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Home , Caterpillar, Parasitoid, Sonia (moth), Sonia Altizer, Thyrinteina

Jaap de Roode Jaap de Roode Sonia Altizer Jaap de Roode Tomato hornworms () are often attacked by Braconid (). larvae eat their way out of the and subsequently spin cocoons on the outside of the caterpillar. After the wasps emerge from their cocoons, the caterpillar may survive for a while, but will ultimately die before realizing its own .

When get Bugs: Top left: A healthy Monarch . Top right: Under natural conditions, Ophryocystis elektroscirrha rarely kills The ABCs of Lepidopteran Disease Monarchs before they turn into free-flying adults. However, captive conditions can result by Sonia Altizer and Jaap de Roode in contamination and a build-up of many parasite spores. When ingest large numbers of these spores, they often die as Introduction: flight muscles warm in the sun. Suddenly, On a cool morning in late winter, sunlight thousands of butterflies take flight in a bright Left: Heavily infected adults may become creeps into a high altitude forest in Central orange explosion, their jewel colored wings stuck to their chrysalis when they are trying to eclose. In such cases, neither the Monarch nor Mexico, illuminating the bodies of millions of stirring the air under a clear blue sky. A its parasites will be able to reproduce. Monarchs draped over the limbs of oyamel small group of tourists gasp with delight and trees. As temperatures warm, sunlight wraps spectacular disbelief. Soon the air is filled with Above: The parasite is spread to the next around motionless clusters of densely packed Monarchs, some landing on the heads and generation of caterpillars when females scatter butterflies suspended high in the forest canopy. shoulders of visitors standing near the colony’s parasite spores (shown at 400x magnification) Monarchs open and close their wings as their edge. onto their eggs. Jaap de Roode 16 American Butterflies,Summer 2010 17 Attack of the but they can also parasitize the larvae, eggs In the movieAlien , larval and pupae of other . Adult female wasps creatures entered human bodies where they have a needle-like ovipositor and can lay 20 or fed, grew, and eventually emerged, ultimately more eggs inside a single . Larvae killing their human host. One of the most of the wasps feed on the living caterpillar’s apparent types of parasites that attack tissues for over a week. The wasps can butterflies and has a life cycle that either pupate inside the host, or in the case follows this same script. Termed parasitoids, of braconid wasps parasitizing the tomato these parasitic and wasps are actually hornworm (see photo, page 16), form dozens a hybrid between a predator and a parasite. of white cocoons peppering the caterpillar’s Tiny as they might be, they pack a powerful back. A tiny wasp will eventually emerge from punch, almost always killing their host after each cocoon, leaving the caterpillar behind to successful infection. And on the scale of things die. that are totally disgusting yet simultaneously Parasitoids can employ incredible stealth fascinating, parasitoids rank pretty high up on in locating their prey, as illustrated the ladder. by two closely related egg parasitoids As one example of a group, ( sp) of Whites and miniscule wasps in the Braconid family European large whites (). Tiny comprise over 1700 North American . female parasitoids (smaller than a dandelion

Richard Bartz The wasps most frequently target caterpillars, seed) hitch a ride on the backs of mated female Adult braconid wasps are tiny, approximately 1/16 inch in length (actual size of wasp is shown by white line), and females have a long, slender ovipositor Parasitoid flies kill many Monarchs protruding from the abdomen.” Left: Adult tachinid flies lay eggs on the outside of Monarch caterpillars, after What the tourists cannot see, however, leaving little trace of infection. In other cases, which the maggot burrows its way into the caterpillar. is that thousands of these Monarchs are insects harboring pathogens might show infected by a debilitating protozoan parasite only subtle outward signs of infection, and Bottom left: After about a week, the called Ophryocystis elektroscirrha (page 17). disease-causing microbes can be hard to detect maggots will have grown large enough Everywhere the infected Monarchs land they without viewing squashed bits of insects under to eat their way out. This particularly scatter hundreds of parasite spores — on the a microscope and using advanced diagnostic unlucky caterpillar harbored no fewer than ground, on tree limbs, and even on people’s guides. 