David S. Hibbett, Ph.D. - US grants

9903835
Hibbett and Donoghue
Homobasidiomycetes include the mushroom-forming fungi and their relatives. About 13,500 species have been described, which is about 23% of all known species of fungi. The fruiting bodies of homobasidiomycetes include conspicuous, developmentally integrated structures such as mushrooms and puffballs, as well as very simple forms such as the crust-like fruiting bodies of "corticioid" fungi. Homobasidiomycetes play major roles in carbon cycling in terrestrial ecosystems; they comprise the majority of wood-decay fungi and ectomycorrhizal fungi (associated with tree roots); and they also include mycoparasites, insect symbionts, lichens, and litter decomposers. Although individual groups of these fungi have been studied intensively, their overall pattern of morphological and ecological evolution has not been determined because to date there has existed no broad well-supported phylogenetic framework for the group.
Over the last few years, considerable progress has been made in understanding homobasidiomycete phylogeny through the use of new DNA sequence characters, particularly the genes encoding nuclear and mitochondrial ribosomal RNA (rDNA). This project will expand the database of nuclear and mitochondrial rDNA sequences and add new data from protein-coding genes under study, such as the tubulins or RNA polymerases. Taxon sampling will target problematic groups such as the polyporoid, hymenochaetoid, and cantharelloid groups of fungi, whose relationships remain weakly supported by the existing molecular data. Conventional protocols for phylogenetic analyses will be employed, and in addition the investigators will explore the use of "supertrees" for combining or collating independent phylogenetic trees with overlapping sets of taxa in order to construct a composite phylogenetic hypothesis. The trees will be used to construct phylogenetic classifications and to study historical patterns of evolution in selected morphological and ecological characters. Specific characters to be investigated include: corticioid versus erect fruiting bodies; brown rot versus white rot modes of wood decay; and ectomycorrhizal versus saprotrophic and pathogenic modes of nutrition. A broad framework phylogeny for the homobasidiomycetes will facilitate, for example by identifying sister groups and nearest outgroups, more intensive analyses of morphological and physiological changes in these fungi.

Abstract Larochelle 0070241

Modern imaging facilities are essential for biological research in cell and developmental biology. The Department of Biology at Clark University is upgrading their imaging facility with a high quality research-grade compound microscope with phase contrast, differential interference contrast, and epifluorescence optics. In addition, a cooled CCD camera, a computer, and software for capturing, processing, and storing images, will be part of the system. Together these constitute a complete system for the generation and analysis of photomicrographs.

Specific research to be carried out with this system will include the microscopic analysis of cells defective in cell division, the examination of nuclear behavior of certain fungi as a means of gaining insights into fungal evolutionary relationships as well as fungal developmental evolution on a cellular level, the analysis of proteins required in the synthesis and regulation of membrane organelles, and the examination of the role of receptor tyrosine kinase signaling pathways in the growth and development of the Drosophila eye.

Six new faculty members (out of nine total) have joined the Department of Biology within the last seven years. With this turnover also comes an updating and improvement of the current facilities. Although the four co-PIs and their associated lab personnel will be the primary users of the facility, this imaging system will be available for research use by all faculty, graduate students, and advanced undergraduate students in the department. The addition of a research-grade microscopy imaging facility will become a cornerstone for the department. Despite its small size, the Department of Biology has been very successful in involving undergraduate students in research and in preparing them for advanced studies. Since 1989 over 350 students that have received their baccalaureate degrees in the sciences from Clark University have gone on to obtain advance degrees in biology or health-related disciplines. This imaging system will contribute greatly to the ability to train students and will be an integral part of the research programs of the four co-PIs.

