TABLE OF CONTENTS

Project Summary

1. INTRODUCTION

2. TAXONOMIC FOCUS FOR MONOGRAPHY

2.1 Tanyderidae

2.2 Rhagionidae

2.3. Aulacigastridae

3. METHODS OF STUDY

3.1 Museum Research Protocols

3.3 Phylogenetic Analyses, Biogeography and Biodiversity Studies

4. TRAINING

4.1 Site Affiliations

4.2 Post-doctoral Fellow

4.3 Graduate Students

4.4 Undergraduate Interns

5. EXPECTED SIGNIFICANCE

5.1 Taxonomy and Monography

5.2 Phylogeny

5.3 Biogeography

6. DISSEMINATION OF RESULTS

7. PLAN OF RESEARCH

BIBIOGRAPHY

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Project Summary

This proposal has three major objectives:

• 1. Production of high quality, modern systematic monographs of 3 understudied taxa of flies (Diptera)
  
• 2. Training 3 Diptera systematists in comprehensive monographic methodology and phylogenetic systematics
  
• 3. Utilizing available technology to create state of the art systematic monographs disseminatedelectronically
  

This proposal requests funding for a five-year term to undertake monographic research on three families of flies (Insecta: Diptera): Tanyderidae, Rhagionidae and Aulacigastridae. Projects have been selected that can be accomplished in a timely manner, as the work will be carried out primarily by PhD students for thesis research and a postdoctoral fellow, under the mentorship of established Diptera researchers at the National Museum of Natural History, Smithsonian Institution. Diverse taxa within the Diptera were chosen for monographic training, rather than training several workers in a single family, so students entering the job market are not researching closely related taxa. However, once students are mentored in comprehensive monography, they should be able to tackle any group within the Diptera in the future. The Smithsonian's collections, libraries, electronic media facilities for taxonomy and ties with two strong local universities with excellent systematics programs are unmatched at any other single working site for the education of highly qualified monographers in Diptera. Monographs of the entire family Tanyderidae, the genera of Rhagionidae and species of the genus Aulacigaster will all contribute significantly to our knowledge of flies. Monographs will be written to include strictly parallel descriptions of taxa, complete treatment of appropriate nomenclature, identification keys, illustrations of critical morphological features, incorporation of distributional information into specimen databases, biogeographic considerations and a complete discussion of known biology. Phylogenetic hypotheses of relationships of these critical, understudied taxa as well as placement of each monographed taxon within the Diptera will be addressed. Electronic products such as specimen and catalog data bases, interactive keys with electronic images, character matrix databases and bibliographies that can be disseminated via the Internet, CD-ROM and other electronic means will be emphasized. Taxa were chosen for monography not only from their intrinsic interest, but also because there are no specialists in these taxa or none using modern approaches, and because advancing our knowledge of these groups will contribute significantly to problems in the phylogeny of Diptera at higher levels.

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1. INTRODUCTION

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The Diptera (flies) contains by far the most important vectors of arthropod borne human diseases (mosquitoes, family Culicidae) and agricultural pests of high significance (e.g. the Med fly, family Tephritidae). Biological control programs have successfully utilized the services of tachinid flies to reduce pest populations in critical agronomic systems (e.g. Woodley 1994 for a summary of one such case). Modern knowledge of genetics is based largely on the paradigm system of Drosophila melanogaster (family Drosophilidae) and another fly, Calliphora vicina (family Calliphoridae) has been extensively used in studies of invertebrate physiology (e.g. Dethier 1976). Willi Hennig was a practicing dipterist and utilized the revolutionary theoretical methodology he developed in his empirical studies of extant and fossil Diptera (e.g. Hennig 1958, 1965). The Diptera are also the exemplary organisms in classic studies of biogeography (Hennig 1960; Brundin 1966) and speciation (Bush 1975; Barton & Charlesworth 1984; Carson & Templeton 1984). Clearly, the Diptera are a highly important group of organisms on many fronts, and fully deserve a steadily increasing knowledge of their systematics, the foundation on which all other biological research rests. The Diptera are a vast order of insects, currently containing about 100,000 described species found from the high Arctic and Antarctic to equatorial regions, which have invaded almost every conceivable habitat (Oldroyd 1964). While a few families are fairly well known, such as the Culicidae because of their medical (and thus economic) importance, taxonomic knowledge for most families ranges from fair to rudimentary. For example, the Phoridae contain about 3,000 described species, but estimates as high as 20,000-50,000 for the family have been predicted (Disney 1983) which translates to 6-15% of the fauna being known. Other families, such as the Stratiomyidae, have received considerable attention of only one or two specialists in this century. The literature of such families gives one the impression that they are fairly well known. However, intensive sampling schemes using Malaise traps and other methods show that they are far more speciose than previously thought (Woodley, pers. com.). The statistics for Diptera are in line with estimates of our knowledge of insects in general, which are believed to be between 2-10% described (Wheeler 1990). In North America we have an excellent assessment of our knowledge of Diptera, and the data clearly indicate that the Diptera are the least known of all North American insects (Kosztarab & Schaefer 1990). Despite the poor knowledge of Diptera,"the future for human resources in Diptera is poor. We have lost by retirement those key teachers, such as Alexander of Massachusetts, Berg of Cornell, Byers of Kansas....who had trained this generation of dipterists. Only Steve Marshall of Guelph, Monty Wood of the Biosystematics Research Centre in Ottawa, and the Maryland Center for Systematic Entomology (MCSE) utilizing the dipterists in Washington, have active programs for training dipterists"(Thompson 1990: 191). As of 1994, we can now add Monty Wood and George Ball to the list of retirements without replacement. Indeed, unless there is some infusion of resources and commitment soon, extinction rates of Diptera systematists will exceed those of the taxa they work on.

This proposal is aimed specifically at training new, young dipterists to continue the efforts of past generations. All of the resources are currently in place for this effort at the Smithsonian Institution (SI). We feel we can train highly competent, modern dipterists that can in turn stimulate and educate the next generation of fly systematists.

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2. TAXONOMIC FOCUS FOR MONOGRAPHY

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For each of three critical, understudied taxa of Diptera we plan to provide modern, state of the art systematic monographs. These will include 1) diagnosing, describing, illustrating, and cataloging the world fauna for each taxon,
2) providing identification aids,
3) determining the phylogenetic placement of each taxon in the Diptera and the cladistic relationships within each, and
4) examining biogeographic patterns of each taxon in a phylogenetic framework.