3 tachinid flymaggots. clothing, cameras and daypacks — leaving But the fact remains that butterflies, E. Sternberg microscopic souvenirs clinging in hidden like most other species, can be Bottom right: A maggot will then form a places. In April when the Monarchs return to infected by many parasites ranging from small brown pupa. their breeding grounds in the southern United viruses and bacteria to parasitic worms and Sonia Altizer States, some of the infected Monarchs will mites. Technically speaking, a parasite is join them, spreading the parasites to a new any that lives within and obtains generation of caterpillars. nutrients from another organism, called Most of us are well aware of the risks the host — usually to the host’s detriment. pathogens pose for human health, in part There are hundreds of parasite species that because of high profile news stories describing can spread among butterflies or caterpillars, outbreaks of emerging diseases like SARS, with major impacts on their development and influenza, Ebola, and West Nile virus. But survival. Examples in this article will illustrate even the most enthusiastic butterfliers don’t some of the ways that parasites can affect the often realize that butterflies can get sick, lives of free-living and captive lepidopterans, too. This is probably because in the wild, and how current human activities might be butterflies and caterpillars that die from altering the spread of butterfly parasites in disease can quickly disintegrate and disappear, some unexpected ways. Jaap de Roode 18 American Butterflies,Summer 2010 19 Alison Hunter (2)

Baculoviruses infect gypsy caterpillars. Left: A healthy gypsy moth caterpillar ingests viral particles (NPV) when feeding on contaminated leaf material.

Right: Within a few days after exposure, the caterpillar dies and rapidly disintegrates, leaving behind a puddle of viral particles and fragments of the caterpillar’s cuticle which disseminate Sonia Altizer Sonia Daniel Winkler viruses to the surrounding leaves. Fungal pathogens of . Above: Moth caterpillars killed by Cordyceps fungus sprout long stroma (finger-like also called nuclear polyhedrosis viruses mix of thread-like fungal hyphae and powder- projections) from their head capsules from (NPVs). These pathogens win the distinction like spores called conidia. The fungal spores which fungal spores are released. of behaving like the Ebola virus of the disperse in a puff of dust that showers the insect world (see photos, this page). Once a vegetation, where they wait to be contacted by Right: A Monarch caterpillar cadaver infected caterpillar unknowingly ingests NPV capsules, another caterpillar. Once a spore germinates with Beauvaria shows the characteristic growth the virus replicates massively inside host cell on the surface of its insect host, the thread- of white fungal mycelium and conidia. nuclei, killing the host rapidly by turning it like hyphae can penetrate the host’s skin and into a sac of jelly, from which millions of replicate inside the animal’s body cavity. viral occlusion bodies (protective capsules Some of the most bizarre-looking fungal butterflies. After the butterfly lays a clutch across North America found that up containing virus particles) then leak out onto pathogens are in the genus Cordyceps. of eggs, the parasitoid jumps off and lays its to 30% of Monarchs die from parasitoid attack surrounding vegetation. In part owing to Caterpillars infected by Cordyceps die after own eggs on or near the host’s. Amazingly, in some regions and years, with up to 10 their rapid lethality and host specificity (most their entire body cavity is filled by the fungal these parasitoids hunt down female Cabbage maggots emerging from a single Monarch NPVs infect a narrow range of Lepidoptera), hyphae; the fungus then sprouts finger-like Whites using an anti-aphrodisiac pheromone pupa. These high rates of parasitism suggest preparations of these viruses have been used projections from the cadavers (opposite page that male butterflies transfer to females upon that tachinid flies could be a major factor as effective biocontrol agents against several top) to help disperse its spores. In traditional mating. This pheromone signals other males to regulating wild Monarch populations. caterpillar pests, including gypsy moths Chinese medicine, Cordyceps has been used keep away from mated female butterflies, but in North America, velvet bean caterpillars for medicinal purposes for almost 1500 years, the same scent lures wasps to freshly mated Viruses and Fungi and Protozoa… Oh My (Anticarsia gemmatalis) in Brazil, and with the goal of maintaining healthy heart females. Although invisible to the naked , microbial