0128925
Hibbett and Binder
The proposed research by Drs. David Hibbett and Manfred Binder at Clark University concerns the evolutionary relationships of homobasidiomycetes, which include the mushroom-forming fungi. There are about 13,500 described species of these fungi. Homobasidiomycetes play important ecological roles as decayers, pathogens, and beneficial symbionts of plants and animals. Traditionally, classifications and evolutionary hypotheses in this diverse group have been based on the morphology of the fruiting bodies, which include forms with a cap and stalk, club and candelabra-shaped forms, puffballs, bracket fungi, etc. Recently, however, DNA sequences have been used to infer evolutionary relationships of homobasidiomycetes. The proposed research will use DNA sequences to study relationships of two groups of homobasidiomycetes: "cyphelloid" fungi (roughly 300 species), which are minute cup-shaped forms, and aquatic homobasidiomycetes (20 species, usually quite rare). Both cyphelloid and aquatic homobasidiomycetes have been taxonomically obscure. Recent molecular evidence suggests that these groups may be closely related, which had not been predicted previously. The proposed research will follow up on the leads generated in recent molecular studies, and will include fieldwork to expand knowledge of the diversity in both groups. The proposed work will increase understanding of the pathways of morphological evolution in homobasidiomycetes. In addition, it will elucidate the ecological transitions from terrestrial to aquatic habitats in fungi.
The proposed research will benefit science and society in two major ways: 1) The proposed research will enhance understanding of biodiversity. In particular, it will highlight an unusual group of organisms that even most biologists are not aware of, but that play important ecological roles. 2) The proposed research will include a significant training and outreach component. A post-doctoral fellow and a Ph.D. student will receive training. In addition, a series of summer workshops in the Worcester, MA, area for local high school students and high school teachers is planned (one student and one teacher will be supported per year). The summer workshops will provide exposure to modern techniques of molecular biology and evolutionary biology. Teachers who participate in the workshops will receive professional development points, which are necessary for annual recertification in Massachusetts, and which Clark University is authorized to award. Teachers and students will be recruited from the Worcester public school system.

0228657
Hibbett
Fungi make up one of the major lineages of life, and include important decay organisms and pathogens of humans, plants, and animals. There are roughly 80,000 described species of Fungi, but the actual diversity in the group has been estimated to be as high as 1.5 million species. This collaborative project by five major investigators and their numerous colleagues worldwide will generate the first global synthesis of the phylogenetic history of the fungi, which is necessary to expand our knowledge of the history of life on Earth as well as the origin of ecosystems and adaptive features that can have a direct impact on human health. Assembling the Fungal Tree of Life (the AFToL project) will significantly enhance our understanding of the fungal kingdom, especially ancient relationships that are not resolved among the chytrids, zygomycetes, ascomycetes, and basidiomycetes. The AFToL project will develop large data sets of molecular and non-molecular (morphological, anatomical, life-history) characters, which will be accessible via the world-wide web in continuously updated databases. Molecular characters to be acquired include DNA sequences for seven nuclear genes sampled from approximately 1500 species, representing all major groups of Fungi. Nonmolecular characters to be sampled include cellular machinery associated with nuclear and cell division and cell wall biochemistry. The AFToL project will be based in five laboratories at four universities with the core responsibilities of the participating laboratories as follows: David Hibbett (Clark University), collection of molecular data from Basidiomycota; Francois Lutzoni (Duke University), collection of molecular data from Ascomycota including lichens and bioinformatics; David McLaughlin (University of Minnesota), collection and databasing of morphological characters; Joseph Spatafora (Oregon State University), collection of molecular data from Ascomycota; and Rytas Vilgalys (Duke University), collection of molecular data from Chytridiomycota and Zygomycota. The AFToL project will involve more than 115 members of the international fungal systematics community in 23 countries for all stages of the project, from selection of taxa to collection and analyses of data. It will be the policy of the AFToL project that all validated data will be released via the web as soon as they have been generated.
Fungi play crucial ecological roles as decayers, mutualistic symbionts, and pathogens, including pathogens of humans. The economic significance of fungi is almost incalculable; they perform vital "ecological services" and their functional roles are the subject of diverse applied disciplines, including agriculture, medicine, and drug discovery, to name just a few. A phylogenetic database for Fungi will facilitate the creation of powerful diagnostic and forensic tools for environmental surveys and other applications, and will enable the discovery of the many fungal species that remain undescribed. Training and outreach activities are an important aspect of the AFToL project and will take several forms including graduate and post-doctoral training, support for visiting graduate students, undergraduate training, summer courses for high school teachers, and website resources for K-12 educators.

