Anurag Agrawal: Monarchs vs. Milkweed

by Anurag Agrawal

Coevolution is a special kind of evolution. And monarchs and milkweeds exemplify this special process. In particular, what makes coevolution special is reciprocity. In other words, coevolution is one species that evolves in response to the other, and the other species evolves in response to the first. Thus, it is a back-and-forth that has the potential to spiral out of control. In some arms races, the two organisms both benefit, such as that between some pollinators and flowering plants. But coevolution is more common among antagonists, like predators and their prey.

When biologists first described coevolution, they likened it to an arms race. An arms race, such as that between political entities, occurs when two nations reciprocally increase their armament in response to each other. So how does an arms race between monarchs and milkweeds, or between cats and mice, or between lions and wildebeest, or between plants and their pathogenic fungi, proceed? When coevolution occurs, it proceeds with “defense” and “counter defense.” And one of the few rules of coevolution is that for every defense that a plant or prey mounts, the predator mounts a counter defense, or an exploitative strategy to overcome the defense.

Once a monarch butterfly lays an egg on a milkweed plant, the natural history of coevolution unfolds. For every defense that the plant mounts, milkweed mounts a counter defense. Once the caterpillar hatches, it must contend with a bed of dense hairs that are a barrier to consumption of the leaf. But monarchs are patient, and have coevolved with the milkweed. So their first strategy is to shave that bed of hairs such that the caterpillar has access to the leaves that lie beneath.


For every defense there’s a counter defense. But next, when the monarch caterpillar sinks its mandibles into the milkweed leaf, it encounters a sticky, poisonous liquid called latex. In this video we will see how the monarch caterpillar deactivates the latex bomb that the milkweed puts forward.

And so the arms race continues, with reciprocal natural selection resulting in coevolution between monarchs and milkweeds. In my book, Monarchs and Milkweed, I outline the third level of defense and counter defense between these two enemies. Milkweed next mounts a remarkable and highly toxic defense chemical called a cardiac glycoside. But, yes, again the Monarch has evolved the means to not only not be poisoned by the cardiac glycoside, but to sequester it away and put it to work in defense of the Monarch itself from its enemies, such as predatory birds. For more on the Monarch – Milkweed arms race see this video, filmed in Ithaca, New York outside of Cornell University where we conduct our research.

AgrawalAnurag Agrawal is a professor in the Department of Ecology and Evolutionary Biology and the Department of Entomology at Cornell University. He is the author of Monarchs and Milkweed: A Migrating Butterfly, a Poisonous Plant, and Their Remarkable Story of Coevolution.

Anurag Agrawal: The oldest butterflies?

by Anurag Agrawal

It’s unclear when humans became humans. Presumably it was a gradual growth of our consciousness over the eons. There are some things, however, that appear to distinguish us from most other animals. For example, our artistic depictions. From the deepest, darkest caves have emerged pictures of humanity from thousands of years ago. And in an Egyptian tomb, that of Nebamun, on a painting called “Fowling in the marshes” (from around 1350 BCE) comes one of the oldest human depictions of butterflies. It happens to be of the African Monarch, Danaus chrysippus, sometimes called the plain tiger, a close relative of our beloved North American Monarch butterfly, Danaus plexippus.

I stumbled on this lovely scrap of history when a friend and colleague, Harry Greene, gifted me a book: Nabokov’s Butterflies (2000), a collection of unpublished and uncollected writings. Some explanation is in order. Harry is an extraordinary naturalist and big thinker in ecology and evolution. Like many senior scholars, his predicament was the lack of shelf-space in his office. And so I was the beneficiary of Nabokov’s Butterflies. Vladimir Nabokov, a Russian-American author, and noted entomologist, was most famous for his writings, for example, Lolita, and his celebrated translation of Pushkin’s novel in verse, Eugene Onegin. His ideas about biology were diverse, he was a passionate lepidopterist, and he often intermixed his literary writing and entomological excursions. Lolita is said to have been written primarily on butterfly collecting trips in the American west. Nonetheless, Nabokov also clung on to other ideas that held little merit in the scientific sphere. Most prominently, Nabokov rejected evolution by natural selection as a driver of certain organismal traits that he deemed ‘coincidental, miraculous, or too luxurious.’


