Date: April 15, 1996
NSF Program: DEB-PEET
NSF Award Number: 95-21925
Period covered by this report: August 11, 1996 - April 15, 1998
PI Name: Michael E. Irwin
PI Organization: University of Illinois at Urbana-Champaign
PI Address: Department of Natural Resources & Environmental Sciences (NRES)
University of Illinois
1101 West Peabody Drive
Urbana, IL 61801
Co-PIs: Brian M. Wiegmann, North Carolina State University
David K. Yeates, University of Queensland
/X/ Continued Funding is Requested
Year 3 NSF PEET Report: Towards a World Monograph of the Therevidae (Insecta: Diptera)
Progress including results obtained to date and their relationship to the general goals of the grant
Summary of work to be performed during the next year of support, if changed from the original proposal; an indication of any current problems or favorable or unusual developments; and any other significan information pertinent to the type of project supported by NSF or as specified by the terms and conditions of the grant.
1A. Use of Electronic Media. This aspect is proceeding much as anticipated in the original proposal. The sheer number of specimens to be processed and verified is still overwhelming but we are becoming more efficient both in data entry and verification.
Networking. The taxonomic community has responded expediently to requests concerning therevid specimens. Many have collaborated with and helped students and PIs during visits to their institutions, and via letters and email. Expeditions conducted during this grant would not have been possible without the cooperation and generosity of countless colleagues and facilitators.
Representatives from other PEET projects (R. Stotler & B. Crandall-Stotler; D. Fautin; P. Arnofsky) and other individuals from scientific meetings (R. Carlson, Ev Schlinger's acrocerid project; V. Moseley, Louisiana State Arthropod Museum) where MANDALA has been demonstrated have received versions and/or training in the use of the databases and their adaptability to other taxa.
Databases. In 1998, the therevid databases, constructed using the cross-platform (Mac & Win95/NT) database engine, Filemaker Pro, finally acquired a real name to reflect their adaptability to a broader scope of taxa than just the Therevidae. Its name, MANDALA, means "interconnectedness," "circle," and "relatedness." Gail Kampmeier, in collaboration with F. Chris Thompson (consultant to our PEET), and with input from all members of the Illinois team, continues to refine MANDALA. The on-going refinements do not preclude data entry by our enthusiastic group of primarily undergraduate students.
The databases were initially demonstrated at the first NSF PEET meeting in Kansas in September 1996. After that meeting and before showing them at the TDWG (Taxonomic Databases Working Group) meeting in Toronto in October 1996, the user interface to the Specimen database underwent extensive revamping, expanding what had been crowded into two layouts into eight, using a 'file folder' metaphor. Also added were layouts for navigation and database management as well as some specialized query layouts. More are planned as researchers refine their search strategies.
Before presenting the databases at the Entomological Society of America's Informal Diptera Conference in December 1996, Gail Kampmeier added the following features:
expanded tracking of multiple taxa determinations and determiners (determnr.fp3);
expanded (unlimited) tracking of museum information, i.e., where specimens are coming from, residing, and moving to (deposit.fp3);
tracking of database users who create or modify records; and
the ability to navigate to related records via a small icon with a magnifying glass.
The following changes were implemented in 1997:
created a "label.fp3" database for typing in label information exactly as it appears without interpretation or additional information. This database speeds up the process of data input (less thought & interpretation required at this step) and provides a place to proof information attached to specimens;
created an illustration database (illus.fp3) to track any type of illustration and link it to specimens, reprints, and taxon names where appropriate;
revamped user interface to lots.fp3 and Reprints.fp3, adding the ability to track collectors and authors respectively in people.fp3 via a join file. This join file is used in the new names.fp3 database (see below) and can handle determiners and illustrators as well so that you can see everything a person has done.
replaced the old flat hierarchy file "taxa" with a new "names.fp3" database, conserving all the old taxon numbers but implementing relationships among taxon names and creating classification hierarchies in "classif.fp3";
added the ability to track multiple associated specimens (assocsp.fp3) and their associations with the specimen in hand;
added the ability to create LIT (literature) numbers as unique identifiers for specimens referred to in the literature that are not associated with physical specimens. Locality/collector information will go into the lots.fp3 db just as for a specimen in hand; etc. Treatment of the specimen in the literature can be tracked using a controlled language (popup selections) in the bioassoc.fp3 database with the addition of fields designating reprint and page number.
Thus far, over 45,000 specimens in over 11,700 lots (unique collecting events) have been recorded, covering nearly all specimens collected from Australia (with the exception of the most recent collections), the generaOzodiceromyia and Cyclotelus, and a smattering of species in other genera.
All specimens of the genus Ozodiceromyia on deposit at the University of Illinois from collections made by our group and from museums worldwide are now in MANDALA, and most specimens have been checked twice for accuracy of data. Students are nearly done with the genus Cyclotelus, which is under revision by Gaimari and Irwin.
The number of records entered into the major databases as of 14 April 1998 (not including material from the 1960s, coded before this project began) is:
Specimens (Specimen.fp3)....................45,143 Labels (label.fp3)..........................16,328 Lots (lots.fp3).............................11,734 Taxa (names.fp3).............................2,680 People (people.fp3)..........................2,214 Museums (museums.fp3)..........................213 Literature citations (Reprints.fp3)............180 Journals (journals.fp3)........................269
PEET student Shaun Winterton visited the Irwin laboratory in April 1997 for an introduction to the use of the databases and returned in March 1998 for more rigorous training with Gail Kampmeier on using MANDALA. Upon returning to Australia he will enter records of specimens of Nanexila andAgapophytus into the database. Therevid specimens collected by the Yeates group during the previous summer in Australia will be sent to the Irwin lab to be entered into the database.
Interactive Keys. Systematists, multimedia designers and programmers at the University of Queensland and the Cooperative Research Centre for Tropical Pest Management in Brisbane have developed the LUCID systema computer program designed for the interactive identification of organisms in a multimedia environment. The system comprises the key shell itself, termed LUCID, and the LUCID builder, a program that allows systematists to quickly write their own interactive keys for use in LUCID. 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 or "hook" once the identification is complete. The program has now been commercially released (Version 1) and funding is now being sought to make Version 2 of the LUCID program compatible with WWW.
