1999 Progress Report

The PEET program has three fundamental goals: 1) training the next generation of taxonomists, 2) electronic information management and dissemination, and 3) monography. We are privileged to have both NSF and Schlinger Foundation support to undertake these intermeshing goals.

1. Training. The training component of this PEET, although multifaceted, is directed primarily towards educating a new generation of Diptera systematics. Those being trained at the University of Illinois at Urbana­Champaign (UIUC) are under the supervision of Irwin, those at North Carolina State University (NCSU) are under Wiegmann, and those at The University of Queensland (UQ) are under Yeates. Graduate students benefit from frequent exchanges among the localities: learning molecular phylogenetics with Wiegmann and morphological and biological aspects with Irwin and Yeates, database training with Irwin, and collecting strategies during field expeditions.

Grant Products


1A. Degree

1B. Informal

1C. Non-degree

1D. Internships

2A. Developing MANDALA

2B. Data Capture & Verification

2C. Output, Data Manipulation, and Report Structuring

2D. Interactive Keys

2E. Linkages to the Internet

2F. Networking

3A. Obtaining Study Material

3B. Cladistic Analyses & Inference

3C. Monographing Taxa

3D. Differentiating Among Treated Taxa

3E. Illustrating Taxa & Character States

3F. Gaining a Knowledge of Therevid Bioecology

3G. Disseminating Knowledge

Grant Products

Degree Training

Our training effort has focused on graduate students, with an emphasis on Ph.D. degrees that train experts in the fields of systematics and Dipterology. Although all of our graduate students (listed below) will probably remain in systematics and Dipterology, none will likely work primarily on the Therevidae once they graduate. For instance, Gaimari is working on Lauxanioidea, Holston will work on Asilidae, Metz on Syrphidae, Winterton on Scenopinidae, Hauser on Stratiomyidae, Yang on higher level phylogenies within the Diptera.

1A Degree

Stephen D. Gaimari completed a Ph.D., Dept. Entomology, UIUC, in June ’98. His study made great headway towards monographing a dominant higher level taxon from the Americas and Asia. A portion of his dissertation is in press, with other parts being prepared for submission. Gaimari received intensive, hands-on training in molecular systematics at NCSU. He accepted a Visiting Lectureship in the Dept. Entomology, UIUC, where he taught Evolution and Classification of Insects (ENT 302), fall ’98 semester. After submitting several manuscripts based on his graduate work, Gaimari will begin a Smithsonian Institution Postdoctoral Fellowship in March ’99, during which he will focus on resolving the basal lineages of the Lauxanioidea (Diptera: Acalyptratae).
Kevin C. Holston successfully defended his M.S. thesis in Nov.’98 on nomenclatural challenges presented by the species-rich genus Thereva. The thesis is currently being crafted into two papers for publication. He will graduate with a Masters of Science in May ’99. He completed all course requirements for a Ph.D. at the Dept. Entomology, UIUC, Dec.’98 and successfully passed the written exams required for advancing to Candidacy for a Ph.D. He will take the oral exams in March ’99. His Ph.D. dissertation will define the clade that contains Thereva and monograph a diverse genus within that species-rich clade. He is on track to receive a Ph.D. in 2000.
Shaun L. Winterton completed a Post-Graduate Diploma in Science (Entomology), July ’97, Dept. Entomology, UQ. His thesis, in press, monographed a new Australian genus of stiletto flies containing 20 new species. His Ph.D. Candidacy began August ’97. He has now defined the clade containing the Australian genus Agapophytus and made exceptional progress toward monographing its complex nominal genus with 70+ species. Feb. and March ’99 were spent at NCSU, where he will work out the phylogenetic relationships among the Australian genera belonging to theAgapophytus­Group using molecular techniques. He is on track to receive his Ph.D. in 2000.
Mark A. Metz completed coursework in May ’99 towards a Ph.D. in Entomology, UIUC. In addition to coursework in entomology, he has taken courses in evolutionary ecology and mathematical modeling. He has completed the written exams and will take the oral exam in mid February for advancing to Candidacy for a Ph.D. He has monographed a clade of 13 species in 3 Lindneria and 2 new genera, one of which is represented by a Dominican amber fossil (submitted). For his dissertation study, he has begun to examine the phylogeny of a species-rich clade within the Therevinae that includes the genus Brachylinga.
Martin Hauser, Diploma, Technical Univ., Darmstadt, Germany, had a two-year traineeship at the Museum of Natural History, Stuttgart. He arrived in August ’98 to begin his Ph.D. studies in the Dept. Entomology, UIUC, accepting the PEET research assistantship just vacated by Gaimari. He has completed the core course in Insect Ecology, proficiencied out of the Evolution and Classification of Insects core course, participated in the fall ’98 ENT 426 seminar (toxicology), and is enrolled in Insect Genomic Analysis. Hauser, with a Diptera background has begun his research into the phylogeny of the Phycinae (Therevidae) and will monograph its included taxa.
Longlong Yang joined the Dept. Entomology, NSCU, in July 1996. His training has involved courses and labs in molecular systematics, genetics, statistics, insect ecology and evolutionary biology. He will take his Ph.D. qualifying exams in May ’99. His dissertation topic focuses on the higher level phylogenetic relationships of Therevidae based on nucleotide sequence data. His research to date includes data collection and phylogenetic analyses for two genes in 45 therevid genera, which form the basis of a manuscript submitted for publication. He is on track to receive a Ph.D. degree in late ’99 or early 2000.
Narelle Power completed a Bachelor’s degree with Honours at UQ in Dec.’98. Her thesis, which is being prepared for publication, analyzed the spatial and temporal distribution of Therevidae in SE Queensland. Power will work as a research assistant in Yeates’ lab during ’99 and will monograph a species-rich Australian therevid taxon for a Ph.D. dissertation starting in 2000.
J. Marie Metz (Jill D. Mullett), a part-time graduate student in Art Education, UIUC, is the illustrator for the therevid project. In training to become a scientific illustrator, she worked with Irwin and graduate students to interpret structures and represent them accurately. Mullett has pursued training in scientific illustration by using therevids as models for class projects and teaching demonstrations, by becoming a member of and attending annual meetings of the Guild of Natural Science Illustrators, and through networking with Guild members. She is becoming efficient at using electronic media to hone her illustrations for publication, has become a resource for other illustrators across the country and around the world, and is building an impressive reputation within the scientific illustration community.

