The Ant Professor--Lin Chung-chi
Coral Lee / photos Chuang Kung-ju / tr. by Scott Williams
March 2006

In high school, Lin Chung-chi dreamed of winning a Nobel Prize. Lin went on to major in entomology at university, where he chose the most unfashionable of unfashionable fields--myrmecology, the study of ants. From his master's degree through his PhD to his post-doctoral research, Lin has been studying ants day and night for 18 years. People paled at 2004's news that Solenopsis invicta, the red imported fire ants that have been plaguing the United States for decades, had invaded Taiwan. As a result, Lin's unmatched knowledge of ants was suddenly in high demand. Once a denizen of an academic backwater, he is now both Taiwan's most sought-after authority on fire ants and the man whose frankness led him to withdraw from a team working on fire-ant control efforts. How did Lin get to where he is today? What charms hide beneath the unassuming exoskeleton of the ant? What role does the ant play in the biosphere? And what does the fire ant invasion reveal about the difficulties faced by ant researchers in Taiwan?
"Typically," says Lin Chung-chi, "the specimens you want to collect don't turn up." Lin, an assistant professor at National Changhua University of Education, is speaking to Hsu Hui-yi, a PhD candidate from Lin's alma mater, National Taiwan University, on an autumn afternoon. The two of them are in the hills behind the campus hunting for the yellow crazy ant (Anoplolepis gracilipes). As a steady stream of people ascends the path, professor and student use spade and trowel in a long and fruitless search beside the steps, around fallen trees and beneath rocks.
Arriving at a level stretch of forest, they discover a volleyball-sized nest hanging from a tree. "The last time I was looking for a black spiny ant [Polyrhachis dives] nest," exclaims Lin, "I couldn't find one!" Yet here one is. As he carefully pulls back an outer layer of the nest, a dozen or two coal-black ants climb out. Everyone gathers round for a closer look at the nest, which is woven out of larval silk, and catches sight of another nest beneath the tree's trunk. "In their search for food," observes Lin, "the spiny weaver ants have probably moved into the long-legged ants' range." He thinks they'll find the ants they're seeking nearby.
Lin soon discovers traces of the reddish-brown long-legged ants under another tree. After turning over a spadeful of earth, he squats down beside the tree. Spotting only a few ants, he uses his hands to clear away earth and stones rapidly but gently. Suddenly, we see ants scurrying about everywhere, and even catch sight of a couple of larvae. Lin quickly pours carbon dioxide into the hole, then shovels the clod of earth into a container. Once he has the clod in the container, he breaks it up in his hands to see if he's collected a queen, periodically moving the loosened earth to a shallow dish for a closer look.

Yellow crazy ants scatter when their nest is dug up from beneath the roots of a tree.
First catch the queen
"Ant queens," says Lin, "usually live deep inside the nest." And queens, which lay the colony's eggs, flee as soon as the nest is invaded. Lin explains that to catch them, you must first stun the ants with gas, then dig up the whole nest. But even this approach doesn't guarantee that you'll catch the queen. You have to examine the nest to be sure you've got one. After a few more hours of digging up nests and picking through them, Lin finally catches sight of a queen and calls it a day.
Collecting ants to raise in the lab is part and parcel of ant research. Hsu Hui-yi, who wants to learn whether yellow crazy ant behavior is cooperative or competitive, will be rearing ants for experimentation and observation for some time to come.
Lin raises all kinds of ants in his own lab. His formicaria are filled with artificial nests of every type and covered with red cellophane to simulate darkness. This allows Lin to observe the ants as they busily search for food, transport captured prey, and care for their larvae.
In addition to the live ants, the lab has 6-7,000 bottles containing preserved specimens. Having collected them himself, Lin can, with just a glance at the label, tell you when and where each specimen was collected, as well as the condition of the ant at the time. In another cabinet, dried ants lie in specimen drawers. Each species is displayed in pairs, one belly up, the other belly down, to facilitate observation of identifying features.

