs in the open air, where this mechanism makes sense. But open-air homes, though convenient for such communication of food location, were vulnerable to predators and also precluded the bees from living in huge areas of the globe, those with cold climates.
Homes in hollow trees allowed the bees to live in areas where they would otherwise be excluded because of cold and/or nest predators. But in such safer homes the combs hung from the roof of a cavity and left no horizontal dancing platform, and additionally the “dance floor” was now in darkness, so bees could not point directly toward the food. Even if they could, they would not be seen. But a breakthrough for indicating horizontal directions on vertical surfaces became possible after some bees started using the hanging flat surfaces of combs as their dancing platform while indicating the sun’s location as the up or “toward” direction in their dance. Additionally, tactile rather than visual orientation became predominant for recruits in reading the code within the nest.
It is amazing enough for an animal to be able to navigate to a location it has never been to before. But some ants do something even more amazing. In North Africa, desert ants live in underground homes where they are protected from the heat. But they must venture out onto the searing surface periodically to forage by scavenging on heat-killed prey. The ants are fast runners that have evolved a very high tolerance for heat. Still, at times it is a matter of life and death even for them to make it back to their cool underground home; they cannot afford to wander on the sand surface for an extended time without access to their shelter to cool down and replenish body fluids. This is where their homing ability comes in; they may have zigzagged in all directions to find a heat-killed insect, but after finding one they must make a straight “ant line” directly back home. This begs the question, Since they are often on a featureless plain and have not kept a steady course, how do they know in what direction to head home?
If one captures bees in one pasture and releases them in another, they usually depart in the direction they would have flown from the original field. That is, they act as one would expect if they do not realize that they have been moved to a new location. Rüdiger Wehner and his colleagues at the University of Zurich came to the same conclusion about desert ant homing in their lifelong experimental studies. The ants use the sun as a compass, but a compass is not enough; the ants, when released from a point they had not themselves traveled to, like the bees caught in one pasture and released in another, apparently got lost.
For homing you must know where you are on “the map” before you head off in the correct direction. The desert ants can return home, but only if they walk to where they find themselves. Wehner concluded that the ants’ homing mechanism involves somehow calculating where they are at all times, probably in measuring distance by keeping a kind of count of their steps, and also keeping track of the angles of their direction from their home relative to the sun’s location. These were not mere speculations, but a hypothesis tested in painstaking experiments that entailed altering the ants’ perception of the sun (holding filters over them that varied the direction of polarized light that they, like bees, use in orientation) and altering their stride length (altering their leg length by gluing on extensions) to find out what information they valued and how they used it. Presumably bees could also have a similar “map sense,” and Randolf Menzel, a neurobiologist in Berlin, was trying to find out how it might work.
Menzel runs the large and active Institute of Neurobiology at the Free University of Berlin, and one of his projects was the burning question of how honeybees seem to find out where they are in order to be able to go where they want to be. Honeybees are suitable animals with which to study this problem because, like ants, you can count on their motivation to return home after they are loaded with food.
We can’t look into a bee’s brain and determine what it knows and what it wants. However, clever experiments based on the bee’s natural history permit inferences. We can determine, for instance, where a bee perceives herself to be relative to her hive. If a bee regularly visits a feeding place, she knows where she is, because she always flies off in a straight line from it back to her hive. If we then remove either the hive or the feeding spot, she circles in the area where her target had been. We know what she is looking for, because when we provide the hive and/or the feeding station within the area where she circles, she quickly finds it. But suppose we capture our bee at the usual feeding station after she tanks up on honey or syrup, put her into a dark box, and then carry her “blind” to a place she has never been. As mentioned, most bees will then make a beeline in the same direction they had normally flown to return to the hive. They will fly as far as before but find no hive there. Yet, they usually eventually do make it back home. How do they find their way? What do they do until they reach home? Until Menzel’s experiments, it had not been possible to track them in flight when they were out of sight out in the field. Menzel had a tool — radar but with a unique twist — whereby he could trace bees’ actual flight paths over a kilometer away by radar and record them on a computer. And he invited me to come see the work in progress.
Bee flight paths. A. A bee’s first trip from a flower patch or bee box back to its bee tree (hive) begins with an orientation flight. B. Later trips are more direct. C. After a bee has been transferred while “blindfolded” to a new spot, she acts as though she perceives herself to be still at the same place as before.
