ENTOMOLOGY-University of Nebraska State Museum-Research

EXECUTIVE SUMMARY
As a result of NSF-PEET support, Team Scarab has built an excellent foundation for training students in systematics, disseminating systematics data electronically, and monographing poorly known pleurostict scarab beetles. These results provide a foundation for continued research and training.

During the past five years, eight students received training in systematics at the MS, PhD, and undergraduate levels. Training included four female students (Aura Paucar, Katie Swoboda, Karla Villatoro, and Stephanie Westergren) and three Latin American students (Federico Ocampo, Aura Paucar, Karla Villatoro), both under-represented groups in systematic entomology. Four graduate students and four undergraduates were trained in scarab systematics. One undergraduate student is now working towards a MS degree in geography with an emphasis in geographic information systems, and another is applying to graduate schools in biology. Two graduate students graduated with MS degrees and are now working as science technicians. One graduate student graduated with a PhD degree and is now a Post-Doc in scarab systematics. One graduate student is continuing his PhD program in our lab and will graduate in May 2003. PEET funding enabled the successful training of members of a new generation of scarab systematists who work as team members and have the capability of using new technologies to further the science of systematics.

A total of 66 peer-reviewed, scientific publications were produced by PEET students and PIs. Several additional publications are in press and in preparation. Among these were 56 papers published on scarab systematics and biology (including nomenclature, cultural impact, fossils, revisions, phylogeny, larvae, taxonomy, biogeography, ecology). The following scientific contributions are of note:

14 Mongraphs of genera
16 Syntheses of higher-level groups
18 Synopses of genera
56 New species described
2 New fossil species described
7 New genera described
4 Larval genera new to science described
7 Larval species new to science described

Published works involved the participation of 17 collaborators (nine of whom were international collaborators from Canada, Czech Republic, Guatemala, Honduras, Mexico, South Africa, and Spain). Results (catalogs, images, keys) for many of these projects are at Scarab Central. As a hallmark of our productivity, an entire issue of the “Coleopterists Bulletin” was devoted to monographs and research papers from our laboratory. PEET funding also contributed to the publication of scientific papers in: weevil systematics (3 publications), study of insect vectors of Chagas disease (4 publications), phylogeny of sea pens (1 publication), study of the endangered insects (2 publications), review of native prairie insects (1 publication), and review of fossil arthropods (1 publication).

In addition to scientific publications, PEET students and PIs presented research findings at national and international conferences (58 presentations) and learned bioinformatics methods resulting in 32 electronic publications. All PEET graduate students gained training in molecular systematics. Two students (Ocampo, Smith) were elected to serve society roles (Executive Council of the Coleopterists Society). PEET students received five awards and 11 scholarships/fellowships. Some of these honors came with cash awards that amounted to $30,850. These awards recognize strong professional achievement and growth. In addition, productivity in our laboratory helped to garner $44,139 in new, internal University funds that were used to further research on the project.

PEET funding facilitated the education of 61 students and faculty in multimedia workshops on biogeography, phylogeny, and larval taxonomy. These workshops involved the participation of 5 other PEET groups and 9 institutions outside the University of Nebraska. Team Scarab hosted a Scarab Larval Workshop with host Dr. Miguel Morón, a Phylogenetics Workshop with host Dr. David Maddison, and a workshop on Biogeographic Theory and Analysis with host Dr. Jorge Crisci.

Team Scarab personnel were instrumental in curation and identification of many scarab collections (Brazil-4, Canada-3, Czech Republic-2, Ecuador-1, El Salvador-3, England-1, France-2, Germany-4, Honduras-3, Netherlands-1, Nicaragua-2, USA-21, Venezuela-1). The U.S. National Museum collection of approximately 300,000 pleurostict scarab specimens was transferred to us for identification, curation, and databasing and is approximately 70% curated. Nine of 13 subfamilies are curated and databased, thus making this international resource once again available to a broad community of users.