1B Informal Training

Degree training we undertake contains aspects that receive no formal credit but greatly influence the potential to construct quality monographs and obtain desired career positions. These fall into two skills categories, scope and breadth in science (electronic media, language, systematics, cladistics, nomenclature, Dipterology, illustration, and fieldwork) and professional development (journal article writing, grant writing, teamworking, networking, oral presentation, resume construction, interviewing).

Systematics Training. Graduate students use PAUP, MACLADE, CLADOS, HENNIG 86, COMPONENT, etc. for cladistic analyses to resolve phylogenetic relationships within specific clades of taxa. All students have access to regular interchange of ideas relating to systematics both among other members of our PEET group and groups reviewing systematics articles on campus. Wiegmann‘s lab is responsible for molecular phylogenetics training, including techniques and interpretation. Gaimari spent 6 weeks in intensive molecular training and Winterton began a three-month internship in ’99. Holston, Metz, and Hauser will join Wiegmann’s lab in spring 2000.
Computer Skills and Techniques. All graduate students and many undergraduates have been trained in a number of computer skills involving, among others, data entry and manipulation of database files; GIS manipulation techniques; software for cladistic analyses (see above); construction of presentations for posters, slides and direct computer output; use of internet-available and CD ROM based gazetteers for refining geographical information; and basic word processing/spreadsheet applications for preparing reports and manuscripts. Some have also been involved in web page construction, scanning and manipulation of print images, and plotting distribution maps.
Scientific Illustration. Although it is important to have a scientific illustrator attached to the project (see Monography section: Illustrating Taxa and Character States), it is also important to ensure that graduate students are capable of and practiced in rendering illustrations for their monographs. To this end, Mullett mentors graduate students in drawing techniques. In April ’98, she conducted a workshop designed to teach 6 of our team to render scientific illustrations professionally and accurately. She is currently helping students produce line drawings that are scanned into the computer and filled with various textured pattern swatches from a library she has created. In Queensland, Winterton is learning illustration techniques fromChristine Lambkin, a professional scientific illustrator with considerable taxonomic experience, who is another Ph.D. student in the Yeates’ lab.
Field Work Skills. We have invested heavily in fieldwork skills for graduate students. These skills provide a foundation for understanding organisms within their environments and allow a better interpretation of morphological and behavioral characteristics upon which a large portion of the systematics is based. We not only insist on fieldwork, we encourage team fieldwork. For instance, Irwin and Webb led a two-week expedition to Guatemala for five graduate students and the PIs, Ev Schlinger, Webb, and 3 graduate students, for two weeks, discussed Diptera ecology and collected therevids while encamped in the Cederbergs of South Africa. Activities like these not only hone field-related skills, they provide mechanisms for cementing team relationships.
Diptera Identification. The UIUC contingent meets Tuesday evenings to sort and identify accumulated Diptera in the Illinois Natural History Survey insect collection. This provides an atmosphere to learn from one another and results in labeled, curated, and classified specimens to be shipped to specialists for monographing and determinations. Visiting dipterists lend their expertise when present Tuesday evenings. This has begun to pick up momentum with additional graduate students joining in, but working on other groups.

1C Non-Degree Training

Twenty undergraduates have obtained training on the database component of this grant, one of which is now a principal databaser with another PEET project. These students now know what constitutes a database, how to enter and manipulate data, and how to obtain further information such as geographic coordinates from USGS sites on the WWW and from Microsoft Encarta™ Global Atlas. Many have taught new students these skills, and one has mastered RangeMapper™ for plotting distribution maps based on coordinate data exported from MANDALA (see species in Chromolepida key).

