The Anthill Chronicles
19
IT WAS TRUE. The Trailhead Queen was dead.
In the first days there had been no overt sign that her long life had ended. There was no fever, there were no spasms, no farewells. She simply sat on the floor of the royal chamber and quietly died. As in life, her body was prone and immobile, her legs and antennae relaxed. Her stillness by itself failed to give warning to her daughters that a catastrophe had occurred for all of them. She lay there in fact as though nothing had happened. She had become a perfect statue of herself.
The deception was the result of the way the bodies of insects decay after death. Where humans and other vertebrate animals have an internal skeleton surrounded by soft tissue that quickly rots away, insects are encased in an external skeleton. Their soft tissues shrivel inwardly into dry threads and lumps, but their exoskeleton around them remains, a knight's armor fully intact long after the knight is gone.
Hence the workers were at first unaware of this mother's death. Her quietude said nothing, and the odors of her life, still rising from her, signaled, I remain among you.
She smelled alive.
The deception was made easier because in life she had never given orders or led them in activities of any kind--even though her brain was fully programmed to perform all of their tasks if she chose.
She was to all purposes a winged wasp who lived in a neutered wingless society. Yet the only initiatives she ever took were all in a burst at the beginning of her adult life--with approximately twenty years left to go--when she left the colony of her birth, abandoning along with it her mother and sisters. She mated then, once for all time--no more sex for her--and started a new colony of her own. Furiously, during a very short period of activity and alone, she performed almost all of the instinctive behaviors of a female of her species, and on top of that the same labor as sterile workers of a developed colony.
The hereditary programs were there, in the sense organs of the Trailhead Queen and in her nerve circuitry, all expressed in the required correct sequence of her actions. These programs faithfully repeated the routine followed by the ancestral wasps that evolved into the first social ants as they crawled among dinosaurs over a hundred million years ago.
Leaving the nest of her birth to start this process, she first spread her four membranous wings and flew into the air. There she joined a swarm of flying males and other virgin queens. One of the males was able to catch her. He clamped his legs around her body, and the couple spiraled down to the ground. On landing, he used large claspers at the rear end of his body to hold their genitalia together and complete the insemination. Within five minutes the act was finished, and the Queen shook the male loose. All of the sperm she received flowed up into a special bag-shaped organ in her abdomen to stay there until called on to fertilize more of her eggs. That might be years into the future. Each sperm was endowed with a potential life-span equal to her own.
In contrast, the father of all her children was programmed to die almost immediately after the mating. The only thing he had ever done was accept meals regurgitated to him like a nestling bird by his sisters, and wait, and wait some more, and finally take the one short flight from his home followed by five minutes of copulation.
The male had started his life as an egg laid by the Queen of the mother colony. The egg hatched into a grub-shaped larva, which was fed by the all-female nurses. Upon reaching full size, the larva metamorphosed into a pupa. This final immature stage was encased in a soft, waxy, temporary exoskeleton with the form of an adult and three body parts--head, thorax, and abdomen. A pair of antennae and three pairs of legs sprouted from the body. Inside, the masses of undifferentiated tissue, protected by the waxy outer skeleton, grew into his final internal organs.
When the transformation was complete, the outer layer was stripped away and eaten by the workers, and the adult male stepped out, complete with wings, large eyes, massive genitalia, rudimentary jaws, tiny brain, and the one big purpose programmed in his tiny brain followed by quick death.
In short, the male was no more than a guided missile loaded with sperm. His life's work would be a single ejaculation. Up to the climactic moment, he had been a parasite in his mother's colony, a layabout fed and groomed by his sisters. He performed no public service. After his world-defining five minutes, he was left with only one instruction that would be enforced if necessary by his sisters. Don't come back here. Just die.
He would not even try to return to the nest. He had no chance at all of survival. A delicate creature, he was not provided with defenses. He had no way to find food, or feed himself if he stumbled across some. He had been issued a one-way ticket. He would die by dehydration, or crushed in the beak of a bird, or chopped into pieces by the jaws of an enemy ant, or, less quickly, pierced by the bloodsucking proboscis of an assassin bug.