Nabokov was a professor at my own Cornell University in the decade following WWII. Although he taught literature and had well-known students at Cornell (including U.S. supreme court justice, Ruth Bader Ginsburg), his entomological interests continued. In fact, after he retired from Cornell in the mid-1960s, Nabokov had sketched out an outline of a book: The Butterflies of Europe. And although the book never came to be, the outline was recapitulated in Nabokov’s Butterflies. Flipping through the book, I stumbled on his entry for Danaus in which he wrote, “This butterfly has the distinction of being the oldest known to have been represented by man. Seven specimens of it (with typical white-dotted Danaus body but somewhat Vanessa cardui like wingtips) are shown flitting over the papyrus swamp…” (page 603).

I later asked another friend, Harvard’s Lepidopterist, Naomi Pierce: did Nabokov have it right? On the money, she independently pointed to the similarity of Danaus chrysippus and the painted lady, Vanessa cardui, wondering if the butterflies on this three thousand year old tomb painting were Danaus or Vanessa. She concluded, as did Nabokov, that the African Monarch ruled. Detailed assessment of the color patterns on the wings were informative to both entomologists. The oldest human depiction of a butterfly? Perhaps not. Naomi mentioned some evidence of butterflies in Minoan artifacts from Crete, a thousand years earlier than Nebamun, and likely in Pyrenees cave paintings, some 10-30 thousand years earlier!

Of course, there is nothing special about being the oldest depiction of a butterfly by Homo sapiens. But suffice it to say, butterflies, metamorphosis, wing patterning, and the beauty of nature have been on our minds for a very long time. Thanks Harry and Naomi! And thanks Nabokov. Who knows what becomes of those side hobbies and obsessions we all hold.


AgrawalAnurag Agrawal is a professor in the Department of Ecology and Evolutionary Biology and the Department of Entomology at Cornell University. He is the author of Monarchs and Milkweed: A Migrating Butterfly, a Poisonous Plant, and Their Remarkable Story of Coevolution.

Anurag Agrawal: Monarch overwintering

by Anurag Agrawal

The estimates of the monarch butterfly overwintering population were announced February 9th by WWF Mexico. The butterflies are so dense at their dozen or so mountain-top clustering sites that overwintering butterflies cannot be individually counted. Instead, the area of forest that is densely coated with butterflies (at about 5,000 butterflies per square meter looking up into the canopy) is estimated as a measure of monarch abundance. Butterflies arrive in Mexico around early November and stay until March.


This winter season (2016-2017), there were approximately 2.9 hectares of forest occupied with dense monarchs (somewhere in the neighborhood of 300,000 million overwintering butterflies). This estimate is down 27% compared to last year. Nonetheless, the previous two years were a 600% increase over the all-time low recorded in the winter of 2013-2014.


Where does this leave us? This year’s population was higher than predicted by many. The season started with a late spring storm that killed an estimated 5-10% of monarchs in March 2016, and many reported low numbers of adults last summer. Nonetheless, the lower numbers this season compared to last are within the range of year-to-year variation, and overall, the population seems to be relatively stable over the past decade. With these 24 years of data, there are various ways to plot and assess the trends. Below I have plotted the four year averages for six periods beginning in 1992. Any way you slice it, the trend has been negative, and the population is not nearly what it once was. Nonetheless, the downward trend seems to have lessened this last period. Is this the new norm? How dangerously low are these numbers? And what can we do to reverse the trend?


AgrawalAnurag Agrawal is a professor in the Department of Ecology and Evolutionary Biology and the Department of Entomology at Cornell University. He is the author of Monarchs and Milkweed: A Migrating Butterfly, a Poisonous Plant, and Their Remarkable Story of Coevolution.