The problem of tracking small objects such as insects from a long distance by radar had always been that radar would “see” too much. You could not isolate and then plot a single specific bee out of all the extraneous noise of echoes bouncing off all objects. The new insect-tracking radar technique started in 1999, when Joe Riley, a British researcher, applied a radar system able to track very small objects over long distances by attaching to the insect a small device that, after receiving the energy of an electromagnetic sound pulse, would respond with a frequency other than that of the transmitted ultrasound. The receiver is then tuned to amplify only that frequency. In this way, it became possible to track the flight paths of individual preselected bees equipped with the appropriate transponders because the echoes from all other objects were filtered out.
The Menzel group’s electronics technician, Uwe Greggers, adopted the Riley system in 1999 and 2001 and got interesting results, but then ran into software problems. Nevertheless, given the promise from the data they did get, the scientists contacted a radar specialist at Emden (north Germany) who agreed to develop the system. The Menzel group then needed to find the right site in which to use it. They needed to locate the experiments at a large flat area devoid of trees in order to be able to record the complete flight paths without interference such as the bees’ trying to avoid objects or being attracted to them. The closest suitable area was an expanse of marshy meadow about a two-hour drive from Berlin. The large, idyllic farmstead near the village of Klein Lübben and land associated with it had accommodations for seven or more helpers, making this site amenable.
One Menzel group experiment in the works when I visited involved training individual bees to expect food at two widely separated feeding stations, but only one station at a time was open to them. I had no idea what to expect, and on my day with the team I was eager not only to watch the bees but also to see the experiment in action.
It was early in the morning when Menzel picked up Greggers and me for our trip to the experiment site in the Brandenburg countryside. We loaded a large, heavy printer that would be used to handle the large-scale printouts of flight paths, and then we were off down the Autobahn. Two hours later we arrived at Klein Lübben, a quiet village of farmsteads that at least in outward appearance has changed little since medieval times. The fields were several kilometers square, flat, and moist — perfect also for frogs, and hence storks which nest there in baskets attached to the tops of red-tiled house roofs. Swarms of starlings swirled through the air, and a pair of white swans paddled serenely down a canal along a dirt road, followed by a line of five still-downy gray cygnets.
At one end of the study field stood a steadily turning radar apparatus with a large round antenna for sending out the signal. A smaller dish antenna mounted directly above it would receive the transformed signal bouncing off the transponder on an airborne bee in the field. On the field sat two blue triangular tents and three yellow ones. They were experimental landmarks for bees that could be made available to them, to find out if they used them, and manipulated for experiments by changing their locations. In the distance sat a beehive, and I noticed a man running from it. He was wildly slapping himself, in an obviously defensive mode. He had been assigned to provide food for the bees close to the hive and then was to gradually move the feeder into the field so that a population of bees from that hive would be available for us to study when we arrived at midmorning. He had come too close to the hive, and at that moment it was he who was getting dispersed over the field, not the bees. Also, as we soon found out, there were no bees coming to the two feeder stations, as they were supposed to have been by now; the student had apparently overslept or been otherwise distracted from his assigned job of luring bees.
The experiment we wanted to do was in doubt. This was serious. Two hundred thousand Euros had already been spent on this study, and the boss was intolerant of negligence. Luckily, bees from hives used previously for another experiment were still coming into the field to search for feeders. He could let some of those bees find the feeders and then train them to come back to specific locations.
For our experiment we needed to establish two feeding stations, A and B, separated by about three hundred meters. Certain bees were already keyed into the routine. When I walked across the field, one bee started following me. It looked most extraordinary: it had a lot of blue and green color, not just the usual plain brown honeybee attire. As soon as Menzel’s helper and I set up our feeder, this specific bee landed on it and immediately started to suck up the rich sugar solution. Now I could examine her more closely: the green was a plastic tag with the number 29 on it that had been glued to her thorax. The blue was a slash of paint that had been daubed onto her abdomen.
Within a few minutes an assembly of several differently color-coded bees was lined up around the edge of the syrup dish. All were sucking up syrup. Some had green on the thorax, some had blue, and still others had yellow tags on their thoraxes, with additional daubs of white, blue, or yellow paint on their abdomens. Uwe Greggers and the unfortunate helper immediately started logging a list of the bees that had shown up in a notebook.
Each bee tanked up quickly, flew off directly toward her hive at the other end of the field, and then came right back to take a next load. Newly recruited (unmarked) individuals were also coming every minute to our site A. At the second feeder (site B) there was a similar flurry of activity, except it involved different individuals.