1D Internships

We have begun an internship program for scientists in countries with severely threatened biotas. Our first intern, Rasolondalao Harin’Hala Hasinjaka (Rin’ha), is attached to the Institute for the Conservation of Tropical Environments (ICTE), Tsimbazaza, Madagascar. He will begin his internship at UIUC and the Illinois Natural History Survey in September ’99. Rinah is a general guide for scientists while they are investigating the biodiversity of Madagascar and has expressed special interest in becoming trained in the science of entomology. Once he returns, he will help curate insects in the national museum, begin a program in monitoring insect abundance and activity throughout Madagascar, and strengthen the national collection of insects.

2. Electronic Information Management and Dissemination. Electronic information management is the keystone of our PEET project. It permeates all aspects of study and structures our efforts. The management component includes the development of MANDALA, our comprehensive database system; entering and verifying data on specimen, label, nomenclatural, and loan information into the system; output, data management, and report structuring; interactive keys; linkages to the internet; and networking. Beyond that, considerable effort has been expended to develop an interactive WWW site that can handle selected portions of the MANDALA database and communicate our activities and findings to multiple client groups.

2A Developing Mandala

Designing our own database using an off-the-shelf engine FileMaker™ Pro was not our original intention when we applied for a PEET grant in early ’95. We were beta-testing Colwell’s Biota, which showed promise, but the released version that might have allowed us the flexibility we needed, came too late; we already had our own. Starting with 5 database files in a quasi-relational but cross-platform (Mac and PC) environment, MANDALA, now comprised of 24 files, is fully relational. Recently, an Apple iMac and FileMaker Pro Server software were purchased as a dedicated server for MANDALA. Now concurrent data entry is possible to authorized individuals using both PC and Mac platforms. A second iMac is a dedicated server for our searchable databases over the web. It is also now possible to obtain a demo version of MANDALA via ftp. Expanding the web presence of MANDALA is a high priority over the next year is coming soon.

2B Mandala: Data Capture & Verification

woman at computerThe on-going refinements do not preclude data entry and verification by our team of undergraduates. Data entry, especially of specimen information, proceeds at a rapid pace. Our strategy has been to enter and verify specimen data of taxa that are currently being monographed so that these data are available for manipulation. As of this writing, nearly 62,000 specimens have been processed.

2C Mandala: Output, Data Manipulation, & Report Structuring

MANDALA has been designed to organize and output data in informative ways. Three broad categories are touched on below.

Internal Use. MANDALA is a research tool. New data are constantly being added; ongoing structural changes are making data more easily accessible. It is composed of data in various stages of verification; literal transcription of the label information and a value added version that fills in knowledge gaps such as geographical coordinates; and constructed not only with valid and invalid taxonomic names but also with working, manuscript, and in press names in mind. For this reason we differentiate between what we offer to the general public on the WWW and what is intended for internal, prepublished use. Specimens in MANDALA have links to taxonomic names, collecting events, illustrations, literature, associated specimens, plant associations, ecological and bioactivity associations via a controlled vocabulary, biogeographic regions, loan and deposit information from collections, determinations, and a wealth of other characteristics about the physical specimen itself, including sex, mounted state, dissections, GENBANK ID, developmental stage collected and in collection, and pupation and emergence dates. The taxonomic names file (NAMES.FP3) allows the user to document all facets of name use, including references verifying that use. Using MANDALA to track museum loans and provide counts of available specimens of each species, Gaimari was able to organize loan returns and additional specimen distributions of some 35,000 Ozodiceromyia specimens.
Client Requests. We have addressed a host of questions about therevids using our MANDALA database from a number of client groups, e.g., generated a report on therevids from Florida (including species, collecting dates, and counties) for Gary Steck (Florida State Collection of Arthropods); generated a list of species and specimens found in the National Parks and National Monuments of Colorado for Virginia Scott (University of Colorado at Boulder Museum). We have also routinely used the database to generate accurate and detailed accounts of material on loan from various museums and material being returned to those museums.
Publication and Poster Layout. Gaimari generated a specimens examined list from MANDALA for his thesis and upcoming publications. He and others have used MANDALA to output the literature cited for publications. Metz and Irwin’s symposium presentation on the Therevidae of Baja California was made possible by data recorded and manipulated in MANDALA. The geographical coordinates data were exported from the database to RangeMapper™ for plotting distributions.

2D Interactive Keys

Systematists (led by Yeates), multimedia designers, and programmers at the University of Queensland have developed LucID, a computer program system designed for building and using interactive keys for the identification of organisms in multimedia environments. The program allows the user to begin the identification of an organism with any character and continue in any desired character order. Still images, video, and sound may be accessed at any stage to increase the speed and accuracy of identification. A galaxy of information, images and other resources may be retrieved using the organism’s name as an index once the identification is complete. Interactive keys to the genus Chromolepida and to the genera of the Cyclotelini were constructed and are being used at UIUC. They will be integrated with other aspects of web-based information once software currently being developed allows LucID to interact with the WWW. We use LucID in demonstrations to students of all ages as an example of how people go about identifying organisms that they have in front of them. LucID is now distributed by CSIRO Publishing (version 1.5 to be released shortly).