A grant has been awarded to Clark University under the direction of Dr. David Hibbett
(PI) with four co-Principal Investigators (co-PIs), titled "Acquisition of a High
Performance Parallel Computing Cluster for the Departments of Biology, Chemistry,
Physics, and Mathematics and Computer Science of Clark University". The proposed
project involves the development of a shared computational faculty for the natural
science programs of Clark University. The instrumentation includes a Beowulf-type clusters, with 32 interconnected "nodes" (individual computers) and associated storage devices, hardware and software. The system will be developed over two years, with one cluster purchased in each of the first two years of the funding period. The proposed system was designed on the basis of an analysis of current and required computing capacity available to the PI and co-PIs.

The PI and co-PIs all have active research programs (including graduate and undergraduate students) that require a high performance parallel computing cluster. The proposed instrumentation will be used in diverse research projects, including: 1) Evolutionary studies in fungi based on DNA sequences (Hibbett, Biology Dept.); 2) Studies of liquid-solid phase changes (co-PI H. Gould, Physics Dept.); 3) Algorithmic analyses of geometric relationships between objects (co-PI L. Han, Mathematics and Computer Science Dept.); and 4) Protein modeling and molecular dynamics (co-PIs R. Bruschweiler and S. Huo, Chemistry Dept.). These research projects are relevant to robotics, conservation of biodiversity, medicine, and other applied disciplines. In addition to research, the PI and co-PIs all participate in the Concentration in Computational Science, which provides a research-oriented background in computational applications for Clark University undergraduates majoring in diverse scientific disciplines. The PI and Co-PIs of this proposal will use the proposed instrumentation in a number of undergraduate courses that satisfy the Concentration in Computational Science, including Computer Simulation Laboratory, Molecular Evolution and Systematics, Quantum Chemistry, Biomolecular NMR, and others. Finally, the proposed cluster will be used in annual workshops on parallel computing and its applications that will be offered for area high school teachers.

The development of a shared computational facility represents a significant interdepartmental initiative in the sciences at Clark. By pooling resources, the proposed project will develop critical resources for research that would be otherwise inaccessible to the individual Departments. In addition to enabling research, the proposed parallel computing cluster will become the cornerstone of the Concentration in Computational Science, which promotes interdisciplinary training for undergraduates in the sciences. The proposed instrumentation will also play a central role in summer workshops in parallel computing technology for local high school teachers. These workshops represent part of Clark University's ongoing community outreach efforts.

Calostoma is a morphologically and ecologically enigmatic genus of puffball-forming fungi. Its bizarre structure has historically left the determination of its relationship to other fungi in dispute. Calostoma's ecological role is also undefined - it has been described as either a litter decomposer or as symbiotic with plants. The purpose of this project is to study the evolutionary relationships and ecological role of Calostoma using a combination of DNA analysis and measurements of nitrogen and carbon isotopes. This study will compare the relationships and ecologies of Calostoma species where they occur in North America, Asia, Southeast Asia and Austrailasia. Fungi are vital to the health of ecosystems, functioning as recyclers of nutrients, or as symbionts of plants by supplying minerals in exchange for sugars from photosynthesis. Understanding the ecological roles of fungi like Calostoma will aid in assessing the health of the ecosystems where these fungi occur. Analyses of the DNA from Calostoma and related fungi will accurately address questions of fungal biodiversity and clarify evolutionary relationships of fungi. The results of this study will be presented at various scientific conferences, published in scientific journals and presented as a web
page for the Tree of Life Web Project.

