Identification of fuzzy insect in Pennsylvania

I live in south eastern Pennsylvania, USA. About a month ago I came across this very peculiar insect. Its body was about an inch long, it was fuzzy, and it had a long proboscis(?) and long orange legs. I've been wondering about it ever since. Can anyone identify it?

This is the nymph of a bug (Hemiptera) from the Genus Arilus, probably Arilus cristatus, also known as "wheel bug".

Here is another image of it, for comparison:


And here is the adult:


Identification of fuzzy insect in Pennsylvania - Biology

Scientific Name: Photuris pennsylvanicus
Common Name: Pennsylvania Firefly

(Information in this species page was gathered in part by Ms. Megan McAuley for an assignment in Biology 220W (Spring semester 2007)).

The Pennsylvania firefly (Photuris pennsylvanicus) (also called the &ldquoLightning bug&rdquo) is a cherished feature of warm Pennsylvania summer evenings. Its pulsing pinpoints of yellow and green light make the dark woodlands, fields, and gardens come alive with movement and possibilities. The firefly was named the state insect of Pennsylvania in 1974.

Classification and Appearance
The &ldquofirefly,&rdquo though, is not really a fly at all. It is a beetle in the family Lampyridae that, along with several hundred other closely related &ldquofirefly&rdquo species, has the remarkable ability, in all of its life stages, to biologically generate light. The adult beetle, which is the form most familiar to people, is ½ to ¾ inch in length, with a flattened body that is predominately black in color with yellow highlights and prominent red spots on the back of its thorax. It has large eyes and long antennae and flies in a gentle, hovering manner. The light generating parts of these adults are in the terminal segments of their abdomens. The adult firefly has long, curved mandibles that suggest a predaceous life style, but only a few species have been shown to actually consume anything other than flower nectar or pollen. The less well known larvae of the firefly, called &ldquoglow worms,&rdquo live in leaf litter and are voracious predators. They eat other insects, mites, earthworms, and even slugs and snails.

Using Light for Communication
The lights of the fireflies represent communication mechanisms. The female fireflies, which are predominantly sessile, perch on the vegetation in its habitat and generate a species specific sequence of light flashes that attract the much more mobile males. The males respond with an answering light sequence and zero in on the females in order to mate. A few species of firefly have been shown to mimic the light sequences of other species in order to draw unsuspecting males to waiting, predaceous females. These females not only gain energy from consuming the males of these other species but also can accumulate chemicals from their prey which help to protect them from their own predators. This behavior is called &ldquoaggressive mimicry.&rdquo

After mating in the late summer, the females lay their eggs one at a time on the surfaces of woody or leaf debris. The eggs hatch in a few weeks and the emerging larvae enter the soil/litter habitat where they actively feed on a wide range of invertebrates. In late fall, the larvae burrow into the soil or under the bark of woody stems where they overwinter. In the spring, they re-emerge and continue to actively feed on their diverse array of prey species. After a few weeks, they re-enter the soil and pupate. They then emerge from their pupal chambers in early to mid summer as adult fireflies.

Light Production
The mechanism for the production of light in fireflies is mediated by the enzyme &ldquoluciferase.&rdquo High energy phosphates generated from food molecules are coupled via luciferase to the direct production of photons of light. This coupling is extremely efficient (90%+) and generates almost no waste energy (heat). The genes that regulate this light generation have been used in cancer research to mark and track metalizing cancerous cells.

There have been many reports of declining numbers of fireflies throughout North America. The widespread use of pesticides and herbicides, the loss of the leaf litter habitat required by larval life stages especially in suburban areas, and drought have all been proposed as factors in the decline of the firefly. It is hoped that this decline can be reversed so that we can all continue to have the pleasure of observing these unique and wonderful organisms.

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1. Spotted Lanternfly

U.S. Department of Agriculture, Flickr // Public Domain

A relative newcomer, the spotted lanternfly arrived in the U.S. from northern China in 2014. Since its discovery in Berks County, Pennsylvania, the insect species has spread to neighboring states in the mid-Atlantic. It’s known for its large, black-spotted gray wings and its destructive behavior. Spotted lanternflies eat sap from more than 70 different plant species, including important crops like grapevines, maple trees, and black walnut trees. Lanternflies can lay up to 200 egg masses on one host plant, and the bugs’ sugary secretions have been known to promote mold growth. According to the Pennsylvania Department of Agriculture, the insect could cost the state $324 million a year if it isn’t controlled.

The Woolly Bear Caterpillar is a furry type of caterpillar with black and orange or brown hairs

The fuzzy black and brown Woolly Bear caterpillar (Pyrrharctia isabella) is one of the most common caterpillars you will see in late summer. You can easily identify this furry caterpillar by the wide brown or orange band around its middle and black ends.

Black and brown caterpillars such as the Woolly Bear aren’t poisonous or a stinging variety. Usually, handling one of these fuzzy worm-like creatures with their spiky tufts of hair may cause skin irritation or contact dermatitis.

One characteristic of the Woolly Bear caterpillar is its defense mechanism. When under threat, the caterpillar rolls up into a spiky ball. When the threat has gone, they quickly crawl away to safety.

