Molecular phylogeny of World Tachinidae

I have been quite lax in providing updates and posts to the Stireman lab website. Hopefully, I will be able to devote time to it in the future. A number of publications have come out of the Stireman lab since I last posted two years ago. Most notable among them is A Molecular Phylogeny of World Tachinidae which was published online in early 2019 (Stiremanetal2019YMPEV6358.pdf), which was the culmination of a number of years’ work by me (Stireman), Jim O’Hara, Pierfilippo Cerretti, Kevin Moulton, and Jeremy Blaschke. I hope that others find this paper interesting and useful. It is by no means the “end of the story” regarding tachinid phylogeny, but it is a significant advancement that we hope will stimulate more research into understanding this evolution and diversification of this fascinating group. Abstract below.



We reconstructed phylogenetic relationships within the diverse parasitoid fly family Tachinidae using four nuclear loci (7800 bp) and including an exceptionally large sample of more than 500 taxa from around the world. The position of the earthworm-parasitizing Polleniinae (Calliphoridae s.l.) as sister to Tachinidae is strongly supported. Our analyses recovered each of the four tachinid subfamilies and most recognized tribes, with some important exceptions in the Dexiinae and Tachininae. Most notably, the tachinine tribes Macquartiini and Myiophasiini form a clade sister to all other Tachinidae, and a clade of Palpostomatini is reconstructed as sister to Dexiinae + Phasiinae. Although most nodes are well-supported, relationships within several lineages that appear to have undergone rapid episodes of diversification (basal Dexiinae and Tachininae, Blondeliini) were poorly resolved. Reconstructions of host use evolution are equivocal, but generally support the hypothesis that the ancestral host of tachinids was a beetle and that subsequent host shifts to caterpillars may coincide with accelerated diversification. Evolutionary reconstructions of reproductive strategy using alternative methods were incongruent, however it is most likely that ancestral tachinids possessed unincubated, thick shelled eggs from which incubated eggs evolved repeatedly, potentially expanding available host niches. These results provide a broad foundation for understanding the phylogeny and evolution of this important family of parasitoid insects. We hope it will serve as a framework to be used in concert with morphology and other sources of evidence to revise the higher taxonomic classification of Tachinidae and further explore their evolutionary history and diversification.

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Adaptive radiation from the top down

Finally, the last paper stemming from Jeremy Heath’s PhD research has been published in The American Naturalist! This is a super-cool paper illustrating the importance of natural enemies (parasitoids in this case) in driving the adaptive diversification in their gall midge hosts. We suggest that such top-down driven diversification (rather than resource-driven) may be much more widespread than is currently appreciated.  I could explain further, but I think the abstract below sums it up nicely.  You can access the paper here.


Aprostocetus tesserus laying an egg in an Asteromyia carbonifera gall. (J.J. Heath)


Most studies of adaptive radiation in animals focus on resource competition as the primary driver of trait divergence. The roles of other ecological interactions in shaping divergent phenotypes during such radiations have received less attention. We evaluate natural enemies as primary agents of diversifying selection on the phenotypes of an actively diverging lineage of gall midges on tall goldenrod. In this system, the gall of the midge consists of a biotrophic fungal symbiont that develops on host-plant leaves and forms distinctly variable protective carapaces over midge larvae. Through field studies, we show that fungal gall morphology, which is induced by midges (i.e., it is an extended phenotype), is under directional and diversifying selection by parasitoid enemies. Overall, natural enemies disruptively select for either small or large galls, mainly along the axis of gall thickness. These results imply that predators are driving the evolution of phenotypic diversity in symbiotic defense traits in this system and that divergence in defensive morphology may provide ecological opportunities that help to fuel the adaptive radiation of this genus of midges on goldenrods. This enemy-driven phenotypic divergence in a diversifying lineage illustrates the potential importance of consumer-resource and symbiotic species interactions in adaptive radiation.


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Mini-review of the “other” parasitoids

fig1Stireman (me) was recently invited to write a mini-review of the community ecology of non-hymenopteran parasitoids for the journal Current Opinion in Insect Science. This was a difficult task as the review was supposed to focus on recent (last 5 years) literature, and was severely constrained in length and number of references. Still, I hope that it provides a decent overview of some of the recent research on the topic (focusing on dipteran parasitoids) and helps to spur further research.

The full text can be accessed here at the publisher’s web site (at least for a while). Enjoy!