The proposed research by Drs. Manfred Binder and David Hibbett concerns the evolutionary relationships of Boletales, a diverse and ecologically important clade of mushroom-forming fungi (homobasidiomycetes). Systematics and ecological studies have been hampered by the lack of a comprehensive phylogenetic classification for the group. The proposed research will use a multi-gene analysis for 52 key species of Boletales and selected outgroups to resolve the major groups of Boletales. Sequences of the internal transcribed spacer region (ITS) and the nuclear large subunit (nuc-lsu) of approximately 700 species sampled worldwide will be assembled into extensive datasets to generate a broad phylogenetic framework and to study character evolution.

This project will contribute to a global classification of the Boletales. The proposed research will provide resources for molecular ecologists and systematists, and will promote the discovery of new species of Boletales. A series of pages in the Tree of Life Web Project will be created for each of the major groups of Boletales to offer resources for a broad audience. In addition, this project will design and implement a four-week learning module on "Molecular Ecology and Systematics of Mycorrhizal fungi" for high school students, in collaboration with Minuteman Regional High School (Lexington, MA). This module will engage students in a discovery-based project that demonstrates the integration of molecular biology, organismal biology, and bioinformatics.

Nitrate transporters deliver nutritional nitrogen to (among other organisms) mushroom forming fungi living in symbiosis with tree roots, yet how nitrate transporter genes evolve and their relative significance to forest ecology is not well understood. This project investigates how nitrate transporters have evolved and diversified in function during a mushroom's adaptation to its ecological niche. Two specific goals of the project are (1) to use DNA and amino acid sequence analysis to develop an hypothesis of the evolution of nitrate transporter genes in the fungi and improve our understanding of fungal taxonomy in some groups and (2) to analyze the patterns of nitrate transporter gene utilization by mushrooms from divergent nitrogen niches under an array of environmental conditions.

This project addresses the importance of environmental nitrogen in the biodiversity and evolution of mushroom forming fungi with a specific focus on a group that forms symbiotic associations with trees. Nitrogen pollution is a broad ecological concern specifically implicated in the recent loss of diversity of these forest fungi. Such a loss threatens the resiliency and viability of the forest ecosystems that depend upon them directly. This project will provide a molecular framework for developing strategies of bioremediation using mushroom forming fungi to counter the impacts of nitrogen pollution.

This project will support a team of investigators who seek to resolve evolutionary relationships within the fungi. The research will enhance understanding of the ancient evolutionary diversification of the fungi by addressing evolutionary relationships among the major groups of fungi. This study will develop integrated molecular and morphological datasets, which will be freely accessible on the Web. Molecular data will include sequences from a target set of 76 genes that were identified from analyses of known fungal genomes. Morphological data will include characters associated with nuclear and cellular division and the morphological organization of hyphae. In addition the study will develop ontologies for morphological characters to promote further sampling and analytical integration across disparate organisms.

Fungi make up one of the major groups of life, with an estimated diversity of approximately 1.5 million species. These organisms play crucial ecological roles as decomposers, beneficial mutualists, and parasites and pathogens, including pathogens of humans. The economic significance of fungi is almost incalculable as they perform essential ecological functions and impact diverse applied disciplines, including agriculture, medicine, and drug discovery. A better understanding of the early evolutionary history of the Fungi is necessary to expand our knowledge of the history of life of Earth and the evolution of its ecosystems. Phylogenomic analyses and databases for fungi will transform the field of comparative fungal biology and will benefit all fields of fungal biology that rely on an accurate understanding of evolutionary relationships and diversity of Fungi. The bioinformatics tools for managing and analyzing phylogenomic data that will be developed will be broadly applicable across the Tree of Life. Training and outreach activities will include graduate and post-doctoral training, undergraduate training, and outreach to K-12 educators.