P.I. Yeates demonstrated LUCID at both the first and second NSF PEET meetings, as well as at the Entomological Society of America meetings in Nashville in 1997. Highlighted at the second PEET meeting was the recently completed builder feature, which allows swift composition of interactive keys.
Communications. The family Therevidae has been on the World Wide Web since April 1996 with periodic updates and additions as time permits by Kampmeier using the input from all of the therevid PEET project. The site details the objectives of our PEET research, profiles therevid research participants, provides minutes of meetings, and recounts therevid hunting expeditions. Although the databases are not yet searchable via the WWW, portions of them will be made available as information on the various groups is published.
1B Training. The training component of this PEET is primarily through the education of graduate students. Our proposal stated that we intended to train five graduate students in the science of Diptera taxonomy. All of the graduate students are actively monographing important therevid genera from around the world. We have added the training of a scientific illustrator to our program since the last report.
University of Illinois. At the Irwin lab, three graduate students are being trained in systematics and taxonomy, and one in scientific illustration. Matching contributions from the Illinois Natural History Survey (INHS) funded a research assistantship for Mr. Stephen Gaimari, which began in September 1995. Mr. Mark Metz and Mr. Kevin Holston began their NSF-PEET supported assistantships in April and May 1996, respectively. The scientific illustrator, Ms. Jill Mullett, is supported by Schlinger Foundation/NSF PEET matching funds.
Mr. Stephen Gaimari will have completed his Ph.D. by 15 August 1998, and has accepted a position as Visiting Lecturer in the Department of Entomology at the University of Illinois to teach insect systematics and taxonomy in the fall of 1998. Mr. Gaimari has also accepted a Smithsonian Institution Postdoctoral Fellowship, with a starting date of 15 January 1999.
Mr. Gaimari has expanded his dissertation project to define the larger monophyletic group (given tribal status) including the genus Ozodiceromyia. This grouping is based on an hypothesized relationship between Ozodiceromyia and Cyclotelus, and has been expanded to include an additional 8 genera, includingBreviperna, Ammothereva, Procyclotelus, and 5 new genera. Their phylogenetic relationships have been estimated using parsimony, and biogeographical relationships have been hypothesized using component analysis. All species for all new genera in this new tribe are being described, and all genera are fully diagnosed, including keys. The large genus Ozodiceromyiais being most fully treated at the species level, with over 100 species delineated and under description. The molecular work in conjunction with Brian Wiegmann's lab deals with the relationships among distinct groups within Ozodiceromyia to relate to the in-progress morphological study.
Mr. Mark Metz has finished two years of course work towards the requirements of the Ph.D. in the Department of Entomology. He will satisfy his course requirements in the Fall 1998 semester with the completion of one seminar course and take both written and oral preliminary examinations for the Entomology Department shortly thereafter. In addition, Mr. Metz has taken coursework ranging in subjects from evolutionary ecology to mathematical modeling to supplement his formal academic training. Informal aspects of training have included, but have not been limited to, field collecting techniques, insect identification, grant proposal composition, and meeting presentation skills. Computer skills have been emphasized and both coursework and on hand experience in databasing, GIS, HTML construction, and other basic computer programs have been acquired. In addition, Mr. Metz has translated 38 species descriptions from the original German texts and is improving his proficiency in German by taking coursework at UIUC.
Mr. Kevin Holston will complete the last of the required Entomology courses in December 1998. He is expanding his background in foreign languages, specifically German, which will aid in his survey of taxonomic and systematic literature related to this project. He has focused considerable attention on nomenclature challenges presented by the genus Thereva by finding and compiling all species names used in combination with Thereva. He is working closely with F. C. Thompson (PEET consultant) and G. E. Kampmeier (PEET collaborator) to develop a systematic database of Thereva names. Under the guidance of Thompson, Mr. Holston spent a week in January 1998 going through literature to develop and improve the database of Thereva names and their associated references. Mr. Holston has worked extensively with Kampmeier on the names.fp3 file of the therevid database system, MANDALA, to improve its ability to handle nomenclatural concerns.
Ms. Jill Mullett is working towards a degree in Art Education at UIUC, where she also has a teaching assistantship. Since joining our lab group in August 1996, she has worked part-time during the school year and full-time during the summers as a scientific illustrator. In training to become a scientific illustrator, Ms. Mullett has had to work closely with the taxonomists and learn to see and interpret structures so their representation is clearer than a mere photograph. Although UIUC does not offer a formal program in scientific illustration, Ms. Mullett has pursued training in this area by using biological models (most often therevids) for many of her class projects and teaching demonstrations, by becoming a member of the Guild of Natural Science Illustrators, building a network of contacts within the Guild via email, attending the GNSI annual meeting in Santa Cruz, CA in August 1997, and persuading the Illinois Natural History Survey to sponsor a visit and seminar by GNSI member, Scott Rawlins on in situ scientific illustration. Whereas her initial illustrations were pen and ink, at the GNSI conference she received training from Elaine Hodges on using the carbon dust technique, which speeds up the production of high quality drawings and has superseded pen and ink for most of her work. Ms. Mullett has completed over 100 illustrations of the genus Anabarhynchus, which have been sent off to Lyneborg (collaborator) for publication in the revision of that genus. Nearly all the illustrations (ca. 165) for theOzodiceromyia (Cylotelus-group) project are complete for a manuscript being prepared by Gaimari and Irwin.
In April 1998, Ms. Mullett led a 3 h workshop at UIUC designed to teach scientists how to render scientific illustrations professionally and accurately. Those attending the workshop (Gaimari, Holston, Irwin, Kampmeier, and Metz from UIUC and Winterton from UQld) were able to complete and understand a tonal scale and to take a sketch of a fly and begin a carbon dust drawing.
Mr. Longlong Yang is continuing his investigation of the higher level phylogenetic relationships of Therevidae based on nucleotide sequence data. Mr. Yang's training has involved both coursework and laboratory experiments in molecular systematics, genetics, statistics and evolutionary biology. During May 1997, Mr. Yang traveled to Guatemala to collect and observe in nature Therevidae with PI Irwin, collaborator Webb, and graduate students from the Irwin and Yeates labs. Yang also traveled to meetings of the North American Dipterists Society and the Entomological Society of America. He prepared poster presentations for the NCSU Entomology departmental retreat and the NSF PEET meeting in Woods Hole, MA. A graduate committee at NCSU consisting of 4 faculty members (3 Entomology, 1 Genetics) has met and approved an official "Plan of Work" and Ph.D. thesis proposal entitled: Higher-level Molecular Systematics of the Family Therevidae (Diptera).