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2.1 Tanyderidae

2.1.1 Specific Objectives for Monograph of Tanyderidae

• 1. Produce comprehensive monograph of the entire family
• 2. Establish monophyly of the family
• 3. Provide data for phylogenetic placement of the Tanyderidae within Nematocera (Diptera)

2.1.2 Past and Present Classification

The tanyderids have long been viewed as a distinct, basal group within the order Diptera (Alexander 1932; Exner & Craig 1976). Osten Sacken (1859) originally recognized the group as a tribe in the family Tipulidae. Revising the familial boundaries of Nematocera, Handlirsch (1909) removed tanyderids from tipulids and placed them in the Ptychopteridae.

For the seventy years since being accorded family rank (Alexander 1919) speculation has surrounded the group's phylogenetic position in the Nematocera. In part, confusion over phylogenetic placement is due to the presence in Tanyderidae of a large number of characters thought to be pleisomorphic in Diptera. In addition, characters from different life stages give incongruent results regarding sister group relationships. For example, Crampton's (1925, 1926) rigorous comparative studies on morphology of the adult thorax contributed significantly to tanyderids being aligned with the Psychodidae and Ptycopteridae. After thoroughly examining the larva of Protoplasa (Diptera: Tanyderidae), however, Crampton (1930b) concluded that no morphological evidence was available to support a close relationship between ptychopterids, tanyderids and psychodids. In Crampton's (1930b) opinion, tanyderid larvae shared far more similarities with Chironomidae. The apparent life stage incongruence observed by Crampton (1930b) led him to reject the evidence provided by larval characters. Alexander (1930b) discussed this ambiguity and observed that each of the known tanyderid life stages possesses characters that suggest a relationship with a separate lineage of primitive flies: larvae - Chironomidae, pupae - Tipulidae, and adults - Psychodidae/Ptychopteridae or Tipulidae. Given the discrepancy among characters, Alexander (1930b) chose to leave the phylogenetic placement of the tanyderids unresolved.

The tanyderids were once again united with tipulids by McAlpine et al. (1981) based on wing venation and condition of the adult pronotum. Under this arrangement, the Tipulomorpha was considered the basal lineage of the order Diptera. Hennig (1981), however, considered the condition of the pronotum and wing venation to be pleisomorphic states for Nematocera. Hennig's classification recognized the clade Ptychopteroidea, composed of tanyderids and ptychopterids, in the superfamily Psychodomorpha. The Ptychopteroidea was supported by one synapomorphy, the ability of males to extend the fifth tarsomere forward, interlocking with specialized setae on the fourth tarsomere. Recent studies by Wood & Borkent (1989) support this arrangement. Their results differed from Hennig (1981) by elevating the superfamily (Ptychopteroidea) to the rank of infraorder (Ptychopteromorpha), and more importantly, removing the tanyderid clade from the Psychodomorpha. This current arrangement places the clade tanyderids + ptycopterids as the sole sister group to the most important pathogen vectors in the Insecta, the Culicomorpha (includes Culicidae, Simuliidae, Ceratopognidae, etc.). Other than the modified male tarsi required to grasp the female in mating, Wood & Borkent (1989) were unable to find additional synapomorphies to support the Ptycopteromorpha. Clearly uneasy with this modest support for the arrangement of Ptycopteromorpha, Wood & Borkent (1989) suggested the ptychopterids alone may be the sister group to the Culicomorpha. However, tanyderids share complex labral brushes with ptychopterids and the Culicomorpha that Wood & Borkent (1989) argue must be homologous across these taxa. The decision to treat ptycopterids as the sister group to the Culicomorpha was therefore deferred till additional support for an alternative arrangement could be found.

Relationships among the species of tanyderids are unknown. One publication (Alexander 1927) has attempted to address the fauna on a worldwide basis. Unfortunately, that generic-level monograph suffers a number of shortcomings. First, less than half of the currently known species were described at the time of publication. Second, the familial boundaries for the tanyderids were based on plesiomorphic characters. This resulted in the inclusion of a number of taxa now recognized as belonging to other nematoceran families, e.g. Bruchomyia (now in Psychodidae) and a larva now recognized as an Axymyiidae (Alexander 1927: Plate 1, Fig. 4). The true immature stages were unknown at that time. Other studies have been confined to regional reviews (Duxbury & Barraclough 1994; Alexander 1981, 1928; Colless & McAlpine 1970; Stuckenberg 1962; Peus 1958; Wood 1952) or species-level descriptions and addenda (Exner & Craig 1976; Hinton 1966; Knight 1963, 1964; Colless 1962; Williams 1933; Alexander 1930a, 1930b; Crampton 1930a).

2.1.3 Diversification and Distribution

The tanyderids are found worldwide, achieving their greatest species diversity in the Southern Hemisphere. Thirty-seven names are currently treated as valid in ten genera (Wagner 1992; Oosterbroek 1989; Hutson 1980, Alexander & Alexander 1973, 1967; Alexander 1965). The group is of particular interest owing to their essentially bipolar pattern of distribution (after Du Rietz 1940). Two of the genera are broadly distributed (Protanyderus, Holarctic and Northern Oriental; Radinoderus, Australian and Oceanian), while the other eight genera are regionally endemic. Six of these genera are monotypic. A world revision and species-level phylogenetic analysis will provide data useful for insights into the complex biogeographical patterns observed in the Pacific and Northern and Southern Hemispheres.

2.1.4 Biology

The tanyderids are poorly known and considered amongst the rarest of Diptera (Alexander et al. 1967). The genera with known larvae occur in two types of aquatic habitats. Larvae of Protoplasa (Alexander 1930b), Mischoderus (Johns, pers. com.), Protanyderus (Exner & Craig 1976, Rose 1963, Knight 1963) and Peringueyomyina (Wood 1952) are free-living and occur in the hyporheic zone of sand bottom streams. Species of Eutanyderus (Colless & McAlpine 1970, Hinton 1966) and the Australian Radinoderus (Cranston, pers. com.) occur in saturated wood along stream margins. Larvae migrate to drier, sandy areas prior to pupation (Knight 1964; Wood 1952; Alexander 1930b). While the adults are frequently reported to be rare, large numbers are collected occasionally in mating swarms or on riparian vegetation (Wood 1952; Alexander 1930b; Crampton 1929). This anomaly suggests some of the records more accurately reflect collecting bias or insufficient knowledge of the life history of the group. The latter was asserted by Exner & Craig (1976) regarding collection of the immature stages. Wood (1952) discusses his search for immatures in which initial collections were tedious and difficult, however, once the habitat was known a subsequent search at a new locality produced high larval numbers in a short period of time.

2.1.5 Available Material

The tanyderid collection at the National Museum of Natural History is unparalleled in coverage, quality and accessibility of material. The foundation of the collection is composed of specimens described by Alexander (Oosterbroek & Theowald 1980), including 21 holotypes and associated paratype series, from all faunal regions. An additional 9 described species are present making exemplars readily available from all genera. Most of the collection consists of pinned specimens, although wings and male terminalia are slide mounted for most species. This collection represents 81% of the currently described fauna. Specimens available in other collections will be borrowed, particularly from the Bishop Museum (Hawaii, USA), The Natural History Museum (Great Britain), CSIRO (Australia), Landcare Institute (New Zealand), MacLeay Museum (Australia), Zoological Institute, Academy of Sciences (Russia) and the Institute of Zoology, Ukrainian Academy of Sciences (Ukraine).