2E Linkages to the Internet

The family Therevidae has been on the WWW since April ’96 with periodic updates and additions. The site details our PEET research, profiles therevid research teamprovides minutes of meetings, and recounts therevid collecting expeditions and museum visits. MANDALA is in the process of becoming accessible via the WWW with a world checklist of taxa; a listing of literature; detailed information on names connected with Thereva; and prototype full exposure to the capabilities of the database using the genus Chromolepida. As other material is verified and published, it will be added to the public view of MANDALA. We have acquired a writer to produce CD-ROMs of our PEET products and the FileMaker Pro Developer package for runtime, cross-platform versions of MANDALA.

2F Networking

Networking has assumed many faces, among which those dealing with MANDALA are highlighted:

PEET Participants
Standing L to R: Gail Kampmeier, Barbara Crandall-Stotler, Ray Stotler,Dave Krayesky, and Jimbo Bray; seated: Bob Pollok (former therevid-PEET databaser, now at SIU)
MANDALA has been adapted for use with taxa other than therevids. Representatives from other PEET projects [Ray Stotler & Barbara Crandall-Stotler (liverworts), Southern Illinois University; Pam Arnofsky (aplacophorans), Woods Hole Oceanographic Institute; Peter Hibbs (snipe flies), Smithsonian Institution] visited our laboratory for training in ’98.
Stotler’s group hired one of our undergraduate student aids in their modifications of MANDALA. Another potential PEET project (on microsporidia, L. Solter et al.) proposes to modify MANDALA for its use.


 Robin Carlson (right, visiting Irwin lab for training) and Sally Regan, part of the Schlinger acrocerid project, continue to use MANDALA to catalog their dipteran spider parasitoids. Gaimari now uses MANDALA to catalog the Lauxanioidea (Diptera). Other individuals (Elden Neal, Southern Illinois University; Ed Dewalt & Kathy Zeiders, Illinois Natural History Survey; Ebbe Nielsen, CSIRO, Canberra, Australia; Lynn Hanson, SIRIC Librarian, INHS; Geoffrey Bowker & Shubha Nagarkar, School of Library Science, UIUC) were given demonstrations at UIUC and clones of MANDALA.  Robin Carlson

Others such as Mark Wetzel (INHS) and individuals from the 4th International Diptera Congress (Dr. Gerhard Baechli, Zoological Museum, Zurich, Switzerland; Marc Pollet, Royal Belgian Institute of Natural Sciences, Belgium; Thomas Pape, Swedish Museum of Natural History, Sweden; Kai Heller, Forschungstelle für Ökozystemforschung, Germany; Matthias Buck, University of Ulm, Germany;) and the ’98 Taxonomic Databases Working Group meeting (Nick Parker & Richard Monk, USGS/Texas Tech University, Lubbock, TX; Georgina MacKenzie, BIOSIS, U.K.; Ian Reid, CSIRO, Canberra, Australia; Charles Hussey, The Natural History Museum, London, U.K.) saw demonstrations and a poster of MANDALA and received demo versions. Two people working at FileMaker, Inc. also requested demo versions of MANDALA (Jeff Gagne & James Gudeli).

3. Monography. Monography needs material for study, and embraces cladistic analyses and inference (including characterizing the lineage of that clade), differentiating among the taxa being treated, and entails illustrating taxa and character states, gaining a knowledge of therevid bioecology, and disseminating knowledge. The last step is highlighted in “Electronic Information Management and Dissemination”” and “Grant Products.” Manuscripts of monographs are just beginning to emerge; the next year-and-a-half should witness the production of several large monographs.

3A Monography: Obtaining Study Material

Insect specimens are the fundamental building blocks for all that is accomplished in monography. Studying primary types is essential for stabilizing nomenclature. Obtaining high quality material in sufficient quantity requires a concerted and consistent effort. We have approached this through three avenues.