UIUC graduate student, Stephen Gaimari, visited PI Wiegmann's lab for 4 weeks in March 1997. Gaimari completed a laboratory rotation project involving amplification and sequencing of the mitochondrial cyctochrome oxidase genes of the genus Ozodiceromyia. Gaimari was trained in PCR techniques, agarose gel fractionation, PCR purification, automated DNA sequencing, and DNA sequence analysis using STADEN package software and GDE 2.2. Nucleotide data was collected for nine Ozodiceromyia species and several outgroup species.
Mr. Shaun Winterton began his studies by monographing a new genus [Australian genus A] with 20 species (Winterton, Irwin, & Yeates, in press), for which he was conferred his Post-Graduate Diploma in Science (Entomology) in July 1997. Shaun started his Ph.D. in August 1997, and has begun monographing the large and complex genus Agapophytus from that same region.
Honours student Ms. Narelle Power has begun a study under the supervision of Co-PI David Yeates at the University of Queensland. Ms. Power will document the spatial and temporal distributions of therevids in three different habitats in south east Queensland. Habitats include coastal heath, open forest and montane rainforest. Malaise trap samples have been collected since August 1997, and this project will continue until August 1998.
Establishing the higher groupings. The PEET proposal stated that a higher level phylogeny for the family Therevidae was needed so that small, monophyletic groupings of taxa could be monographed. The strategy for establishing a higher level classification of the Therevidae entailed two aspects. The first was a strong cladistic analysis based on morphological evidence, which would develop hypotheses of monophyletic units. This was to be followed by a molecular cladistic study to test those hypotheses. We felt that this would provide the most rigorous test of monophyletic units in the Therevidae.
Morphological progress. PI Irwin continues to work with Co-PI Yeates on developing a morphologically based higher classification of the Therevidae. Characters on the female terminalia and head have already been found and their states defined and polarized. Irwin is currently searching for characters on the thorax, wing, legs, abdomen, and male terminalia. These characters are being polarized and scored for all 50 plus genera and five outgroups currently under study. Preliminary trees based on parsimony and maximum-likelihood approaches will be generated using Hennig 86 and PAUP and examined using MacClade. The shortest resulting cladogram(s) will form hypotheses to be tested by molecular studies.
Mr. Gaimari and Dr. Irwin have delimited a new tribe of Therevinae, with a New World-Asian distribution. This tribe includes Ozodiceromyia, Cyclotelus, Breviperna, Procyclotelus, Ammothereva, and 5 new genera. All genera have been fully reviewed and diagnosed, and new synonymies and combinations have been established. Additional morphological information has been provided for all genera, and are included in a key. A monograph on the tribe is ready for submission for publication.
Molecular progress. Brian Wiegmann and Longlong Yang have completed amplification and sequencing of 1.3 kb of the 28S rRNA and 1 kb of Elongation Factor- 1a genes in 40 species representing 30 genera of Therevidae and outgroups. These data were obtained by direct PCR for 28S and RT-PCR for EF1a, followed by automated DNA sequencing on an ABI 377 autosequencer. Sequence electropherograms were edited and base calls checked by inspection in the program GAP4 of the STADEN package. Sequences were aligned in GDE 2.2. Phylogenetic analyses were carried out in PAUP* 4 test versions (D. Swofford pers. comm.). The following questions are being addressed in phylogenetic analyses of these data:
Does the 28S ribosomal RNA gene contain enough phylogenetically informative variation to reliably reconstruct higher-level relationships in the Therevidae?
Is the family Therevidae a monophyletic group?
What is the sister-group of Therevidae? Do molecular data support morphological evidence implicating the Scenopinidae as the closest relative of the Therevidae?
What is the phylogenetic status of the genera Apsilocephala andClesthentia?
What are the major groupings of Therevidae below the family level? Do the molecular data support the current morphological classification?
Current results of molecular phylogenetic analyses: High bootstrap values (>60%) support the monophyly of the Therevidae (excluding Apsilocephala) and monophyly of Therevinae. The monophyly Phycinae + Ruepellia-group is supported by parsimony analysis, but bootstrap values for this clade are low. Our results differ from the preliminary 5-subfamily classification except that the Therevidae is divided into two major groups, Phycinae and Therevinae that may represent two reasonable subfamilies. The controversial genus,Apsilocephala, is placed outside the Therevidae within the Scenopinidae.
Within subfamilies, the monophyly of Therevinae is highly supported by higher bootstrap value (>90%). However, the relationships amongAgapophytus-group genera are not well resolved by the data from 28S rRNA gene. The Agapophytus-group is paraphyletic on our trees and branch lengths for individual clades within it are short, indicating that 28S rRNA gene alone may not provide sufficient phylogenetic information to recover relationships below the subfamily level. Additional molecular data will be required to gain strong support for relationships at lower level in the family and within subfamilies and to place controversial genera such as Apsilocephala.
Elongation factor -1alpha (EF-1a) has been sequenced for most of these same taxa. This gene shows higher variability within therevid subgroups, suggesting that it is a suitable candidate for lower level phylogenetic analysis within the Therevidae. Phylogenetic analysis of 908 aligned bp of EF-1a for 17 taxa is concordant with results found for 28S rDNA. Like 28S, EF-1a supports the monophyly of the Therevidae excluding Apsilocephala, the monophyly of the Phycinae + Ruepellia-group clade, and the Therevinae.
Nucleotide sequence data have also been collected for 28S rDNA and phosphoenolpyruvate carboxykinase to examine the position of the Therevidae within the Asiloidea and resolve relationships between lower brachyceran subgroups. Sequencing of 28S rDNA for 85 brachyceran taxa has been completed.
A review of higher-level phylogenetic evidence and research on Diptera was completed for the serial Annual Review of Entomology by PI's Yeates and Wiegmann. This paper summarizes recent support for dipteran phylogeny from morphological and molecular studies covering the past 20-25 years. The manuscript was completed while PI Yeates was on sabbatical to the laboratory of PI Wiegmann at NCSU.