African armyworms (Spodoptera exempta) in function, energy, positive outlook, immune Given that parasitoids are common and pathogens, including viruses, bacteria, and Tanzania. health and even sexual stamina. Current highly lethal, they can have major impacts on protozoa, can commonly infect butterflies. Fungi outnumber all other butterfly consumption in China and other countries has caterpillar survival in the wild. For example, These pathogens most often attack caterpillars pathogens by a factor of about 4:1. Instead driven down the abundance of wild-collected Monarchs in the are frequently (as opposed to other life stages), and routes of causing the to disintegrate into Cordyceps and led to increasing cultivation of attacked by the generalist parasitoid Lespesia of invasion tend to be through the accidental goo, fungal pathogens cause a light and fluffy this fungus by humans. It is interesting that a archippivora, a tachinid fly that has been ingestion of capsules or spores. One of the growth over the outside of the animal, giving pathogen highly lethal to caterpillars is viewed reported to infect caterpillars from 14 different most common and lethal groups of caterpillar it a marshmallow-like appearance (see photo, as promoting the health of humans! families (see page 19). One study of fly infections are caused by baculoviruses, middle of opposite page). That white fluff is a 20 American Butterflies,Summer 2010 21 Scott O’Neill faced a 2-4 fold higher probability of extinction than uninfected populations due to the impacts of this pathogen during the invasion phase. A practical guide to preventing disease when Why do Parasites Cause Disease or Kill rearing butterflies at home their Hosts? Because of the disease they cause and the Many people enjoy bringing caterpillars killing they do, parasites are not often seen from their gardens indoors to pupate as friendly creatures. But from the parasite’s and emerge as adult butterflies. But point of view, all that matters in life is to occasionally, these caterpillars harbor produce offspring that can infect new hosts - infectious parasites including bacteria, even if this requires killing its current host. In viruses or protozoa. this regard, it makes sense for baculoviruses Sick caterpillars might appear to turn caterpillars into the sticky goo that lethargic, pale, and may stop eating contains millions of virus offspring. This is or fail to molt properly — although because the virus particles that ooze onto leaf some animals harboring low levels material can be ingested by new caterpillars, of infection could appear identical to causing the next generation of infections. And healthy caterpillars. parasitoids can usually only grow big enough Importantly, many parasite species Transmission electron micrograph of Wolbachia bacteria developing within an insect cell. to form a cocoon by eating so much caterpillar can produce millions of propagules tissue or blood that the caterpillar will die under the right conditions, and it often before completing its own metamorphosis. takes only one of these to cause a Wolbachia: Harmless Symbiont or Sexual bolina. In an interesting twist of events, But not all parasites kill their hosts. The new infection. In nature, the chances Sabotage? between 2004-2006, scientists discovered a protozoan Ophryocystis elektroscirrha, for of parasite propagules reaching Bacteria in the genus Wolbachia are some dramatic recovery of males on some of the example, spreads to new hosts when female new caterpillars are low, but captive of the most common microbes infecting islands, indicating that these butterflies had Monarchs lay eggs onto which the parasite conditions often facilitate the spread of Lepidoptera and other insects. These evolved resistance to Wolbachia’s male-killing spores are scattered. These parasites are parasites: parasites can easily build up maternally-transmitted bacteria don’t kill properties. eaten by caterpillars hatching from these in rearing chambers and thereby cause their hosts directly, but can manipulate Given the widespread nature of eggs. Because this parasite needs its host to unwanted infection of caterpillars reared insect reproduction. Depending on the host these endosymbiotic bacteria and their reach adulthood and lay eggs, killing the host side-by-side with an infected animal. species affected, Wolbachia can destroy ancient associations with insects and other prematurely would be a bad idea, and this Because some parasite spores can male embryos, turn males into females, invertebrates, it is tempting to assume that parasite rarely kills Monarch caterpillars or live for many months after a butterfly or cause mating