The weakness of the SI's collection is the absence of immature stages, reflecting their relatively recent discovery. Many of these immatures have been recovered in intensive aquatic surveys and are currently not described (Cranston, pers. com.; Johns, pers. com.). Judd will collaborate with other investigators and participate in field work at known sites to supplement data on the immature stages (see attached documentation).

In addition to the Alexander specimens, his personal correspondence, notebooks, and library are available at the National Museum of Natural History. We feel this material will be invaluable to the reexamination of the family.

2.2 Rhagionidae

2.2.1 Specific Objectives of Monograph of Rhagionidae

• 1. Produce comprehensive monograph of"rhagioniform"genera
• 2. Establish monophyly of the family(ies) of rhagioniform genera
• 3. Provide data for phylogenetic placement of the rhagioniform genera within the lower Brachycera and Tabanomorpha.

2.2.2 Present and Past Classification

The Rhagionidae have been recognized as a distinct, family-level group of Diptera since the early part of the 19th century (Latreille 1802), and the formal family name we use today dates from Samouelle (1819). The early concept of the family is based almost entirely on the genera Rhagio Fabricius and Chrysopilus Macquart.

Despite the long-term recognition of the Rhagionidae, the limits of the family have not been stable. A number of genera such as Bolbomyia Loew, Glutops Burgess, Pseudoerinna Shiraki, Vermileo Macquart, Dialysis Walker, and Atherix Meigen have been placed in the Rhagionidae (James 1965) until relatively recently. Hennig (1972) discussed the importance of rhagionids in phylogenetic studies of brachycerous Diptera. Woodley (1989) summarized cladistic argumentation for the relationships of the 4 families of Tabanomorpha, a well-corroborated infraorder to which the Rhagionidae belongs, and reviewed recent literature discussing placement of a number of genera. Stuckenberg (1973) convincingly showed that Atherix and a few other"rhagioniform"genera are really more closely related to Tabanidae than to the rhagionid residue, and placed them in the new family Athericidae. Nagatomi (1975) suggested Dialysis belonged in the Coenomyiidae [=Xylophagidae], strongly confirmed by discovery of its larva (Webb & Lisowski 1983). Nagatomi (1982) reviewed the genera of Rhagionidae and Pelecorhynchidae and proposed a"supposed phylogeny"in the form of a tree (Nagatomi 1982: fig. 1) but omitted any discussion of character support for its structure. Woodley (1989) placed Glutops and Pseudoerinna in the Pelecorhynchidae following Teskey (1970, 1981), but this scheme was not followed by Majer (1988) who left them in the Rhagionidae in the latest Palaearctic catalog.

Because the Rhagionidae lack a modern, cladistic definition (Hennig 1973; Woodley 1989), it remains a catchall taxon for flies exhibiting a suite of plesiomorphic character states. The rhagioniform genera may be paraphyletic or even polyphyletic with respect to other Tabanomorpha. Lack of recent, critical morphological study of the adult flies has resulted in poor definition of a number of critical genera. While there is convincing evidence that the Tabanomorpha is monophyletic, its relationships to other major clades of lower Brachycera are unresolved (Woodley 1989; Sinclair et al. 1994). This is due in large part to our poor knowledge of the Rhagionidae, the most plesiomorphic lineage of the Tabanomorpha. A thorough, comprehensive monograph of the rhagioniform genera is critical to further knowledge in this area.

2.2.3 Diversification and Distribution

The Rhagionidae are found worldwide. There are about 520 described species placed in 21 genera. Rhagio (about 270 spp.) and Chrysopilus (about 110 spp.) are by far the largest genera, followed by Atherimorpha White (38 spp.), Symphoromyia Frauenfeld (32 spp.), and Ptiolina Zetterstedt (22 spp.). The remaining genera each have fewer than 10 included species. Collecting in southern South America and Australia is likely to increase the number of species, but not overwhelmingly so, as suggested by Stuckenberg's (1956, 1960) South African field work. A number of monotypic genera, such as Alloleptis Nagatomi & Saigusa, Desmomyia Brunetti, Neorhagio Lindner, Solomomyia Nagatomi, and Stylospania Frey may simply be autapomorphic species that could be included in other genera if the genera were rigorously defined in a cladistic framework.

There are three general distribution patterns in the Rhagionidae. 1) Widespread genera found virtually worldwide (e.g. Chrysopilus, Rhagio), 2) north temperate genera (e.g. Athroceras Williston, Symphoromyia, Ptiolina), and 3) south temperate genera (e.g. Austroleptis Hardy, Atherimorpha, Spaniopsis White).

2.2.4 Biology

The biology of the Rhagionidae is very poorly known. Adults are commonly collected sweeping in moist, shady, often forested areas. Leonard (1930) and Paramonov (1962) stated that adults were predators, at least occasionally, on other insects, but the fragile morphology of most species suggests that other feeding habits predominate. Females of at least two genera, Spaniopsis (Paramonov 1962) and Symphoromyia (Turner 1974), feed on vertebrate blood. Males of Symphoromyia form conspecific mating swarms (Turner 1974), and sexual dimorphism, particularly in eye structure, suggests that other genera also have this habit.

Immature stages are poorly known. Larvae of those few taxa that are known have been collected in moist soil rich in organic matter (Turner 1974). Paramonov (1962) and James & Turner (1981) indicated that they are predators on a variety of invertebrates, but gave very little confirming evidence. Paramonov also inferred that some were saprophagous or rarely coprophagous. The wide dispersion of individual larvae in their habitat (Turner, pers. com.) does seem to indicate a predaceous habit.

2.2.5 Available material

The Smithsonian collection of Rhagionidae is excellent, with representatives of most genera and with good diversity of species in the larger genera. We anticipate borrowing further material, especially those genera that have been recently described from little material (e.g. Solomomyia Nagatomi) from the Bishop Museum, The Natural History Museum (London), Natal Museum (South Africa), CSIRO (Australia), and the Museo Nacional (Santiago, Chile). Further supplementation will be made by field work, especially in southern South America and Australia, to obtain fresh material. No rhagionid specialist has collected extensively in the south temperate regions except Stuckenberg in South Africa. Holotypes will be borrowed from a variety of institutions as necessary.