Status of Agapophytus. This large speciose genus is being monographed by Mr. Shaun Winterton. Original descriptions and revisions of the 13 valid species have been compiled. Type material for most of these species has been borrowed, and arrangements are being made to visit collections holding types unavailable for loan. Most specimens required for this study are on loan with a large collection of approximately 70 to 100 undescribed species accumulated and sorted awaiting description. All specimens for study have been entered into the database by the Irwin lab and labeled with loan information and unique specimen numbers. Most of the specimens required for molecular studies have been collected and are presently stored at the University of Queensland.
Status of Anabarhynchus. The monograph, by Dr. Lyneborg, PI Irwin, and Co-PI Yeates, for this speciose Australian genus is nearly finished. The three met in Copenhagen in 1996 to work on the manuscript. Species descriptions have been completed and PMTs of over 100 illustrations by Jill Mullett were sent to Denmark in Fall 1997.
Status of Brachylinga. Mr. Metz has gathered specimens for a monographic treatment of the genus Brachylinga from the world collections. Mr. Metz is now sorting specimens to species from his previously delimited sorting groups mainly using dissected male genitalia. Four holotypes ofBrachylinga species are currently on hand. Most remaining holotypes will have to be examined in European museums.
Status of Cylotelus. This genus is being treated by Mr. Gaimari and Dr. Irwin, as part of ongoing studies of the newly erected tribe, Cyclotelini including Ozodiceromyia. All of the specimens of Cyclotelus on hand have been entered into the database system, and preliminary sorting has yielded 26 species in the South and Central American fauna, and at least 3 species endemic to Greater Antillean islands. The North American fauna has yet to be sorted fully, but numerous species have been recognized, and working concepts for species-groups are being refined.
Status of Lindneria. Starting September 1997, Mr. Metz and Dr. Irwin initiated a treatment of the small Neotropical genus Lindneria. Represented by few specimens in collections, this genus is currently monotypic, but will include an additional species moved from the genus Psilocephala. Both holotype specimens for the monographic treatment are on loan. Illustrations of included taxa are underway. Recent study of a fossil specimen from Dominican amber of Oligo-Miocene age has suggested that it is not contained within the Lindneria genus concept, but may be the sister genus. The fossil specimen will be described as the type of a new genus.
Status of Ozodiceromyia. This large genus is under study by Mr. Gaimari. Approximately 30,000 specimens have been sorted into 112 species. All specimens have been input into the MANDALA database system. After studying all the types of the known species, names have been associated with their proper species, and synonymies and new combinations have been established. Of the 36 names from Irwin and Lyneborg (1981), 27 are valid, including new combinations. The known species have been fully analyzed using parsimony to provide a phylogenetic framework for the remaining species. The additional 85 species are currently being described and placed within this phylogenetic context. In addition, the genus is placed in a phylogenetic context within a new tribe including 10 genera. This new tribe has been fully treated phylogenetically, and biogeographical analyses have yielded an interesting account of their New World and Asian distribution.
Status of Pandivirilia. This genus contains 17 species from the Nearctic and Palaearctic regions that had previously been placed in the generaDichoglena, Pandavirilia, and Viriliricta. Three species are new to science. The species were revised by Don Webb and Mike Irwin, a phylogeny is hypothesized, and keys to the species are provided. This revision is currently undergoing in-house review.
Status of Tabuda and Tabudamima with the description of a new genus. The genus Tabuda, which contains two Nearctic species and four Palaearctic species (two of which are new to science), has been revised. The species Tabuda borealis was determined to be phylogenetically distinct fromTabuda and has been placed in a new genus. The genus Tabudamima, which contains one Nearctic species was included in this revision for the purpose of generic comparison. The species were revised by Don Webb (collaborator) and Mike Irwin, a phylogeny is hypothesized, and keys to the genera and species are provided. This revision is currently undergoing in-house review.
Status of Thereva and Acrosathe. These two genera, both found in the Nearctic and Palearctic regions, are being revised by Mr. Holston. Therevaincludes over 200 currently valid species, and over 200 additional species names are relevant to the nomenclature of this genus. By comparison,Acrosathe includes only 14 valid species names with nine additional species names relevant to the nomenclature of Acrosathe. With the guidance of F. C. Thompson, Mr. Holston has compiled these species names into a systematic database, which will be completed by Fall 1998. Name records from this database are being incorporated into the therevid database, MANDALA, names.fp3 file. Concurrently, Mr. Holston will compose a historical review of the family Therevidae with an emphasis on the type genus, Thereva. This nomenclatural work will constitute the majority of his Master's thesis, which will be completed by Fall 1998. Mr. Holston has examined the available specimens of Nearctic Thereva from museum collections and has begun morphological study on Thereva material from other biotic regions, received primarily from European museums.
Status of [new Australian genus A]. This genus of endemic Australian therevids was monographed by Mr. Shaun Winterton and is presently in press with Invertebrate Taxonomy (CSIRO publications). Of the twenty species in this genus 19 are new to science. The genus has been placed in theTaenogera-Group of genera.
Relationships of male & female genitalia. Mr. Metz continues to analyze the relationship of male and female Ozodiceromyia nanella (Cole) genitalia. He has two years of data on a population from a site in New Mexico and has written two small proposals to continue this work in 1998.
Parasitic mites on Diptera. Mr. Metz has also initiated the study from the same site in New Mexico of the association of the families of Diptera with parasitic mites.
Long-term Malaise trapping survey. Honours student, Narelle Power is conducting a long-term malaise trapping survey near Brisbane, Australia using 6 m Focks Malaise traps. Three traps are stationed in each of three habitats: coastal dunes (alt 5m), sclerophyll forest (200 m) and upland rainforest (ca. 600 m). They have been monitored since the beginning of August 1997. As of 20 March 1998, 1100 specimens of 47 species of therevids have been collected in 17 genera
Genera are as follows:Agapophytus .......................(4 species) Bonjeania .........................(1 species) Squamopygia (Pipinnipons)..........(2 species) Acraspisa..........................(1 species) Anabarhynchus.............(at least 7 species) Johnmannia.........................(1 species) Taenogera..........................(2 species) Ectinorhynchus............(approx. 12 species) Acatopygia.........................(2 species) Acupalpa ...........................(4 species) Australian genus A.................(3 species) Australian genus B.................(2 species) Australian genus C.................(1 species) Australian genus D.................(1 species) Australian genus E.................(1 species) Undescribed genus..................(1 species) Parapsilocephala...................(2 species)
Standardized terminology. As part of his Ph.D., and in collaboration with Mr. Stephen Gaimari and Co-PI's David Yeates and Michael Irwin, Mr. Shaun Winterton has proposed a standardized terminology for body vestiture and male genitalia. The former developed using scanning electron microscopy facilities at the University of Queensland. This terminology will be adopted in all monographs produced by the therevid group.