Once he has a nest, Lin checks it carefully to see if he has collected a queen.
A barren waste
Ant research has long been unfashionable in Taiwan. According to Professor Wu Wen-jer of the Department of Entomology at National Taiwan University, entomology got a later start than botany and vertebrate zoology and has fewer people working on even as basic an area as taxonomy. The number of known species of insect in Taiwan currently stands at more than 200,000, but this number is thought to be less than 10% of the actual total. Taxonomy is fundamental to biology. At both the macro-scale of ecological research and the micro-scale of molecular biology, the field relies on comprehensive records of plants and animals. Taiwan's dearth of taxonomic data has slowed the pace of insect-related scientific research on the island.
Myrmecology is a particularly desolate subfield of the already neglected field of entomology. Throughout the Japanese occupation, Western and Japanese myrmecologists surveyed local ants and collected specimens, ultimately recording 140-some species of ants on Taiwan. In marked contrast, in the more than 50 years since the ROC government came to Taiwan in 1949, only one scholar has conducted basic surveys and research.
"When I was a student," recalls Lin, "NTU didn't yet have an entomology department; there was only an entomological group in the Department of Botany and Plant Pathology." In those days, he says, the department was split into two areas: economic applications and basic research. Because he was interested in the phenomenon of life, he naturally chose the theoretical group.
"I've been fascinated by the worlds glimpsed through microscopes and telescopes since childhood," says Lin. He recalls that when he was young, he used to squat on the ground for hours at a time to stare at the fish and shrimp in aquaria or to watch ants transport things. Later, when his mother bought him a microscope, he became absorbed by the mysteries of the tiny living things he saw under the glass. It therefore seems only natural that when it came time to choose a research subject at university, he rejected large insects like butterflies and beetles in favor of ants. The day after he asked Professor Wu Wen-jer about the possibility of pursuing myrmecological research, Wu brought him a stack of articles from around the world and noted that ants had provided important material for life sciences research. With Wu's encouragement and backing, Lin hitched his wagon to an ant.

Black spiny ants are often seen in Taiwan at low elevations. Here one is gathering nectar.
Dances with ants
Lin first produced "A Comparison of Taiwan's Ants," a survey, taxonomy and specimen collection.
In 1990, Japanese scholars had collated all of the myrmecological literature on Taiwan and had found 197 species. Lin has spent more than ten years in the field building on this foundation and has increased the number of known species to 276. He has also collected specimens of 90-some species which had previously been identified but not collected.
Though it generally isn't difficult to collect ant specimens, there are a few unusual species that can drive a collector up a wall. Lin had to search for one such species for more than seven or eight years before finally collecting it.
"One ant of the genus Trocanilla," says Lin, "was previously unrecorded on the island. But it had been found in Japan, China and the Philippines, suggesting that it should exist in Taiwan as well." Lin explains that the ants of this species are very difficult to collect--not only do they spend their entire life hidden in the earth, but are now also facing extinction. While still a doctoral candidate, Lin stumbled across one on the Huisun Farm, but couldn't find the nest in the earth nearby. He made over a dozen trips to the farm over the next several years, but came away empty handed each time. It wasn't until much later, on a research expedition in Ilan's Fushan Botanical Park, that he finally stumbled across a nest. Lin discovered the deep-dwelling ants where they had come up close to the surface after a heavy rain. It was then that he learned that they usually live at a depth of 10-15 centimeters.
"You have to be able to bear loneliness to do basic entomological research," says Lin. Wu Wen-jer, who has been working with Lin since Lin was a student, says that taxonomists spend an incredibly long time preparing to enter their field. To have a sufficiently broad perspective, they must have a solid understanding of theory, including genetics, evolution and taxonomy. They also need a tremendous amount of experience in the field. A really good taxonomist is, without doubt, a modern naturalist, familiar on the one hand with the ecology and the environment, and on the other with genes and proteins.