To escape the same fate, the newly mated future Queen of the Trailhead Colony, full-brained and powerfully muscled, hurried to find shelter. She had to get back underground as quickly as possible after receiving her sperm load. First, however, she had to take a few minutes to shed her four wings. To do that she simply bent her middle legs forward, pressed them against the base of the wings, and snapped them off. This mutilation caused no injury to the rest of her body. It caused no pain. From the start the wings had been lifeless films and struts of chitin, joined to the body in a way that made them easy to break off painlessly and then discard.
The Queen was a parachutist that slipped her harness upon landing. Now she could move more quickly to avoid ants, spiders, and other predators hunting around her in the grassroots jungle. She was fortunate to come upon an open space between grass clumps, a small, ant-sized clearing at the Lake Nokobee trailhead. By luck she had found it an ideal site. If she could build a nest there, it would be her home for, perhaps, twenty years. She set out at once to dig a vertical tunnel in the sandy clay soil. Her movements were swift and precise, and within minutes she had deepened the shaft to more than her body length. It provided some degree of protection, but needed to be completed as quickly as possible. She had to hurry. Her life remained in constant danger; there was not a minute to lose.
At a predetermined depth, which she measured by the time it took her to climb up and down the shaft, the young Queen turned to the side at the bottom and began to excavate a wider space. She continued until she had fashioned a round chamber about three times wider than the vertical shaft. Her safety was now enhanced but far from ensured. Predators and marauding ants could still climb down the shaft to attack her. At least now the enemies would be confined to a narrow space by the walls of the shaft and forced to confront the young Queen's thrusting sting and snapping jaws head-on before they could reach her vulnerable body.
With this much achieved, and as the shadows of the surrounding pines lengthened across the trailhead, she had beaten odds of about a hundred to one. For every hundred young queens leaving the mother nest in order to be mated and start a new colony, only one at the end of the day now sat in the bottom chamber of an incipient nest.
Yet in spite of this huge achievement, and no matter how securely she had constructed her home, the odds against final success were still stacked against her. The chance of progressing from being the architect of a nearly excavated nest to being the mother of a large, mature colony was also about one in a hundred. Thus the Trailhead Queen would be statistically the single one in ten thousand who flew from the mother nest and went on to finish the entire process. She alone would enjoy a long life in the deep royal chamber of a mound nest. Defended by an army of fierce daughters, she would be as safe as any insect in the whole world could expect to be.
Even with the excavation of the first chamber complete, the Trailhead Queen still had heavy work ahead. First, she laid a small cluster of eggs on the earthen floor. These tiny objects she was compelled to lick continuously. It was an urgent task, because to the peril from enemies above was now added the threat of bacteria and fungi teeming in the soil all around her. If the eggs were not regularly cleaned and coated with antibiotic saliva, they would be soon overgrown by an invading mold and consumed. And from a single bacterium in the soil excavated by the Queen, millions could proliferate on any ant tissue left unprotected.
Other young queens around the Trailhead Queen were digging in on their own. All failed: one by one predators killed them. No sister worked side by side with the Trailhead Queen, no worker had yet been born that could support her. Outside the sisterhood of the mother colony that produced her, nature remained a battleground in a pitiless and total war every minute of the day. Intruders winkled the survivors out of their little nests and ate them. Other risks were present inside the nests themselves: the eggs might not be properly inseminated, or the sperm might be genetically defective.
But the dice fell right time and time again for the future Queen of the Trailhead Colony. As tiny larvae hatched from her eggs, she fed them highly nutritious food secreted from a large gland that partly filled her head and emptied through her mouth. This baby food was manufactured from masses of fat stored in the rear segment of the Queen's body. It was also created by metabolism of her now-useless wing muscles.
From the reserves of her own body the young Queen reared a dozen workers. All were female. They were tiny and weak, barely able to perform the work necessary for the little colony to survive. By necessity they came into the world as midgets. If each were larger in size, fewer of them could have been raised. The number would have fallen below the level necessary to provide adequate labor for the survival of the newborn colony.