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Ph.D. student opportunity: ecology, evolution, and systematics of tachinid flies

I am currently seunknownParerigoniiniAUS064eking a Ph.D. student to join my laboratory studying the evolution and ecology of parasitoid flies. While the specific focus of the dissertation research is negotiable, the research assistantship will require contributing to a collaborative, NSF/Brazil(FAPESP) funded Dimensions of Biodiversity project focused on “Chemically mediated multi-trophic interaction diversity across tropical gradients.” My laboratory’s role in this international collaborative project is primarily focused on tachinid parasitoids. This includes identifying and documenting species, studying how they influence and are influenced by hosts and their host-plants, analyzing population- and phylo-genetic/genomic patterns and processes, and revisionary taxonomy and species description. Students will also have the opportunity to contribute to other aspects of this large and multi-disciplinary project . The successful applicant will develop a thesis research project on tachinid ecology, evolution and/or systematics employing ecological, phylogenomic, taxonomic, and comparative methods. The student will also have the opportunity to visit and participate in field research in Brazil as well as other Latin American Countries. Latin American students are particularly encouraged to apply. Click here for more information.

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Master Karen Pedersen

Congratulations are in order! WSU master’s student Karen Pedersen has successfully defended her thesis research on: LIMITATIONS OF HOST PLANT USE IN TWO ANDEAN ALTINOTE (NYMPHALIDAE, HELICONIINAE, ACRAEINI), BUTTERFLIES, FROM A TRITROPHIC PERSPECTIVE. In this research project, Karen tried to assess the determinants of host plant use in two co-occurring species of Altinote butterflies in Ecuador. Her research took place at Yanayacu Biological Station, and is an extension of our work there on plant-caterpillar-parasitoid interactions. Karen produced several short videos focused on her study system system that you can check out:

Catching Butterflies in Ecuador!!!

Egg to Adult in two Altinote butterflies

hyperparasitoid wasp easily overcomes caterpillar defenses

Finding a Host-Plant

Dysschema sp? caterpillar dances


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Ghosts of the cloud forest – 11 new species of Erythromelana (Tachinidae)

Although MS student Diego Inclan successfully defended his thesis more than two years ago, the focus of his research was published this year:

Inclan, D.J. & Stireman, J. O. III. 2013. Revision of the genus Erythromelana Townsend, 1918 (Diptera: Tachinidae) with description of 11 new species and analysis of their phylogeny and diversification. Zootaxa. 3621:1-82.(Abstract here)

In this lengthy manuscript Diego describes 11 new species (as you can tell from the title) of a little known, small bodied and somewhat gracile genus of tachinids in the tribe Blondeliini that occur in the Neotropical region (Northern Argentina

to Southern Mexico). Most of the species are found in cloud forest habitats in the Andes.  Diego also examines their phylogenetic relationships using morphological and molecular approaches and examines potential modes of diversification for the genus. With only three previously described species, Diego has more than quadrupled the number of known species in the genus.

Why focus on this group of small, poorly known cloud forest tachinids?

First,  exploring, documenting and describing all of Earth’s organisms is a worthy goal in itself. Each has many fascinating biological “stories” or “lessons” to tell with its unique evolutionary history and ecological niche.  Describing these species is a first step towards elucidating these lessons, which may be far reaching (e.g., I always like to point out the vast scientific knowledge we have gleaned from studying the small and nondescript species Drosophila melanogaster).

Erythromelana cryptica Inclan, one of the new species described in this study

Erythromelana cryptica Inclan, one of the new species described in this study


A species of the diverse geometrid genus Eois, hosts of Erythromelana

However, there is another reason that we focused on this genus. It just so happens that Erythromelana species are parasitoids of small geometrids in the genus Eois. We know this because we reared several species (first known rearings for the genus) from Eois in the cloud forests of Ecuador (see post Ecuador Expedition). Eois is a very diverse genus of geometrids that feed on plants in the hyper-diverse genus Piper (Piperaceae), and this tritrophic system of plants, herbivores, and parasitoids has been the subject of a number of studies by collaborators of our “Caterpillars and Parasitoids of the Eastern Andes” biological survey project (e.g., see Wilsonetal2011). One goal of Diego’s paper was to see if Erythromelana exhibited evidence of co-evolution and host-associated differentiation with their Eois hosts. Although the degree of specificity of Erythromelana species is difficult to evaluate due to the many undescribed and cryptic species of Eois that we know of, our rearing records suggest considerable overlap in host use among these tachinids.  Thus, it would appear that perhaps geographic isolation may play a greater role in Erythromelana diversification than host associations.