Acquiring Specimens from Existing Collections. Museums have been combed for specimens of the family Therevidae. The worldwide systematic community has responded expediently to requests, and many have collaborated with and helped students and PIs during visits to their institutions and via letters and email. We have specifically borrowed material within the species-rich genera that we are monographing from over a hundred museums. We estimate that we now have on loan about 100,000 specimens from museums in all over the globe.
Examining Types and Other Material at Museums and Private Collections. We lack many primary types and material that remains unsorted in collections. We have visited many museums around the world to study primary types and uncover unsorted therevids. Trips have been taken to St. Petersburg, Paris, Vienna, London, Oxford, Copenhagen, Budapest, Madrid, Berlin, Eberswalde, Wiesbaden, Stuttgart, private collections of Geller-Grimm, Kassebeer, and Schmid-Egger in Europe; Tel Aviv in the Middle East; Pietermaritzburg, Cape Town, Windhoek, and Nairobi in Africa; Canberra, Sydney, Brisbane, Perth, Melbourne, Adelaide, Launceston, and Hobart in Australia; Tananarive in Madagascar; Auckland in New Zealand; Sao Paulo, Rio de Janeiro, Santiago, Concepcion, La Serena, Lima, Quito, La Paz, and Buenos Aires in South America; San Jose (INBio), Guatemala City, and Zamorano in Central America; Mexico City, Ottawa; and many of the important collections in the United States in North America.
Increasing the Knowledge Base of Therevidae Through Expeditions. Because many areas of the world are poorly collected and because, even when abundant, therevids are difficult to collect, there are enormous taxon and geographic gaps in material in collections, and specimens of key taxa needed for molecular studies have not been properly preserved. For these reasons, we undertook a number of collecting expeditions to remote areas of the globe over the past 3 years, which were made possible by the cooperation and generosity of countless colleagues and facilitators for, among other things, hospitality and obtaining collecting and exit permits.

Our expeditions had to meet three criteria: the potential to

1) gather specimens of critical taxa for both morphological and molecular studies;
2) increase the holdings of specimens from areas lacking representation in the world’s collections; and
3) explore areas of expected radiation but that have not been collected.

Expeditions thus far have included Australia (Queensland, NSW, Victoria, ACT,Tasmania), New Caledonia, New Zealand, South Africa (Natal, Cape), Namibia, Chile,ArgentinaMadagascar, Israel, Denmark, Guatemala, Mexico (Baja Calif.), USA (ColoradoNew Mexico, ArizonaTexasCalifornia, and Utah).

Moreover, a number of Malaise traps, which are excellent for gathering therevid material, were left with colleagues in many of these countries and have produced a steady stream of material over several seasons, allowing us not only to increase our holdings, but to understand the flight phenologies of therevids in specific areas of the world. Material other than therevids collected during our expeditions has been shared with many museums and taxonomists. Wharton and Wooley (PEET project, Texas A&M Univ.) received Hymenoptera Parasitica; the National Pollinating Insect Collection, Logan, received bees; The Bohart Museum, Univ. Calif., Davis received a wide diversity of wasps; the Illinois Natural History Museum collection has been the depository of misc. material.

3B Monography: Cladistic Analyses & Inference

Although monographing clades of Therevidae is a major goal of our project, as predicted, difficulties arose early in defining discrete monophyletic units. The family Therevidae had no internal hierarchy except for subfamilial lineages when the project was initiated. An early objective, therefore, was to clarify basal and delineate near-basal clades so that monophyletic groups could be established and monographed. To this end, much progress is being made on a major manuscript by Irwin and Yeates, using 94 adult morphological characters, to generate a phylogeny for 55 in-group taxa (chosen from a world-wide sampling of genera) with 5 out-group taxa. Independently, using a large subset of the same taxa, Wiegmann and Yang produced a phylogeny based on nucleotide sequences of two genes (28S rDNA and EF1a). Dopa decarboxylase and PEPCK genes are being amplified to determine their potential to differentiate among therevid genera and asiloid taxa. The cladistic analyses from morphology and molecular data sets, for the most part, support one another, especially at the basal and near-basal nodes. These two sets of information now permit the selection of discrete monophyletic units to monograph. Not all lineages have been resolved (this proposal points to lineages that remain poorly differentiated and need clarification through added taxa and character analyses).

Students engaged in monographic revisions of specific genera also defined the clade to which that particular genus belonged. No easy task, the genera selected for monography are species-rich, and their near relatives are unknown and potentially from any biogeographic region of the world. Two students hit paydirt in hypothesizing that discovered synapomorphies would define inclusive clades. Gaimari has defined the Tribe Cyclotelini, containing the species-rich genus Ozodiceromyia, and Winterton defined the “Agapophytus Group” within which lies the diverse nominal genus. Holston and Metz are taking the same approach with Thereva and Brachylinga.

3C Monographing Taxa

We are attempting to monograph a wide range of clades during this grant. The approach has been to select for revision a species-rich genus that seems to be within a cluster of related genera. Through this strategy, we hoped to begin piecing together the world Therevidae. A number of genera have been worked on during the three years of our project. Following are the genera for which manuscripts have not yet been submitted followed in parentheses by the number of species involved: Acrosathe (20), Agapophytus (>70), Anabarhynchus (>200), Bonjeania(20), Brachylinga (45), Cyclotelus (>60), Lindneria Group (11), Ozodiceromyia(120), Pandivirilia (25), Thereva (250), and several Australian genera with manuscript names (25). Spiriverpa, a Holarctic genus, is under study.