Also called the Isabella Tiger Moth caterpillar, this spiky looking insect feeds on herbs, tree leaves, and other plants.

Identifying features

Short spiky tufts of brown/orange and black hairs cover this species of furry caterpillar.

One of the larger types of black furry caterpillars that grows up to 2.3” (6 cm) in length.

Identification of fuzzy insect in Pennsylvania - Biology

Common Name: Tree of heaven, Chinese sumac, stinking sumac, varnish tree

Scientific Name: Ailanthus altissima


Phylum or Division: Magnoliophyta
Class: Magnoliopsida
Order: Sapindales
Family: Sunaroubaceae

Identification: Ailanthus altissima is the only species introduced into North America of the Quassia family, which includes five native trees and two native shrubs. It is a large deciduous tree that grows up to 80 feet rapidly from a straight, gray trunk. The leaves are pinnately compound, with up to 41 leaflets spaced alternately on the one-to-four-foot leaf veins (see illustration). At the base of each leaflet are one or two teeth. Green above and silvery below, the leaflets are fuzzy when young oil glands at the base produce a foul smell when the leaves are crushed. Seedpods are reddish brown, produced in late summer. Each is between 1 ½ to 2 ½ inches long and twisted like a propeller, each with one seed. Bark is gray, smooth to bumpy and it becomes fissured with age. Like the leaves it is hairy when the tree is young. Trees are unisexual and bisexual with small, half-inch five-petaled yellow-green flowers in dense terminal clusters. Male flowers have a disagreeable odor, which has led to one of the plant’s common names, stinking sumac. Twigs are also hairy and produce a round, wide, open crown. Both the bark and wood contain astringent chemicals.

Original Distribution: Northern and central China

Current Distribution: In the United States, 41 continuguous continental states, except Minnesota, Montana, New Hampshire, North Dakota, South Dakota Vermont, and Wyoming, as well as Hawaii.

Site and Date of Introduction: Introduced into Philadelphia, Pennsylvania, by William Hamilton in 1784 for his garden. On the West coast it was also introduced for medicinal purposes by Chinese immigrants in the Gold Rush of the 1850's.

Mode(s) of Introduction: Hamilton brought a specimen from England by ship for ornamental purposes the Chinese also brought their samples with them by ship.

Reason(s) Why it has Become Established: A. altissima is a "classic weed" species (Stobel, 1991): It reproduces at a rate of 325,000 seeds per tree per year. Its winged seeds disperse easily, often taking advantage of the wind-tunnel effect of roadways, but the plant also spreads by aggressive suckers, or runners. The root system can push into cracks in sidewalks and building foundations (see photo). It can establish itself in the poorest of ecological environments. This capacity led to its becoming a metaphor for the resilience of poor families in large cities, as expressed in A Tree Grows in Brooklyn by Betty Smith. Not only does it grow in the poorest soils, but it also tolerates air pollution well.
The tree of heaven grows swiftly to become a large tree, growing four feet per year and blocking the light for native species beneath it. In addition, its leaves are toxic to over 40 native species of plants, and it is unpalatable to herbivores. Ailanthus can overwinter in northern climates, and is resistant to both frost and drought, giving it a clear competitive edge over native species.
Because of these attributes, A. altissima has been dubbed the "tree of hell" by Pennsylvania state botanists.

penetrating building foundation City block invaded by Ailanthus

Ecological Role: A. altissima prevents erosion, and it provides shade and roosts for nesting birds. As a pioneer species, it grows in environments where other species of plants cannot.

Benefit(s): The tree of heaven is believed to have medicinal value in Asian traditional medicine as a remedy for asthma, worm infestation, vaginal discharge, diarrhea, and mentrual cramps. In Africa, it is used as a treatment for heart problems, seizures, and menstrual discomfort. In France, the leaves of the tree of heaven are used instead of mulberry leaves for feeding silkworm moths. The wood of A. altissima may be used in crafts and woodworking. The toxin produced in the leaves, bark, and wood is currently under study as a natural herbicide source.

Threat(s): The tree of heaven is a dominant tree that crowds out native species by its fast reproduction and aggressive range expansion its suckers choke out native seedlings, greatly reducing local biodiversity. It invades urban spaces, blocking sunlight, views, and breaking up pavement and building foundations and blocking plumbing and sewer systems. In fact, in cities, this weed will grow virtually everywhere.
In the countryside, it is currently seen in fields, woodland periphery. and untended open spaces, preventing the propagation of native species. In agricultural environments, the tree of heaven pollutes fodder and damages farm equipment. It proliferates in sections of forest opened by gypsy moth predation.
Despite threats to native species, Alianthus seeds are still readily available in nurseries and seed catalogs in the United States.

Control Level Diagnosis: A. altissima is classified as an invasive, nonnative plant in many states.