Lin carefully peels back the outer layer of a black spiny ant nest so his students can see how the ants have woven it together out of larval silk and leaves.
A long road
Keeping research going requires long-term financial support. But funding from research bodies such as the National Science Council tends to go either to topics of global interest or topics that have practical applications, in areas such as medicine or electronics. Grants for basic research are limited and competition is fierce.
For projects that span several years and for which no long-term grants are available, funding can be piecemeal. Lin says that during the 11 years in which he was conducting his doctoral and post-doctoral research, he needed from NT$600,000 to NT$1 million per year to fund experiments and cover travel expenses. Without Wu's support, much of that research would never have been completed.
Why would people devote themselves to the study of a creature that is less than one millionth the size of a human being? "Humans have existed for about seven million years," says Lin. "The ant has been around for more than 100 million." The more Lin studies the ant, the more awed he is by its ability to survive and thrive.
Within the biosphere, ants serve as food for a variety of other insects and arachnids, as well as themselves disposing of dead creatures. Ants also spread plant seeds, and their earthmoving activities are crucial to the circulation of soil nutrients. In short, ants have been important to the evolution of all of the Earth's plant and animal species and still play a significant role in their survival.

Here, the common Monomorium floricola sucks sweet secretions out of a cocoon.
A cornerstone species
"From the perspective of the food chain," says Lin, "ants are a cornerstone species." He explains that in a food web in which 20-30 species depend upon one another, the disappearance of eagles from the top of the web or a small plant from the bottom has less impact than the disappearance of a species from the middle. Ants exist in great numbers and great variety, and consequently have a greater effect on the food web.
Take the army ants of the Amazon jungle, for example. Each species of army ant is the natural enemy of a particular jungle pest. They therefore play a crucial role in controlling these pests. Perhaps less obvious is that many birds depend upon army ants for their survival.
Tropical rainforests have evolved over millions of years. Birds can't see insects moving under the thick layer of leaves that covers the ground. But when army ants go out foraging in their hundreds of thousands, they make a humming noise that flushes other insects from their cover. This, in turn, gives the birds an opportunity to eat. Consequently, each species of army ant has symbiotic relationships with about ten bird species. With the jungle being cut down, the army ants need an area of at least 100 hectares of protected forest if they are to survive. But the protected areas are too small for the birds. With the ants disappearing from outside such protected areas, the birds face extinction.

Tapinoma indicum, a house ant often found in the vicinity of Taiwanese homes, can store energy and water in its body.
Strength in numbers
The number and breadth of distribution of ants is startling. A single worker ant weighs only one to five milligrams, yet the total weight of all the ants in the world is equal to that of all the people. From the tropics to the forests of the Arctic Circle, from deep in the soil to the top of the forest canopy, from the pathways of urban parks to homes and factories, ants are everywhere.
"The ant's extreme physical and behavioral specialization has enabled it to adapt to all kinds of environments," says Lin, who finds their distinctive behavioral patterns a fascinating research subject.
As everyone knows, ants exhibit social behavior that involves a distribution of labor: Some worker ants are responsible for foraging for food while others care for the larvae; soldier ants defend the nest; the queen lays eggs; the males fertilize the eggs. Within the colony, each class fulfills its responsibilities, cooperating ceaselessly with the others. This leads to the high degree of efficiency that is the ant's survival advantage.
Many ant species have also developed amazing behavior patterns that have enabled them to adapt to hostile environments.
For example, among the various species of "honeypot" ants, a worker functions as a living larder. These workers have a crop in their abdomen that can, when filled with honeydew, swell to several times their normal body size. When food is scarce, they regurgitate this honeydew to feed the other ants in the colony. Interestingly, in order to prevent the formation of the "bedsores," these obese living larders hang from the ceilings of their nests rather than lie on the ground.
"In the animal kingdom," says Lin, "probably only humans and ants gather and store water." He says that the many "altruistic" and "self-sacrificing" behaviors of the ant have been crucial to making Formicidae one of the world's dominant animal families.