3D Monography: Differentiating Among Treated Taxa

Dichotomous keys have been constructed to the therevid taxa being monographed. Furthermore, considerable effort has been exerted to develop tools to interactively identify organisms. Yeates’ team (Univ. Qld.) has developed the software package LucID, an electronic, interactive key that includes a builder module, is compatible with DELTA, and is being adapted for WWW interface. We have tested the capabilities of this key format by building a prototype LucID key to the species within the genus Chromolepida Cole and to the genera of Cyclotelini. A web-based dichotomous key to the Chromolepida is also now available.

3E Monography:  Illustrating Taxa & Character States

Illustrations are indispensable for monographs. Obtaining sufficient high quality illustrations for our monographic studies has been an obstacle during the course of this project. To overcome this, we have used a number of techniques and continue to explore new ones as they present themselves.

Anabarhyncus tristis  Habitus Photographs. A photographic library of habitus poses of therevids, developed by Yeates and Anthony O’Toole, is maintained at the Univ. Queensland. Fourteen species in 9 Australian genera have been photographed while living. Other photos displayed are used by permission from the photographer.

Light Imaging Systems. Two PIs and 3 graduate students visited Sabine Huhndorf’s (PEET grant, fungi) lab at the Field Museum of Natural History, Chicago, to observe computer image capturing and manipulation techniques. Although more optimal for smaller, flatter objects, we realized the applicability to some of our imaging needs and submitted a proposal to the Illinois Natural History Survey’s request for future equipment (year 2000), and we plan to adapt this digital imaging system to a Wild MZ12 if INHS funds become available.
Environmental Scanning Electron Microscope. This past year, the Beckman Institute at UIUC spearheaded a proposal to NSF for a field-emission environmental scanning electron microscope. Irwin, a collaborator on the proposal, was interested in using the ESEM (no coating, no vacuum, no special mounting needed; i.e., one can even use holotypes or live larvae and pupae) for imaging small external anatomical features of therevids to complement the illustrations provided by other means and to investigate microstructural features as possible characters for phylogenetic and taxonomic studies. The proposal was funded and an XL30 ESEM-FEG (FEI Co., Hillsboro, OR) has been purchased along with a special, just released LF-GSED detector (large field, low magnification gaseous secondary electron detector). Irwin’s group is being trained in its use and will take advantage of this extraordinary piece of equipment over the next 5 years.
Scientific Illustrator. Perhaps the most important, if traditional, way to obtain clear, accurate illustrations is through a highly skilled scientific illustrator. J. Marie Metz, the illustrator on our project, uses mostly carbon dust but also uses pen and ink, mixed media, and acrylic air brush (for color) techniques. She acquired studio space in the Dept. Art Education for airbrush paintings at no cost to our project. To date, she has produced >100 illustrations (mostly internal spermathecae and associated sacs) for the mss. on the genus Anabarhynchus, >165 drawings, primarily in mixed media, for a submitted mss. on the Cyclotelini, 20 drawings for the adult cladistic mss., and >40 drawings for the Lindneria mss. She has mastered electronic modification of her illustrations and is scanning each illustration to store them electronically and cataloguing them in the illustrations portion of MANDALA.

3F Gaining a Knowledge of Therevid Bioecology

A number of ecology studies of stiletto flies have been initiated. Power’s thesis onspatial and temporal distributions of Therevidae in southeast Queenslandrepresents one of the most extensive and intensive quantitative surveys of Therevidae yet conducted. We now have a much greater knowledge of the form of the therevid bioscape in southeastern Queensland.


Namib Desert, Namibia
Namib Desert, Namibia

 A year-long sampling of therevids along theKuiseb River of the Namib Desert, Namibia, has shed considerable light on the flight phenologies of the 4 species that were captured. Yet another study at Sedgewick State Park near Santa Barbara, California, is providing altitudinal richness and flight phenologies of the stiletto flies.

A similar study, undertaken in the central foothills of Chile has provided an excellent account of the timing, relative abundance, and species richness of therevids in that part of the world. An intensive study of the coastal dunes of central California revealed habitat boundaries of 5 stiletto fly species inhabiting that dune system.

3G Dissemination Knowledge

Taxonomy for Tots A number of knowledge dissemination media have been used in our project: oral and poster presentations at meetings and congressespublications in journals and other print media, and by theInternet . In addition, a number of team members have interacted with members of the public, particularly children, in Insect Expos (see Mark Metz left with “Taxonomy for Tots”), and otherspecial events for outreach. Several team members have also taught classes or courses involving the principles of insect systematics at the university level.


