■ Phylogeny of World Tachinidae Project

The information given here has been extracted from an article in The Tachinid Times:

Stireman, J.O. III, O'Hara, J.E., Moulton, K., Cerretti, P. & Winkler, I.S. (2013) Progress towards a phylogeny of world Tachinidae. Year 1. The Tachinid Times, 26 , 4-9.

Introduction by John Stireman

Department of Biological Sciences, 3640 Colonel Glenn Highway, 235A, BH, Wright State University, Dayton, Ohio 45435, USA. E-mail: john.stireman@wright.edu

John Stireman

Developing a structural phylogenetic framework for the family Tachinidae has been an interest of mine since I first started working on tachinids as a graduate student. It was then that I developed a new phylogenetic perspective with which to view the world, and many of the questions I wanted to ask of tachinids depended on some level of phylogenetic knowledge of them. I was surprised at the time that no one had attempted a broad quantitative phylogenetic analysis of the family, and a publication from my thesis work on the Exoristinae (Stireman 2002) became the first such study that I know of. Ever since that initial work, I have been seeking ways to continue working towards the development of a broad and robust phylogeny of Tachinidae (e.g., Stireman 2005, 2010).

About ten years ago, Kevin Moulton (Univ. of Tennessee), developed the first full proposal for a molecular phylogeny of Tachinidae and invited me to participate as a postdoc. Unfortunately this, and a subsequent proposal, failed to garner interest from U.S. governmental funding agencies, and we put it "on the back burner" and pursued other research projects. Obtaining a professorship position in 2005 at Wright State University allowed me the resources and opportunity to begin collecting tachinid samples and sequence data "on the side" while conducting other work. We were not in a big rush to get rejected yet again, so we put off re-submitting a revamped proposal until 2010, when we invited Jim O'Hara (CNC) to be a collaborator. In this revised proposal, we sought to make the tachinid phylogeny project more of a collaborative effort as you can see from my 2010 article in The Tachinid Times exhorting aid from tachinid workers. This same year, Tachi and Shima (2010) published their paper focused on the phylogeny of Exoristinae, a vast improvement over my initial attempts. This paper demonstrated the potential insight that phylogenetic analysis of tachinids could provide, and spurred our efforts. Again, our proposal was rejected. But we were not so easily deterred; we revised it and resubmitted it twice more, each time expanding our goals and proposed products. We were finally awarded a grant in 2012 from U.S. National Science Foundation to pursue our goals of developing a broad phylogenetic framework for Tachinidae. This initiative, "The Phylogeny and Evolution of World Tachinidae", was designed to be a collaborative project, involving many tachinid researchers from around the world. The primary collaborators are John Stireman (me, Wright State Univ.), Jim O'Hara (CNC), Kevin Moulton (Univ. of Tennessee), Pierfilippo Cerretti (Univ. of Roma), and Isaac Winkler (who recently joined my laboratory as a postdoc working on the project). I also have a Ph. D. student, Zachary Burington, working in my lab on this project (see the Student News section of this newsletter for a note from Zach).

In addition, we have solicited the involvement of many additional tachinid researchers. Thus far, we have received advice and assistance from a number of colleagues, including but not limited to Monty Wood, Ashley Kirk-Spriggs, Dan Janzen, Daniel Whitmore, Takuji Tachi, and Hiroshi Shima.

Jim O'Hara

Canadian National Collection of Insects, Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, Ontario, K1A 0C6, Canada; E-mail: james.ohara@agr.gc.ca

Jim O'Hara

My contributions to this study are intertwined with my own research program funded by Agriculture and Agri-Food Canada (AAFC) that supports myself and my technician Shannon Henderson. My research over the past decade has shifted away from traditional taxonomic revisions to gathering together information for broader use; e.g., regional catalogues, an ongoing world taxonomic database, and involvement in the Tachinidae chapter of the Manual of Afrotropical Tachinidae (led by Pierfilippo Cerretti). If the present study leads to a better understanding of tachinid relationships and an improved classification, then this will be good for the projects I am pursuing on behalf of AAFC.

For the NSF project, Shannon and I will be expanding the coverage of the Tachinidae in the TachImage Gallery. I will be supplying Pierfilippo with specimens from the Canadian National Collection of Insects for use in an interactive key to the Tachinidae of America north Mexico and assisting with its development. I will participate in the interpretation of results and the writing of papers, and the collection of specimens for molecular analyses. The last is a vital part of the project and, from my perspective, one of the more enjoyable aspects of it. I started collecting specimens for molecular analyses several years ago in the hopes that the project would be funded. In 2012 I collected specimens for this project in Arizona and New Mexico and also participated in the group trip to South Africa.