Field research and collecting in Latin America were used to train students in obtaining permits, collecting methods, and international collaboration (see Research and Collection Trips). Team Scarab conducted field and collections research in Ecuador, Venezuela, Honduras, Nicaragua, and El Salvador. In addition, we conducted collections research in Brazil, Canada, France, Germany, Netherlands, Czech Republic, and England.

The project has successfully launched interactive, on-line keys to scarab genera
(Generic Guide to New World New World Scarab Beetles), a World Directory of Scarab Workers that enables collaboration, catalogs and checklists of scarabs (e.g., Checklist of the Scarabaeoidea of the Nearctic Realm and Catalog of the New World Melolonthinae), and on-line species-level and specimen-level databases of scarab collections (UNSM Entomology Databases). “Generic Profile” web pages (e.g., Epectinaspis Blanchard 1850 and Brachysternus Guerin-Meneville 1831) and “Species Profile” web pages (e.g., Epectinaspis opacicollis Bates 1888 and Trizogeniates foveicollis Ohaus 1922) add to the operability of the key to genera of scarabs. The PEET project also influenced the development of the on-line University of Nebraska Division of Entomology Type Database and the Nebraska Dragonfly Database (see http://129.93.13.109/default.htm).

PROJECT ACTIVITIES AND FINDINGS
Systematics research produced 14 monographs of genera, 16 syntheses of higher-level groups, and 18 synopses of genera. In these publications, Team Scarab described 56 species, 2 fossil species, and 7 genera new to science. Seven species of larvae and 4 genera of larvae were described for the first time. Descriptions of new adults, larvae, and fossils greatly expand our understanding of the biodiversity of this speciose group of insects (over 35,000 species of scarabs are known).

Phylogenetics studies that accompanied each of the monographs identified sister taxa and provided a better understanding of character evolution and biogeography. For example, phylogenetic analysis of the Eucraniini (Scarabaeinae) based on morphology and molecular data provided evidence that: 1) overturns highly contentious hypotheses of sister-group relationships; 2) allows for interpretation of food relocation behavior in this unusual group of dung beetles; 3) and creates a foundation for addressing the conservation of this endemic, flightless, Argentinian group. Similar findings that illucidate our understanding of classification, land management, biogeography, and behavior were discovered in each monograph.

Working in collaboration with David Hawks and John Heraty (University of California, Riverside), molecular phylogenetic data based on 350 exemplars of Scarabaeoidea have greatly increased our understanding of scarabaeoid relationships, classification, and biogeography. Based on these data, some major clades are obviously paraphyletic and polyphyletic, thus revolutionizing the way in which we view evolution and classification of scarabaeoids. Examples include paraphyly in the southern hemispheric pleurostict tribe Anoplognathini (Rutelinae), thus promoting a re-interpretation of biogeographic events; paraphyly between the pleurostict subfamilies Dynastinae and tribe Anomalini (subfamily Rutelinae), thus causing a re-interpretation of classification; and polyphyly in the basal scarabaeoid family Lucanidae, a topic that is targeted for systematics research in our PEET2 grant.

Our on-line “Generic Guide to New World Scarab Beetles” includes keys to genera that are used by many people engaged in agriculture, habitat surveys, conservation studies, biodiversity research, collection management, evolutionary studies, taxonomy, and comparative biology. Keys, catalogs, images, maps, and biology that are produced for monographic research are integral to this guide. We have received positive feedback from people who are using this resource for identification, nomenclature and classification, and distributional data for species/genera. Our on-line “Scarab Beetle Database” provides access to 30,578 specimen-level records from multiple institutions and 3,595 species-level records. It integrates data from all our scarab databases (including the database of Smithsonian Institution higher scarabs, databases created for biotic inventory research [DEB 9870202 and DEB9200760], and databases for monographic research), thus elevating the power of an online search.