Control Method: The entire area colonized by the tree of heaven must be treated over a long period of time for effective eradication. Young seedlings must be dug up, root suckers must be destroyed by removal or herbicide cutting alone is ineffective as this action stimulates the stump to send out new suckers and sprouts. Careful application of plant poison to the bark in late winter is often effective on trees less than six inches in diameter specific herbicides are listed on the National Park Service website.
Any method must be diligently and continuously pursued to eradicate this invasive weed. For the occasional stubborn individual tree, the Division of Forestry recommends flame-throwers and bulldozers as a means of permanent removal.
Natural control methodds are limited. The tree of heaven is preyed upon by very few insects due to the chemicals in its wood and bark. The fungus Verticillium albo-antrum kills A. altissima, but unfortunately the fungus spores remain in the area of the dead tree and will kill many species of native trees that might germinate in the infected location.

Photo Credits: Mary Knight illustration by Patricia J. Wynne

Author: Patricia J. Wynne
Last Edited: December 9, 2002

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Insect Conservation Biology

Up until the mid-1980's, New York's state insect, the native ladybird beetle, Coccinella novemnotata (C-9) was the most common lady beetle (coccinellid) in the northeastern U.S. This relatively large (5-7 mm) species ranged across the U.S. and through Southern Canada and was an important biological control agent in gardens and crops in the northeast. Collections of C-9 declined through the mid to late 1980's. The latest reported collection in the northeastern U.S. was 1992, although C-9 may have persisted beyond this date in low densities. C-9 could eat many different species of aphid and could live in many different crops including alfalfa, clover, corn, cotton, potatoes, soybeans and arboreal habitats (Harmon et al 2007).

The historically broad geographic range of C-9 stands in stark contrast to its current range. An extensive USDA APHIS coccinellid survey in 1993 found no C-9 in eleven Northeastern states. This cooperative study focused on 100 counties and was based on comprehensive fieldwork and data from personal collections. Based on the latest records in the literature, C-9 was last collected in Maryland in 1986, Pennsylvania in 1987, Delaware in 1988 and Maine in 1992. A coccinellid study several years in duration in Kentucky found no C-9 in corn, tobacco, tomatoes and soybeans. Declines in C-9 populations in Alabama and Mississippi have been recorded since the early 1990's, so there is little reason to believe that C-9 will not continue to disappear from its current range.

There are many questions to be answered on the disappearance of this native ladybird beetle, including the possibility that introduced ladybirds have excluded it from habitats that it once favored. The timing of the extirpation of this native species overlaps with the arrival and establishment of its congener, Coccinella septempunctata or C-7. Making a definitive statement about the effects of C-7 on C-9 is difficult as no data were taken as C-7 expanded its range and C-9&rsquos contracted. Since then, three other introduced species have established, Harmonia axyridis, Propylaea quatuordecimpunctata and Hippodamia variegata. These species are particularly voracious, but it is difficult to quantify this factor in an ecologically realistic setting. Coccinellids are frequently parasitized by Dinocampus coccinellae, a braconid wasp. Although this wasp is native to the Northeast, it is possible that introduced species carried in more wasps upon introduction, potentially altering parasitoid-host dynamics for native species. Cropping patterns and loss of agricultural land may have played a prominent role as well.

Fortunately, although C-9 has declined precipitiously some recent discoveries indicate that it continues to persist. Jilene (age 11) and Jonathan (age 10) Penhale found a nine spotted ladybug near their home in Virginia in October 2006 (read more about this discovery). This is the first C-9 seen in the eastern U.S. in 14 years. Their finding confirmed that the species is not extinct and gave specialists a place to start some intensive hunting. Recent sitings were also recorded from Alberta, Canada (Ladybugs of Alberta, John Acorn, 2006) and Nebraska in 2007 (Scott Black, Xerces Society, 2007). A new citizen science project has been launched at Cornell (the Lost Ladybug Project) to educate the public on the importance of biodiversity and conservation and to recruit them to join in the effort to document the current status of C-9 and other rare ladybug species.

Resources: See Harmon et al. 2007 (J. Insect Cons. Bio. 11:85-94) and essay in Wings

Selected Publications

Calvin, D. D., Keller, J., J. Rost, B. Walsh, D. Biddinger, Hoover, B. Treichler A. Johnson, and R. T. Roush. 2021. Spotted lanternfly (Hemiptera: Fulgoridae) nymph dispersion patterns and their influence on field experiments. J. Econ. Entomol., in revision.

Tan, C-W., M. Peiffer, A. Jones, J. G. Ali, R. J. Schilder, Hoover, C. Rosa, and G. W. Felton. 2021. Stung by a wasp: Multitrophic effects of a parasitoid in a non-permissive host. Nature Communications, in review.

Uyi, O., J.A. Keller, and Hoover. 2021. Performance and host association of spotted lanternfly (Lycorma delicatula) among common woody ornamentals. Sci. Reports., in revision.

Mason, C.J., Felton, 2021. Effects of maize (Zea mays) genotypes and microbial sources in shaping fall armyworm (Spodoptera frugiperda) gut bacterial communities. Sci. Reports

Mason, C.J., K. Rubert-Nason, D. Long, R.L. Lindroth, J. Shi, and Hoover. 2021. Salicinoid phenolics reduce adult Anoplophora glabripennis (Cerambicidae: Lamiinae) feeding and egg production. Arthropod-Plant Interactions 15(1): 127-136.