Menzel then instructed us to move our food station A one hundred meters closer to the second one, B. Bee numbers 29 and 30 green, both with blue tails, number 2 yellow with white tail, and number 39 green with red tail (who had all been present at A) then almost immediately started showing up at B, the new location. When crowds of bees had done the same, we removed one station and put the remaining one into the middle, between the two original sites. Next we moved our feeder to site B. Most of the bees, such as 30 green with blue and 39 green with red, who had been at the previous site, showed up. That is, we had trained bees who had been at one site to come to the second site, so we knew they now knew two sites and could potentially use either as a reference site to return home.
For the planned experiment it was important that the bees forget the intermediate sites that had been instrumental in getting them to go to the two widely separated sites. So, for the rest of the day, we alternately fed the bees, first at site A, then at site B, and monitored which individuals were showing up at both sites (most of the individuals continued to forage at either one or the other site).
We were now, near the end of the day, finally ready to move from training to trials. The experimental plan was to select one of the bees who knew both sites. This bee would, after feeding at one site and getting ready to leave, be captured in a dark box and thus “blindfolded” and then brought to a third feeding site where she had never been before. Here she would be released after being equipped with a radar-tracking transponder. We presumed that she would do at least one of three things: she might recognize where she was and fly straight home; she might instead fly off in her original (now wrong) direction; or she might immediately know that she was at an unknown location and search until she found one of her two feeding sites and from there take a direct beeline home. Knowing her exact flight path would allow us to distinguish among the alternatives, which would be essential to ultimately decoding her homing mechanism. Setting up this experiment had taken a long time, but I would now, possibly, be treated to an exciting demonstration of bee homing, one I could never have imagined possible.
Menzel picked up his walkie-talkie to call the radar station: “Mike, we’re now going to put a transponder on a bee — are you ready?” Mike had spent some years in the army where he was trained on radar, and he was now working part-time while getting a university degree in computer technology. He replied yes, he was ready. Menzel then took me to feeding station A, where a whole lineup of bees was coming and going.
“Which one do you want?” Menzel asked me. I wanted a bee that I had gotten to know over the course of the day, so I chose 39 green with red-tipped abdomen. We waited for her to arrive and let her feed for a while. As planned, Menzel then held a glass vial over her while she was distracted sucking up syrup. When she was full, she walked up into the vial, and Menzel corked it shut and darkened it by wrapping his hand around it. We then took her to a site distant from both feeders, a place where she had not previously fed and from where we would now release her.
The vial holding “39 green” had a plunger at the bottom with a wide-mesh screen at the top. Menzel gently pushed the plunger in and forced the bee up against the screen, held her there, and picked up a tiny transponder (a wire holding a diode with a sticky pad at one end). With fine tweezers, he deftly removed the protective paper from the sticky pad and glued the transponder onto the top of the bee’s thorax. “Ready?” he radioed Mike.
“OK.”
Menzel removed the plunger and held the vial with the open end up, for the bee to crawl up. She hesitated at the lip of the glass, groomed her antennae, and then lifted off. She showed no strain in flight. (The transponder’s weight is twenty milligrams, and a bee can fly with double her body weight, carrying a hundred-milligram load of nectar in her honey stomach plus two pollen packets on her hind legs.) However, she flew only two or three meters before dropping down into the grass, stopping to preen herself some more. But a couple of minutes later, she finally took off again. Mike, who was now monitoring her flight, radioed us. At intervals we heard: “She is heading south-north-east-north-west-south.” Then, finally, Mike continued: “Now her path is straightening out — now she is heading directly for her hive!”
She had suddenly oriented correctly. This was the crucial point: she had apparently recognized something that had “placed her on the map,” so that she then “knew” in what direction to fly to reach home. Assuming she had taken a path she had never taken before, did her successful homing suggest a “map sense”?
I ran over to the radar tent where Mike showed me the radar screen and the dots where the three-second successive readings traced the bee’s path. A computer screen, where software had converted the time and directions of the bee’s flight path into different-colored images for easy reading, showed that the bee’s original flight direction was toward where the hive would have been had we not moved her from her feeding spot. In other words, she acted as though she didn’t know where she was when we released her. As expected, however, after she reached the area where her hive would have been, she flew loops in several directions. Then, after she had flown ever-farther away from both her real and “would-have-been” hive locations, she suddenly seemed to orient and flew directly toward the hive. Amazingly, it was along a route that had not been her normal foraging route from her two feeding sites. Had she perhaps seen a blue or a yellow tent and, having learned their relationship to each other during previous orientation flights, transposed that information to fix her new location? Only more bees could tell.
