Source: MICHIGAN STATE UNIV submitted to NRP
FLORISTICS OF MICHIGAN AND THE ROLE OF FLOWER FORM AND FUNCTION
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
0180833
Grant No.
(N/A)
Cumulative Award Amt.
(N/A)
Proposal No.
(N/A)
Multistate No.
(N/A)
Project Start Date
Apr 1, 2009
Project End Date
Mar 31, 2014
Grant Year
(N/A)
Program Code
[(N/A)]- (N/A)
Recipient Organization
MICHIGAN STATE UNIV
(N/A)
EAST LANSING,MI 48824
Performing Department
Plant Biology
Non Technical Summary
The importance of plant collecting in cataloguing biodiversity cannot be understated: specimens are the very basis of floristic and plant taxonomic science, and as such provide the foundation of plant nomenclature, the basis for identification, the common reference for communication, the vouchers for floras, and the tools for teaching. All fields of biological science from systematics and evolution to ecology and even physiology and molecular biology are ultimately dependent on collections, not just for application of names, but as the basis for referencing all aspects of biodiversity. The fact that the U.S. flora is not completely known has been well-documented, if not widely appreciated. Given the high rate of habitat destruction and other threats to native species, it is of utmost urgency to fully catalogue our flora. Non-native species now constitute a considerable fraction of our flora. Non-native organisms can cause tremendous economic losses and are the second largest threat to biodiversity loss in the U.S. (after habitat destruction) for plants as well as for all species. After preventing introductions, the best strategy for reducing this threat is early detection and elimination so it is obvious that we must continually monitor even the most botanically well-known regions in the U.S. Documenting extinction has particular relevance today because 5090 species, or one third of the total number of plant species in the U.S., are of conservation concern, i.e. at risk of extinction. The number of plant species determined to be at risk is an order of magnitude larger than that of any other group of organisms. The best evidence for extinction is clear documentation (i.e. specimens) of the former existence of a species that is no longer present, based on extensive and well-documented field searches. Expansion and contraction of distributions are the phenomena that contribute the most to the dynamic nature of our flora. In modern times, range contraction is pervasive and is largely driven by habitat destruction, although range contractions and expansions are also natural processes. Without excellent data (i.e. specimens) documenting plant occurrences and continued collecting over time, we will lack sound information on distributional changes. Without studies concerning how and why distributions change, we will never be able to build good predictive models on how our flora will change in the future. Evolutionary biology is the organizing principle at all levels of life as well as the unifying concept among all fields of biology. Phylogeny reconstruction, the estimation of an evolutionary tree, is central to evolutionary biology and provides the foundation for many types of biological inquiry. The Mint (Lamiaceae) and Phlox (Polemoniaceae) families provide excellent opportunities for study in this regard because the floral morphology is notoriously labile and the phylogenetic history of these families have been shown to be relatively tractable, though not without some difficulties.
Animal Health Component
(N/A)
Research Effort Categories
Basic
(N/A)
Applied
(N/A)
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
1362420107020%
1362499107020%
2062420107020%
2062499107040%
Goals / Objectives
GOALS: This project will focus on understanding the Flora of Michigan and the patterns of evolution of flora form that contribute to the reproduction and persistence of plants. This information well help us understand how and why components of the flora changes and the role of floral morphology in those changes. The specific foci of this project are to: 1) Better understand the flora of Michigan a) Collect baseline data of the flora of Michigan b) Make existing data more readily available to those who need it c) Collect and use empirical data to learn how the flora is changing 2) Better understand evolutionary patterns in floral form a) Investigate the phylogeny of select groups in the Mint and Phlox families b) Develop systematically appropriate, useable taxonomies for these groups c) Document floral lability and the evolution of floral form in these groups
Project Methods
To better understand the flora of Michigan, standard methods will be used. Specimens will be collected at select localities at Michigan and will be deposited in the MSU Herbarium. These specimens will be preserved for posterity by standard archival preparation and curation. The data from these specimens and from existing specimens will be added to the MSU Herbarium database. This database has been established using SPECIFY software which is continuously developed with funds from the U.S. National Science Foundation. The database is hosted on a server owned by the MSU Herbarium and operated in the Plant Biology computing facility. Portions of this database are currently being served publicly. Currently the lichen collection is publicly available. In the near future the vascular plants should be made publicly available, though only about one quarter of the collection has been included in the database. Data entry is ongoing. We have initiated a project at Baker Woodlot where we will collect new data on species presence and absence, validated by new plant collections, to compare to historical studies to learn how the flora is changing. To better understand evolutionary patterns in floral form we will estimate phylogenies standard phylogenetic techniques will be used, as described below. Markers. Sequences of low-copy nuclear genes, nrITS, and cpDNA regions will be used for phylogenetic estimation. Because of the widespread applicability of ITS, as well as its demonstrated utility in related plants, it will be used as a marker in many of these studies. Low-copy nuclear genes can provide more variation than ITS in cases where resolution from ITS is poor. Because low-copy nuclear genes are less susceptible to concerted evolution, it is possible to clone each homolog from each parent. These data should provide the history of the parental lineage of each allopolyploid species. DNA extraction, PCR amplification, cloning and Sequencing. Total DNA will be extracted from fresh or silica dried material. DNA extraction, purification, amplification, cloning and sequencing will be performed according to standard protocols. Modified ITS primers will be used to amplify the ITS1, 5.8S, and ITS2 regions from nuclear ribosomal DNA. Previously designed Adh primers are also being used. Phylogenetic Analyses. Edited sequences will be aligned manually. Gaps will be treated as missing data in one analysis, and scored using the simple indel coding procedure. Parsimony will be implemented using PAUP*. Support for monophyletic groups will be assessed using 10,000 bootstrap replications with 100 addition sequence replicates per bootstrap replicate. Interpretation of Lability. Recently developed methods to compare levels of homoplasy will be employed, as well as standard measures of homoplasy and character mapping.