Homology of musculature and sclerotized portions of male genitalia. Co-PI Yeates has collaborated with Dr. Olga Ovtchinichova (Zoological Institute, St. Petersburg) in a study of the homology of musculature and sclerotized portions of the male genitalia of Therevidae. This has resulted in a manuscript in press in The Australian Journal of Entomology.
Morphology & histology of a novel structure in the female therevid reproductive system. In collaboration with Dr. David Merritt and Anthony O'Toole (University of Queensland), Mr. Shaun Winterton, Co-PI David Yeates and PI Michael Irwin have made detailed studies into the morphology and histology of a novel structure in the female therevid reproductive system. The studies include histological sections of virgin and mated females to determine the possible function of this structure.
Photographic library of live therevids. In collaboration with Mr. Anthony O'Toole (University of Queensland), Mr. Shaun Winterton and Co-PI David Yeates have begun to accumulate a photographic library of live therevids collected from the field. One of these photographs (of an undescribed genus and species) recently won first prize in the photographic competition at the 1997 annual general meeting of the Australian Entomological Society in September.
Yeates, D. K., and M. E. Irwin. 1996. Cladistic reappraisal and biogeography of the Apioceridae (Diptera: Asiloidea). Zoological Journal of the Linnaean Society. 116:247-301.
Irwin, M. E. 1997. Therevidae. In: Solís, A. (ed.), Las Familias de insectos de Costa Rica. INBio. <http://www.inbio.ac.cr/papers/insectoscr/Texto438.html>
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) <http://www.inhs.uiuc.edu/chf/pub/surveyreports/sep-oct97/peet.html>
1D Expeditions to Increase the Knowledge Base of Therevidae
During our PEET year-one meeting, we determined the criteria our expeditions should meet in order to be most productive. First and foremost was the need to gather specimens of critical taxa for both morphological and molecular studies. Second was the need to increase the holdings of specimens from areas lacking representation in the world's collections. The third criterion was to explore areas of expected radiation but that have not been well collected.
Eleven expeditions and one trip to European museums were supported by Schlinger Foundation funds since the last PEET report in August 1996: southern Africa; Guatemala; Chile and Argentina; southwestern U.S.; Texas; Utah; California; Europe and Russia; Australia; Baja California, Mexico; Madagascar. Further details on many of the expeditions can be found on the expeditions page.
Europe and Russia [Sept.Oct. 1996]. In September and October of 1996, Steve Gaimari visited several important insect collections in Europe to study type materials of Therevidae. These collections included the Zoological Institute of the Russian Academy of Sciences in St. Petersburg, the Naturhistorisches Museum Wien, in Wien, Austria, the Museum National d'Histoire Naturelle in Paris, France, and the Natural History Museum in London, England. Vadim Zaitzev was very kind in providing our project with a synoptic collection (including paratypes) of nearly all determined Therevidae in the Zoological Institute's collection (about 75 species), with very important material from Asia, Mongolia, and the Russian Far East. Also very helpful in locating materials for study were Ruth Contreras-Lichtenberg in Wien, Loïc Matile in Paris, and John Chainey in London.
Australia. Coolola National Park, Queensland, Australia [October 1996]. In early October, Co-PI Yeates led a one week field expedition to Coolola National Park, a large sand and heath area near Frazer Island, Queensland. The area has many undescribed and, undoubtedly, unknown species of therevids. Accompanying Yeates was Greg Daniels, curator of the insect collection at the University of Queensland, and graduate students Shaun Winterton, Chris Lambkin, and others from the University of Queensland. Asiloidea was the target group of organisms, and this provided Mr. Winterton with an excellent initial opportunity to observe and collect therevids in an area where they abound.
During September 1997, Mr. Winterton, with other postgraduate students from the University of Queensland Entomology department, traveled on a one week trip to far western Queensland to collect near the town ofBirdsville. Several new species of desert Therevidae were collected.
The following month, Mr. Winterton, with other postgraduate students from the University of Queensland Entomology department, traveled for one month through Victoria, South Australia and New South Wales. Collections were made in Wyperfeld, Flinders Ranges, Dutchmans Stern, Warrumbungle and Gammon Ranges National Parks. Approximately 40 species from close to 25 genera were collected during the trip, with greatest diversity in Gammon Ranges and Warrumbungle National Parks.
Southern Africa [mid Octoberlate November 1996]. A large-scale, six-week expedition was carried out in late 1996. The following specific localities were included: Kalahari Desert, Namib Desert, Namaqualand (western Cape Province of South Africa), and N'dumu (north eastern Natal Province of South Africa). This expedition involved rotations of all PIs, the Schlingers, Webb, and students during its various phases. The area is extremely rich in endemic taxa.
Texas [March 1997]. Collaborator Webb led a one week trip during UIUC spring break to southern Texas with graduate students Holston and Metz. Because the weather remained cold even in southern Texas, fewer taxa were collected than anticipated.
Guatemala [May 1997]. A two-week expedition was conducted in Guatemala by Irwin, Webb, the five graduate students, and their guide, Ms Maria Lucia Prinz. On the first full day in Guatemala City, they went to the insect collection at the Universidade del Valle de Guatemala, sorted their unsorted Diptera to family, and curated several families of flies. At Monterrico, the coastal dunes yielded many therevid specimens, including several Megalinga bolbocera (Osten-Sacken), numerous specimens ofBrachylinga, and Penniverpa, and a single female of "Psilocephala" breviventrisKröber. Megalinga was collected at the lights at night, and on the walls of the hotel in the morning, and occasionally in the vegetation. All other genera were collected perching on coastal vegetation, or were swept from grasses slightly farther in from the coast. They also put malaise traps at the very base of the Sierra de las Minas, in the very dry thorn tree zone at an elevation of about 250-400 meters. These traps proved to be very productive, collecting numerous specimens of Ataenogera abdominalis Kröber and Chromolepida pruinosa (Coquillett), as well as a species ofOzodiceromyia. Nearby they also visited a local hot springs, Quebrada de Agua Caliente, where numerous larval therevids were collected from the litter and sand under large trees. Many of these larvae have pupated and emerged, and belong in the genera Brachylinga and Penniverpa. Many malaise traps were also placed in the transition zone below the cloud forest in the Sierra de las Minas Biosphere Reserva at sites from 1600 to 2400 meters elevation. Very different insects were associated with each elevation. Therevids were abundant in a few of the traps, including many specimens ofOzodiceromyia schroederi (Kröber) and Ozodiceromyia obliquefasciata(Kröber). In the cloud forest itself, Irwin collected a singleton male of a new and very strange species of Phycus from low-lying vegetation.