TRAINING AND DEVELOPMENT
Eight students received training in systematics, including four female students (Aura Paucar, Katie Swoboda, Karla Villatoro, and Stephanie Westergren) and three Latin American students (Federico Ocampo, Aura Paucar, Karla Villatoro), both under-represented groups in systematic entomology. Four graduate students (2 MS and 2 PhD) and four undergraduates were trained in scarab systematics. One undergraduate student is now working towards a MS degree in geography with an emphasis in geographic information systems (Brett Ratcliffe), and another is applying to graduate schools in biology (Katie Swoboda). Due to her high quality thesis on scarab systematics, Swoboda graduated “With Distinction”. Two graduate students graduated with MS degrees (Aura Paucar and Karla Villatoro) and are now working as science technicians. Both continue to work on projects in the lab in their spare time, and one student (Aura Paucar) intends to work towards her PhD in the near future. One graduate student (Andrew Smith) graduated with a PhD degree and is now a Post-Doc in scarab systematics in our laboratory. One graduate student (Federico Ocampo) is continuing his PhD program in our lab and will graduate in May 2003.

Students were successful in publishing peer-reviewed papers and learning techniques for disseminating data electronically, illustration and imaging, databasing, molecular methods, research presentations, grant-writing, and presentations for the public (see Team Scarab PEET products). In recognition of high achievement, leadership, instruction, and community service, PEET students received five awards and 11 scholarships/fellowships. Examples include a one-year fellowship from UNL to Ocampo (the most prestigious award to a graduate student at UNL), the J.H. Comstock award from the Entomological Society of America to Smith (the highest graduate student award from the Society), the Ernst Mayr fund from Harvard to Villatoro, the Outstanding Research and Creativity Award to Ratcliffe (the most prestigious research award for faculty at UNL). Some of these honors came with cash awards that amounted to $30,850.

Three undergraduates (Mitch Wellman, Ian Ratcliffe, Katie Swoboda) were trained in curation, databasing, and scarab systematics as part of an REU. Two undergrads (Katie woboda, Stephanie Westergren) had independent research projects on scarab systematics as part of the UNL honor’s program. Swoboda earned a B.S. “With Distinction” for her high achievement and thesis on scarabs.
PEET funding facilitated the education of 61 students and faculty in multimedia workshops on biogeography, phylogeny, and larval taxonomy. These workshops involved the participation of 5 other PEET groups and 9 institutions outside the University of Nebraska.

PEET graduate and undergraduate students were actively involved in identification, curation, and databasing of the University of Nebraska scarab collection, the National Collection of Scarabs from the Smithsonian Institution, and numerous other institutions from countries previously listed. The U.S. National Museum collection of approximately 300,000 pleurostict scarabs specimens was transferred to us for identification, curation, and databasing and is approximately 70% curated. Nine of 13 subfamilies are curated and databased, thus making this international resource once again available to a broad community of users.

OUTREACH ACTIVITIES
PEET personnel are actively involved in educational outreach activities that range from informal presentations on insect diversity and scarabs to formal workshops on evolution, biogeography, and phylogeny. To increase their communications skills, some PEET students have volunteered to teach introductory courses (Federico Ocampo, Karla Villatoro, Aura Paucar). Due to the success of our “Biogeography Workshop”, Ocampo was invited to present a similar workshop at the New York Botanical Gardens (2002) and California Academy of Sciences (2002). Outreach activities also range from exhibits on scarab diversity and systematics research (e.g., “Treasures of the Museum” 2002-2003) to participation in “Bug Bash”, a public educational event on insects.

CONTRIBUTIONS WITHIN DISCIPLINE
Within the disciplines of entomology and systematics, products of this PEET research are far-reaching. Monographs and electronic data provide the foundation for understanding and interpreting the diversity of a speciose group. The on-line “Guide to New World Genera” provides a means of identifying the higher genera scarabs (approximately 320), and created the foundation for continuation of PEET research in our lab on all New World genera of scarabs (approximately 600). The Guide provides a means of identification and includes keys to taxa, overviews of taxa, generic profiles, species profiles, distribution maps, temporal distribution, and images. It is our hope that this Guide will be the foundation for an on-line guide to all world scarabaeoids as well as a guide to all world larvae. Databases that resulted from this research allow for sharing of data. Specimen-level databases enable the examination of past and present diversity. Results of on-going phylogenetic analyses provide a basis for a new, stable classification of scarabaeoids that reflect evolutionary patterns.

