David Grimaldi - US grants

The abundant deposits of Oligo-miocene amber from southern Mexico and Hispaniola have yielded a great diversity of exquisitely preserved biological inclusions, the large majority being arthropods and especially Diptera. The amber specimens afford an unparalleled opportunity to address questions on the relationships and origins of the Caribbean fauna, but critical information is lacking on the distributions of most extant Diptera in the caribbean, and the phylogenetic relationships of the amber fossil taxa to living species. This research proposes to examine selected groups represented in the amber collections the AMNH and the NMNH: Asteia (Asteiidae), Aulacigaster (Aulacigastridae), Beckeriella (Ephydridae), Beebeomyia (Richardiidae), Carnus/Meoneura (Carnidae), Lygistorrhina and Proceroplatus (Mycetophilidae), Diathoneura and selected Drosophila groups (Drosophilidae), Odinia (Odiniidae), and a species group within Pholeomyia (Milichiidae). Taxonomic revisions of each will be based on species-level treatments using adult morphology and cladistic methodology to examine phylogenetic relationships. Study will resort to pinned museum specimens but requires extensive sampling using several general and specific collecting techniques in lowland tropical forests, which is proposed for Hispa ola, Cuba, and Costa Rica. Gas chromatography/mass spectroscopy will be done by a collaborator on selected amber specimens to more accurately determine their geological provenance. Aims of the research are the following: define species and their distributions for the above groups in the Netropics; test for repeated, congruent patterns of distributions; and place the fossils among the phylogenetic system of extant species. Implications of these results are that distributions can be examined for correspondence to island origins (vicariance), and the minimum, fossil age of a clade may provide permissive evidence for or refute a particular geological explanation. Also, the role of geographical extinctions in modifying distributions can be assessed, which is directly applicable to vicariance methodology.

9870097 Krishna The order Isoptera (termites) comprises about 2,300 described and several hundred undescribed species (living and fossil), classified into 6 families, 12 subfamilies, and 290 genera. In spite of its comparatively small size, because the order has generated great interest, a massive body of literature has been produced on all aspects of termite biology. Aside from their notorious inportance as insect pests, termites, because of their intricate social biology and their critical importance in the ecosystem as recyclers of cellulose, have attracted the attention of a wide range of researchers in many fields: evolutionary biology, ecology, behavior, physiology, economic biology, genetics, cytology, and other areas of investigation. The aim of this project is to create a taxonomic database of the termites of the world. The database (and the printed volume derived from it) will provide complete and up-to-date references to all systematic, zoogeographical, and biological information available for each species of Isoptera. The database will be put on line on the World-Wide Web, where it will provide comprehensive and current information to biologists around the world and will serve as a global data bank for the future.

DEB- 0075360
David A. Grimaldi

Drs. David Grimaldi and Valerie Schawaroch of the American Museum of Natural History have been awarded a grant to study the anatomy, habits, distribution, and evolutionary relationships of a group of fruitflies used as an experimental foundation by many researchers in genetics and developmental biology. The common laboratory fruitfly, Drosophila melanogaster, is probably better known than any other animal species on earth. Serious study of the fly began in 1910, when it was first used as the model organism for genetics. Drosophila melanogaster actually belongs to a group of 175 named species, the melanogaster species group, all of which live naturally in tropical forests of Africa, Eurasia, Australia, and Pacific Islands. Some species, like D. melanogaster, breed in a great variety of fruits and vegetables, and have become widespread (even agricultural pests) wherever people have settled. The diversity of species and habits, their ease of lab culture, and unique knowledge of the model species, has made the melanogaster species group an ideal subject for all aspects of comparative biology.

Nevertheless, accurate identification of species by specialists and especially other biologists like geneticists is very difficult. Distributions of individual species are poorly documented, and introductions of exotic species are therefore difficult to detect. Collections of major natural history museums already harbor new species, awaiting complete description, naming, and classification. The goal of the PIs is to rectify deficiencies in the basic systematics of the melanogaster species group, by production of a hard copy and digital synthetic monograph which will treat the anatomy, habits, distribution, and relationships of each species in detail, including newly described ones. This monograph will stand as a major reference, and will be accompanied by an interactive, lavishly illustrated CD-ROM for identification of species. The package will be of widespread use to students, laboratory technicians, quarantine entomologists, and various biologists, and will greatly facilitate all aspects of comparative biology wherever these flies are used, from genetics and physiology to development and evolution.

The Department of Entomology at the American Museum of Natural History houses 13,500 specimens of fossil insects, including 624 types. The main strengths are fossils in amber, Cretaceous fossils, and sizeable historical collection of insects in Oligocene shales from Florissant, Colorado, that has received virtually no curation for 80 years. This proposal seeks to complete preparation and cataloguing of several thousand amber and compression fossil specimens; to identify approximately 2000 specimens; to house the compression fossils in new drawers; to prepare all types and other important amber specimens for long-term conservation; and to create high-resolution Computed Tomography images of selected specimens. In addition, the project will investigate new technology such as High-Resolution Soft-Xray imaging as a means to generate data from damaged and degraded amber sources.