Fungi play important ecological roles and affect society through their activities as decayers, mutualists, and pathogens of plants and animals. Their enzymes are critical to biofuel production. Despite their importance, there is little known about how many fungi obtain food. Mycorrhizal fungi obtain sugars from plants in exchange for helping their host plants take up nutrients, and saprotrophic fungi obtain their energy and nutrients from decomposition of dead organic matter. Whether fungi switch between nutritional strategies is a key question in the evolutionary ecology of fungi. Recently, measuring carbon-12, carbon-13, and carbon-14 isotopes has been shown to reliably assess nutritional strategy, and therefore provides new opportunities to examine the evolution of nutritional strategies in fungi. This study will use archived fungi from Australia, Asia, and the US to conduct a broad survey of fungal nutritional strategies. A site-specific study at the Harvard Forest research site in Massachusetts will also be used to examine whether fungi can switch nutritional strategies. These studies will greatly expand knowledge on the links among fungal function, phylogeny, and ecology. The broader scientific impacts of the proposed research will result from integrating functional (isotopic) methods and genetic methods in a novel collaboration that should stimulate interest within the larger community of evolutionary biologists, ecologists, and mycologists. These methods could be readily expanded to Europe and Africa to include the global distribution of mushroom-producing fungi. This project will also incorporate current scientific research into specific educational activities designed to inspire future scientists including high school (fungal forays), college (Clark University's ACE Summer Institute, Research Experience for Undergraduates at Harvard Forest, the UNH Undergraduate Research Conference), and graduate students.

The Polyporales is a diverse group of fungi that includes "bracket fungi," crust-like forms, gilled mushrooms, and others. Recent years have seen much new information on how the major groups of polypores are related to each other, but genus-level classification remains poorly resolved, and it is likely that many species remain undiscovered. To understand the diversity of Polyporales, classical taxonomic approaches must be combined with morphology and modern molecular methods. Unfortunately, however, few mycologists have expertise in both areas, and many of the world's experts are retired or are nearing retirement. The goals of this project are: 1) To train the next generation of fungal taxonomists with expertise in morphological and molecular methods; 2) To produce comprehensive monographs of selected genera of Polyporales; 3) To create informatics resources for mycologists and the general public; 4) To promote development of an international community of Polyporales taxonomists; 5) To encourage participation in science by members of underrepresented groups; and, 6) To raise awareness about mycology among high school students and educators. The proposed research and training activities will be conducted by an international team of four "Principal Collaborators," and five "Trainees," including three PhD students and two early-career post-doctoral researchers.

As decayers and pathogens, Fungi have profound impacts on human affairs. They also function as beneficial symbionts, provide food, drugs, and other useful biochemicals, and serve as model systems for genetics and molecular biology. However, it is estimated that only 5-10% of the species of Fungi have been described. The Polyporales contains many wood-decay fungi, as well as timber pathogens. They play an important role in the carbon cycle, and produce diverse enzymes that break down lignin and cellulose. Monographic studies in Polyporales will improve understanding of biodiversity and will provide resources for applied research into biofuel production, bioremediation, and other "green" technologies.

The tree of life links all biodiversity through a shared evolutionary history. This project will produce the first online, comprehensive first-draft tree of all 1.8 million named species, accessible to both the public and scientific communities. Assembly of the tree will incorporate previously published results, with strong collaborations between computational and empirical biologists to develop, test and improve methods of data synthesis. This initial tree of life will not be static; instead, we will develop tools for scientists to update and revise the tree as new data come in. Early release of the tree and tools will motivate data sharing and facilitate ongoing synthesis of knowledge.

Biological research of all kinds, including studies of ecological health, environmental change, and human disease, increasingly depends on knowing how species are related to each other. Yet there is no single resource that unites knowledge of the tree of life. Instead, only small parts of the tree are individually available, generally as printed figures in journal articles. This project will provide the global community of scientists who study the tree of life with a means to share and combine their results, and will enable large-scale studies of Earth's biodiversity. It will also create a resource where students, educators and citizens can go to explore and learn about life's evolutionary history.