Some of these pioneers, guided entirely by instinct because no one existed to teach them, set out to forage for food. Others took care of the Queen and reared the next generation of workers to maturity. Still others devoted time to enlarging the nest. Failure to perform all these tasks with exactitude would mean death for the colony. The young Queen could help no more. On the contrary, she desperately needed help herself to continue living. Her expendable body tissues were depleted. They had almost all been fed to the larval daughters, and now she was starving. Her body was a shell of chitin containing only tissues necessary for her own life. The first foragers venturing timidly away from the nest were able to bring back a few scraps of food. Their prizes included a fallen mosquito, a bit of shed caterpillar skin, and a newly hatched spiderling, which were enough to keep the colony alive and allow the Queen to regain some of her weight and strength.
Workers of the next generation, raised on food harvested from terrain outside the nest, were somewhat larger in physical size than the first generation, and stronger. They began to dig out many more tunnels to accommodate the growing population. As the colony and their habitation grew, the endangered home assumed the form of a labyrinth of chambers and connecting galleries. It became an enemy-proof fortress. A mound of excavated soil formed above it, reinforcing the roof and capturing the warmth of the sun.
As the months passed, the Queen, growing heavy with eggfilled ovaries, retreated ever deeper, distancing herself from the still-dangerous nest exterior. She had become an extreme specialist. She alone laid eggs, she alone was the growing tip of the burgeoning colony. The workers performed all the labor needed to raise her offspring, their sisters. They were the Queen's hands and feet and jaws, and increasingly they replaced her brain. They functioned together as a well-organized whole. They were altruistic toward one another, and they divided labor without regard to their own welfare. The Trailhead Colony came to resemble a large, diffuse organism. In a word, it became a superorganism.
By the time the colony reached its full mature size two years after the nuptial flight of the Queen, it contained over ten thousand workers. It was then able, in the following year, to rear virgin queens, and males, and from them to give birth to new colonies. By that time the Queen was producing eggs at the average rate of one every fifteen minutes. Heavy and torpid, she lay in the royal chamber at the bottom of the subterranean nest, five feet below the surface, a distance of four hundred ant lengths. By human scale the ant city was the equivalent of two hundred underground stories. The mound of excavated soil capping the nest added another fifty stories aboveground.
The Queen may not have been the leader of this miniature civilization, but she was the fountainhead of all its energies and growth. She was the key to its success or failure. The metronomic pumping out of fertilized eggs from her twenty ovaries was the heartbeat of the colony. That it should continue strong and true was the ultimate purpose of all the workers' labor. Their careful construction of the nest labyrinth, their readiness to risk life in daily searches for food abroad, their suicidal defense of the nest entrance, all their sacrifices were for her and for the creation of more altruistic workers like themselves.
One worker, or a thousand workers, could die and the colony would go on, repairing itself as needed. But the failure of the Queen, if not corrected, would be fatal.
Now after twenty more years that catastrophe had occurred. The death of the Queen was the greatest challenge the colony had faced since the days of its founding. Yet the workers could not take action until they learned for certain that the Queen was dead. They knew that something was not right, that something unnamed had settled upon them, but they did not yet realize the extent of her problem. The signs were not yet strong enough. So the Trailhead Colony thrummed on for a while longer with bustle and precision. Like a large ship at sea, it could not be easily turned from the shoals coming at it.
The reason for the continued momentum of the Trailhead Colony lay in the way ants communicate. Because they live most of their lives in underground darkness, they cannot speak to each other with sight or sound. Instead, they are forced to communicate with chemical signals. Human beings think in sound and vision. Ants, forced to be pheromonal, think only in taste and smell. No human can understand the chemical sensations that crowd the brain of a worker ant. We have no understanding of the entities she conceives, or the tones, the accounts, and the blends that course through her mind. While the Trailhead Colony may have been silent to unaided human perception, it was thunderous with pheromonal chatter among the ants.
The Trailhead Colony communicated using about a dozen chemical signals. The retinue of workers crowded around their dead mother were locked to her by several of these pheromones still oozing from her body. Translated into a human voice, they whispered a ghostly command: Come here, gather around me, stay close to me.
The attendants licked her body lasciviously with their pad-shaped tongues. They continued eagerly to clean her, picking up substances from her to pass on to others outside the retinue. The pheromones that triggered their intimate care spoke through taste and smell: Wash me, eat the substances you clean from my body, share them with your sisters.