INSECT AND VERTEBRATE FAUNAS OF A UNIQUE LATE TRIASSIC FRESHWATER ECOSYSTEM IN EASTERN NORTH AMERICA

Nicholas C. Fraser and David A. Grimaldi
EAR-0106309

The end of the Triassic period (230 -200 mya) was a pivotal time in the evolution of modern terrestrial ecosystems. There is a good global record for tetrapods, including the first mammals, crocodiles, turtles, lissamphibians (frogs and salamanders), sphenodontians, and, by extension, their sister taxon, the lizards. By contrast the record for the most significant group of modern day animals, the insects, is considerably poorer. Only a handful of rich insect-producing Triassic sequences are known worldwide, and the majority of fossils from these localities comprise incomplete specimens such as isolated wings and elytra.

A single site in the Newark Supergroup of eastern North America has been found to contain numerous complete Late Triassic insects. The Solite Quarry is situated on the Virginia-North Carolina state line, and it has produced the oldest global records for water bugs, thrips and caddisflies. Other fossils found to date include a unique gliding reptile and numerous specimens of a small aquatic reptile that frequently preserve details of their soft part anatomy. These have yet to be fully described. Only a relatively small portion of the fossiliferous sequences has been excavated and PIs will open large-scale excavations in the most productive quarry units. The next phase of the work will be to conduct detailed studies of all the finds and to compare them with the two major assemblages that share the most in common: the Madygen Formation of Kirghizistan and the Molteno Formation of South Africa. The exquisite preservation of the Solite insects will permit a complete re-evaluation of the extensive Russian and South African insect collections and thereby allow for a better understanding of the mode and tempo of insect evolution and radiation at this critical time.

For a long time the Solite quarry was regarded as being largely unfossiliferous. Certainly the microscopic insect fossils can be readily overlooked under normal lighting conditions. The search image is critical to the PIs success, and using the same methods they have identified other localities nearby that also contain insect remains. Together with the diverse plant and vertebrate remains, they shall reconstruct a detailed picture of marginal freshwater ecosystems in the Late Triassic.

Insects are the most diverse class of organisms in the 3.5 billion years of life on earth, and one (but not exclusive) explanation for this is that insects radiated in the Cretaceous as plant feeders and pollinators along with the radiation of the dominant form of plant life, the angiosperms. In other words: insects may be exceptionally diverse because plants speeded their rate of origin or speciation. This theory will be addressed by studying diverse species of insects fossilized in ancient resin (amber) from around the world, including hundreds of specimens from 3 collections in the United States, 2 in Canada, and 5 in the UK, Spain, France, Russia, and Germany. Specimens range in age from 120 to 70 million years old, and study will focus on the megadiverse orders Diptera (flies) and Hymenoptera (ants, bees, wasps), with additional work on the Lepidoptera (moths) and termites. The amber will be meticulously prepared and detailed microscopic study will be made of the insect inclusions, along with photomicrographs, detailed illustrations and descriptions, and the specimens will then be compared to living and other extinct species to determine phylogenetic relationships. If angiosperms affected the evolution of insects, then groups of insects that today are intimately associated with angiosperms (like gall midges and gall wasps) will show rapid diversification in the Late Cretaceous and Tertiary, whereas the diversification of predatory and scavenger insects should not be timed to that of angiosperms. A low rate of extinction can also increase diversity, and this study will also address whether insects were less affected by extinctions at the Cretaceous-Tertiary boundary 65 mya that killed off other major groups of terrestrial life.
The intricate, life-like preservation in amber provides unique scientific and educational opportunities. Virtually all morphological characters found in living species are visible in amber fossils, thus allowing profoundly accurate measures of the amount of change over vast expanses of time and discerning the relationships of extinct life forms. With this, scientists can directly confront criticisms about the incompleteness of the fossil record for revealing evolutionary change. Also, the esthetic appeal of fossils preserved in amber engages the public with insect diversity, fossils, evolution and the history of life in general, as well as with the interface between biology and earth history. Besides the formal education of students in systematics, entomology, and paleontology, this research will promote informal science education through a new web site and popular publications.

A Time Scale for Scales: Basal Phylogeny and Divergence Times in Coccoidea (Hemiptera: Insecta)

Scale insects are plant pests of considerable significance in agriculture and forestry, causing billions of dollars in damage annually. For this reason, the taxonomy and identification of species is quite sophisticated and specialized, but relationships among the larger lineages are still poorly explored. With this grant, research will be expanded on the evolutionary relationships among the basal families of scale insects, using anatomical structures of the adult females and males as well as DNA sequences. The results will allow informed predictions to be made about groups whose biology (e.g., plant hosts) is poorly known. Since scale insects are so diverse in ambers 130 to 20 million years old, they are a model group to examine how plant-feeding insects have evolved with respect to the flowering plants (angiosperms). Scales preserved in ambers from around the world will be studied, and the anatomical data from the fossils in a large data set with the modern species analyzed. Select amber specimens will be scanned at the Advanced Photon Source, Argonne National Laboratory, to visualize obscured microscopic structures. The project will allow to determine when the families of scale insects diverged in geological time, and assess if their evolution coincided with that of the flowering plants.