2.3. Aulacigastridae

• 1. Produce a comprehensive monograph of the species of Aulacigastridae
• 2. Determine the sister group of the family
• 3. Provide data for the phylogenetic placement of the Aulacigastridae within the Asteiioidea

2.3.2 Past & Present Classification.

The Aulacigastridae are a small, curious, discrete and uncommon group of acalyptrate Diptera that was first accorded familial status by Duda (1924). Until Hennig (1969, 1971) recharacterized Aulacigastridae, adding four more genera (Cyamops Melander, Planinasus Cresson, Stenomicra Coquillett, and Schizochroa Hennig), the family comprised a single species, Aulacigaster leucopeza (Meigen). Hennig's precedent was followed by most dipterists for almost two decades (Griffiths 1972, Teskey 1987). David McAlpine (1983, 1985) described two new genera, Nemo and Ningulus, in the family and proposed a division of Aulacigastridae into two subfamilies: Nemininae with his new genera, and Aulacigastrinae with Aulacigaster Macquart (Schizochroa Hennig was synonymized with the latter). McAlpine's recharacterization of the family excluded Cyamops, Planinasus and Stenomicra, which he placed in the family Periscelididae. This classification and characterization was adopted by most subsequent workers (Colless & McAlpine 1991, Mathis & Papp 1992) with the exception of Freidberg (1994), who accorded familial status to Neminidae, described a new genus, Nemula, and enumerated 18 differences between Neminidae and Aulacigastridae sensu stricto. The result again is a monobasic concept of Aulacigastridae: Aulacigaster, consisting of 11 species worldwide. The pared down concept of the family is unquestionably monophyletic with numerous synapomorphies, including the peculiarly shaped and colored head and other morphological characters (D. K. McAlpine 1983, Mathis & Freidberg 1994). Although the higher classification for the Aulacigastridae is not entirely resolved, there is general agreement that the family is allied with the families Neurochaetidae, Periscelididae and Asteiidae in either the superfamily Asteioidea (Crosskey 1980, D. K. McAlpine 1978, Colless & McAlpine 1991) or the superfamily Opomyzoidea but under the suprafamily Asteioinea (J. F. McAlpine 1989). These differences largely represent issues of nomenclature and taxonomic rank rather than cladistic relationships.

2.3.3 Diversification and Distribution.

Although once considered to be species poor, recent sampling of Neotropical Diptera has revealed the Aulacigastridae to be far richer in species than was previously indicated (Mathis & Freidberg 1994). A single, three-week field trip to Peru, for example, resulted in nearly 20 species from a remote ranger station on the Manu River (Pakitza, 11¿56.75'S, 71¿17'W). We soon discovered additional species from other localities, nearly all undescribed. Over 40 species are preliminarily sorted thus far. The richness of these collections and subsequent attempts to identify them has prompted our proposal of this family for training in monographic work.

The aulacigastrids occur worldwide in temperate and tropical climes, but with considerable unevenness in their distribution. The Old World is species poor with only three species: Palearctic, 1 sp. (Papp 1984); Afrotropical, 2 spp. (Barraclough 1993). The New World is comparatively species rich with nearly 40 spp. (Mathis & Freidberg 1994), including eight that were previously described. Most New-World species are undescribed, and their distribution is also asymmetrical, with three species in the United States and Canada and the remainder in the neotropics.

2.3.4 Biology.

Flies of the family Aulacigastridae are peculiar to weeping wounds and sap fluxes of deciduous trees (Robinson 1953, Teskey 1976). Near or within these habitats, the species breed and undergo most of their life cycle. Eggs are laid singly or in small groups of two or three, usually directly in or on the flux, which apparently is the primary source of larval food. Larvae have three instars, the first being metapneustic, the second and third being amphipneustic. The third-instar larva, which along with the puparium was first described by Dufour (1846), is quite distinctive, with a long respiratory tube and a long, filamentous, prothoracic spiracle. Adults frequently occur on or near the flux or they hover within centimeters of its surface.

2.3.5 Available Material.

The largest and most diverse collection of Aulacigastridae, both in terms of new species and those previously described, is in the National Museum of Natural History (SI). Other significant collections are now available from the following institutions: American Museum of Natural History, New York; California Academy of Sciences; Canadian National Collection; Museo Zoologico"La Specola", Florence, Italy; private collection of Dr. Norman E. Woodley; Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania; and Tel Aviv University, Tel Aviv, Israel. We are in frequent communication with D.A. Barraclough, who will be cooperative with Afrotropical species, including holotypes.

3.1 Museum Research Protocols

A significant portion of data collection will be museum oriented and involve standard morphological procedures that need not be elaborated. We anticipate extensive use of SEM for character surveys.

Monographs attempt to synthesize all available information on a given taxon. Systematic monographs concentrate on identification aids (keys and illustrations), diagnoses and descriptions with associated illustrations, nomenclature, phylogenetic analyses and classification, information on distributions and biogeographic analyses, life histories, and a comprehensive bibliography. To prepare a monograph the monographer must study all available material germane to the taxonomic focus, which can be extensive. Thus, management of specimens and associated data is critical.

To properly and efficiently handle specimens used in research, the Diptera Unit at SI has developed and implemented a system of bar-coding specimens while storing label data in a relational database (Microsoft ACCESS). Basic label data (locality including coordinates, date, collector, collection depository) associated with bar-coded specimens will be accumulated for new material that is accessioned, SI material studied, and that obtained on loan. The bar codes and data elements conform to the appropriate community standards (Thompson 1990, 1994). The unique bar code will also be used to associate observations and measurements made while developing character taxon matrices for the specimens used (see Section 6).

Characters and information on their states within taxa will be stored in the DEscriptive Language for TAxonomy (DELTA) files (Dallwitz 1980, Pankhurst 1991). Use of the DELTA standard will insure efficient generation of parallel descriptions and keys, consistent terminology, and wide dissemination of results in different formats.

Students will be able to query the Databases of World Diptera at SI, which provides basic nomenclatorial data on described taxa. An interim nomenclature (="interim taxonomy" of Farrell & Erwin 1988) wherein unique species identification numbers are used in place of scientific names for unpublished taxa will insure that the various databases (tables) are synchronized. Interim nomenclature and character taxon matrices in digital formats will allow for rapid generation of useful products, such as specialized identification systems for All-Taxa-Biodiversity-Inventories (ATBIs) and national biological surveys, long before the final monographs are published or made available electronically.

The Diptera Unit maintains a species level inventory of its holdings (Diptera Unit 1994) which are curated according to community standards (McGinley 1993). The unit is developing a system of bar-coding to maintain the integrity of this inventory. Thus, all specimens used for the proposed monographs will be cared for according to the highest community standards.

3.2 Field Work

Field studies are required to supplement existing collections, document species distributions, and gather data on life history and behavior. All collecting during the proposed study will be in compliance with local, state, national and international laws and permit requirements. Material collected will be deposited in collections specified by collecting permits of the host country. In those instances where a depository has not been identified material will be placed in collections at SI.