Kevin Moulton

Department of Entomology and Plant Pathology, University of Tennessee, 2431 Joe Johnson Drive, 205 Ellington Plant Sciences Bldg., Knoxville, Tennessee 37996-4560, USA. E-mail: jmoulton@utk.edu

Kevin Moulton

Myself and my Master's student Jeremy Blaschke at the University of Tennessee, Knoxville are primarily responsible for collecting molecular data for the subfamily Phasiinae, collection of sequences for the gene MAC, and aiding Stireman's group in the collection of sequences for newly developed genes. The phasiines are an agronomically important group of heteropteran parasitoids and many genera have been used with varied success as biocontrol agents. From personal collecting trips and additional material sent from collaborators, we have been able to extract DNA from 35 phasiines representing 28 genera from 8 of the 9 collectable worldwide tribes. We are sequencing five different genes: CAD, MAC, LGL, MCS, and OPA or TUFT. These genes have varying evolutionary rates and when combined into a single data set will hopefully be able to reconstruct subfamilies, tribes, and beyond. We have sequences for CAD and MAC from all 35 available taxa, 33 for LGL, 16 for MCS, 8 for OPA, and 4 for TUFT.

As mentioned earlier, we are responsible for providing MAC, LGL, and MCS sequence data for a preliminary investigation of gene utility using a sparse though representative sample of genera. This initial study will consist of representative genera from important tribes within each subfamily as well as several outgroup taxa from each oestroidean family. We want to include around 32 taxa in this phylogeny and of those we have 32 for MAC, 30 for LGL, and 24 for MCS. We hope to have sequence data for all genes by the end of spring semester 2013.

Some interesting notes from collecting efforts in 2012. We have been baiting flight intercept traps with pheromones from Halyomorpha halys (Stål) (brown marmorated stink bug). The pheromones have successfully doubled (sometimes more) our catch rate of several phasiine genera. We will be exploring heteropteran pheromone baiting in the upcoming collecting season. Lastly, we have discovered what might be two distinct morphotypes of the important biocontrol species Trichopoda pennipes (Fab.) in the eastern United States. Further morphological and molecular research will be done to determine if these entities are valid species.

Pierfilippo Cerretti

Department of Biology and Biotechnology 'Charles Darwin', University of Rome 'La Sapienza', Piazzale A. Moro 5, 00185, Rome, Italy; and Centro Nazionale Biodiversità Forestale - Corpo Forestale dello Stato, via Carlo Ederle 16/a, 37100 Verona, Italy. E-mail: pierfilippocerretti@yahoo.it

Pierfilippo Cerretti

My focus on tachinid systematics dates back to 1998, when I first met Jim O'Hara and Monty Wood at the International Congress of Dipterology in Oxford (UK). Soon after the meeting Monty kindly introduced me to Benno Herting and Peter Tschorsnig (Stuttgart). Since 1999 I have visited the Tschorsnig lab several times and have learned a lot from Peter about tachinid systematics and biology. In more or less a decade I revised the whole Italian fauna to species level (Cerretti 2010), producing also an interactive key to Palaearctic genera in collaboration with Peter and Massimo Lopresti (Verona) (Cerretti et al. 2012). At the same time I started studying the Afrotropical fauna in order to revise the tribal and generic level classification and eventually extend the interactive key to this huge fauna. It is because of my interest in the African fauna that Ashley Kirk-Spriggs in 2009 invited me to lead the tachinid chapter of the upcoming Manual of Afrotropical Diptera that will be co-authored by Jim O'Hara, John Stireman and Monty Wood. In this project Jim is leading a new revised version of the Afrotropical tachinid catalogue.

Scoring morphological characters and developing interactive key applications is now one of my main activities and the reason why John Stireman and Jim O'Hara kindly asked me to join the "Phylogeny and Evolution of World Tachinidae" project proposal in 2010. As mentioned already by John, the project has several ambitious goals and these are so interesting to me that I simply could not turn down their request to collaborate. In this context my main responsibilities are:

  1. to score morphological characters to infer the first phylogenetic reconstruction of the Tachinidae using a cladistic approach (we are confident that a manuscript with preliminary but interesting results can be submitted in spring of 2013),
  2. to lead the development of an interactive key to Nearctic genera, and
  3. to supply John's and Kevin's labs with specimens for molecular analyses.

Of course, to study the evolution of morphological and biological traits over trees generated by molecular data will probably be the most exciting challenge for all of us!