Baja California Norte [June 1997]. The primary destination of Mike Irwin and Ev Schlinger was Parque Nacional de Sierra San Pedro Martir, a massive mountain range that divides Baja California longitudinally. June 9 was spent placing malaise traps at strategic waterholes at elevations above 2300 m. It was a memorable day because they discovered thousands of therevids at one of the higher waterholes, most belonging to one species,Ozodiceromyia nanella (Cole). That evening they met the expedition organizers, Dr. Ernesto Franco, Department of Crop and Soil Sciences, Michigan State University; Dr. Horacio de la Cueva, Centro de Investigacion Cientifica y de Educacion Superior de Ensenada (CICESE); and Dr. James Berry, state ecologist, Department of Parks and Recreation, Sacramento, CA, who were returning from a two-day hike. During this nine day trip, seventeen sites were sampled.
California. Following the expedition to Guatemala, Mr. Holston spent a week observing the coastal dune therevids of Oso Flaco Lake, Guadalupe-Nipomo Dunes Preserve, California. Mr. Holston was accompanied and otherwise supported by PI Irwin, Dr. Evert Schlinger, and Dr. Ebbe Nielson during the first three days of the trip, after which he spent the rest of the time on his own at the dunes. He recorded behavioral observations of adult therevids and collected therevid larvae, pupae, and adults. Over 40 specimens determined as species of Acrosathe, Tabudamima, and Therevawere reared later in the Irwin lab. Detailed notes of the habitat in which these specimens were found suggest differences in therevid habitat utilization across areas of the dunes. Morphological study of the larvae and pupae has provided more data for upcoming taxonomic reviews and keys.
Southwestern U.S. In June/July 1997, Mr. Metz, returned to the southwest U.S. to collect data for a second year of morphometric analysis of the genitalia of Ozodiceromyia nanella (Cole). In connection with this trip, he also collected in the extreme southern borders of Arizona and New Mexico. Mr. Metz returned with over 1500 specimens in 10 genera of Therevidae. Some specimens from this trip may become holotypes of new Ozodiceromyiaspecies in Mr. Gaimari's upcoming monographic treatment of the genus. Mr. Metz' trip was supported in part from a $1000 (Canadian) from the North American Dipterists Society.
Utah [21-25 July 1997]. Irwin and Webb traveled to Logan County, UT, joining Dr. Frank Parker and Dr. Wilbur Hansen in a short collecting trip. The primary objective of this trip was to find larval Apsilocephala, adults of which had been found in earlier collections from the area. Unfortunately, they were unable to find any larvae of Apsilocephala, but did return with many adult therevids and a Thereva larva, which Mr. Holston successfully reared to adulthood.
Chile [27 Sept.-Oct. 9; 25 Oct.-21 Nov. 1997] A two-month expedition was carried out as planned for areas in this taxa-rich country of southern South America. All PIs, Thompson, Webb, and the Schlingers rotated in and out on two-week cycles during this expedition. The timing was very good for the northern portion of the trip, especially around La Serena and Santiago, but was too early to gather many specimens in the southern part of the country. There are many undescribed genera and species in Chile and this expedition was focused on obtaining specimens in 95% ethanol of many of these groups for molecular and morphological study. Our expectations were achieved, and we returned with a wealth of specimens preserved in several ways for future study. In addition, funds were found to provide support for Chileans to maintain several of the malaise traps during the rest of the summer. These were finally taken down in late April. Gerardo Barria, who was in charge of these traps, recently reported that he has over 5 kilograms of preserved specimens to send us from the samples. He plans to continue the trapping effort next summer (September 1998 to April 1999).
Argentina [9-25 Oct. 1997]. This expedition was sandwiched between two-week cycles during the Chile expedition. Dr. Frank Parker joined PI-Irwin, who were escorted and guided to several areas to the north of Mendoza by Dr. Sergio Roig of the Unidad de Zoología y Ecología Animal, Instituto Argentino de Investigaciones de las Zonas Aridas (IADIZA), Centro Regional de Investigaciones Cientificas y Tecnologicas (CRICYT). The two-week expedition proved very successful. Numerous species representing several species in a wide aray of genera were collected and many of the taxa were preserved in 95% ethanol. Many rare and previously unknown groups of taxa were taken. For instance, a species of Phycus was collected that had previously only been known from Mexico to Costa Rica in the New World. Its African roots are now much better explained by the population collected in northwestern Argentina.
Madagascar [8-25 April 1998]. PI Irwin and Dr. Schlinger accompanied members of the Entomology Department of the California Academy of Sciences on a three-week expedition to several areas in Madagascar. This is a difficult place for an expedition and required an initial trip to develop the appropriate links before a full-blown expedition can occur. We focused on Ranomafana National Park in the rainforest near the eastern coast of the country, but had time to sample in the coastal beach vegetation to the east of the park and westward to Tulear and the dry spiny forest area of Madagascar. Collecting was relatively poor because we were not there during the peak of the season. We hope to return to Madagascar in November of 1999 during the peak season. However, we did manage to gather about 16 species of therevids representing three or four genera and at least four species during this past trip. Three of these taxa are preserved in 95% ethanol for molecular and internal morphological studies.
Irwin and Yeates presented a paper at the North American Dipterists Society informal conference at the Entomological Society of America meeting in Louisville, Kentucky, in early December 1996. The paper was entitled, Spermathecae and associated tissues of the female terminalia hold promise for constructing a higher-level classification of the Therevidae.
Michael Irwin moderated the North American Dipterists Society meeting at the National meeting of the Entomological Society of America, Louisville, KY, December 1996. Brian Wiegmann organized and moderated the conference in 1997 in Nashville, TN.