Other bee homing experiments with hundreds of bees were ongoing. And in the group’s final publication two years later, the thirteen-author research team headed by Menzel concluded that honeybees incorporate information for flight direction from both their previously learned flights as well as landmarks and from the flight directions learned from hive mates within the hive. But they can redirect their flight vectors to and from the hive and perform novel shortcut flights between the learned and the communicated vectors.
“The” homing instinct, recognized and traded on by every American beeliner to get honey, and used by von Frisch to decipher the bee language, is a source of fascination and mystery still. Von Frisch had likened it to a “magic well” from which the more you take, the more runs back in. The “well” is still doing that, three-quarters of a century after his prophetic pronouncement.
GETTING TO A GOOD PLACE (#ulink_a1addad7-22a9-51d5-ad06-01f378599df8)
THE TENT CATERPILLAR MOTH, MALACOSOMA AMERICANUM, is common in North America. It emerges from its light yellow silk cocoon in late summer, and the female is then ready to deposit her batch of over a hundred eggs. She searches for an apple or a cherry tree, and somewhere out on a thin twig of just the right diameter — about a half centimeter — she exudes her eggs along with sticky foam to form her egg mass into a ring that wraps around the twig. The foam dries and hardens, encasing the clutch of eggs and gluing them solidly to the branch where they stay through the coming winter. But the larvae develop inside the eggs during the summer and, while confined in their eggs through the winter, hatch at almost precisely the day, about nine months after the egg-laying, when their tree breaks its buds.
The moth is named for the conspicuous communal homes of silk, called “tents,” that its caterpillars make, and in the spring of 2013 I found a just-made tent on a young black cherry tree next to my Maine cabin. Like nearly everyone else in this part of the country, I was long familiar with these caterpillars but had not deemed them worthy of a closer look. The tents act, I learned, like miniature greenhouses and warm the new caterpillars at a time when nightly frosts are still common. But, despite its advantages, to have any home is to incur costs: it has to be made, and it takes time, energy, and expertise to make, and the wherewithal to travel to and from it. For the time being, I wanted to know where the caterpillars making this home had come from. To my surprise, the ring on the twig with the now-emptied eggs I was looking for was almost a meter from the tent. How had the many hatchling caterpillars “decided” or been able to stay together and then coordinate to make their tent? Squinting against the sun, I could see a glistening trail of fine silk leading from the emptied egg-case ring to their home, so here was at least a hint as to how they crawl together to end up at the same place.
On the second day after I found the tent, May 1, there was still snow on the ground in the woods. There was as yet no sign of fresh green anywhere. But I wrote in my journal, “Black cherry buds ready to pop leaves.” These trees are the first to leaf out, and the caterpillars could not have fed yet. What would they do? An hour after the sun came up, the tiny caterpillars emerged from their tent and massed on its sunny side. An hour later they started milling about, and then a few started crawling, seemingly aimlessly, several centimeters up and down the trunk and branches of the cherry tree.
As I had anticipated, some of the tiny caterpillars started to crawl back onto the same branch they had come from, possibly following their previously made silk trail. But they went only six centimeters before turning back. Others went down the trunk of the tree. Always some would turn back, and then the others followed one behind the other in a line. Finally, by 7:30 a.m., a contingent of about twenty of them had progressed nine centimeters down the tree trunk, although two were coming back up. Then more started to leave the tent, and eventually all were in one long line, going only down the trunk and then angling up another branch. In half an hour the leaders had traveled seventy-three centimeters and reached a bud. The rest were strung out all the way to the tent, but their two other travel-direction options had been abandoned. All were eventually massed at the same cherry bud, three-quarters of a meter from their tent, and in an hour and a half they had all returned to their tent, one following the other in a long train.
The young black cherry tree showing relative locations of a tent caterpillar moth egg cluster (C) from which the clutch of just-hatched caterpillars emerged and traveled to start making their home (H) in a crotch of the tree, and their first travels as a group (T) to feeding places
At noon they came out and crawled onto the outside of their tent, waving their heads back and forth, apparently weaving silk from their salivary glands to enlarge it. Another hour later they were again all massed inside the tent and perched, immobile, tightly against the bark, where they were barely visible through the thin gossamer veil of silk.
The caterpillars stayed in their tent through the night, and I expected them to go at sunup to the same branch where they had been the day before. But instead, this time they all followed an entirely different path, going directly up the tree instead of down as on the previous day, and without taking another side branch. I could not detect any silk on their so-far two different foraging trails, and this time they went even farther — a distance of 130 centimeters. After their one meal the day before, they were already noticeably larger. A few were the same size as the day before, but most had probably doubled in weight. There were many tiny fecal droplets in their web. So they had fed, even though it seemed hardly possible that they had anything to feed on at the barely opening bud.