The substances commanding the retinue were held in the forward chamber of the gut of each of the ants. They mingled there with liquid food. The ants smelled each other constantly by sweeps of their antennae, the "feelers" on their heads, the equivalent of the human nose. An ant that was well fed, with lots of food resting in her gut, said to a less well-fed nestmate, Smell this, and if you are hungry, eat. If the ant approached and was in fact hungry, she extended her tongue, and the donor ant rewarded her by regurgitating liquid directly into her mouth.
The exchanges among the sisters continued in this way. The combined intelligence of the colony listened to the flood of crosstalk among its members. They spoke in pheromones in all the messages they were programmed to send and receive. The colony exchanged information within itself in the same way the body of one ant, one human, or any other single organism exchanges information within itself by hormones. The superorganism pheromones suggested, begged, and commanded.
Outside the nest, not far from the Trailheader mound, a wood thrush flew by one day carrying a grasshopper to her own nest. Part of the crushed insect broke off and fell to the ground. In less than a minute a patrolling worker found it, triggering a chain of action of the kind followed countless times by Trailheaders before. She examined the grasshopper, tasted it briefly, and ran back to the nest entrance. On the way, she touched the tip of her abdomen to the ground, laying down a thin trail of chemicals. Entering the nest, she rushed up to each nestmate she passed, brushing her face close to theirs. The odor-sensitive antennae of the nestmates detected both the trail substance and the smell of grasshopper. The signals now proclaimed, Food, food. I have found food, follow my trail!
Soon a mob of ants ran out. They followed the trail, and gathered around the delicious haunch of grasshopper. Some of the first to arrive ran back to the nest, laying trails of their own, reinforcing the message, saying, Come on, come on, we need help.
The ants working on the grasshopper piece began to drag it toward the nest entrance. A catbird perched on the branch of a tree nearby saw the activity and swept down to investigate. She pecked at the grasshopper, scattering the ants and injuring several. The ants expelled a pheromone from a gland that opened at the base of their jaws. A chemical vapor spread fast. It shouted, Danger! Emergency! Run! Run! Get out of here!
And so the business of the Trailhead Colony was conducted by a vocabulary of odor and taste. Pheromones were emitted, occasionally reinforced by touch. Messages were created, sometimes with a single chemical substance, sometimes with the same substance at different concentrations, and on occasion two or more in combination. Meanings were changed according to where the substances were delivered. The vocabulary grew. Different messages were delivered.
Here, let me lick and clean you.
Get to work, do what others are doing here.
This is my caste, and this is my condition.
Let us lay down territorial pheromones, announcing to rivals our dominion over this land.
We don't have enough soldiers; raise more in the nursery.
We have too many soldiers; raise fewer.
Who is leading the struggle to become our new Queen?
The members of the Trailhead Colony lived every second of their lives by instructions in the clouds and torrents of pheromones around them. Some signals such as the alarm pheromones spread and faded fast, drawing the attention of many nestmates as needed locally, but not holding on long enough to create panic throughout the colony. At the opposite extreme, some odors spread slowly and lasted a long time. Among them were the royal pheromones of the Trailhead Queen. Even as her body began to decay, the pheromones she had manufactured in life persisted in the minds and bodies of her colony.
The royal presence had been woven into the pheromone life of the Trailhead Colony a second way. Her secretions were blended with other substances to create an odor unique to the colony as a whole. The odorants were absorbed into the waxy cuticle that covered the body of every member of the colony. Each colony of ants had a personal bouquet that all its members shared and learned and to which they remained absolutely faithful. When two ants met, regardless of their origins, they both swept their antennae back and forth over the other. The movement was too swift for the unaided human eye to follow, but the brain of each ant almost instantly processed the information. If the two ants shared the same odor, the message was, She belongs to my colony, no other. The two then either continued on their way without pause, or paused to groom each other, or exchanged food. If, on the other hand, their odors differed even slightly, the message was, Different colony, look out! Like strange dogs meeting in the street, two ants of separate origin stopped to examine each other more carefully. They then either launched an attack or ran from each other.
No words, no signals of motion were used to establish an ant's tribal identity. None was needed. The Trailhead Colony was united simply and entirely by possession of the same smell. If that were to disappear, the superorganism would quickly dissolve into a mob of disoriented organisms. They would fight among themselves. Enemies would scatter them. Predators would close in for an easy meal.