Fungi represent a major group of organisms that have profound economic and ecological impacts as decayers, pathogens (of plants and animals, including humans), and mutualists. Fungi are important sources of industrial bioproducts (antibiotics and enzymes) and they are critical to the fermentation industry, including biofuel production. However, much of the basic biology of fungi remains unknown, including knowledge on how fungi develop. This project will address the historical patterns and genetic mechanisms of developmental and morphological evolution in Agaricomycetes, which is a large group of fungi that includes gilled mushrooms, polypores, puffballs, and other complex forms. The results of this research will provide baseline information on the mechanisms of fungal development, as well as gene regulation and other processes of general significance for basic and applied fungal biology. This project will provide training for a PhD student, a Postdoctoral Fellow, and undergraduates. Education and outreach activities will make connections between art and science. Undergraduates with interests in art and science will engage in joint classes that develop observational skills and raise awareness about fungal biodiversity. The project will also create educational posters appropriate for high school students that illustrate fungal biology and general evolutionary principles. Posters and associated lesson plans will be made available for free download on a project web site and will be distributed through workshops at professional conferences for science educators.

Gene sequences for thousands of Agaricomycetes are available and complete genomes are available for over 90 species and increasing. However, these data have not been combined and queried to understand processes of fungal developmental evolution. The proposed research includes (1) macroevolutionary analyses, (2) phylogenomic analyses, and (3) developmental analyses. Macroevolutionary analyses will employ large-scale phylogenies (up to ca. 4000 terminals), drawing on publicly available data. Comparative methods will be used to perform ancestral state reconstruction, address directional trends, and test key innovation hypotheses. Phylogenomic analyses will identify shifts in copy number in gene families, such as transcription factors, hydrophobins, lectins and cell wall biogenesis enzymes, that may be correlated with evolution of complex forms. Developmental analyses will focus on Lentinus tigrinus, typically a gilled mushroom, but there is also a puffball-like "secotioid" form that is conferred by a recessive allele at a single locus (Sec). Two genome sequencs of L. tigrinus are available, one carrying the Sec+ allele and the other with the recessive sec allele. Bulk segregant analysis will be used to determine the identity of the secotioid locus. Pileus induction in L. tigrinus requires light and transcriptomic analyses will be used to compare gene expression patterns in vegetative mycelium and fruiting bodies produced in light or in darkness.

Wood (lignocellulose) is one of the most abundant carbon pools in terrestrial ecosystems. Fungal decomposition of wood is a critical component of the carbon cycle, impacts soil productivity, and has the potential to be exploited in the production of biofuels and other 'green' technologies. The major wood-decaying fungi are mushroom-forming (Agaricomycetes). The physical characteristics and chemical composition of wood vary among plant species, and most wood-decaying Agaricomycetes have characteristic wood substrate ranges. However, it is not understood why individual species of wood decaying fungi tend to occur on specific plant hosts. This project will investigate the mechanisms of substrate specificity and substrate switching in wood-decaying Agaricomycetes using a combination of analyses of genes expressed during decay, and physical and chemical characterization of the decay process. Fungal enzymes that are involved in wood decay or degradation have potential applications in emerging bioprocesses, such as energy-related bioconversions, including biofuel production, and bioremediation. Enhanced understanding of the mechanisms that allow different species of fungi to exploit particular wood substrates could help guide development of genetic resources that could be used for such applied purposes. This project will support one Postdoctoral Fellow and two graduate students, and will provide training to undergraduates at three collaborating academic institutions in the US. To bring information about fungi and wood decomposition to a wide audience, this project will create interactive exhibits and develop accompanying public programs at the Worcester EcoTarium (http://www.ecotarium.org/), a science and nature museum which serves over 140,000 visitors annually including large numbers of students from regional public schools. Exhibits will illustrate fungal diversity, the decay process, and the role of fungi in the carbon cycle.