The PI and co-PI are actively involved with informal science education, and the funded work will be incorporated into it. The Co-PI has designed Entomology sessions of the biodiversity course for the NSF-sponsored Science Research Mentoring Program with which the AMNH Education Department is preparing the production of web material, and will be available through this online portal. The PI mentors two high school students in this program, working on amber. The Co-PI will also be involved in informal public education, including hands-on laboratory experiences for middle and high school students in the Sackler Educational Laboratory. Lastly, a product of this research will be an online interactive identification key available on the co-PI?s research website.

Fossil insects provide a unique deep-time record of ecological and evolutionary response to past environmental changes and therefore are invaluable for understanding the impacts of climate change on the current biodiversity crisis. Given current models of future climate change and the important role that insects play in human society (biodiversity, pests, pollination, vectors of disease) the ability to access these data and make predictions about future insect populations becomes even more urgent. The Fossil Insect Collaborative will make available all the major collections of fossil insect specimens in the United States by creating electronic specimen records consisting of digital images and associated collection data.

The digitized fossil insect collections will be made broadly accessible to the research community, K-16 education, government and industry, the general public, and the media through the project website and a central site integrating all the paleobiological Thematic Collections Networks called iDigPaleo. Mobile apps and activities that allow a wide variety of users to experience and interact directly with the collections data will be developed. This award is made as part of the National Resource for Digitization of Biological Collections through the Advancing Digitization of Biological Collections program and all data resulting from this award will be available through the national resource (iDigBio.org).

Researchers at the American Museum of Natural History are using 20 to 100 million year old amber specimens (fossilized tree resin) from deposits around the world to understand how ants evolved from an ancestral solitary wasp into the diverse and abundant group of social creatures they are today. Fossil and modern specimens in the AMNH, as well as in museums in Russia, England, France, and Brazil are being studied using X-Ray-based 3d reconstruction and traditional light microscopy. These data are being combined with published DNA sequences from modern ants and analyzed to produce a clear pattern of evolution through time and the major physiological, behavioral, and ecological features that aided in diversification.

Ants are one of the greatest success stories in nature. At present there are over 12,000 species of ants, with an estimated biomass at least equal to all humans. This project seeks to fundamentally inform other studies relating to behavior, ecology, systematics, and developmental biology by detailing relationships among these model organisms in a novel way. In addition, because ants are very familiar and recognizable to the public, there is tremendous potential for outreach. The co-PI has and will continue to educate K-12 students and the museum-going public using ants as a model for understanding biological concepts at large.

This project will synthesize more than 30 years of research on the evolutionary history and classification of the "higher" flies, including robber flies, bee flies, flower flies, fruit flies, carrion flies, bot flies, and tsetse flies, many of which are economically and ecologically important. It will integrate exciting new fossil discoveries as well as anatomical and new genomic data. A unique feature of the fossil record for these flies is that many diverse higher flies are preserved with life-like fidelity in fossilized resin (amber) that is from 17 to 130 million years old. This profoundly enhances the accuracy with which the timing and amount of evolutionary change can be measured. With these data, this project can address whether higher flies are diverse because their speciation is so rapid, or because their extinction rate is low, or both. It also can address whether higher flies were affected by mass extinctions that occurred 66 million years ago, when most dinosaurs became extinct, and whether the largest group of higher flies, which represents the largest radiation of all life in the past 66 million years, also are the fastest evolving. Besides these broad questions, this study will also address the rate of evolution of fruit flies (Drosophila), based on several hundred specimens in amber. Since fruit flies are a model system for genetic and genomic research, this will provide a unique context for research in genomics and evolution. The project also will provide research training and experience for high school students and develop a museum exhibit on the evolution of flies.

This project will construct a comprehensive evolutionary framework through the analysis, study, and interpretation of over 30 years of accumulated data on the phylogeny, biogeography, and general evolution of the "higher" flies (Brachyceran Diptera). It will build upon the principal investigator's experience from hundreds dissections of exemplar living species, thousands of measurements and digital images of living and fossil species, and writing descriptions for over 70 fossil species to first assemble a data matrix of 592 morphological characters from 526 taxa, including both fossil and extant species. This morphological and molecular data will be used to compile a monograph on the morphology, fossil record, and phylogeny of the Brachycera, emphasizing the Mesozoic Era. This project also will produce a detailed descriptive study on diverse new Drosophilidae preserved in amber (42-17 million years ago) and two papers that address divergence times of lineages in lower Brachycera and in the Ephydroidea, especially Drosophila, analyzed with morphological and genomic data. Outreach will include mentoring high school student interns in the amber laboratory and developing a traveling or permanent exhibit at the American Museum of Natural History, "Flies."