Anurag Agrawal on Monarchs and Milkweed

AgrawalMonarch butterflies are one of nature’s most recognizable creatures, known for their bright colors and epic annual migration from the United States and Canada to Mexico. Yet there is much more to the monarch than its distinctive presence and mythic journeying. In Monarchs and Milkweed, Anurag Agrawal presents a vivid investigation into how the monarch butterfly has evolved closely alongside the milkweed—a toxic plant named for the sticky white substance emitted when its leaves are damaged—and how this inextricable and intimate relationship has been like an arms race over the millennia, a battle of exploitation and defense between two fascinating species. Check the PUP blog each Monday for new installments in our “Monarch Monday” blog series by Anurag Agrawal.

What makes monarchs and milkweeds so special?

AA: Monarchs and milkweed are remarkable creatures, they’re on a wild ride! From the monarch’s perspective, its only food as a caterpillar is the milkweed plant. This makes them highly specialized, highly evolved, and very picky eaters indeed. They’re actually not that unique among butterflies, but they are extreme. Milkweed does everything in its power to defend itself against being eaten by monarchs. They make sandpapery leaves, toxins that can stop a human heart, and a thick poisonous goo that can glue an insect’s mouth shut. Again, although milkweed is not unique among plants, it is extreme. In what is called a coevolutionary arms race, monarchs and milkweed have been continually evolving over the eons to keep up with each other. As such, they have a lot to teach us about the way nature works, the way plants and animals interact, and about the various paths that evolution can take different species. And this is all to say nothing of the monarch’s spectacular annual migration, often over 3,000 miles flown by individual butterflies, using the sun to navigate, and having stored away milkweed’s poisons to protect themselves from being eaten by birds. Monarchs and milkweeds are royal representatives of all interacting species.

Why did you write this book?

AA: After studying monarchs and milkweed myself for over 15 years, I felt like I had a lot I wanted to share, especially with non-scientists and nature lovers. Monarchs and milkweed are such fascinating organisms, and yet so much of their beautiful biology is not widely known. I also wrote the book because there are areas of my own knowledge about monarchs and milkweed that I wanted to immerse myself in, but that I had not yet done any research on. So as an author, getting to visit the overwintering sites in Mexico, to study the population decline of monarch butterflies, and to understand their mating rituals were all fascinating detours from my everyday research life at Cornell University. The book was incredibly fun to write, and getting to work with artists and historians made it all the more rich. I hope that anybody that has an appreciation for nature, an interest in science, or just a curiosity about the ecology of plants and butterflies will enjoy this book. Working on this project has surely altered the course of my own research, the classes I teach, and how I see the natural world.

Why have you highlighted some of the personalities of the scientists studying monarchs and milkweeds in this book?

AA: One of the most amazing things about monarchs and milkweeds is the scientists who have studied them. They were such remarkable characters, especially those pioneering studies back in the 1950s: tremendously creative, sometimes competitive, and with some of their discoveries worthy of a Nobel prize. Getting to know them, both from their discoveries and their personalities, and how they interacted, has enriched my appreciation for how science is done. It also highlights the meandering and sometimes serendipitous nature of discoveries. I wanted to share the thrill of science, its ups and downs, and the process by which it is done with the curious reader.

Can you share one of your ah-ha! moments from studying monarchs and milkweeds?

AA: One of my favorites was from when I was an assistant professor at the University of Toronto. One day I was eating lunch by myself in a small downtown garden. Just by chance, I happened to sit on a bench beneath a very tall milkweed plant that had a very large monarch caterpillar feeding away. Without giving away all the details, that one hour encounter, in the middle of a city with 3 million people, changed my perspective on monarchs and milkweed forever. It was so unlikely an event, perhaps 1 in a 1,000 that a butterfly had been flying by and happened to lay an egg on this Toronto milkweed, and then a further 1 in 100 chance of that egg hatching and surviving to be that large caterpillar that I could watch it. And probably a 1 in a million event that I would happen to be eating lunch there, that day, to observe the events. In biology one has to work hard, be patient, and occasionally get very lucky! Throughout my studies on monarchs and milkweed, I have had tremendous luck in encountering wonderful biology that has had profound consequences.