Grant Products


  • Irwin, M. E. 1997. Therevidae. In: Solís, A. (ed.), Las Familias de insectos de Costa Rica. INBio.
  • Irwin, M. E. and G. E. Kampmeier. 1997. PEET: A training effort that is paying dividends. Illinois Natural History Survey Reports 347: 2. (September/October).
  • Kampmeier, G. E. 1997. Therevid databases: Reinventing the Wheel or Prudently Hitching a Ride? Entomological Collections Network 1996 annual meeting, Louisville, KY.
  • Kampmeier, G. E. and M. E. Irwin. 1998. Database gathers taxonomic information. Illinois Natural History Survey Reports, 353: 4, 8.
  • Ovtshinnikova, O. G. and D. K. Yeates. 1998. The male genital musculature of Therevidae and Scenopinidae (Diptera: Asiloidea): Structure, homology, and phylogenetic implications. Australian Journal of Entomology 37: 27-33.
  • Yeates, D. K. and B. M. Wiegmann. 1999. Congruence and controversy: Toward a higher-level phylogeny of the Diptera. Annual Review of Entomology 44: 397-428.
  • Irwin, M., P. Raven, J. Oliver, E. Wilson, E. Schlinger, R. May. Draft Entomological Society of America Position Statement: Insects and Biodiversity. ESA Newsletter 22 (1): 1, 4. [contains statement on importance of PEET program.]

Dissertations and Theses

  • Winterton, S. L. 1997. Systematics of [a new genus] (Diptera: Therevidae) from Australia. Postgraduate Diploma in Science. The University of Queensland, Australia. 99 pp. + 185 figs.
  • Gaimari, S. D. 1998. Phylogeny, classification, and biogeography of the cycloteline Therevinae (Diptera: Therevidae). 1998. Ph.D. Dissertation, University of Illinois at Urbana-Champaign. 273 pp.
  • Power, N. 1998. Diversity and Spatial Distribution of Therevidae in South East Queensland. Thesis, Bachelor of Science (Honours), Department of Entomology, The University of Queensland, Brisbane, Australia. 169 pp.
  • Holston, K. C. 1999. Answering the Nomenclatural Challenge with a Database ofThereva Names (Diptera: Therevidae). M.S. Thesis, University of Illinois at Urbana-Champaign. 150 pp.

In Press

  • Gaimari, S. D. and M. E. Irwin. 1999. Phylogeny, classification, and biogeography of the cycloteline Therevinae (Diptera: Therevidae). Zool. J. Linn. Soc.
  • Winterton, S. L., M. E. Irwin, D. K. Yeates. 1999. Systematics of [a new genus] (Diptera: Therevidae) from Australia. Invertebrate Taxonomy.
  • Winterton, S. L., M. E. Irwin and D. K. Yeates. 1999. Systematic revision of theTaenogera Krober genus-group (Diptera: Therevidae), with descriptions of two new genera. Australian Journal of Entomology.


  • Gaimari, S. D. and M. B. Mostovski. A new genus for Psilocephala electrellaCockerell from burmite (Diptera: Asiloidea). Bulletin of the Natural History Museum (London). [special volume on burmite geology and paleontology].
  • Metz, M. A., and M. E. Irwin. 1999. A new genus from Dominican amber and its phylogenetic relationship with the extant Neotropical genus Lindneria Krober (Therevidae). Annals Entomol. Soc. Amer.
  • Webb, D. W. and M. E. Irwin. 1999. A revision of Tabuda Walker and TabudamimaIrwin and Lyneborg with the description of a new genus [B] Webb and Irwin (Diptera: Therevidae: Therevinae). Annals Entomol. Soc. Amer.
  • Winterton, S. L., J. H. Skevington, M. E. Irwin, and D. K. Yeates. 1999. Phylogenetic revision of Bonjeania Irwin and Lyneborg (Diptera: Therevidae)Systematic Entomology.
  • Winterton, S. L., D. J. Merritt, A. O’Toole, D. K. Yeates and M.E. Irwin. 1999. Morphology and histology of the spermathecal sac, a novel structure in the female reproductive system of Therevidae (Diptera: Asiloidea). International Journal of Insect Morphology and Embryology.
  • Yang, L. L., B. M. Wiegmann, D. K. Yeates, & M. E. Irwin. 1999. Higher level phylogenetic reconstruction of the Therevidae based on 28S ribosomal and elongation factor-1alpha gene sequences. Molecular Phylogenetics and Evolution

In Preparation

  • Gaimari, S. D., and M. E. Irwin. Phylogenetic relationships within the cycloteline genus Ozodiceromyia Bigot (Diptera: Therevidae: Therevinae). Thomas Say Miscellaneous Publications.
  • Gaimari, S. D., and M. E. Irwin. Phylogenetic revision of the mexicana-group ofOzodiceromyia (Diptera: Therevidae). Annals of the Entomological Society of America.
  • M.E. Irwin and D.K. Yeates, Phylogeny of the Therevidae based on adult morphological evidence.
    Tsaur, S.C., B.M. Wiegmann, B.K. Cassel and D.W. Webb. Monophyly and relationships of the Tabanomorpha (Diptera: Insecta).Annals of the Entomological Society of America.
  • Wiegmann, B.M., S.C. Tsaur, and D.K. Yeates. Synthesis of molecular and morphological evidence on phylogeny of the brachyceran Diptera (Insecta).Systematic Biology.
  • Wiegmann, B.M. D.K. Yeates, J.L. Thorne, and H. Kishino. A revised time-scale for fly evolution: Divergence time estimates without a molecular clock. Proc. Nat. Acad. Sci. USA.