Isaac Winkler

Department of Biological Sciences, 3640 Colonel Glenn Highway, 235A, BH, Wright State University, Dayton, Ohio 45435, USA. E-mail: isw971@gmail.com

Isaac Winkler

I have been helping to coordinate and assemble the molecular data for the overall tachinid tree (except phasiines). We are focusing initially on getting sequence for CAD and 28S. Our student technician, Beth Stayrook, has also been doing a lot of lab work, and we now have some sequence data from 19 tribes and nearly 200 genera.

Also, we have some transcriptome sequence data now from eight tachinid species (two from each subfamily). These each consist of about 30 million short reads, assembled into sequences for about 20,000 genes. It has been interesting and fun learning how to assemble and analyze this genomic data. By summer of 2013 we should have a phylogenomic data set put together and analyzed. The data should also be helpful for designing primers, studying specific tachinid genes, and even for learning about genome evolution in tachinids.

Graduate Students

Jeremy D. Blaschke

Department of Entomology and Plant Pathology, 2505 E. J. Chapman Drive, 370 Plant Biotechnology Building, University of Tennessee, Knoxville, TN 37996 , USA. E-mail: jeremy.blaschke@gmail.com

Jeremy Blaschke

As a Masters student in the Moulton Lab at the University of Tennessee, I am studying the phylogenetic relationships within the agronomically important tachinid subfamily Phasiinae I am contributing to the larger "Phylogeny and Evolution of World Tachinidae" project by providing sequence data for the phasiines and several specific outgroup taxa. Phasiinae are the smallest tachinid subfamily but include a remarkable degree of diversity within the various tribes and genera. They mostly attack hemipteran hosts and that preference remains one of the few unifying characteristics of this subfamily. My research has three main goals:

  1. determine if Phasiinae are a monophyletic group,
  2. resolve basal relationships within the Phasiinae, and
  3. resolve intertribal relationships within the Phasiinae.

I also hope to confidently resolve the position of the enigmatic genera Strongygaster, Euthera, Epigrimyia, and Catharosia within (or outside of) Phasiinae. I will be using over 8kb of sequence data from four single copy nuclear coding genes (MAC, MCS, CAD and LGL). As many worldwide genera as possible will be analyzed, with representative genera from each tribe included. I am hoping for at least 50 genera (we currently have 38). At least 35 intra- and interfamilial outgroup taxa will also be included. Maximum parsimony and maximum likelihood inference methods will be used to reconstruct phylogenies in separate analyses of each gene and all data together as a combined dataset.

Zachary L. Burington

Department of Biological Sciences, 3640 Colonel Glenn Highway, 235A, BH, Wright State University, Dayton, Ohio 45435, USA. E-mail: keroplatus@gmail.com

Zachary Burington

I am a PhD student at Wright State University working under Dr. John Stireman on the "Phylogeny and Evolution of World Tachinidae" project. My background is mainly in the taxonomy of Trichoptera and other aquatic insects, but following my M.Sc. at Clemson University in 2011 I became interested in true flies. I had plans to work on fungus gnats for my Ph.D., but when I saw this opportunity at WSU I eagerly applied. My focus in the phylogeny project will be the tribe Blondeliini, especially looking at the relationships of those members with keeled abdomens and piercing ovipositors. This complex of genera, including Eucelatoria Townsend, Blondelia Robineau-Desvoidy, Compsilura Bouché, Vibrissina Rondani, and others, has been treated as a natural group based on morphological characters. I plan to infer a rough phylogeny of the tribe and determine the singular or multiple origin of the "keeled piercer" character. I also (tentatively) plan to revise the large and difficult New World genus Eucelatoria. This will include a redescription and key for the Nearctic species, as well as descriptions of new species from Central and South America. The rough Blondeliini phylogeny will be used to determine the scope of the tribe and any genera that clearly do not belong in relation to the core group. The third portion of my dissertation will be devoted to a longitudinal comparison of tachinid diversity between the Neotropics and the Nearctic temperate region. This will use a year length time series of species and morphospecies from tropical, subtropical and temperate sites, including Yanayacu Biological Preserve in Ecuador and the Wright State University Forest.


Cerretti, P. (2010) I tachinidi della fauna italiana (Diptera Tachinidae) con chiave interattiva dei generi ovest-paleartici. Cierre Edizioni, Verona. Volume I: 573 pp. Volume II: 339 pp. + CD ROM.

Cerretti, P., Tschorsnig, H.-P., Lopresti, M. & Di Giovanni, F. (2012) MOSCHweb – a matrix-based interactive key to the genera of the Palaearctic Tachinidae (Insecta, Diptera). ZooKeys, 205, 5–18.

Stireman, J.O. (2002) Phylogenetic relationships of tachinid flies in subfamily Exoristinae (Tachinidae: Diptera) based on 28S rDNA and elongation factor-1α. Systematic Entomology, 27, 409–435.

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