Mason, C.J. A. St. Clair, M. Peiffer, E. Gomez, A. Jones, G.W. Felton and Hoover. 2020. Diet influences proliferation of gut bacterial populations in herbivorous caterpillars. PLoS ONE 15(3): e0229848.

Keller, J., J. Rost, K. Hoover, J. Urban, H. Leach, M. Porras, B. Walsh, M. Bosold , and D. D. Calvin. 2020. Dispersion pattern and sample size estimates for spotted lanternfly, Lycorma delicatula (Hemiptera: Fulgoridae) egg masses. Environ. Entomol. 49(6): 1462-1472.

Keller, J.A., A.E. Johnson, O. Uyi, S. Wurzbacher, D. Long, and Hoover. 2020. Dispersal of Lycorma delicatula (Hemiptera: Fulgoridae) nymphs through contiguous, deciduous forest. Environ. Entomol., DOI: 10.1093/ee/nvaa107.

Uyi, Osariyekemwen, J. Keller, A. Johnson, D. Long, B. Walsh, and Hoover. 2020. Spotted lanternfly can complete development and oviposit without access to the preferred host, Ailanthus altissima. Environ. Entomol. DOI:10.1093/ee/nvaa083.

Paudel, S., P.A. Lin, K, Hoover, G.W. Felton and E.G. Rajotte. 2020. Asymmetric responses to climate change: Temperature differentially alters an herbivore salivary elicitor and host plant responses to herbivory. J. Chem. Ecol. DOI 10.1007/s10886-01201-6.

Trotter, R.T., S. Limbu, K. Hoover, H. Nadel and M.A. Keena. 2020. Region-specific agent-based phenology models for the Asian gypsy moth (Lymantria dispar asiatica and dispar japonica) in East Asia ports. Special Collection: Geospatial Analysis of Invasive Insects (J. Morisette and K. Macaluso, eds.). Annals of the ESA,

Mason, C.J., S. Ray, I. Shikano, M. Peiffer, A.G. Jones, D.S. Luthe, K. Hoover and G.W. Felton. 2019. Plant defenses interact with insect enteric bacteria by initiating a leaky gut syndrome. PNAS 116(32): 15991-15996.

Tan, C-W, M. Peiffer, K. Hoover, C. Rosa and G.W. Felton. 2019. Parasitic wasp mediates plant perception of insect herbivores. J Chem. Ecol. .

Pan, Q., I. Shikano, G.W. Felton, T-X. Liu, and K. Hoover. 2020. Host permissiveness to baculovirus influences time-dependent immune responses and fitness costs. Insect Science https://DOI:10.1111/1744-7917.12755.

Jones, A.G., K. Hoover, K. Pearsons, J.F. Tooker, and G.W. Felton. 2019. Potential impacts of translocation of neonicotinoid insecticides to cotton (Gossypium hirsutum [Malvales: Malvaceae]) extrafloral nectar on parasitoids. Entomol. DOI: 10.1093/ee/nvz157.

Mason, C.J., S. Ray, I. Shikano, M. Peiffer, A.G. Jones, D.S. Luthe, K. Hoover and G.W. Felton. 2019. Plant defenses interact with insect enteric bacteria by initiating a leaky gut syndrome. PNAS 116(32): 15991-15996. (PDF)

Jones, A.G., K. Hoover, K. Pearsons, J.F. Tooker, and G.W. Felton. 2019. Potential impacts of translocation of neonicotinoid insecticides to cotton (Gossypium hirsutum [Malvales: Malvaceae]) extrafloral nectar on parasitoids. Environ. Entomol. DOI: 10.1093/ee/nvz157. (PDF)

Hoover, K. 2019. Editorial overview: Insect behavior and parasites: From manipulation to self-medication. Current Opinion in Insect Science 33: vi-viii.

Hoover, K. (guest ed.). 2019. Behavioural Ecology: Insect behavior and parasites: From manipulation to self-medication. Current Opinion in Insect Science 33: 1-69 (Denlinger and Casas, eds.). (PDF)

Pan, Q., I. Shikano, K. Hoover, T-X Liu, G.W. Felton. 2019. Pathogen-mediated tritrophic interactions: Baculovirus-challenged caterpillars induce higher plant defenses than healthy caterpillars. J. Chem. Ecol. 45: 515-524. (PDF)

Hall, L., A.J. Myrick, F. Graves, K. Hoover and T. Baker. 2019. Labial and maxillary palp recordings of the Asian longhorned beetles, Anoplophora glabripennis, reveal olfactory and hygroreceptive capabilities. J. Insect Physiol., (PDF)

Mason, C.J., A.M. Campbell, E.D. Scully and K. Hoover. 2018. Bacterial and fungal midgut community dynamics and transfer between mother and brood in the Asian longhorned beetle (Anoplophora glabripennis), an invasive xylophage. Microbial Ecology 77: 230-242. (PDF)