On the third day the buds had opened and the tree was replete with new small leaves pushing out of the buds. But it had been a cool night — there was again frost on the ground at dawn — and the caterpillars made a slow start.
The pattern soon became clear: the caterpillars spent most of the night and most of the day when they were not feeding in their home. The time spent on tree branches was brief, and it could not have been just to keep warm that they stayed in their home because they went back inside just as quickly after feeding regardless of temperature or time of day.
Having found and watched the caterpillars of one tent, I then observed others for more clues to their homing behavior. One of the surprises to me was that as they grew larger, they foraged independently of one another, no longer going to and from feeding areas in groups. Furthermore, after they were about half grown they left their tents, not to return at all but still to continue feeding before eventually searching for a spot in which to spin their flimsy cocoon. Tent caterpillars usually choose a bark crevice to pupate, although commonly they also choose the cracks in the sides of buildings. But why were the young caterpillars strongly homebound and the older ones not?
I suspect the young ones’ web-making behavior may have evolved in part as an anti-predator response. The tents were visited by red wood ants, Formica rufa, and right after the caterpillars hatched, these ants often loitered alongside them on their trails. I tore a nest open on one side to find out if it served as protection. It must have, because ants entered, though frequently wiping their antennae as though irritated. Nevertheless they tarried inside the damaged nest, and I saw them grab and walk off with caterpillars. No ants entered an intact nest of the several I watched, each of which consisted of several successive layers of silk. Thus, the webbing of the tent acts as a deterrent to predators such as ants. Staying inside the home most of the day and night, as these caterpillars appear to do when they are small, probably reduces mortality from parasitic flies and ichneumon wasps as well. When they are larger, the caterpillars are probably protected from the ants, as well as from most birds, by a layer of fine spines. They pupate without having to bury themselves to escape frost, because the adult emerges long before there is any frost.
Because these caterpillars are protected from predators in the summer homes they build and by the spines they wear, because they mature early enough in the summer for the pupae to avoid the cold of winter (by early emergence of the moth), and because the eggs and young larvae are immune to freezing because of the antifreeze they contain, “everything” in the life of the tent caterpillar moth may be found within a few meters. The adults that emerge in late June are not far from the apple or cherry trees where the parent left her eggs, and their life cycle can be completed without their having to go far from home, unlike some other insects which traverse a continent to be able to satisfy all their needs.
Monarchs. Of all the insects, the travels of the monarch butterfly, Danaus plexippus, are perhaps most famously spectacular in both scale and scope. Dr. Lincoln P. Brower of the University of Florida in Gainesville (now at Greenbriar College), who has studied this butterfly and its migration for over forty years, records the rich history of the emergence of our knowledge of monarch migrations. Early naturalists saw “immense swarms” in the prairie states where the caterpillars fed on the leaves of the many native species of milkweed (Asclepias) and the adults fed on the nectar of their flowers. Monarchs declined when later industrial agriculture destroyed many of their food plants, but in the nineteenth century they resurged in the East due to land clearing and the spread mainly of one milkweed, A. syriaca. Millions of them were seen passing for hours, even in Boston. This was a phenomenon that is hard to imagine now and it ignited much interest then. Charles Valentine Riley, the entomologist who first hypothesized that these butterflies engaged in a birdlike migration, cites people seeing them in the fall in swarms that extended for kilometers and obscured the sun, “blurring day into night.” Huge lines of them passing Boston in 1880 were described as “almost beyond belief.” Now, with reforestation, plowing, and then the use of Roundup and other weed killers that eliminated their food plants in agricultural fields, the monarch is but a shadow of what it was. In the past several years in the East, it seems to have almost disappeared. For the first time, I saw not a single one in late summer of 2013. But our knowledge of the scope of the monarch migration has blossomed.
Monarchs migrate on their own power for thousands of kilometers, and, unlike many other insect migrants, the population (though not the individuals) has a regular two-way migration, although as with the other insect migrants, the individuals that come back are not the same ones that left.