The Smithsonian has a number of collaborative programs throughout the world, with emphasis on Latin America. These include field stations and programs in Guyana, Brazil, Peru, Belize which can be used to facilitate collecting in other remote areas. The Diptera Unit at SI also has strong ties with INBio (Costa Rica; see section 4.1.1).

3.2.1 Tanyderidae

Knowledge of immatures is critical to further understanding of the Tanyderidae. Immatures are known for Protoplasa (Crampton 1930a, b; Alexander 1930b), Protanyderus (Nearctic only; Exner & Craig 1976; Knight 1963, 1964; Rose 1963), Eutanyderus (Hinton 1966; Cranston, pers. com.), Radinoderus (Australian only; Cranston, pers. com.), Mischoderus (Crosby, pers. com.) and Peringueyomyia (Wood 1952). With the exception of Protoplasa, none of the immatures have been reared. In regions where the potential for species overlap exists, doubt remains as to the identity of the immatures collected.

Field work for the tanyderid monograph will emphasize immature stages. Travel to Australia-New Zealand, Chile-Argentina, Japan, and western North America is planned for collecting material. Life history patterns for the group make it impossible to sample all regions in a 2 year study. We have therefore chosen sites that reflect recent activity in the group and answer pertinent questions in systematics and biogeography. Methods for collecting immatures will follow descriptions provided by Exner & Craig (1976) for free-living larvae. Larvae associated with saturated wood adjacent to stream banks (Australian species of Eutanyderus, Radinoderus) will be collected by washing techniques. Where successful, the nearby gravel substrate will also be sampled to insure a thorough sampling regime and recover any pupae in the vicinity. Since there is only one record of a tanyderid rearing (Alexander 1930b) there are no explicit protocols. We are familiar with rearing various other nematocerous Diptera from a variety of habitats including saturated wood, knowledge that will be useful in attempts to rear tanyderids. Field studies will also supplement existing collections of adults. Most of the species have narrow windows of emergence so availability is often limited. Fresh material is required to supplement those taxa known from only type material.

An additional justification for field work is to expand and verify behavioral information. Hennig (1981) discussed modifications of the 4th and 5th tarsal segments of males as a synapomorphy for the clade uniting Ptychopteridae and Tanyderidae which he presupposed to be an adaptation in males insure successful copulation with females. Reservations were expressed by Wood & Borkent (1989) on the strength of this character. Two of the three genera in Ptychopteridae do not possess the apomorphic condition, and no one has examined the distribution of this character throughout the Tanyderidae. Field observations would verify the function of this character.

3.2.2 Rhagionidae

Field collecting for Rhagionidae will emphasize collecting adult material of poorly known genera. This will be primarily by Malaise trap collecting and hand sweeping. Travel to survey the south temperate faunas of South America and Australia is planned because no specialized collecting for rhagionids has been undertaken in these regions, and diversity is likely to exceed present knowledge.

A minor effort will be made to find and rear immatures. Unfortunately, larval rhagionids tend to be widely dispersed in soil habitats so that collection has a considerable chance element (Turner, pers. com.). Although information on immatures of enigmatic genera such as Austroleptis would be extraordinarily valuable for phylogenetic characters, the effort necessary to find them is likely to exceed the time frame of this proposal.

3.2.3 Aulacigastridae

As noted above (section 2.3.3), the Aulacigastridae have proven to be quite diverse at a single Neotropical locality. Field work for this monograph will therefore concentrate on sampling at additional locations to insure adequate geographic coverage and to assess the total diversity of the family. We have selected Costa Rica, Ecuador, and Bolivia as being important field sites for this purpose. Collecting will be by Malaise trap sampling and hand netting, especially near weeping wounds and sap fluxes that serve as adult habitats.

3.3 Phylogenetic Analyses, Biogeography and Biodiversity Studies

Characters will be selected primarily from comparative morphology, but behavioral and ecological data will be used if available. Morphological characters will be chosen and homologies interpreted based on extensive comparative study. Characters will be coded for phylogenetic analyses as either binary or qualitative multistate. All characters will be treated initially as unweighted (Hennig86 and PAUP). Characters will be treated as additive or nonadditive depending on the results of prior analyses (Wood & Borkent 1989; Woodley 1989; J. F. McAlpine 1989). Multiple heuristic searches will be done for each data set changing the addition sequence of taxa to insure finding the most parsimonous tree sets. Following initial parsimony analysis characters will be weighted using successive approximation (Farris 1969) and the homoplasy concave function discussed by Goloboff (1993b).

Selection of taxa is dependent on the focal taxon. Outgroup choice is discussed in the specific sections (3.2.1-3).

Biogeographic studies will follow methods described in Nelson & Platnick (1981), Humphries & Parenti (1986) and Humphries et al. (1988). Names for terminal taxa will be substituted with the area(s) in which they exist. We will explore the relationships of these areas with Hennig86, PAUP, COMPONENT (Page 1993) and TAS (Nelson & Ladiges 1992).

Information on biodiversity of taxa, including identifying areas of endemism, character and species richness will be assessed using the program WORLDMAP 3 (Williams 1994). Information of this type is then used to identify critical areas for future studies in biodiversity and conservation.

3.3.1 Tanyderidae

Phylogenetic analyses will be examined at the species level. All taxa will be included in the analyses. Morphological character systems from larvae, pupae and adults will be examined. The species placed in the Tanyderidae currently are united only on sympleisomorphies. For example, tanyderids are the only flies with four radial sector veins reaching the wing margin and a discal cell (Hennig 1973). Dipterists have tended to recognize a maximum complement of veins and the presence of a discal cell as the ground plan venation. These sympleisomorphic characters, however, are not informative in a phylogenetic context. We feel that a thorough, rigorous, comparative monographic study of all life stages will provide new character systems to establish the monophyly of tanyderids, resolve relationships within the tanyderids and confirm their phylogenetic placement in relation to other Nematocera. We have identified three preliminary character systems potentially useful in establishing the monophyly of the tanyderids: structure of the male aedeagus, structures of the adult compound eye, and structures of the posterior abdominal segment in larvae.

The choice of outgroup taxa will be critical for determining character state polarity within tanyderids and for establishing monophyly of the family. The outgroup taxa should be adequate to represent the diversity of character states of the ingroup, a complex issue if more than a one life stage is included in the analysis. Selection difficulties also arise due to various methodologies (or lack thereof) implemented by previous investigators. Two higher classification studies of Nematocera used explicit methods (Wood & Borkent 1989; Hennig 1981), while other studies did not have these techniques available (Wood 1952; Crampton 1930b, 1926, 1925). The latter investigations, however, were thorough examinations of character systems for the life stage under study and are valuable contributions to dipterology. Therefore, outgroups will be chosen based on evidence from all of these previous studies as a first approximation for assessing the phylogenetic relationships in Tanyderidae. Exemplars from the following families will used as outgroups: Ptychopteridae (1 each from Ptychopterinae and Bittacomorphinae), Psychodidae (1 from Bruchomyiinae), Dixidae (1 from Dixini), and Tipulidae (1 each from Hexatomini and Eriopterini).