Pan, Q., I. Shikano, K. Hoover, T.-X. Liu, and G.W. Felton. 2018. Enterobacter ludwigii, isolated from the gut microbiota of Helicoverpa zea, promotes tomato plant growth and yield without compromising anti-herbivore defenses. Arthropod-Plant Interactions 13: 271-278. (PDF)

Shikano, I., Q. Pan, K. Hoover, and G.W. Felton. 2018. Herbivore-induced defenses in tomato plants enhance the lethality of the entomopathogenic bacterium, Bacillus thuringiensis var. kurstaki. J. Chem Ecol. 44: 946-956. (PDF)

Shikano, I., E.M. McCarthy, J.M. Slavicek and K. Hoover. 2018. Jasmonic acid-induced plant defenses delay caterpillar developmental resistance to a baculovirus: Slow-growth, high-mortality hypothesis in plant–insect–pathogen interactions, J. Invertebr. Pathol. 158: 16-23. (PDF)

Wang, J., M. Yang Y. Song F.E. Acevedo K. Hoover R. Zeng G.W. Felton. 2018. Gut-associated bacteria of Helicoverpa zea indirectly trigger plant defenses in maize. J Chem Ecol 44: 690-699. (PDF)

Tan, C.-W., M. Peiffer, K. Hoover, C. Rosa, F. E. Acevedo, G. W. Felton. 2018. Symbiotic polydnavirus of a parasite manipulates caterpillar and plant immunity. PNAS 201717934 DOI: 10.1073/pnas.171793411. (PDF)

Ash tree species likely will survive emerald ash borer beetles, but just barely

UNIVERSITY PARK, Pa. — “Lingering ash." That’s what the U.S. Forest Service calls the relatively few green and white ash trees that survive the emerald ash borer onslaught. Those trees do not survive by accident, and that may save the species, according to Penn State researchers, who conducted a six-year study of ash decline and mortality.

The research shows some ash trees have varying degrees of resistance to the strangely beautiful, invasive beetle from Asia. The study is unique because it took place at a plantation of ash trees planted on Penn State’s University Park campus in the mid-1970s.

“We found that genetic variation exists in trees from around the country, and through time — especially as the emerald ash borer population collapses because host trees are rapidly disappearing — the resistance that we observed will likely ensure the survival of the species,” said Kim Steiner, professor of forest biology, College of Agricultural Sciences.

Genetics moderated the rapidity with which emerald ash borers injured and killed trees, researchers learned. This suggests that some ash genotypes, especially on favorable sites, will survive.

Steiner, who also is director of The Arboretum at Penn State, collected seeds from wild green ash trees in 27 states and Canadian provinces in the fall of 1975. He grew the seedlings for two years before methodically planting 2,100 of them, all 12 feet apart, in a seven-acre plot. Mixed in were a small number of white ash trees.

Steiner conducted a provenance trial — moving trees that had evolved in different climates to one location and carefully monitoring their growth and other characteristics — with the goal of understanding how species adapt to their environments. Over the last few decades, researchers maintained the plantation to study the effects of climate change on trees.

This little-known ash plantation off Porter Road near Penn State's Swine Research Facility — the largest collection of green ash germplasm in one location in the world — may play a role in saving the species.

“We began measuring the decline in 2012, shortly after emerald ash borers arrived in the plantation, and we measured it every year through 2017,” said Steiner. “The effect of the insect was devastating. As of August of this year, only 13 trees remained of the 1,762 that were alive when the emerald ash borer arrived.”

Researchers began measuring the decline of ash trees in the Penn State plantation in 2012, shortly after emerald ash borers arrived there, and they measured it every year through 2017. The effect of the insect was devastating.

Although final destruction was nearly complete, genetics moderated the rapidity with which emerald ash borers injured and killed trees, noted Lake Graboski, Steiner’s assistant, who earned a master’s degree in ecology at Penn State.

“This suggests that some ash genotypes, especially on favorable sites, will survive with lower densities of emerald ash borer beetles on the landscape,” he said.

The fact that some trees survived longer means there are heritable genetic differences among trees from different populations and seed parents, Steiner added.

“For the first time, this study demonstrated that there is genetic variation that could be captured in a breeding program to improve resistance to emerald ash borer in both white ash and green ash species," he said.

There are three kinds of resistance to insects commonly exhibited by trees, Steiner explained, and more research will be needed to determine which ones the ash trees may be deploying. One is avoidance, when a tree doesn't attract the adult females that are flying between the trees as they look for a place to lay their eggs. A tree may accomplish this by not emitting a chemical signal the insects are homing in on.

The second is surviving attack. Adult insects lay eggs on a tree, the larvae hatch and the insects grow into adulthood, all the while causing damage, but the tree is vigorous enough to withstand that injury.

In this 1977 photo, taken off University Drive near where the Forest Resources Building sits today, Kim Steiner, professor of forest biology, surveys a makeshift nursery where he grew the seedlings that were later moved to his ash plantation.

The third mode of resistance involves the tree producing compounds — or alternately, not producing compounds — that reduce the likelihood the larvae will survive to adulthood, either by actively killing the larvae or by not offering the nourishment they need.