incompatibility between Wolbachia do not pose a significant risk pupas under natural conditions. This is not to is reared and released, a single infected males and females with unrelated strains. for wild insect populations. However, these say that O. elektroscirrha does not damage caterpillar could continue to infect These manipulations give the bacteria a parasites can reduce host reproduction during Monarchs. Indeed, the parasite needs to multiple generations of butterflies. distinct advantage in spreading through host the invasion phase, and can also lower host produce millions of offspring to successfully One remedy to deal with such populations, and also cause problems for some genetic diversity due to extreme sex ratio bias. transmit spores to Monarch eggs, and this contaminations is to regularly sterilize butterfly species. In African populations of This risk was recently evaluated for reproduction means that infected adult (with 10% bleach solution) rearing the White-barred Acraea (Acraea encedon), ‘Karner’ Melissa Blues. Wolbachia that Monarchs don’t live as long or fly as well as chambers, tools and gloves that are used for example, 70-100% of the butterflies were closely related to strains from the non- uninfected butterflies. Moreover, in captivity, for rearing caterpillars or housing pupas are infected with Wolbachia. At sites with endangered western populations of Melissa the parasite can often build up to high numbers and butterflies. It is also wise to quickly high infection rates, the proportion of males Blues were found to be common in the (unless careful measures are taken to prevent remove any sick-looking caterpillars is exceedingly low, most females will die western range of ‘Karner’ Melissa Blues, transmission) and thereby cause damaging and individuals to avoid the maturation before mating, and females have taken to suggesting that a recent transmission event infections in captive-bred Monarchs. of parasites and their subsequent spread. mobbing the few remaining males. Extreme occurred between the two sub-species. A To facilitate their own transmission, loss of males has also occurred in Polynesian computer analysis showed that Wolbachia- many parasites have evolved the ability populations of the butterflyHypolimnas infected ‘Karner’ Melissa Blue populations to manipulate their hosts’ behavior.

22 American Butterflies,Summer 2010 23 E. Sternberg directly. Parasites that are too large to be might affect their interactions with parasites engulfed by immune cells, such as parasitoid and other natural enemies. In general, at eggs, can be encapsulated by aggregations of least three types of human activities might hemocytes and through melanization, which impact butterfly-parasite interactions: habitat involves the deposition of melanin pigment, destruction, climate warming, and releases of effectively producing a dark capsule around farmed butterflies into the environment. foreign material (see photo, opposite page, top First, global climate change may impact right). butterfly diseases by affecting pathogen Insects can also employ physical and development, survival rates of hosts and behavioral defenses to avoid parasites. parasites, disease transmission processes, and Gypsy moth caterpillars, for example, can host susceptibility. Because harsh winters can actively avoid feeding on leaves contaminated cause the die offs of pathogens that might with virus particles and hence avoid hang out in the environment, a question raised

Sonia Altizer Sonia infection. There is also evidence that arctiid in recent years is “will a warmer world be moth caterpillars can increase their body a sicker world?” With less seasonality and Butterfly immune defenses temperature by basking in the sun, and this warmer temperatures, will pathogens survive may help them fight their parasitoid flies. better and be transmitted over longer periods Above left: Butterflies have hemocytes (immune cells) in their blood; shown here are Much like humans have mastered the of time? plasmatocytes, one type of immune cell that helps to encapsulate foreign material. art of herbal medicine, moths and butterflies Moreover, habitat destruction and also seem able to use nature as an apothecary fragmentation could crowd some insect Above right: To measure encapsulation activity, sephadex beads (shown against their bugs. African armyworms, for populations into smaller habitat patches, unencapsulated as red bead) were implanted into a Monarch caterpillar. The next example, can increase their dietary intake of potentially increasing disease transmission, day, beads were extracted, showing the deposition of black pigments onto them. proteins, which