3.3.2 Rhagionidae

Phylogenetic analysis will be at the genus level. Exemplars of each genus, once determined to be monophyletic, will be chosen for character scoring. Characters will be mostly or entirely adult morphological characters.

Outgroup taxa will be selected from basal lineages of Xylophagomorpha, Stratiomyomorpha (Pantophthalmidae), and Vermileonidae because these groups are by current estimations most likely to contain the sister taxon to the Tabanomorpha (Sinclair et al., 1994). We realize this is a first approximation, and that some Nematoceran group should probably also be included. However, as Griffiths (1994) stated, the Nematoceran sister group to the Brachycera remains highly speculative.

3.3.3 Aulacigastridae

Selection of an outgroup will be from families and/or other higher-level taxa within the superfamily Asteioidea (sensu Colless & McAlpine 1991; = suprafamily Asteioinea within the suprafamily Opomyzoidea of J. McAlpine 1989), with Neurochaetidae, Periscelididae and Neminidae being likely candidates. The family Neminidae, which is a plausible outgroup, was initially proposed as a subfamily within the Aulacigastridae (D. McAlpine 1985), and the latter was the"out group"in Friedberg's (1994) comparative study of Neminidae.

The Diptera Unit at the Smithsonian has a well documented record for training successful dipterists, most of whom have produced exemplary monographs. These individuals entered our program as pre- or postdoctoral fellows, finished their research, and went on to secure positions at prestigous institutions. Recent fellows include: Brian Brown (Natural History Museum of LA County), Martha Condon (Hofstra University), Greg Courtney (Grand Valley State University), Amnon Freidberg (Tel Aviv University), John Gelhaus (Academy of Natural Sciences, Philadelphia), Thomas Pape (Swedish Museum of Natural History), and Brian Wiegmann (University of North Carolina).

Diptera researchers based at the Smithsonian (see biographical sketches) are strongly committed to overseeing student education. We envision the following activities to educate dipterists:

1) We have agreed to"team teach"the two students funded by this proposal in dipterology at SI as a graduate-level special topics course (e.g. Biology 398, Advanced Reading & Research at GWU). This course will focus on taxonomy at the familial rank, and include taxonomy, biology and life history patterns, nomenclature, and phylogeny. Practical skills of importance to monographers will also be addressed, such as field collection methods, specimen preparation, construction and use of identification keys, biological illustration and curation. These skills are necessary to successfully complete the field and museum components for dissertation research;

2) A Diptera sorting group, composed of fellows and research scientists, is currently in place at the Museum. We sort a wide diversity of Diptera from Malaise trap samples and canopy fogging programs. Students will be expected to actively participate in sorting, as this is a means to see an incredible diversity of flies rapidly, which will assist in developing"an eye"for character variation in the order; and 3) Experts-in-training will be required to curate the Smithsonian collection of their focal taxon. This will include all facets of collection management (see section 3.1).

A second component to training competitive scientists is to have a system available to mentor those under stewardship. Mentoring is instrumental in building self-confidence, comradeship, honing communication skills, and the avoidance of unnecessary obstacles to progress. We are in the unique position of having a stable, cohesive critical mass of scientists all focusing on taxonomic issues in Diptera. However, existing as a graduate student is a difficult period in one's career regardless of the perceived advantages of a particular program. Our proposal is designed to maximize available mentorship. The post-doctoral trainee (Judd, section 4.2) will be a recent PhD familiar with the intricacy of living in Washington and being based at the Smithsonian Institution. This will allow the post-doc to assist the first graduate student in settling into the SI routine. At the end of two years, we expect the first graduate student to have completed all course work and be starting full-time research. Thus the first graduate student will be fully ensconced at SI and will be a mentor to the incoming second student.

A third important component in our proposal is our ability to evaluate progress prior to project completion. The post-doctoral trainee will expedite the evaluation process. We anticipate our first expert-in-training (Judd) will have produced a monograph on the Tanyderidae by the end of the second year. This will allow us to determine the success of our methodologies (section 3), training (section 4), and the choice of taxon focus (e.g. was the group correct size for the time requested, etc.). Graduate students will also contribute to the evaluation process although their comments will be more difficult to use since both are projected to finish near the end of the proposal term.

The success of our training can be assessed in other ways as well. A good monographer should be able to apply acquired skills to other taxa. Our post-doctoral trainee (Judd) will have done significant research on two separate and quite different taxa after finishing her work on tanyderids, which will be an actual example of this philosophy. We expect our trainees will thus have developed broadly useable skills that they can apply to any group of Diptera, and be very competitive in the job market.

4.1 Site Affiliations

The SI's National Museum of Natural History epitomizes cooperative research, particularly with units from other federal agencies (USDA, NOAA, NBS, DOD etc.), but also with departments in local universities (GWU, MCSE; see section 4.1.2). Within the SI there is an extraordinary network of satellite organizations, programs and facilities that will be accessible to participants in this proposal. The latter includes:

(1) an exceptional research collection that is one of, if not the finest in the world (5,300 drawers of pinned material, 470,000 slide-mounted specimens, and 17 quarter cases of specimens in alcohol) (Diptera Unit 1994);
(2) the largest community of scholars in biological systematics;
(3) access to outstanding library facilities (SI, Library of Congress, National Agriculture Library);
(4) the SI's facilities and field stations in the New World (Panama, Peru and Belize) and Old World (Aldabra and Kenya);
(5) excellent scanning electron microscopy facilities and photographic services;
(6) access to illustrators for advice and illustration supplies;
(7) collection related equipment (cabinets and drawers) and supplies (pins, labels, traps etc.); and (8) the extensive programmatic offerings of the SI's Biodiversity Program, such as the Biological Diversity in Latin America Project. The Diptera Unit also has very strong ties with the Costa Rican biodiversity program (INBio), where we have provided funds, basic literature resources, training for the INBio Diptera curator, and expertise in Diptera systematics. Students will be involved in this endeavor.

Training of students and dissemination of knowledge, which complement the objectives of PEET, are critical missions of SI and its Biodiversity Program. Furthermore, the SI's Department of Entomology is now recruiting for a collection-based research entomologist and anticipates additional recruitments, given the size of the Department, its responsibilities within the Museum and to the community, and its current age structure. There is and will be a need for well-trained entomologists.

4.1.2 University Affiliations

Two outstanding local universities strongly support our PEET proposal (see attached letters).