Abstracts in Proceedings

  • Gaimari, S. D., and M. E. Irwin. 1998. Systematics and biogeography of the cycloteline Therevinae (Therevidae). Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 58-59.
  • Gaimari, S. D., M. E. Irwin, and G. E. Kampmeier. 1998. Importance of collections and specimen level databases for dipterological research. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 59-60.
  • Irwin, M. E., and A. H. Kirk-Spriggs. 1998. Species composition and flight phenology of stiletto flies and window flies (Asiloidea: Therevidae and Scenopinidae) occurring along the Kuiseb River, Gobabeb, Namibia. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 89.
  • Irwin, M. E., and D. K. Yeates. 1998. The internal hierarchy of the Therevidae and biogeographic patterns of extant higher taxa (Brachycera, Asiloidea). Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 90.
  • Kampmeier, G. E., M. E. Irwin, and F. C. Thompson. 1998. MANDALA: a database weaving together information on specimens, nomenclature, and literature for systematics research. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 98-99.
  • Metz, M. A., and M. E. Irwin. 1998. A new genus from Dominican amber and its phylogenetic relationship with the extant Neotropical genus Lindneria Krober (Therevidae). Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 148.
  • Metz, M. A., and M. E. Irwin. 1998. The relationship of parasitic trombidiid mites with the Diptera in an arid ravine in New Mexico, USA. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 148-149.
  • Winterton, S. L., A. O’Toole, M. E. Irwin, D. Merritt, and D. K. Yeates. 1998. Morphology and ultrastructure of the “spermathecal sac,” a previously unknown structure in the reproductive system of female Therevidae. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 247.
  • Winterton, S. L., J. H. Skevington, M. E. Irwin, and D. K. Yeates. 1998. Revision of the bizarre genus Bonjeania Irwin & Lyneborg (Therevidae) from Australia. Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 248.
  • Yang, L. L., B. M. Wiegmann, and M. E. Irwin. 1998. Multiple molecular datasets resolve the phylogeny of the stiletto flies (Therevidae). Fourth International Congress of Dipterology, Oxford, England, Abstract Volume, p. 250.

Meetings & Seminars to Communicate Advances, Plan Future Activities and Network

  • IVth International Congress of Dipterology. All graduate students, PIs, most collaborators, and Thompson actively participated in this Congress, Oxford, England, 6-11 Sept.’98. Wiegmann and Yeates were elected to the International Council of Dipterology, and Yeates has agreed to host the 5th International Congress of Dipterology in 2002, at the University of Queensland, Brisbane, Australia.
  • University of California at Davis Department of Entomology Seminar. Irwin gave a lecture, entitled, “Dune revisited: Exploring the lives and times of sandworms,” in April ’97.
  • UIUC Department of Entomology Seminar. Gaimari lectured on the “Phylogenetics and biogeography of a Laurasian group of Therevidae (Diptera),” March ’98.
  • North American Dipterists Society Meetings and Field Trips. Our PEET team has been very active in this organization. Irwin chaired the meeting in Louisville KY in 1996; Wiegmann chaired it at Nashville TN in ’97, and Gaimari chaired the meeting at Las Vegas NV in ’98. Wiegmann organized and will lead the ’99 field excursion to the Great Smoky Mountains.
  • Entomological Collections Network. Our team has participated in these activities for the past three years. Wiegmann lectured on the advances the Therevid PEET project had made during the meeting at Mesquite NV in ’98.
  • Taxonomic Databases Working Group (TDWG) Meeting. Kampmeier attended these meetings Toronto, Canada, in ’96 and in Reading, England, in ’98. She has been appointed to the TDWG Executive Committee to help facilitate involvement of zoologists as this group expands its scope.
  • Therevid/PEET Workshops. Each summer of the grant, our entire team assembles at UIUC to discuss the year’s accomplishments and plan for the coming year. See details of the 19961997, and 1998 workshops.
  • NSF PEET Workshops. Irwin, Yeates, and Kampmeier attended the first workshop at Lawrence KS in ’96. Kampmeier demonstrated the databases and Yeates demonstrated LucID to the group. All PIs, the five Diptera graduate students, Kampmeier, and Thompson participated in the second at Woods Hole MA in ’98. Irwin, assisted by Thompson, moderated a session on monography; Kampmeier presented during the database workshop; Holston participated in the student panel. Yang, Winterton, Gaimari, and the Therevid PEET team all presented posters. Metz is on the organizing committee for the next NSF PEET Workshop in 2000.
  • Guild of Natural Science Illustrators (GNSI). Mullett participated in this conference at Santa Cruz, CA in ’97 and at Ames IA in ’98. Her artwork was selected for the professional juried art competition in Ames.
    Missouri Botanical Garden Systematics Symposium. Irwin, Gaimari, Holston, Hauser, and Metz attended the 45th annual symposium St. Louis, Missouri, Oct. ’98.