they use to fuel their immune and exposure to or other Caterpillars can use such pigments to encapsulate parasitoid larvae and other types of response against bacterial pathogens. And environmental stressors could increase parasites. See the article on page 27 to learn how many wasps avoid this defense. woolly bear caterpillars can extract toxic susceptibility to infection. In terms of habitat chemicals from their host plants to fight changes, for many butterfly species, humans their parasitoid flies. Recent studies in our have converted habitats in open meadows or own labs have also shown that Monarch prairies or coastal areas into suburban/urban Baculoviruses and fungal pathogens can Butterflies Fight Back caterpillars that feed on certain milkweed developed areas or crop systems. cause summit disease, a syndrome that makes Caterpillars and butterflies can warn or trick species containing high levels of cardinolides, This is especially true with the increasing caterpillars climb to high vegetation. Here the predators by using camouflage, false body pathogens will kill the host and produce spores parts and warning coloration, but how do or capsules that can be carried far and wide by they fight off microbial diseases? When the wind to infect new caterpillars. One of the people get sick, our immune systems mobilize “global climate change may impact butterfly diseases ” most bizarre examples of host manipulation rapidly to fight off infection, and humans comes from a braconid wasp parasitoid of the can engage in behaviors like hand washing geometrid moth Thyrinteina leucocerae. In and covering our mouths during a sneeze such as tropical milkweed ( popularity of roundup ready crops that are this case, most parasitoids eat their way out or cough to limit disease transmission. Like currasavica), experience less severe infections genetically engineered to be herbicide tolerant, of the caterpillar when they are ready to form people, butterflies and other insects can deploy by the parasite Ophryocystis elektroscirrha. so that agricultural weeds that butterflies a cocoon, which they do on the vegetation several lines of defense to protect themselves Interestingly, female butterflies appear able might otherwise use for food or shelter are outside of the caterpillar. However, a number against infectious agents. In terms of immune to take advantage of these curative plants much more effectively eliminated. Butterflies of wasps stay behind inside the caterpillar defenses, antimicrobial proteins float around by preferentially laying their eggs on them that are crowded into remnant habitats could and apparently manipulate it to become in insect blood, and attach to (and mark or to reduce the disease symptoms of their encounter more for resources — a bodyguard of the cocoons: the infected attack) extracellular pathogens. Insects also offspring. and possibly higher spread of parasites. caterpillar fends off predators of the wasp have immune cells called hemocytes that As a final issue, the growing popularity of cocoons with violent head-swings and finally operate similar to our white blood cells (see Butterfly Diseases in a Changing World raising and releasing insects such as ladybird dies when the wasps have emerged from their photo, top left). Some of these hemocytes can Humans are currently changing the climate , , and butterflies has resulted in cocoons and no longer need the caterpillar. engulf (or phagocytize) microbial parasites and habitats of butterflies in ways that the rising commercial sales of many insect 24 American Butterflies,Summer 2010 25 species. Butterflies in particular are sold in nature. At the same time, biologists have for educational activities or for release at uncovered only a miniscule percentage of the weddings and other special events. In the diversity of infectious from natural United States alone, as of 1998, more than 60 host communities, and nowhere is this gap and Parasitoids: commercial butterfly farms were in operation, more extreme than for pathogens from the vast with an estimated tens of millions of dollars majority of butterflies. Not only is describing The Ultimate Virus-Host Relationship spent annually on butterfly sales. Introducing butterfly pathogens important for inventories large numbers of captive raised insects into of biodiversity, but baseline data on the by Don Stoltz the wild raises concerns for the spread of distribution and prevalence of insect pathogens