The Department of Biological Sciences at George Washington University (GWU) and the Maryland Center for Systematic Entomology at the Department of Entomology at the University of Maryland (MCSE) have a firm commitment to education in systematics. Departments and curators at the Smithsonian have actively participated with these institutions in training systematists. We believe students funded by this proposal will be provided the best available education and training in collection-based, comparative biology in the United States.

Students will be given the option of enrolling in GWU or MCSE. However, course work and the expertise available will not be limited to any single university or institution. The Consortium of Universities of the Washington Metropolitan Area allows students to enroll in courses throughout the District of Columbia (e.g. Georgetown University, Howard University, American University, etc.). This cooperative agreement presents a unique opportunity for students and advisors to tailor a custom program.

GWU is located in Washington, DC within fifteen minutes of the Smithsonian. A private, nonsectarian institution, the School of Arts and Sciences has an annual enrollment of 1500 graduate students. GWU has recently demonstrated its commitment to systematics by creating three new positions within the last year (mammals-Dr. Marc W. Allard, paleontology-Dr. James M. Clark, and entomology, selection-in-progress).

Students at GWU are expected to be knowledgable in three to four areas related to the focus of their dissertation. Graduate courses in systematic theory, evolution, molecular systematics, paleobiology, methodological-oriented techniques, and topical weekly seminars are currently being taught. Laboratories are associated with lecture courses and students receive extensive experience with software programs for phylogenetic analysis. The department is expanding its curriculum and courses in molecular evolution, conservation genetics, coevolution, biogeography, and insect phylogeny will be offered over the next three years. Currently there are eleven graduate students in systematic biology at GWU.

The University of Maryland is a land-grant institution located 8 miles from the Smithsonian with easy access by subway. MCSE is a long-standing consortium between the University of Maryland, the Systematic Entomology Laboratory (USDA), and the Smithsonian Institution (SI) for training graduate students. MCSE has two systematists on the Maryland faculty (Dr. Charles Mitter and Dr. J. Schultz) and six graduate students.

Systematic students at MCSE take core courses including insect classification, evolution, comparative development, population genetics, and physiology giving them a strong backround in entomology. A team-taught course in systematics is offered that covers modern systematic theory and practice and addresses practical issues related to species descriptions, curation, and publication. The MCSE program also requires that students attend a weekly discussion group on current papers in systematics and evolution.

In addition to opportunities provided by GWU, MCSE, The Consortium of Universities of the Washington Metropolitan Area and the Smithsonian, students will be able to compete for positions in courses offered through the Organization for Tropical Studies (OTS). The Smithsonian is a member of OTS and our curators are active participants as coordinators and resource scientists. Issues critical to training future scientists, such as project design and implementation, statistics, cooperation, biodiversity, and written and oral presentations, are addressed. We consider this program to be vital and expect our students to be highly competitive for available slots.

For students entering the current job market, knowledge and competency in molecular biology is required. GWU, MCSE, and the Laboratory of Molecular Systematics at the Smithsonian have established high quality research and training programs in molecular systematics. Participants from all these programs attend regularly scheduled discussions of current research papers. Courses are available at the identified universities to provide an understanding of the complex issues related to evolution, structure, and phylogenetic anaylsis. We expect to produce students that are competent to analyze molecular data and assess current issues in molecular biology. The post doctoral trainee has prior experience and course work in this area of study.

4.2 Post-doctoral Fellow

4.3 Graduate Students

A PhD student working on Aulacigastridae will begin in Year 1 and continue through Year 5. A potential graduate student, Alessandra Rung de Paula Baptista (Rio de Janeiro, Brazil), has expressed an interest in working on Aulacigastridae and seeking a Ph.D. A biographical sketch is attached. 4.3.2 Rhagionidae

The second PhD student will begin studies in Year 3 and we anticipate completion at the end of Year 5. No student has been identified so we anticipate a recruitment (see below). Given the duration of funding, preference will be given to a candidate possessing a Master's degree in systematic entomology. Final receipt of the NSF traineeship will be dependent on acceptance by GWU or MCSE. The Smithsonian has committed to additional funding for this student if the project is not finished by the end of the 5th year of this proposal (see budget justification).

We will recruit the second PhD student as follows. Upon notification of funding we will prepare an announcement to be placed in Fly Times, a newsletter of the North American Dipterist Society issued quarterly and mailed to approximately 175 individuals throughout North America with various interests in dipterology. An more detailed announcement will be placed in national publications (e.g. Newsletter of the Entomological Society of america, Science), at the national meeting of the Entomological Society of America, and sent to university departments during Year 1. Prospective experts-in-training will be interviewed by Smithsonian and GWU and/or MCSE staff with final selection by agreement between the PI and the appropriate faculty. SI will provide funds for related expenses.

4.3.3 Advising Graduate Students

Each student in this proposal will have a committee that will consist of the PI, a primary faculty adviser at either GWU or MCSE, a minimum of one other dipterist, and other individuals deemed critical to the successful completion and enhancement of the dissertation. Expenses for external examiners will be provided by SI. The purpose of the committee will be to produce dipterists broadly trained in comparative biology and capable of competing successfully in a highly competitive future job market.

4.4 Undergraduate Interns

The SI, through its National Museum of Natural History Research Training Program, annually sponsors and hosts a highly competitive and successful program for undergraduate interns. In 1994 over 502 candidates applied for the 26 positions that were funded. Last year Mr. Navin Rao (University of Puget Sound) worked with the PI and produced a revision of the shore-fly genus Rhysophora (Rao & Mathis 1995). We hope to utilize this program to expose talented young students to systematics, especially through field work in North America.

5.1 Taxonomy and Monography

The completion of the work in this proposal will result in three highly trained monographers with extensive museum, field, and analytical experience. Our experts-in-training will able to:
1) prepare high quality specimens,
2) manage specimen and character data and be familiar with curation of major collections,
3) recognize and understand homologies in the Diptera,
4) characterize and identify taxa
5) understand nomenclature, and
6) make predictive, stable classifications based on phylogenetic principles. These monographers will have produced three outstanding, comprehensive monographs on taxa of Diptera that are poorly known and in which there are no current workers using modern methodology. Each of these scientists will have a broad, fundamental knowledge of Diptera, allowing them to pursue further systematic research on any taxon within the order. They will also be able to train future dipterists in a highly competent manner.

5.2 Phylogeny

The importance of the Tanyderidae rests in their phylogenetic position which is currently unresolved. They are undoubtedly one of the basal lineages of the Diptera. As a basal lineage, the character systems are of critical importance for determining homologies and assessing character state polarities for the entire Diptera. Characterization of all life stages are inadequate and no recent studies have addressed the fauna worldwide. The absence of knowledge for this group subsequently affects our ability to evaluate operational hypotheses in behavior, vector potential, and ecology for the Nematocera.