Student training invests in the future of systematics and provides critical mass in our lab. This new generation of scarab systematists is available to train additional systematists and coleopterists. The broad training that students received prepared them to be competitive in areas of traditional and molecular systematics with emphasis in entomology. This generation of systematists works effectively in a team and has the capability of using new tools that will advance the science of systematics to a new level. PEET students influence our disciplines because of their representation in nationally elected offices (Federico Ocampo and Andrew Smith are members of the Executive Board for the Coleopterists Society). The PEET PIs also influence disciplines: Brett Ratcliffe and Mary Liz Jameson are subject editors for the Annals of the Entomological Society of America; Brett Ratcliffe is Secretary of the Coleopterists Society; Mary Liz Jameson is president elect of the Coleopterists Society and Central States Entomological Society.

Our Scarab Workers Directory provides substantial information to the field and allows for collaboration and communication. The Scarabs-L listserve (initiated by Andrew Smith) provides a quick and easy means of communicating among the community (including amateur collectors, collection managers, agriculturalists, and ecologists).

Authoritatively identified and well-curated collections that have resulted from this research allow a broad community of users to use material for studies beyond entomology and systematics. The field of collections management directly uses products of our research in retrieving, organizing, databasing, and identifying scarab collections.

CONTRIBUTIONS TO OTHER DISCIPLINES
Results of PEET research influenced many disciplines outside of entomology and systematics. To name a few, the fields of biotechnology and geography will mine data from specimen-level databases for tests of geographic information systems, examination of biodiversity patterns, and predicting patterns of diversity. The on-line “Guide to New World Genera” is useful in agriculture, habitat surveys, and conservation studies. Phylogenetic classifications furnish the scientific context for understanding the processes of ecology, behavior, and conservation patterns. Cultural anthropologists and paleontologists will use data generated from published papers. Smith and Paucar (2000) recorded use of scarabs by indigenous peoples in Ecuador as a food source. Publications by Ratcliffe (e.g., 2001) and Ocampo (e.g., 2002) provide data on scarab fossils in amber as well as insect trace fossils.

This research influenced the publication of scientific papers in weevil systematics (3 publications), Chagas disease (4 publications), phylogeny of sea pens (1 publication), endangered insects (2 publications), review of native prairie insects (1 publication), and review of fossil arthropods (1 publication).

CONTRIBUTIONS TO HUMAN RESOURCE DEVELOPMENT
Our PEET program provides opportunities for our students to conduct research and teaching. In so doing, they improve their performance and skills. Members of our team are from under-represented groups in systematics: Latin American (3 participants) and female (4 participants). All of our Team is developing and disseminating new educational materials (lab manuals, internet resources, seminars, teaching at university and K-12 levels, exhibits). The K-12 involvement provides exposure to science to young people and non-scientist members of the public. All published papers included Spanish or Portuguese abstracts and/or keys that enabled our audience to use our publications. The on-line “Guide to New World Genera” will include keys in English and Spanish.

CONTRIBUTIONS TO RESOURCES FOR RESEARCH AND EDUCATION
Databases, authoritatively identified collections, DNA collections, On-line Guides, publications are all useful to a broad audience of users (see “Contributions to other Disciplines” and “Contributions to Human Resource Development”). The results of our PEET program have leveraged additional lab space in order to conduct our activities.

CONTRIBUTIONS BEYOND SCIENCE AND ENGINEERING
Scientific information that is mined from our databases, presentations, publications, and on-line data will be used by resource managers to help monitor areas and to justify conservation measures for areas. Education (formal and informal) on systematics and biodiversity will positively impact students and the general public by providing the data necessary to make wise decisions for the future of the planet. Products of this research will positively impact the field of bioinformatics, making additional data available with which to test diversity models.
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