In the article “When Butterflies get Bugs” over the past 40 years in my laboratory and in “The time is ripe to reexamine regulations on by Sonia Altizer and Jaap de Roode, on page the laboratories of other scientists who became 16 of this issue of American Butterflies, you interested in this fascinating system. commercial growers who sell live butterflies for learned about the many diseases of butterflies. When a braconid wasp encounters the release into the environment.” It might be a surprise for most people to right kind of caterpillar, it tries to inject its learn that for every lepidopteran species, eggs into the caterpillar using a specialized there exists one or more species of parasitic structure called an ovipositor (many of you pathogens, especially because rearing animals are essential to detecting future changes in wasp (parasitoid) which attacks it. Here, we will have experienced the effects of a certain at high densities in commercial operations prevalence or novel pathogen introductions take a look at a very unusual relationship modification of the ovipositor, namely the could increase their exposure to some diseases. that might result from anthropogenic change. between viruses belonging to the family, stinger!). If it succeeds, it also injects some Although the USDA regulates the This is especially true for butterfly populations Polydnaviridae, and certain species of wasp fluid, and this fluid contains interstate shipment of live butterflies, existing subject to habitat loss and exposure to belonging to the well-known wasp families, particles. The recognition that virus particles permits do not track the number of butterflies environmental stressors that could make and . were injected along with the eggs was made in transported nor do they require the butterflies species more susceptible to disease-mediated Those of you who keep a garden, and the mid 1970s in my laboratory. to be screened for disease. Both NABA and declines or extinction. avoid the use of chemical pesticides, are likely In a strange twist, however, the virus the Xerces Society have proposed a ban on the to have observed Cabbage White caterpillars particles that have been examined don’t environmental release of commercially reared festooned with small white cocoons — these seem to contain any viral DNA; rather they butterflies, with the spread of parasites and will have been spun by larvae of the braconid contain wasp DNA. Once the polydnavirus diseases listed as one major concern. Given For Further Reading parasitoid, Cotesia rubeculae. Wasps in the particles are within the caterpillar, the wasp the growing popularity of butterfly releases, Tanada, Y., and H.K. Kaya. 1993. Insect Pathology. genus Cotesia belong to a lineage (termed genes contained within them spring into lack of required screening for parasites, and Academic Press, San Diego, California. the microgastrine lineage), comprising more action and prevent the caterpillar’s immune potential for cross-species transmission in than 10,000 species worldwide. As far as system from destroying the wasp eggs. But, operations where multiple butterfly species Federman, A. 2008. All aflutter: The flap over the we can tell, every species within this very the DNA released into the caterpillar cells are reared together, the time is ripe to mail order butterfly industry. Earth Island Journal, large and successful lineage of wasps carries from the polydnavirus particles doesn’t reexamine regulations on commercial growers Fall 2008. http://www.earthisland.org/journal/ a polydnavirus. Polydnaviruses are also replicate. In the 1990s, the DNA packaged index.php/eij/article/all_aflutter/ who sell live butterflies for release into the associated with at least two ichneumonid into certain polydnavirus particles was Highly recommended! environment. subfamilies. Polydnaviruses carried by completely sequenced. One of the results of It is important to note that before scientists Hurst, L.D. and Randerson, J. 2002. Parasitic sex braconid wasps are called bracoviruses while this work was the realization that the genes can predict how butterfly pathogens will puppeteers. Scientific American. April 2002. Pp. those carried by ichneumonids are called required to make progeny virus particles respond to future environmental changes, we 56-61. ichnoviruses. In terms of morphology, the were not packaged into said particles! So, need baseline information on how common bracoviruses and the ichnoviruses are very the polydnaviruses are in the unique position butterfly pathogens are in wild populations different (see photos, pages 28 and 29), of being unlike any other kind of virus, to — and how butterfly diseases are affected by however their life cycles are essentially the point that some questioned whether they factors like climate and habitat characteristics. identical and remarkable! should legitimately be called