According to Wu Wen-jer of the Department of Entomology at National Taiwan University, Taiwan's research environment is not supportive of basic biological research. It takes a very long time to develop the skills necessary for such research, so those who wish to pursue it must be very comfortable working alone.
Queenless ants
In recent years, much of Lin's research has focused on the extremely interesting behavior patterns of the cannibal ant (Cerapachys biroi). "Only five of the world's more than 15,000 ant species exist without a queen," notes Lin. And the cannibal ant, which is endemic to Taiwan and Okinawa, is one of the five. In place of the typical queen model, cannibal ants share egg-laying responsibilities among all the worker ants, thereby increasing their chances of survival. Cannibal ants are also known to wipe out colonies of neighboring ants of other species by eating their larvae.
How can cannibal ants reproduce without a queen? This question leads to others that fall within the realm of biotechnological research: What applications might this kind of asexual cloning have? Could genetic material with medical applications be inserted into cannibal ants, be cloned in large volumes by the ants, then extracted from the ant larvae?
Cannibal ants' behavior raises another question: Ant larvae are usually heavily protected. "How," asks Lin, "are cannibal ants able to overcome the powerful defenses of other ants and strike at the heart of their colonies?" In fact, this is another of Lin's research interests. He says that cannibal ants secrete a chemical compound that makes rival ants believe they have already been defeated--they surrender and even offer up their larvae to the invaders.
With the arrival of fire ants on Taiwan, Lin has been thinking about cannibal ants even more. Can, for example, cannibal ants' need to feed on the larvae of other ants be turned against fire ants?

Lin Chung-chi's first love was the jungle, teeming with life. To his students, it is a paradise to be explored and researched. The photo shows the Fushan Botanical Park.
A fight between bullies
Lin notes that the cannibal ant is an indigenous ant. The local biosphere is a system in balance, and the ant, which has no natural enemies, sits near the top of that system. Its numbers are limited by the quality of its environment and the availability of resources. Lin argues that because the cannibal ant is a native species, raising it in the lab and training it to eat fire-ant larvae would be a more appropriate means of biologically controlling Taiwan's fire-ant population than would importing the fire ant's natural predators.
"The fact that cannibal ants reproduce without a queen," says Lin, "and produce a new generation every 40 days, makes it possible to breed large numbers of them. This is the only way to deal with the red imported fire ant, which owes its own success to its reproductive speed and numbers." According to Lin, after nearly a year of selective breeding, he has developed a strain of cannibal ants that, in the lab, have shown a taste for fire ant larva. When they reach the point of preferring fire ant larvae 99 times out of 100, he can begin raising them in large numbers and releasing them in areas that are seriously threatened by fire ants. Lin believes that this will help control fire ant populations and allow other ant species that have been driven out of their ranges by the fire ants to recover. When the numbers of nests in these areas return to their original levels, the cannibal ant population will also fall because there won't be enough food to support it, thus restoring the ecological balance.
"The government has supported me for more than ten years," says Lin. "Now that fire ants have come to Taiwan, its only right that I pay that back." To that end, Lin has spent the last two years running all over the island. He has conducted surveys, appraised the situation, and developed methods and plans for dealing with the fire-ant invasion. In the process, he's also become a focus of media attention. Though he doesn't like having his work interrupted, he says he talks to the media because he feels it is his responsibility to do so. Asked about the vacillation that has been evident in the government's fire-ant control policy, he says bluntly that it is on the verge of missing its decade-long window of opportunity.
Away from the media and back at his academic post, Lin is also concerned that the outstanding students of the last few years are even less interested in basic research than the students of his day. Lin hopes that he can pass on to them the research bug that Wu Wen-jer passed on to him and his classmates. At the same time, he remains very happy in his lab exploring the mysteries of the world of ants.

Specimens are a necessity when doing taxonomic research. Dried specimens like those in the picture--one ant prone, the other prostrate--allow researchers to identify unique characteristics.

The cannibal ant is one of the few queenless ant species. Here, workers are taking larvae of other ant species to feed their own larvae. The translucent larva in the lower left of the picture is that of another species. Those to the right and at the top are cannibal ant larvae.