Michael Irwin presented a Department of Entomology seminar at the University of California at Davis, intitled Dune revisited: Exploring the lives and times of sandworms in April 1997.
A Year-Two working group meeting of our PEET grant took place in Urbana, Illinois, from 21-23 September 1997. Present were Dr. and Mrs. Schlinger, PI Irwin, Co-PIs Wiegmann and Yeates, Collaborators Webb and Kampmeier, graduate students Gaimari, Metz, Holston, and Mullett (UIUC).
Gaimari presented "Phylogenetic and biogeographic history of the cycloteline Therevinae" at the NSF/PEET Therevidae working group annual meeting, Urbana, IL in September 1997.
Kampmeier presented a seminar demonstrating MANDALA to the systematics collections group at the Illinois Natural History Survey in December 1997.
Gaimari was invited to present "Using cladistic methodology for phylogeny estimation and zoological classification, using an example from the Diptera (Insecta)" as a Quantitative Psychology seminar, University of Illinois, Urbana, IL in November 1997.
All PIs, the five Diptera graduate students, Kampmeier, and Thompson attended the second NSF PEET conference in Woods Hole, MA in March 1998. Irwin moderated a session on monography with help from Thompson; Kampmeier demonstrated MANDALA at the database session; Holston was a member of the student panel on training; Winterton ("Monographs of the world Therevidae: Australia's endemic fauna"), Yang ("Higher level Molecular Phylogeny of the Stiletto Flies"), and Gaimari ("Systematics and biogeography of the cycloteline Therevinae (Diptera:Therevidae)") each submitted posters on their research; and the therevid PEET team put together a poster complete with sound bites that gave an overall view of our project .
Gaimari presented his Ph.D. seminar to the Department of Entomology at UIUC entitled "Phylogenetics and biogeography of a Laurasian group of Therevidae (Diptera)" on 30 March 1998.
To Be Done
Summary of work to be performed during the next year of support, if changed from the original proposal; an indication of any current problems or favorable or unusual developments; and any other significan information pertinent to the type of project supported by NSF or as specified by the terms and conditions of the grant.
Networking. Several people have expressed interest in MANDALA, adapting all or portions of it for their taxonomic needs. Pam Arnofsky, postdoc working with Ami Scheltema, will spend a few days with Gail Kampmeier in late May working on this. Others have requested clones (structure, no records) of the databases, which can be sent via email or on diskette. Ms. Kampmeier will also be demonstrating MANDALA at the Taxonomic Databases Working Group meeting in Reading, England, in Sept. 1998.
Databases. Future plans for MANDALA include:
making the lots.fp3 database internally relational in a manner similar to the metamorphosis of the former taxa db to names.fp3
link a geographic region database to specimen locations,
implement the tracking of determiners (in determnr.fp3) and illustrators (in illus.fp3) via peo_join.fp3 (this side enabled already)
Interactive Keys. Over the next two years, we will adopt LUCID for keys of therevid taxa and use it in demonstrations to students of all ages as a example of how people go about identifying the organism they have in front of them.
Communications. We would like to make CD-ROMs of MANDALA available for distribution on a periodic basis. This will either require inclusion of a runtime version of FileMaker Pro (purchase of a license is required) on the CD-ROM or current access to FileMaker Pro 4.0 by the end user.
We also will have some parts of MANDALA web-accessible within the next year. Updates to the therevid WWW pages were recently made.
University of Illinois. Mr. Metz and Mr. Holston will finish taking courses towards a Ph.D. from the Department of Entomology at the University of Illinois in the fall semester of 1998. Both Mr. Metz and Mr. Holston will spend the majority of the following spring semester in Wiegmann's lab learning molecular techniques using material collected in alcohol on numerous expeditions and attending a formal course in systematics taught by Wiegmann. PI Irwin, co-PI Yeates, Mr. Metz, Mr. Holston, and Mr. Gaimari will attend an informal workshop in May 1998 with Dr. Sabine Huhndorf at the Field Museum of Natural History in Chicago to learn computer image capturing and manipulation techniques. Ms. Jill Mullett hopes to attend the annual GNSI meeting in Iowa this summer and improve skills relating to computer-assisted illustration.
North Carolina State University. Mr. Yang hopes to take summer courses on biological technology at molecular and cellar level. He will also take Insect Ecology and Systematic Botany next year. His advisor, Brian Wiegmann, will teach him various methods of analyzing molecular data using different software. Mr. Yang will also increase his proficiency in the use of other pertinent software during the coming year.
University of Queensland. Mr. Winterton will continue his revision of the large and speciose genus Agapophytus, concentrating on the morphological characteristics of the species groups within the genus. In January 1999, he will spend several months in the Wiegmann lab (North Carolina) to work on the molecular systematics of Agapophytus and hone his skills in this important technology.
Morphological progress. Irwin's lab is continuing to gather together specimens, enter data into the databases, and make critical observations on therevid morphology. Meanwhile, specimens of critical taxa are being collected during expeditions and preserved in 100% ethanol for molecular studies.
While Co-PI David Yeates is on sabbatical leave for 2 months starting May 1, 1998 with Irwin, they will complete a comprehensive cladistic analysis of the Therevidae based on over 100 morphological characters. This analysis will help resolve the relationships between, and identity of, the therevid subfamilies, tribes, and genus-groups of the world.
A comprehensive treatment of the higher level classification of the Therevidae and its close relatives will be published based on external and internal morphology. This will set up the hypotheses to be tested by the molecular approach in Wiegmann's lab.
Molecular progress at North Carolina State University will continue towards building a stable phylogenetic classification of therevid genera. Dopa decarboxylase and PEPCK genes will be amplified for a subset of the larger therevid dataset. Manuscripts will be prepared for publication, including results of the following current project-funded studies:
Analyses of 28S rDNA and EF1a in Therevidae;
Higher level phylogeny of the lower Brachycera;
An improved time-scale for dipteran diversification;
Molecular phylogeny of the therevid genusOzodiceromyia.
Status of Brachylinga. The species-group concepts is being refined while Mr. Metz continues to sort specimens to species-group and to species using both external morphology and genitalic characters. A study of the holotypes will illuminate the current nomenclature of the genus and necessitate a trip to European collections that are reluctant to loan type material. Monophyly of the genus will be established before a descriptive treatment of the species is to be considered. A preliminary treatment of "Brachylinga group" genera may be necessary before the Brachylinga species are treated. A paper on the higher level groupings of Brachylinga will be submitted for publication.