The irony of addressing a modern-day ecological disaster, such as the emerald ash borer invasion, with research done in a 43-year-old experimental plantation intended to serve an entirely different purpose, was not lost on Graboski.

“Dr. Steiner planted those ash trees long before I was born, and the ultimate fate of the ash species may not be decided in my lifetime because the trees must evolve to survive attacks by the invasive beetles," said Graboski. "That is just the reality of working with trees.”

Tachinid Flies

Parasitism, a reproductive strategy not typically associated with flies, serves as a key trait uniting the large and diverse fly family Tachinidae, better known as tachinid flies.

With a world wide distribution of some ten thousand species, the tachinids have been loosely organized according to common parasitic strategies along with common physical traits. For example, the smallest subfamily of Tachinidae, the Phasiinae, usually get classified as the least hairy tachinids that parasitize true bugs (order Heteroptera).

Entomologists generally converge on the idea that most Tachininae species, along with other tachinids, adopt a parasitic reproductive strategy that targets members of the Lepidoptera order, butterflies and moths, as the host for their larvae.

Furthermore, because so many moth larvae are classified as agricultural pests, tachinids that parasitize them often get classified as beneficial insects. The use of tachinid flies to control gypsy moth infestations of forests serves as one of the most prominent examples of their use as biological control agents.

The proven and potential utility of Tachinidae gives them a prominent place in present and future entomological research.

North America hosts over five dozen different tachinid genera. As a rule of thumb, tachinids can be identified by their hairy and/or colorful abdomens.


Articles and book chapters

* = senior author
** = MSc student

Araújo, J., and D.P. Hughes (2016) Diversity of Entomopathogenic Fungi: Which Groups Conquered the Insect Body? Advances in Genetics Vol 93

Loreto, R and D.P. Hughes (2016) Disease Dynamics in Ants: A Critical Review of the Ecological Relevance of Using Generalist Fungi to Study Infections in Insect Societies Advances in Genetics Vol 93

Hughes, D.P. Araújo, J. Loreto, R., Quevillon, L., de Bekker, C. and H.C. Evans (2016) From So Simple a Beginning: The Evolution of Behavioral Manipulation by Fungi Vol 93 Genetics corrected proof Vol 93

Hughes, D. P. (2015). Behavioral Ecology: Manipulative Mutualism. Current Biology 25:R806-R808.

de Bekker, C., R. A. Ohm, R. G. Loreto, A. Sebastian, I. Albert, M. Merrow, A. Brachmann, and D.P. Hughes (2015). Gene expression during zombie ant biting behavior reflects the complexity underlying fungal parasitic behavioral manipulation. BMC Genomics 16:620.

Quevillon, L. E., Hanks, E. M., Bansal, S., & Hughes, D. P. (2015). Social, spatial, and temporal organization in a complex insect society. Nature Scientific Reports 5:13393

NESCent Working Group on the Evolutionary Biology of the Built Environment: Martin, L.J., Adams, R.I, Bateman, A., Bik, H.M. Hawks, J. Hird, S.M. Hughes, D.P., Kembel, S.W., Kinney, K., Sergios-Orestis, K., Levy, G., McClain, C., Meadow, J.F., Medina, R.F., Mhuireach, G., Moreau, C.S., Munshi-South, J., Nichols, L.M., Palmer, C., Popova, L., Schal, C., Täubel, M., Trautwein, M., Ugalde, J.A. & R. R. Dunn (2015) Evolution of the indoor biome, Trends in Ecology & Evolution, Volume 30, Issue 4, April 2015, Pages 223-232, ISSN 0169-5347.

Biron, D. Panek, J. Chetouhi, C. El Alaoui, H., Texier, C., Langin, T. de Bekker, C., Bonhomme, L. Urbach, S. Demettre, E. Misse, D. Holzmuller, P., Hughes, D.P. Zanzoni, A., Brun, C. (2015) Cross-talk in host-parasite associations: what do past and recent proteomics tools tell us? Infection, Genetics and Evolution Jul33:84-94. doi: 10.1016/j.meegid.2015.04.015.

Alisha Quandt, C. Kepler, R.M. Gams, W., Araújo, J. P., Ban, S., Evans, H. C., Hughes, D.P.Humber, R. Hywel-Jones, N. Li, Z, Luangsa-ard, J.J. Rehner, S.A. Sanjuan, T. Sato, H. Shrestha, B. Sung, G.H. Yao, Y. Zare, R and J. W. Spatafora (2014). Phylogenetic-based nomenclatural proposals for Ophiocordycipitaceae (Hypocreales) with new combinations in Tolypocladium. IMA fungus, 5(1), 121.

de Bekker,C, Quevillon, L., Smith, P. Patterson, A.D, Flemming, K., Ghosh, D. and D.P. Hughes Species-specific Ant Brain Manipulation by a Specialized Fungal Parasite BMC Evolutionary Biology 14.1 (2014): 166.

Loreto, R. G., Elliot, S. L., Freitas, M. L., Pereira, T. M., & Hughes, D. P. (2014) Long-Term Disease Dynamics for a Specialized Parasite of Ant Societies: A Field Study PloS One, 9(8), e103516.