viruses. Parasites and infectious diseases comprise a Here is a somewhat simplified description Inevitably, the debate centered on the of the entwined life histories of the wasp and question of origins, for which there were two major fraction of the biodiversity on Earth, July 5, 2009, Manion Corners, Ontario, Canada. Same individual as photo 23. with half or more of all species being parasitic the virus. Much of this story was worked out possibilities: one, certain wasps had evolved 26 American Butterflies,Spring 2010 27 Contributors , then in his own lab at Emory. Until he knew that butterflies Sonia Altizer is an carry interesting parasites, Jaap never used a Associate Professor butterfly net. in the Odum Leslie Ries is Don Stoltz is School of Jeffrey Glassberg’s brief biographical sketch a conservation originally from at the University appeared in the Spring 2010 issue of American biologist who southern Ontario, of Georgia. Butterflies. has worked where as a boy he She received largely on collected all sorts her Bachelor’s Kimberly Pegram the impacts of things, including degree from Duke is a graduate of landscape insects. He later University and a student working structure on pursued a career in Ph.D. from the in Ron Rutowski’s habitat quality. science, and in that University of lab at Arizona All of her past context has been Minnesota, where she was co-advised by State University. field work able to indulge all of Karen Oberhauser. For the past 15 years, She currently has focused his favorite interests (entomology, virology Sonia has studied Monarch behavior, ecology, studies the evolved on butterflies. Leslie was an undergraduate and microscopy). In 1974, he moved to and interactions with a protozoan parasite, design features of at University of Maryland (Zoology). She Dalhousie University, Halifax, where he began asking how seasonal migration of these warning coloration received an M.S. in Ecology, Evolution and to characterize some unusual parasitoid- butterflies affects parasite transmission. In using Pipevine Behavior from Iowa State University and associated viruses; this work eventually led to 2006, she launched the citizen science project Swallowtails. a Ph.D. in from Northern Arizona the establishment of a new family of viruses MonarchHealth, now in its 5th year, and also Kimberly graduated from the University of University. She is currently a post-doctoral (Polydnaviridae). Now officially retired, he’s maintains a webpage dedicated to Monarch Mary Washington in Fredericksburg, VA associate at the University of Maryland where nonetheless more likely to be found in his lab parasites (www.monarchparasites.org). Sonia where she did undergraduate research on she is working on the NABA 4th of July than anywhere else. In his spare time, Don is a founding member of the MonarchNet Painted Ladies with Joella Killian. Also as Butterfly Count dataset. takes delight in stalking neotropical insects steering committee, a group dedicated an undergraduate, she carried out surveys of with a camcorder. to integrating, analyzing and publicizing butterflies on Plummers Island with John W. Ron Rutowski has Harry Zirlin’s Monarch monitoring data. Brown at the Smithsonian’s National Museum been a Professor of fascination of Natural History as part of their Research Biology at Arizona with butterflies Training Program. State University since and other Jaap de Roode 1976 and has authored insects dates works at Emory Mike Reese updates over 80 publications to his early University, where the NABA Recent that have helped childhood. he runs a research Sightings web keep butterflies in He travels lab on parasites and pages. He enjoys the mainstream of extensively teaches ecology and photographing behavioral ecology. in the United evolution. Born in wild flowers, birds, His research has States, studying and photographing butterflies. the Netherlands, dragonflies, and, of focused on vision and He supports this passion by working as an he studied course, butterflies. visual signals in butterflies especially in the attorney in the environmental area at the New population biology He is an educator in context of mate choice and mate detection. York law firm of Debevoise & Plimpton. at Wageningen Wautoma, Wisconsin University. He then moved to the University and has been recording of Edinburgh to study malaria, which sparked and documenting the butterflies that are found a keen interest in disease. In 2005 he moved there for over 15 years. He also maintains a to the United States to study the parasites website on the Butterflies of Waushara County, of Monarchs, first with Sonia Altizer at the Wisconsin. 48 American Butterflies,Summer 2010 49