Status of Lindneria. Specimens in and near Lindneria are being examined for character analysis. Monophyly of the genus will be tested and a number of closely related genera may require study and description. All taxa will continue to be illustrated for inclusion in the final monograph that we expect to have in press by this time next year.
Status of Ozodiceromyia. The large number of new species are currently being described and placed into a phylogenetic context. This work will likely be completed by the end of 1998. All specimens are sorted to species, and new species have been assigned manuscript names and are currently being described. Through parsimony analysis, the genus will have a fully phylogenetic classification.
Status of Cyclotelus. Specimens of this genus are currently being sorted to species, and names based on study of types will be applied to sorted species. New species will be described and old species will be redescribed and placed within a phylogenetic framework. This genus should be completed by this time next year and a draft monograph developed.
Status of Thereva and Acrosathe. Having studied specimens of Thereva from the Nearctic in detail, graduate student Holston will expand the scope of his morphological study to Thereva of other biotic regions. An attempt will be made to divide the genus into monophyletic groups based on morpohology.
- Gaimari, S. D. and M. B. Mostovski. A new genus for Psilocephala electrella Cockerell from burmite (Diptera: Asiloidea). Bulletin of the Natural History Museum (London). [special volume on burmite geology and paleontology], submitted.
- Gaimari, S. D. and M. E. Irwin. Phylogenetic revision of the cycloteline genus Ozodiceromyia Bigot (Diptera: Therevidae)
- Gaimari, S. D. and M. E. Irwin. Phylogeny, classification, and biogeography of the cycloteline Therevinae (Diptera: Therevidae).
- Irwin, M. E. and D. K. Yeates. Towards an internal classification of the family Therevidae.
- Winterton, S. L., M. E. Irwin, D. K. Yeates. Systematics of [Australian genus A] gen. nov. (Diptera: Therevidae) from Australia.Invertebrate Taxonomy, in press.
- Yeates, D. K. and O. Ovtchinichova. The male genital musculature of Therevidae and Scenopinidae (Diptera: Asiloidea): Structure, Homology, and Phylogenetic Implications. The Australian Journal of Entomology, in press
- Yeates D. K. and B. M. Wiegmann. Congruence and controversy: Towards a higher-level phylogeny of the Diptera. Annual Review of Entomology, submitted, in review.
After attending the International Congress of Dipterology in Oxford in September 1998, Mr. Winterton will visit various museums throughout Europe and the United Kingdom to examine type specimens.
Mr. Holston plans to spend one or two weeks with Dr. Leif Lyneborg (PEET collaborator) in October 1998 to benefit from Dr. Lyneborg's experience withThereva and Acrosathe, particularly with the Palearctic species. Following his stay at the lab of BMW in the spring of 1999, Mr. Holston plans to visit several European museums in Fall 1999 to complete his study of the Therevatype specimens and examine other museum specimens.
Caribbean Islands [Winter 1999]. Mr. Metz anticipates an expedition to the islands of the Caribbean to increase his knowledge of island biogeography and the major radiation of the genus Brachylinga. A month-long expedition would be necessary for a comprehensive representation of both lesser and greater Antillean island fauna. The few specimens of Therevidae collected from these islands indicate the need to mine this wealth of diversity.
Gobi Desert of China and Inner Mongolia [summer 1999?]. A month long expedition is planned for this under-collected xeric zone. Very few taxa have been taken from this area but many are anticipated. Graduate student Yang will begin organizing this expedition in the near future. Several PIs and others associated with the project will rotate in and out during this expedition.
Madagascar [November 1999]. The preliminary expedition in May 1998 established essential contacts and determined that the optimal timing for a future expedition to this island would be in November 1999. It is expected that several PIs and others associated with the project will rotate in and out during this expedition.
The next therevid PEET meeting is currently planned for mid-August 1998 in Santa Ynez, CA.
All graduate students, PIs, Thompson, Kampmeier, and Mullett (as accompanying person), are planning to attend the 4th International Diptera Congress in Oxford, England in September 1997. Each will present posters or papers on their research.
Kampmeier will attend the Taxonomic Databases Working Group meeting in Reading, England following IDC4 in Oxford. She will present MANDALA and exchange ideas with other participants. She will represent not only our therevid PEET group but the Illinois Natural History Survey at this meeting.
All of our PEET group attending the national meeting of the Entomological Society of America in Las Vegas, NV in November 1998, will attend the North American Dipterists Conference held jointly at that time. Steve Gaimari is organizer and moderator of this meeting.
Members of the therevid PEET team have contributed heavily to outreach efforts in entomology. The Entomology Graduate Student Association at UIUC helps sponsor events such as Insect Expo each fall and several grad students and collaborators were involved in creating and running various activities. Mr. Metz displayed "Taxonomy for Tots", which takes kids through a visual dichotomous key, and Mr. Gaimari helped run this booth; Ms. Kampmeier ran the "I am an Insect/I am not an Insect" booth; Ms. Mullett worked with rubber stamps of insects in ecological settings; Dr. Webb set up and ran a forensic entomology booth. Many of these displays were also used at the Insect Expo in Louisville, KY at the national Entomological Society of America meetings in December 1996, in which we all participated. Mr. Holston, Mr. Metz, and Ms. Mullett also participated in a mini-Insect Expo in Danville, IL in April 1998.
Webb has lectured on forensic entomology to Dr. May Berenbaum's honors biology and Insects and People classes.
Kampmeier has also been active in programs to get and maintain young women in the sciences and mathematics, using entomology as a model. She and Ms. Mullett participated in a program in Springfield, IL on Feb. 28, 1998 "Expanding your horizons in Science & Mathematics" and in a career day for all students at the Urbana Middle School on April 9, 1998. Ms. Kampmeier also participated in the GEMS program (Girls in Engineering, Math, and Sciences) on April 15th, which also targeted middle school aged girls.
Many of the UIUC team will be involved in the CYCLE program on May 2, 1998 with an inner city youth group from Chicago. The presentation will include explanations of "What is taxonomy" and "What do taxonomists do and why is it important?" and allow the youth to use the computerized key LucID to key out various insects to order.
Steve Gaimari will be teaching "Classification and Evolutionary History of Insects" (ENT 302) in Fall 1998 at UIUC as a Visiting Lecturer.