Why ‘ghosts of the cloud forest’? This somewhat dramatic title refers to the rarity of Erythromelana species. Total parasitism of Eois (from over 5000 rearings) is less than 0.5% and we only recovered a single specimen from a pan trapping effort at Yanayacu that collected over 2000 other tachinids. Several of the described species are known from just a handful of specimens. Still, these ‘ghosts’ can be observed, if one knows where to look. On a few occasions they have been seen, flitting about sunlit leaves over small streams in deep cloud forests of Costa Rica and Ecuador.

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Yard insects

Just a few recent photos from my yard (click for larger images)


The Blue Dasher (Pachydiplax longipennis: Libellulidae)


Gymnosoma sp. (Tachinidae)

a bee fly (Bombyliidae: Phthirinae)

a bee fly (Bombyliidae: Phthirinae)

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It’s not just what’s in the field, it’s what’s around it

I have taken a long hiatus from posting any news or other items on here (for no particularly good reason) and I thought I might try to do some catching up on news from the lab. In the next several posts I will highlight some recent publications and other events over the last half year or so.OLYMPUS DIGITAL CAMERA

First, I would like to highlight a not-so-recent publication (LetBothStire2012 – actually published in 2012) led by collaborators at UC Santa Cruz, PhD student Sara Bothwell Allen and Deborah Letourneau, her advisor. For Sara’s thesis she studied the effects of surrounding land use on parasitism levels of pests and parasitoid diversity in organic agricultural fields in California. She set out “sentinel” letourneaufigspest caterpillars, confined to their plants, for one week to examine parasitism and she set out Malaise traps for two days (twice per season) to sample parasitoid diversity. She then characterized the surrounding landscape cover using GIS. Because they got quite a few tachinids, Sara and Deborah invited me to collaborate on an analysis of how landscape variables outside the fields affected tachinid diversity and parasitism inside the fields. Interestingly we found effects of many landscape variables on the tachinid fly com

munities in the agricultural fields, but the strongest pattern was an increase in tachinid abundance and diversity with increasing coverage of semi-wild perennial vegetation in the landscape (see graphs below). These results argue for that current trends towards agricultural intensification -plowing fence line to fence line are counterproductive.  Semi-wild corridors and fragments can act as refugia for parasitoids, which may reduce the need for pest control measures in organic or conventional agriculture.

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Pivotal Pervasive Pigments: Carotenoids in insect ecology

Carotenoids are nearly ubiquitous organic compounds involved in all sorts of important

Diverse carrots colored by carotenoids

Diverse carrots colored by carotenoids

functions across all major groups of organisms. They play key roles in photosynthesis in plants, they function as antioxidants, and they provide many of the bright yellow, orange, and red coloring we see in plants, fungi, and animals. For example, the familiar orange coloration of carrots and red coloration of tomatoes is derived from carotenoids.

In a newly published paper in the journal Arthropod-Plant Interactions, Jeremy Heath (former PhD student), reviews the many varied functions of carotenoids in insects, with emphasis in how carotenoids and their derivatives influence interactions between insect and their environments (notably plants).

A stylized, schematic representation of the various known and hypothesized functions of carotenoids in insects that mediate ecological interactions.

A stylized, schematic representation of the various known and hypothesized functions of carotenoids in insects that mediate ecological interactions.

He briefly reviews the structure and biosynthesis of these molecules and then discusses their roles in cryptic and aposematic coloration in insects, their importance in vision, photoperiodism and diapause, their function as antioxidants, and their role in signaling. He also explores the possible functions of carotenoid derivatives such as strigolactones and volatile apocarotenoids in mediating interactions between insects and plants (and fungi), and between insects and their parasitoid enemies. Contact Jeremy ( or myself ( for reprints.

Heath, J.J., D. Cipollini, and J.O. Stireman III. 2013. The role of carotenoids and their derivatives in mediating interactions between insects and their environment. Arthropod-Plant Interactions 7:1-20.

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Ecuador Expedition 2012

I (Stireman) recently returned from a trip to Ecuador. The primary reasons for the trip were 1. To collect tachinid flies for systematic and biodiversity research, and 2. To get graduate student, Karen Pedersen, set up for her studies on the multitrophic determinants of host specificity in Altinote butterflies (Nymphalidae: Acraeinae). There was also the added benefit of meeting up with some of my old grad school buddies, Harold Greeney (Tropical Ornithologist and Entomologist), Caleb Gordon (bird fanatic), Matt Kaplan (high through-put moleculoid master and lizard man), and Margy Green (Optics engineer and herp/bug photographer) and several of their biophile colleagues.