Recognition of importance of basal taxa has also been cited in studies of conservation biology and biodiversity (Stiassy 1992). Vane-Wright, Humphries & Williams (1991) use an index of phylogenetic position (where older and less speciose taxa are given higher values) to make decisions on which areas should be preserved given limited funding. Results of this study, as well as that on the Rhagionidae, will provide valuable data for studies in biodiversity.

5.2.2 Rhagionidae

Completion of a monograph of the genera of Rhagionidae will resolve a number of phylogenetic issues. It will establish the monophyly of the family or suggest an alternative phylogeny that might recognize more than one family from the rhagioniform genera. Knowledge of this basal clade in the Tabanomorpha is critical to understanding the relationships within that lineage, and its placement with respect to the other lower Brachycera (Woodley 1989). Furthermore, this phylogenetic resolution will enable other biological information to be placed in context, For example, is vertebrate blood feeding in rhagionids of monophyletic or polyphyletic origin? Or, is haematophagy primitive to the Tabanomorpha (it occurs also in Athericidae and Tabanidae) and subsequently lost in multiple lineages?

5.2.3 Aulacigastridae

A monograph of the Aulacigastridae will advance knowledge of the cladistic relationships in acalyptrate flies. We anticipate that phylogenetic placement of this group with respect to other periscelidid and related flies will be resolved.

5.3 Biogeography

The distribution observed in the Tanyderidae is one of the most interesting and puzzling patterns in biogeography. Holarctic in the North, Gondwanan distribution in the South and a smattering of taxa in the Pacific. Various groups from plants (Du Rietz 1940) to crustaceans (Ekman 1953) to sardines (Regan 1916) display a similar pattern. The geological models thus far proposed to interpret land mass distribution have been less than satisfactory for explaining this type of distribution for floral and faunal components (Nelson 1985). Our data will allow us not only to examine current geological models (Nur & Ben-Avraham 1981; Carey 1983; Sluys 1994; Shields 1991), but supply empirical data to hotly debated issues on observed patterns of distribution among taxa (Holzenthal 1986; Craw 1983, Humphries 1983; Fleming 1979).

5.3.2 Rhagionidae

A thorough understanding of rhagionid phylogeny will provide data for assessing transantarctic biogeographic patterns in the family. The family has been used in compilations of taxa showing such distribution patterns (Hennig 1960), but until the genera or higher taxa are shown to be monophyletic, their support for biogeographic concepts is equivocal.

5.3.3 Aulacigastridae

Recently collected specimens indicate that the center for aulacigastrid diversity is the New World tropics, where 90% or more of the fauna occurs, mostly undescribed. The accelerated loss of biodiversity in the neotropics is a compelling argument that motivated our proposal of Aulacigastridae for monographic study and training. A cladistic knowledge of these flies may also provide data for examining theories of hypothetical refugia and centers of radiation in northern South America and their importance in conservation efforts (Haffer 1974; Barrowclough 1992; Erwin 1991).

The Smithsonian Contributions to Zoology is a peer-reviewed primary journal devoted to taxonomic and monographic studies. We view this as the primary publication resource for the monographs produced in this proposal. SI has committed to publish the three monographs produced by the work in this proposal.

The future of monography is undoubtedly in electronic formats. CD-ROM technology and Internet access is increasing at breath-taking speed. Dissemination of taxonomic information will be cheaper and in a more efficiently usable form with these media. The user will access the information through query options of database programs, expert systems (identification) or graphic viewers. The electronic hardware, software, and technological expertise currently exists at the Smithsonian to produce electronic monographs. We envision the translation of a traditional monograph into electronic format as follows:

Title Page: This is the initial (logon) screen that the user sees when they access the digitial medium via gopher (top menu), World Wide Web (WWW; home page) server, or a program loaded from CD-ROM.

Table of Contents: This translates into the menu of options that the user can select from. This willfrequently be a hierarchical series of menus, the first of which is part of the logon screen.

Introduction: This will be an ASCII text file (or series of text files) marked up according to the appropriate"hypertext"markup language (see below).

Acknowledgments: This will also be an ASCII text file.

Keys: These will be replaced by an expert identification system. They use DELTA datafiles (ONLINE). The datafiles for the identification program may be separate from those of the phylogenetic program (see below) as not all characters are useful for both functions. While markup languages allow for the traditional key to be used in a digitial environment, the hypertext format still restricts the user to a predefined path through the characters (if you don't know the first or subsequent characters, you are stuck at that point). Hence, this approach will only be used in those environments (Internet) where more interactive programs are not supported.

Phylogeny & Biogeography: The analyses will be included in marked-up ASCII text files, but the supporting data will be included as character taxa (phylogenetic) or taxa area (biogeography) matricies. The preferred cladograms and maps will also be stored as images.

Taxonomic descriptions: Descriptions will be derived from the character taxa matricies that are available from the previous sections. Descriptions can also be stored as marked-up ASCII text files.

Nomenclature: Will be derived from a database. Predefined queries will generate the traditional formated list of names and nomenclatorial details (literature citations, type material, status, etc.).

Distribution: Will be derived from a database of specimen label data. Mapping programs will be included to generate graphic images of distributions.

Life histories: Will be summarized in marked-up ASCII text files, but data on associates, phenology, etc., will be stored in databases.

Figures: Illustrations of characters, species, distributions, relationships (cladograms) and any other necessary figures will be image files.

Bibliography: Will be in the form of a database.

Index: Will not be necessary!

Given the current status of the digitial environment, we would disseminate our proposed monographs on CD-ROM and over the Internet. The CD-ROM version would consist of a relational database (Microsoft ACCESS) and expert system (Pankhurst's ONLINE) linked by a batch program. These would be transferred to the user's computer by an INSTALL program (Knowledge Dynamics), but the data and image files would be resident on the CD-ROM. The Internet version would consist of the traditional printed copy marked up (HTML) for a WWW server. However, we believe a more interactive version will be possible by the time the first monograph is finalized.

• Year 1: Tanyderidae expert-in-training starts research; collecting trips to Japan, Australia, New Zealand, North America. Aulacigastridae expert-in-training starts coursework.
• Year 2: Tanyderidae expert-in-training continues research; collecting trip to Chile-Argentina; project completion. Aulacigastridae expert-in-training finishes course work; collecting trip to Costa Rica.
• Year 3: Aulacigastridae expert-in-training starts full-time research; collecting trip to Ecuador. Rhagionidae expert-in-training starts course work and research; collecting trip to Chile-Argentina.
• Year 4: Aulacigastridae expert-in-training continues research; collecting trip to Bolivia. Rhagionidae expert-in-training finishes course work and begins full-time research; collecting trip to Australia.
• Year 5: Aulacigastridae expert-in-training completes project. Rhagionidae expert-in-training completes project.

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