Hughes, D.P. (2014) On the origins of parasite extended phenotypes Integrative and Comparative Biology 54 (2): 210-217

de Bekker, C., Merrow, M., and D.P. Hughes (2014) From behavior to molecular mechanisms: an integrative approach to parasitic host manipulation Integrative and Comparative Biology (2): 166-176.

Smith, R. A., & Hughes, D.P (2014). Infectious Disease Stigmas: Maladaptive in Modern Society. Communication Studies, 65(2), Special Issue on Stigma.

Lachaud, J. P., Lenoir, A., & D.P. Hughes (2013). Ants and Their Parasites 2013. Psyche: A Journal of Entomology Vol 2013 1-5

Loreto RG, Hart A, Pereira TM, Freitas ML, Hughes DP, Elliot SL (2013) Foraging ants trade off further for faster: use of natural bridges and trunk-trail permanency in carpenter ants. Naturwissenschaften Vol 100 Issue 10 pp 957-963

de Bekker, C. Smith, P. Patterson, A.D and D.P. Hughes (2013) Metabolomics reveals the heterogeneous secretome of two entomopathogenic fungi to ex vivo cultured insect tissues PloS One 8(8): e70609. doi:10.1371/journal.pone.0070609 (Article)

Maure, F. Brodeur, J. Hughes, D.P. and F. Thomas (2013) How much energy should manipulative parasites leave to their hosts to ensure altered behaviours? Journal of Experimental Biology 2012 216: 43-6. (Article)

Hughes, D.P. (2013) Pathways to understanding the extended phenotype of parasites in their hosts Journal of Experimental Biology 216:142-147 (Article)

Hughes, D.P. Parasites and the Superogranism (2012). In Host Manipulation by Parasites Edited by David P. Hughes, Jacques Brodeur, and Frédéric Thomas

Andersen SB, Ferrari M, Evans HC, Elliot SL, Boomsma JJ, and D.P. Hughes (2012) Disease Dynamics in a Specialized Parasite of Ant Societies. PLoS ONE 7(5): e36352. doi:10.1371/journal.pone.0036352

Andersen SB and D.P. Hughes (2012) Host specificity of parasite manipulation –zombie ant death location in Thailand vs. Brazil Communicative & Integrative Biology 5:2, 1–3 March/April

Harry C. Evans, Simon L. Elliot and David P. Hughes (2011) Ophiocordyceps unilateralis: A keystone species for unraveling ecosystem functioning and biodiversity of fungi in tropical forests? Communicative & Integrative Biology 4:5, 598-602 (Article)

Hoover, K., M. Grove, M. Gardner, D.P. Hughes, J. McNeil and J. Slavicek. (2011) A gene for an extended phenotype. Science 333: 1401. This paper has been designated a Faculty of 1000 Must Read Factor 8

Hughes, D.P., Andersen, S.* Hywel-Jones, N.L. , Himaman, W., Bilen, J and J.J. Boomsma. Behavioral mechanisms and morphological symptoms of zombie ants dying from fungal infection BMC Ecology 2011, 11:13doi:10.1186/1472-6785-11-1 (Article)

Evans, H.E., Elliot, S.L, and D.P. Hughes (2011) Hidden diversity behind the Zombie-Ant fungus Ophiocordyceps unilateralis: Four new species described from Carpenter ants in Minas Gerais, Brazil PloS One. 2nd March 2011

Semenova, T,** Hughes, D.P. Boomsma, J.J. & Schioett (2011) Evolutionary patterns of proteinase activity in attine ant fungus gardens BMC Microbiology. BMC Microbiology 11:15

D.P. Hughes (2011) Recent developments in sociobiology and the scientific method Trends in Ecology and Evolution Vol 26 (2) 57-8

D.P. Hughes, Wappler, T, & C. C. Labandeira (2010) Ancient death-grip leaf scars reveal ant fungal parasitism Biology Letters 18th August doi:10.1098/rsbl.2010.0521

D.P. Hughes (2010) A philosophical view of biology Trends in Ecology and Evolution Vol 25 (7) 384-385

Andersen, SB, ** S. Gerritsma, ** K.M. Yusah, D. Mayntz, N.L. Hywel-Jones, J. Billen, J.J. Boomsma & D.P. Hughes* (2009). The life of a dead ant - the expression of an extended phenotype. American Naturalist 174: 424–433

Pontoppidan, M-B, ** W. Himaman, N.L. Hywel-Jones, J.J. Boomsma & D.P. Hughes* (2009). Graveyards on the move: The spatio-temporal distribution of dead Ophiocordyceps infected ants. PloS One 4(3): e4835 doi:10.1371/journal.pone.0004835

Hughes, D.P. (2009). Altruists since life began: the evolution of the superorganism (book review). Trends in Ecology and Evolution 24(8): 417–418

Hughes, D.P., H.C. Evans, N.L. Hywel-Jones, J.J. Boomsma & S.A.O. Armitage (2009). Emerging fungal diseases in complex leaf-cutting ant societies. Ecological Entomology in 34: 214–220