Is the monarch butterfly going extinct?

AA: The answer to this very important and timely question is both simple and complex. On the simple side, there is no way the monarch butterfly is going extinct anytime soon. Having said that, the butterfly, and especially the long-distance migration that occurs every fall from Southern Canada and the USA, all the way to Mexico’s highlands in Michoacán, is indeed declining at a rapid pace, and we should all be worried about the sustainability of the annual migration. There’s so much information and misinformation floating around in the news these days about the causes of the monarchs decline. What I’ve tried to do in the book is outline the best knowledge that we have to date and to examine the facts critically, so we can really understand what might be going on. Unfortunately, we don’t have all the answers, but we can reject some of the most prominent explanations for the population decline of the monarch butterfly. As I argue in the book, planting milkweed certainly won’t hurt, but it is unlikely to save the monarchs annual migratory cycle. It is perhaps ironic that I spend eight chapters of the book discussing and detailing the importance of milkweed for monarchs, and nothing could be more true than their intertwined and intense evolutionary battle, but at this stage, and thinking about their conservation, it does not appear that milkweed is what is limiting the monarch’s population. Monarchs will persist for a very long time, but given that they are migratory butterflies that taste their way across North America, their declining population is something we must try to understand. Much more than the monarch is at stake, these butterflies are sentinels for the health of our continent!

Anurag Agrawal is a professor in the Department of Ecology and Evolutionary Biology and the Department of Entomology at Cornell University. He lives in Ithaca, New York. He is the author of Monarchs and Milkweed: A Migrating Butterfly, a Poisonous Plant, and Their Remarkable Story of Coevolution.

Sunny, Spring Book List

The Bee
Say goodbye to cold, dreary winter and hello to spring. Welcome the season by checking out Princeton University Press’s selection of natural history books. Wondering what bird you hear chirping? Download BirdGenie Backyard Birds East and West and find out. Some choices to get you out of the house and into nature:


The Bee:
A Natural History

Noah Wilson-Rich
With contributions from Kelly Allin, Norman Carreck & Andrea Quigley
“The natural history of solitary, bumble, honey and stingless bees is as gripping as our lengthy alliance, as urban beekeeper Noah Wilson-Rich and contributors show in this charming compilation. They cover evolution, biology (including a unique proboscis made of two organs), behaviours (such as honey bee ‘quacking’), the causes of catastrophic die-offs, and more.”–Barbara Kiser, Nature



Birds of Australia:
A Photographic Guide

Iain Campbell, Sam Woods & Nick Leseberg
With photography by Geoff Jones
“Written primarily for the visiting birder, reading a copy of this book will fire up anyone’s desire to get out into the field to see more of our fabulous birds.”–Sean Dooley, Australian Bird Life


bookjacket Britain’s Butterflies:
A Field Guide to the Butterflies of Britain and Ireland

Fully Revised and Updated Third edition
David Newland, Robert Still, Andy Swash & David Tomlinson
“The images in this pocket-sized photo-guide are excellent and include pictures of eggs, chrysalids and caterpillars of all breeding species. Comparing very similar species can be difficult, but computer mock-ups helpful place specimen in situ. Clear text and page design make the book easy and fun to use.”BBC Wildlife magazine


bookjacket Britain’s Hoverflies:
A Field Guide

Revised and Updated Second edition
Stuart Ball & Roger Morris
Praise for the previous edition: “The latest field guild from the excellent Wildguides. . . . Beautifully and clearly laid out.”–Charlie Moores, Talking Naturally