Unlike most of the other North American butterflies and moths, which overwinter in New England as eggs, larvae, pupae, or adults, monarchs cannot survive there through the winter in any stage. The population that normally now graces fields all along eastern North America overwinters at around three thousand meters’ elevation in dense fir groves on the southwest slopes of volcanic mountains thousands of kilometers to the southwest, near Mexico City. The monarchs find shelter in those fir stands from rain, hail, and occasional snow. It is not cold enough for the butterflies to freeze there, but it is cool enough for them to conserve the energy resources that they have accumulated on their way south.
The monarch butterfly adult, caterpillar, and chrysalis
In the summer, the monarchs fly in what look like random zigzag patterns over the New England fields as they stop here and there to sip nectar. Occasionally you see a mated pair, the female doing the work of flying, the male dangling passively with folded wings while attached by his genitals. After the prolonged mating (and/or technically “mate guarding,” since it prevents mating by other males), the female glues her delicately patterned green eggs with gold markings, one at a time, to the undersides of milkweed plants. In a few days, the flashy yellow-black-white larvae hatch and start chomping. After about fifteen days (depending on the temperature), the caterpillars have increased their weight to 1.5 grams (2,780 times the hatchling weight). The caterpillar attaches itself to a support such as the underside of a leaf by a clasping organ at the hind end of its abdomen to hang upside down. It will then molt into the bright green pupa (chrysalis) with the shiny golden spots that is surely familiar to almost all school kids. In a few days, the chrysalis starts to turn dark, and the outlines of the orange-patterned wings are visible through the now-transparent cuticle. When the chrysalis splits, along a predetermined line of weakness in the back, the limp adult slips out and expands its wings, and in two or three hours hormones will have instigated a biochemical process that hardens its body armor and stiffens its wings. The butterfly is ready to fly. Where will its wings take it?
Thanks to the monarch studies initiated in 1935 by Dr. Fred A. Urquhart and his wife, Norah Urquhart, from the Zoology Department of the University of Toronto and continued to the present day with the input and cooperation of thousands of amateur volunteers, there is now an amazing story to tell. The Urquharts noted in the late 1930s that the monarchs they saw in late May and early June in Canada had tattered wings, and they knew that this species would not and could not overwinter in Canada, so they suspected that they may have come a very long way. Monarchs fly in a southwesterly direction in the fall, but nobody had a clue where they ended up. To get some idea of the butterflies’ movements, these researchers in 1937 began gluing paper tags onto monarch wings with this message: “Please send to Zoology University Toronto Canada.” Monarchs weigh almost half a gram and the wing tags only 0.01 gram, so the tags were not likely to hamper the animals’ movements. Similar tags, used today, have pressure-adhesive backing and can be folded in half and glued over the leading edge of the forewing (after the scales are removed).
The idea from the inception of the monarch-marking studies was to try to find out if the butterflies migrated — an idea that at the time, as Urquhart noted, “was considered quite impossible.” But the question of where the butterflies might be going to and coming from grabbed the imagination, and anyone seeing a tagged butterfly would be sure to try to catch it. Sure enough, tags were returned over decades that suggested a migratory pattern. Individual tags were returned from huge distances, up to 1,288 kilometers. One monarch that was tagged in Ontario in 1957 was recovered eighteen days later in Atlanta, Georgia, 1,184 air kilometers distant. Clearly, when the butterflies left Canada in the fall, they headed south.
Still, nobody knew what happened to the mass of butterflies. Then, in January 1975, Cathy and Ken Brugger of Mexico City found them — a dazzling, shimmering, orange display of an estimated 22.5 million monarch butterflies on one 2.2-hectare site (which turned out to be only one of ultimately thirteen overwintering sites in the mountains of Mexico). The millions of monarchs were festooned in the trees in the mountains of Michoacán near Mexico City. The Urquharts excitedly traveled to see the site and on January 18, 1976, listened to “the sound of the fluttering of thousands of wings [that was] like that of a distant waterfall.” As they stood awestruck by this dazzling display, a pine branch broke off from the sheer weight of butterflies attached to it, and it crashed to the ground right in front of them. Fred Urquhart had been posing for a National Geographic photographer surrounded by these just-fallen butterflies when, incredibly, he saw a tagged one among them. When he traced its origin, he learned that it had been tagged on September 6, 1975, by Jim Gilbert, from Chaska, Minnesota. Urquhart, who had encountered countless tagged butterflies in his career, said it was “the most exciting one I have ever experienced.”
The picture that has now emerged from decades of study is that individual butterflies migrate all the way from Ontario to Mexico in the fall, arriving there at their overwintering sites in a torrent during October. They spend most of the winter in Mexico in a cooled low-energy state but soar around on warm days to drink water and replenish on nectar. In early spring, when their sex urge awakens, there is a mating orgy followed by a mass exodus. Most of the females mate before leaving, and their “compasses,” which were set to take them south in late fall, are now “reset” to take them in a northerly direction.