Lefevre, T., S. Adamo, D.G. Biron, D. Misee, *D.P. Hughes & F. Thomas (2009). Invasion of the Body Snatchers: The Diversity and Evolution of Manipulative Strategies in Host–Parasite Interactions. Advances in Parasitology 68: 45–83

Cremer, S., L.V. Ugelvig, F.P. Drijfhout, B.C. Schlick-Steiner, F.M. Steiner, B. Seifert, D.P. Hughes, A. Schultz, K.S. Petersen, H. Konrad, C. Stauffer, K. Kiran, X. Espadaler, P. d'Ettorre, N. Aktaç, J. Eilenberg, G.R. Jones, D.R. Nash, J.S. Pedersen, J.J. Boomsma (2008). The Evolution of Invasiveness in Garden Ants. PLoS ONE 3(12): e3838

Hughes, D.P., N.E. Pierce & J.J. Boomsma (2008). Social insect symbionts: evolution in homeostatic fortresses. Trends in Ecology and Evolution. 23(12): 672-677

Hughes, D.P. (2008). The extended phenotype within the colony and how it obscures social communication. (book chapter in Sociobiology of Communication eds. P. d'Ettorre and D.P. Hughes)

Hughes, D.P., D.J.C. Kronauer & J. J. Boomsma (2008). Extended Phenotype: Nematodes turn ants into bird-dispersed fruits. Current Biology. 18: R294–R295

Sánchez, M.I., F. Ponton, A. Schmidt-Rhaesa, D.P. Hughes, D. Missé & F. Thomas (2008). Two steps to suicide in insects harbouring hairworms. Animal Behaviour, 75(5): 1621–1624

Sánchez, M.I., F. Ponton, D. Missé, D.P. Hughes & F. Thomas (2007). Hairworm response to notonectid attacks. Animal Behaviour 75(3): 823–826

Hughes, D.P. & S. Cremer (2007). Plasticity in anti-parasite behaviours and its suggested role in invasion biology. Animal Behaviour 74(5): 1593–1599

Lefevre, T., M. Sanchez, D.P. Hughes & F. Thomas (2007). Virulence and resistance in malaria: who drives the outcome of the infection? Trends in Parasitology 23(7): 299–302

Ponton, F., C. Lebarbenchon, T. Lefevre, D.G. Biron, D. Duneau, D.P. Hughes & F. Thomas (2006). Hairworm anti-predator strategy: a study of causes and consequences. Parasitology 133(5): 631–638

Hughes, D.P. & J.J. Boomsma (2006). Muscling out malaria. Trends in Ecology and Evolution 21(10): 533–534

Ponton, F., C. Lebarbenchon, T. Lefevre, D.G. Biron, D. Duneau, *D.P. Hughes & F. Thomas (2006). Parasite survives predation on its host. Nature 440: 756–756

Kathirithamby J. & D.P. Hughes (2006). Description and biological notes of the first species of Xenos (Strepsiptera:Stylopidae) parasitic in Polistes carnifex (Hymenoptera: Vespidae) in Mexico. Zootaxa 1104: 35–45

Hughes, D.P. & J. Kathirithamby (2005). Virulence under low extrinsic mortality: the benefit of parasitizing social insects? Oikos 110: 42–434

Hughes, D.P. (2005). Parasitic Manipulators: a social context. Behavioural Processes 68(3): 263–266

Hughes, D.P., J. Kathirithamby & L. Beani (2004). Prevalence of the parasite Strepsiptera in adult Polistes wasps: field collections and literature overview. Ethology, Ecology and Evolution 16: 363–75

Johnston, S.J., L. Ross, L. Beani, D.P. Hughes & J. Kathirithamby (2004). Tiny genomes and endoreduplication in Strepsiptera. Insect Molecular Biology 13(6): 581–585

Hughes, D.P., J. Kathirithamby, S. Turillazzi & L. Beani (2004). Social wasps desert the colony and aggregate outside if parasitized: parasite manipulation? Behavioural Ecology 15(6): 1037–1043

Hughes, D.P., P. Pamilo & J. Kathirithamby (2004). Horizontal transmission of Wolbachia by strepsipteran endoparasites? A reply to Noda et al. 2001. Molecular Ecology 13(2): 507–9

Hughes, D.P., G. Moya-Raygoza & J. Kathirithamby (2003). The first record among Dolichodernae (Formicidae) of parasitism by Strepsiptera. Insectes Sociaux. 50 (2): 148–150

Hughes, D.P., L. Beani, S. Turillazzi & J. Kathirithamby (2003). Prevalence of the parasite Strepsiptera in Polistes as detected by dissection of immatures. Insectes Sociaux 50(1): 62–68

Hughes, D.P. (2002). The value of a broad mind: some natural history meanderings of W.D. Hamilton. Ethology, Ecology and Evolution 14 (2): 83–89

Kathirithamby, J. & D.P. Hughes (2002). Caenocholax fenyesi Pierce (Strepsiptera: Myrmecolacidae) parasitic in Camponotus planatus Roger (Hymenoptera: Formicidae) in Mexico: is this its original host? Annals of the Entomological Society of America 95 (5): 558–563