The Trailhead Queen lay in state. It could not last forever. Eventually parts of her body were eaten and the rest carried away to the ant cemetery. For a week the pheromones licked off the remaining fragments continued to broadcast her existential message. Thereafter the chemicals gradually dissipated, and at last the message faded away.
A few chemical signs appeared early, but it was on the third day after she died that the Queen's pheromones began to be overlaid by the faint evidence of death. Her overall odor became ambiguous, and with it the posthumous messages she sent. Still, nothing mattered in the recognition of her corpse other than the odor of decay. Visual appearance and the cessation of movement meant nothing. The Queen could have lain on her back with her legs held rigidly up in the air. She could have turned any color: red, black, metallic gold, or any other hue or shade, it would not matter. Instead, the Queen had to smell dead in order to be classified as dead. And not from the blends of substances in corpses repellent to the human nose--not, for example, from the loathsome skatole and indole that distinguish human feces, nor the trimethylamine that rises dramatically from spoiled fish. Such chemicals, when encountered alone, would cause alarm in the ants and repel them. The same was true of other volatile toxic substances. Only oleic acid and its ester, which are decomposition products of fat, were effective messengers of death. They mean little or nothing to the human nose. But they mean dead to an ant. When encountered on the corpse of a nestmate, they caused the ants to pick it up and carry it away for disposal.
Within a week, the constant licking of the royal corpse in the Trailhead Colony started to break it into pieces. One by one the fragments, reeking of the oleic compounds, were carried out of the royal chamber. Unknowingly the ants bade farewell to their mother. No ceremony was performed. Instead the workers bearing the body parts wandered alone through the nest galleries in search of the Trailheader cemetery. This special place was not marked by ceremonial trappings. It had no special shape, nor did it contain any token of remembrance, even for a queen. It was merely a chamber at the periphery of the underground nest. The ants dumped all kinds of debris into it, including discarded cocoons shed by newly emerged adults, inedible parts of prey, and deceased colony members.
When the corpse carriers came close to the refuse chamber, they turned their burdens over to cemetery workers. These specialists were ants who constantly rearranged and added to the refuse piles. They stayed close to their work and were for the most part avoided by their nestmates.
In cemetery work and all other activities, the Trailhead Colony organized its labor by altruistic rules of labor specialization. Everything they did was restrained by some degree of self-sacrificial altruism. Above all, the workers had given up the chance to reproduce, at least so long as the Queen was alive and healthy. They accepted service in foraging, soldiering, and other dangerous occupations that increased their risk, often to the point of certain early death. The dominance of the Trailhead Colony over its individual member ants was total. The welfare of the superorganism was paramount, and a worker's life story was programmed to be subordinate to the superorganism's needs. If a worker died, the loss weakened the colony to some measurable but relatively inconsequential extent. The deficit could be quickly made up by rearing another worker in the nursery. If, on the other hand, a worker behaved in a selfish manner, consuming for a good part of its life more resources than it contributed, it weakened the colony far more than if it just had the decency to desert or die.
The decency of ants was, in disability, to leave and trouble no more. The self-sacrifices that led to the success of the Trailhead Colony were evident in every task performed by all of the worker force in all circumstances. The sick and injured received no care. In fact, they avoided such attention, moving on their own to the outermost nest chambers. The disabled were among the colony's most aggressive fighters. Dying workers often left the nest completely, thereby avoiding the spread of infectious diseases.
Older workers that stayed healthy but were approaching the end of their natural life-span also emigrated to the nest perimeter. From there they were prone to become foragers, leaving the nest to search for food, which exposed them to a much higher risk from enemies. When defending the nest, elders were among the most suicidally aggressive. They were obedient to a simple truth that separates our two species: where humans send their young men to war, ants send their old ladies.
The Queen had been the exclusive reproducer of the colony, the mother of the entirety of its inhabitants and the new colonies they were able to produce. All the sacrifices offered by the workers were made to protect her life and to enhance her fertility. Whether the Trailhead Colony would now live or die depended upon its ability to replace the mother Queen. That would require all their skill in retaining their strength until a new Queen was installed.