As the tide of butterflies advances northward, the females stop to lay their eggs on milkweed. Some of the butterflies from Mexico make it all the way to the north, and others (their offspring) that grow from the eggs laid along the way arrive later. Those of the first generation have slightly tattered wings when they arrive in the north, while those that arrive later have untattered wings. (However, not all monarch populations migrate, and not all that do, travel in the same directions as the populations of northeastern North America.)
One of the mysteries that puzzled Fred Urquhart was how the butterflies home. In Urquhart’s 1987 book on the monarch, he speculated that the butterflies perhaps use the Earth’s magnetic lines of force, although different populations of the butterfly migrate in different directions, so they could not all be orienting to it in the same way.
A potentially even more puzzling question is the ultimate (evolutionary) one of why these butterflies migrate in the first place. Urquhart simply suggested what he admitted was a “perhaps far-fetched” idea: that “twice each year it [Earth] passes through an area rich in some sort of radiation that could impinge upon animal life [that] might affect in some manner the cells of the body causing reproductive organs to abort in the fall and develop in the spring and initiate the migratory response.” This is an unlikely theory, though, mostly because it depends on a mechanism that is not adaptive in evolutionary terms. Instead, more current thinking about the adaptive reason why the phenomenon has evolved focuses on energy economy and maximization of resource use under the expected evolutionary constraints from the monarch’s having evolved in the tropics, meaning it was not able to survive northern winters. (Monarchs belong to the family Danaidae, an otherwise strictly tropical group.) Migration to the north in the spring opens up the milkweed crop over a major swath of North America as a food base for the larvae. In addition, the journey is probably not costly to the monarchs, either in terms of predation (since they are chemically protected from predation by poisons they sequester from their food plants) or in terms of energy costs, since their energy intake along the way more than makes up for the energy expended for travel. Indeed, unlike most birds that may deplete all their fat reserves on migration, these butterflies instead fatten up on their journey and may consist of about 50 percent body fat by the time they arrive in Mexico, where their overwintering fast begins.
Butterflies and moths experience tremendous selective pressure, and undoubtedly there are constant readjustments of survival strategies. Weather affects the populations, not only through flight activity and flight range as well as growth rates of larvae, but perhaps also indirectly by influencing virus infections. But Urquhart noted that each female monarch butterfly lays up to seven hundred eggs, and he calculated that the “biotic potential” — the number of individuals if there are no deaths — of one female after only four generations (that is, at the end of one summer) is 30,012,500,000 adults. Luckily for the planet, animals’ reproductive potentials are never naturally realized, for long. The limit is quickly reached when the population uses up its food base, in this case milkweed. In some years a virus decimates most of the monarch population over North America, but then several years later it rebounds. But the population cannot rebound from some things: in recent years there have been massive declines of the monarch population that cannot be reversed, because they are due to unnatural causes — the massive conversion of land to crops, and the introduction of genetically modified crops that tolerate herbicides, which have allowed the elimination of milkweed that formerly grew between rows of corn.
The flight performance of monarchs is spectacular, but like the hordes of cluster flies from the surrounding fields and woods that overwinter in my cabin, they are traveling to a specific place for overwintering where they have never been before. Such homing movements are diverse, but common. Robert D. Stevenson and William A. Haber of the University of Massachusetts, Boston, found a regular seasonal migration of about eighty percent (250 species) of butterflies living in the dry lowlands of the Pacific Slope of Costa Rica that migrate to wetter forests of the east. Distances traveled range from ten to a hundred kilometers.
In North America as well as in Europe, the cosmopolitan painted lady, Vanessa cardui, a mostly orange and black butterfly with white spots and pink and blue “eyes” on its under-wings, at times appears in large numbers and then is not seen again for years. Usually the individuals are seen crossing a road, and almost all will be heading in the same direction. The painted lady regularly migrates north from Mexico, from where it originates, after heavy rains in the deserts have created an abundance of food plants, primarily thistles. A friend told me of one migration while he was in Arizona when his windshield wipers “soon became useless” because of the huge numbers of painted ladies plastered onto them as he was driving. I see them regularly in Vermont and Maine, but seldom in large numbers (the summer of 2012 was one of the exceptions).
Red admiral butterfly larva, adult, and chrysalis. The larva makes a shelter for itself by pulling leaves together and holding them with silk, while then feeding on the leaf.