The proposed research will focus on four species of Agaricomycetes (with distinct substrate preferences) with available genome sequences. The fungi will be cultured on solid wafers of four different tree species (two conifers and two hardwoods). Transcriptome profiling will be performed with the Illumina HiSeq 2000 platform, focusing on genes encoding enzymes known or suspected to be involved in wood decay, as well as co-expressed genes of unknown function. Phylogenomic analyses will make use of all available genomes of wood-decaying Agaricomycetes, focusing on gene families that are likely to play a role in decay, as informed by expression profiles. Physical and chemical characteristics of decay will be addressed using microscopy, mass spectrometry, and chemical analyses of colonized substrates. The proposed research is a collaboration between researchers with expertise in fungal systematics and molecular evolution, wood decay and forest products pathology, fungal genomics, and genetics and biochemistry of decay systems. This project will capitalize on recent advances in genomics of wood-decaying basidiomycetes. It will provide insight into the functioning and evolution of fungi in forest ecosystems, and create a framework for future studies aimed at understanding the specific plant-derived compounds that affect expression of particular fungal genes.

Most mushrooms are classified in the fungal group Agaricomycotina, comprising about 22,000 described species. Members of the Agaricomycotina have profound impacts on the functioning of ecosystems through their activities as decayers, pathogens, and beneficial symbionts. Researchers who study members of the Agaricomycotina include medical mycologists, plant pathologists, biofuels researchers, ecologists, and evolutionary biologists. However, such activities are often hampered by the lack of a coherent evolutionary and taxonomic framework for Agaricomycotina. In addition to professional scientists, many members of the public are interested in mushrooms, which they may collect for food, dyes, or as art objects. Amateur mycologists use field guides that are organized according to the physical forms of the mushrooms (e.g., gilled mushroom, polypores, coral fungi, etc.), but these often do not reflect evolutionary relationships. Consequently, amateur mycologists fail to appreciate the genealogical groupings of Agaricomycotina, or the rationale for taxonomic name changes when they occur. In this project, 25 years of work on the Agaricomycotina will be synthesized to produce two books on the subject. One will be a technical publication on "Systematics of Agaricomycotina" for professional scientists and graduate students. "Systematics of Agaricomycotina" will enable scientists from diverse backgrounds to gain an overview of the diversity and evolutionary relationships of mushroom-forming fungi, and promote comparative perspectives in basic and applied fungal science. The other book will be a popular text, titled "Mushrooms in the Tree of Life" for amateur mycologists. "Mushrooms in the Tree of Life" will show how evolutionary trees are estimated and help amateur mycologists make connections between field guides and the technical literature in fungal systematics.

This work will result in a comprehensive scientific summary text and a popular text for the public. The first, entitled "Systematics of Agaricomycotina", will include five chapters on general topics, including (1) History of phylogenetics of Agaricomycotina; (2) Overview of phylogeny and diversity of Agaricomycotina; (3) Character bases of systematics of Agaricomycotina; (4) Morphological and developmental evolution in Agaricomycotina; and (5) Ecological diversity and evolution of nutritional modes in Agaricomycotina; as well as eleven chapters (6-16) describing individual groups of Agaricomycotina. The popular text, "Mushrooms in the Tree of Life," will be organized in three parts: (1) "Reconstructing the tree of life", which will contain a concise primer of fungal phylogenetics, with minimal jargon; (2) "Mushrooms in the tree of life", which will present a higher-level phylogenetic framework for mushroom-forming fungi, emphasizing groups that are familiar to amateur mycologists; and (3) "Stories from the tree of life", which will contain a series of evolutionary vignettes about mushrooms, illustrating historical narratives that can only be addressed with a phylogenetic perspective.