This was my favorite trip to Ecuador thus far (and I’ve been there a few times). We explored a wide diversity of habitat types and were able to travel to some magnificent areas. We started out at Yanayacu Biological Station, epicenter of my research program in

The view form Yanayacu

The view from Yanayacu

Ecuador, in montane cloud forest on the East slope of the Andes (~2000 m). This the best place to find tachinids that I have ever seen. They are everywhere, in an amazing variety of forms. They are thick as, well…flies. You hardly have to try to look for them. My pet hypothesis for their amazing abundance and diversity here is a combination of the almost infinite opportunities for habitat specialization and semi-isolation provided by the Andes, the ridiculous diversity of potential hosts, and the relative lack of ants. Perhaps I will elaborate sometime…

From Yanayacu, we travelled to the high paramo, reaching elevations as high as 4300 meters (Like 14400 ft). It is remarkable habitat of cushion plants and club mosses, but it’s awful chilly when the sun isn’t out (as it was not when we went…which explains the poor showing of tachinids).

Paramo near Papallacta

Paramo near Papallacta

From there some of us split off to visit a small reserve called Jatun Sacha. This is a lowland reserve at about 450 m with nice accommodations and some attractive old growth forest

A tree frog at Jatun Sacha (see, I like vertebrates too)

A tree frog at Jatun Sacha (see, I like vertebrates too)

(It’s also easy to get to). The group rejoined to visit another small reserve in the Andean foothills called Bigal River Biological Reserve that adjoins the huge Sumaco National Park. This reserve, operated by a French couple (Marion and Thierry, who cooked us super-delicious food) required a little bit of walking on a treacherously muddy old logging road to get to (luckily with donkeys packing in most of our gear). Though this is a fairly new reserve, the forest is old, diverse, and

Lycaenid at El Bigal

Lycaenid at El Bigal

relatively untouched. I spent most of my time walking along the road (good place for flies) and peeing on select plants to attract a diversity of flies, stingless bees, and butterflies. I cannot overstate the utility of urine as a tachinid (and other fly) attractant in the wet, salt poor lowland forest. I even saved jars of pee for this express purpose.

Finally, we traveled to the heart of the Ecuadorian Amazon to a fantastic lodge called Shiripuno, located on the Huaorani (native peoples)  Reserve and not far from the “intangible zone”, where there are people living that have no, and want no, contact with western civilization. Being in this place, it is easy to forget that there is any threat to the lowland rainforest; the forest seems endless, there are parrots and toucans everywhere,

boating down the Shiripuno

boating down the Shiripuno

there are 10 species of monkeys, and the insect diversity is amazing. The naturalist Fernando (Vaca), as well as several Huaorani guides, led us on a number of eventful outings on the river and on trails, but we were also free to explore the trails on our own. I spent a lot of time in around the edges of the main clearing of the lodge, monitoring my pee and sugar-water trapline for flies – and even this was awesome.

I should note that my good friend and scientific collaborator Harold Greeney arranged all these trips, and I am very thankful he did. I should also note that my tachinid collecting was conducted under the auspices of two major research programs the Stireman lab is involved with: A long term biotic survey and inventory of caterpillar-plant-parasitoid interactions in Ecuador (led by Lee Dyer at U. Nevada) and the World Phylogeny of Tachinidae project led by our lab (see previous post). I was able to collect quite a diversity of taxa (many of which will be deposited in the Museo Nacional del Ecuador in Quito) that will be extremely useful for our ongoing investigations of tachinid  ecology, diversity, and phylogeny. By the way, I left Karen (my student) down there at Yanayacu for four months to work on her studies of plant-butterfly-parasitoid interactions. I’m a little jealous.

Endless forest seen from the Mirador trail at Shiripuno

“Endless” forest seen from the Mirador trail at Shiripuno

One of the few tachinid shots I took (probably a blondeliine. Phyllophilopsis??)

One of the few tachinid shots I took (probably a blondeliine. Phyllophilopsis??)

Mesembrinella (Calliphoridae sensu lato) on rotting banana

Mesembrinella (Calliphoridae sensu lato) on rotting banana

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