One of the butterflies that not only migrates as an adult but also hibernates in some parts of its range is the red admiral, Vanessa atalanta. It is (as are all butterflies!) beautifully colored. It sports a wide red stripe across each dark forewing ornamented with white spots, and its larvae feed on nettles. I wrote in my journal on May 11, 1985, near my home in Vermont: “In the afternoon from around 2:30 to 4:30 PM, as I was jogging along on an 18-mile circular loop I counted 512 red admiral, crossing the road in front of me. All but 5 of these were flying in a northeasterly direction. At 5:00 PM, after I was home, I take compass readings of butterflies flying over a plowed field where they funnel onto it through a valley. I can see them to take a bearing for at least 50 paces — 250 feet. All 22 that I observed flew in NE direction. At 6:00 PM activity almost stopped. The breeze is slight, from northwest.” In the summer of 2001 and again in the spring of 2010 I saw large numbers of red admirals. They fed on freshly opened apple blossoms, and later all the nettle plants in a neighbors’ sheep pasture had an abundance of their caterpillars.
Moth migrations are perhaps more spectacular than those of butterflies. Jason W. Chapman and colleagues report one recent ten-year study involving radar tracking of about one hundred thousand owlet (Noctuid) moths, primarily the silver Y moth, Autographa gamma, migrating south in the fall from northern Europe, and then north from the Mediterranean in the spring. Like the butterflies, these insects breed along their migration route. Also like the butterflies, the moths partially correct for crosswinds, to maintain specific directions. Most surprising perhaps is the moths’ windsurfing; they choose the most favorable wind currents corresponding to their respective spring or fall migratory directions. If the wind shifts about twenty degrees from the favorable direction, they adjust their flight to accommodate and maintain the correct direction. If the wind shifts ninety degrees, though, they stop and wait for a favorable wind. Millions of them fly together in the dark of night, and, like the monarchs’, their compass directions are likely tuned to the Earth’s magnetic fields. Some studies of radio-tagged green darner dragonflies, Anax junius, suggest that these insects also migrate hundreds to thousands of kilometers from north to south with those that return being a different generation.
These behaviors get the animals to a good place (for overwintering or for reproduction). Like the long-range movements with specific endpoints on the map, homing to a good place is not always easily distinguished from moving out of a bad place. The behavior is a mechanism with deep evolutionary roots. Indeed, insect wings (and metamorphosis) may themselves have been an original adaptation for dispersal, to colonize temporary pools, animal carcasses, or other temporary resources. The first individuals to reach the resource won the competition to use it and multiply there, and these were more likely to be the ones that flew, and flew far and wide, rather than those that walked at random.
Wings and metamorphosis have lesser value in constant conditions. Some insects are able to respond in real time to the changes in conditions they experience (especially crowding), in that when they don’t “need” to disperse they either don’t grow wings (some aphids) or the muscles to power the wings are broken down and the amino acids from the protein are used instead to make more eggs (in some Hemiptera bugs). Often there are discrete “dispersers” versus “non-dispersers” in any given insect population, and the percentage of each depends on the quality of the home habitat and hence the relative cost/benefit ratio of moving versus staying.
Dispersing to “anywhere but here” generally applies to nonmigratory species that have no encoded or learned directions to go to but may have innate instructions to move in more-or-less straight lines rather than potentially going in circles in order to achieve distance. In Africa, dung-ball-rolling scarab beetles race away from their often thousands of competitors at a dung pile at night by using the swath of stars of the Milky Way galaxy as a reference. Swarms of insects feeding at dung and carcasses also attract predators, and as soon as they finish feeding, many distance themselves from those predators. I’ve observed blowfly larvae at animal carcasses keeping to almost perfectly straight lines in their getaway at dawn, by steering directly toward the direction of the rising sun. Mass movements sometimes observed in some rodents, such as lemmings and gray squirrels (as in 1935 in New England) following a population explosion after a superabundance of food, may be another example of dispersal to get to a better place, though not necessarily a predetermined one.
On the other hand, “true” migrants are able to utilize ideal conditions in two places, provided they vary predictably. Arctic terns, Sterna paradisaea, breed throughout the Arctic, then fly to Antarctica to escape winter when food availability declines and to arrive in spring and food again, a round-trip distance of nearly seventy-one thousand kilometers. Gray whales, Eschrichtius robustus, also feed in the Arctic in the summer but then travel eight thousand kilometers along the coastline to Mexico to bear their calves in warm waters.