Progress 04/01/09 to 03/31/14

Outputs
Target Audience: General Public: We have floristic databases of our non-vascular and vascular plants. This is an area of rapid growth over the term of the project, as well as in the near future. Currently all of our specimen data are served online via the Michigan State University Herbarium Database portal (http://herbarium.lib.msu.edu/). The non-vascular database is also accessible via the Global Biodiversity Information Facility (http://www.gbif.org/) and has more recently been made available via the Consortium of North American Lichen Herbaria portal (http://lichenportal.org/portal/) and the Integrated Digitized Biocollections portal (https://www.idigbio.org/portal). Soon the vascular plant database will also be available via the Global Biodiversity Information Facility and Integrated Digitized Biocollections portals. Collectively these portals serve data from hundreds of millions of records and are the most important source of biodiversity data available to the general public, the scientific community, and those agencies and organizations that focus on resource management. We have also focused on the general public in a number of biodiversity presentations to lay audiences including groups such as the Red Cedar Chapter of the Michigan Botanical Club, the East Lansing Chapter of the Wild Ones, Alumni groups on international travel tours from various universities (but always including Michigan State University), and the Merry Mates of the First United Methodist Church in Birmingham, MI. The local area public was the focus of our participation in the Michigan State University Science Festival by hosting a public open house for over 160 visitors. MSU Science Festival is a free, public, educational outreach event aimed at families living in Central Michigan. Stations were arranged around the herbarium by 3 graduate and 5 undergraduate students to discuss plant preservation methods, invasive species, tropical plants, lichen, and poisonous plants. Visitors were able to mount their own plant to take home, observe lichen under microscopes, and view many ecologically important specimens from our collection, with the focus of the exhibits as a whole focusing on biodiversity. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? Three graduate students and several undergraduate students have been invovled in the research peformed as part of this project. They have all received training in research and also learned about data entry and data management in publicly available databases. The investigator organized, coordinated, or hosted six workshops focused on issues realated to data associated with biological specimens and how to best make those data availble over the course of htis project. He particapted in nine additional workshops. Two of the graduate students invovled have received their degrees and the third is continuing. Each of them attended workshops and scientific conferences related to this project. How have the results been disseminated to communities of interest? The results have been disseminated through a combination of talks to lay audiences (7 over the reporting period), posters or talks at scientific meetings (5), participation or organization of workshops for scientists or educators (15), non-peer reviewed reports to agencies or groups (6), peer-reviewed papers in scientific journals (3), and public dissemation of data via online resources. What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? As part of our efforts to better understand the flora of Michigan, we made great strides in two specific areas. First, we received several small grants to make the data on plant biodiversity in Michigan more accessible to those who might benefit from it. These helped us to make 22,272 additional specimen records from the Michigan State University Herbarium available online through multiple collaborative portals. Access to these data makes us a leader in herbarium digitization. We also made high resolution images of our entire collection of type specimens, over 2300, and made them available through the JSTOR Plants website. These efforts will continue into the future. Second, we have a better understanding of how the flora of Michigan is changing because of a targeted study of baseline species changes to a natural area, Baker Woodlot, on the Michigan State University campus. This multi-year floristic project was begun on the natural area, a 70+ acre parcel that was the subject of a botanical survey about 40 years ago. The primary goal of this study is to detect changes in the flora of this parcel and determine the extent of change during this interval. Some potentially invasive species, such as Garlic Mustard, have established populations on the site. Many species that were present 30 years ago have not been relocated, suggesting that they may have been locally extirpated. Many new native species have been recorded, suggesting that migration onto this site from nearby areas is ongoing. Overall, the total number of species at the site increased by more than 20% and the proportion of non-native species is about 30%. This is comparable to the proportion of non-native plants in the state as a whole. Interestingly, the proportion of native to non-native species has not changed significantly over time because the number of native and non-native taxa have both increased over the time interval. This is contrary to the pattern found in other studies across eastern deciduous forests in the U.S. The efforts of my research group to better understand the evolution of floral morphology have resulted in several significant findings. A phylogeny ofCantuawas estimated using parsimony analyses of DNA sequences from the nuclear ribosomal ITS region and the cpDNA spacerstrnT-tmL, tmL-trnF,and partialndhFcoding region to better understand the phylogeny of this group, including patters of floral evolution. This result, in combination with our earlier work, suggests that floral morphology is highly variable within this group and is associated with a diversity of pollination syndromes. Closely related species are often very different in terms of flower shape and size. One of the two species formerly included inHuthiawas included in these analyses and is shown to derived within Cantua. This work was performed in collaboration with Anna Monfils. Poliomintha(Lamiaceae) is a small genus of shrubs from the southwestern United States and northern Mexico. The circumscription of this group, and its close relatives, has been confounding for decades because, in part, high levels of homoplasy in floral morphology. Sequence data from the nuclear ribosomal ITS and two plastid regions,trnL-trnL-trnF and rpL32-trnL, were used to objectively test morphologically-based hypotheses of relationship withinPoliominthaand among other closely related genera.Based on these analyses, Poliominthaactually comprises at least two lineages and so is not a tenable genus, taxonomically. This work comprises the thesis of former student Grant Godden. Monarda subgenus Cheilyctis (Lamiaceae) is a group of 17 morphological complex species that has been shown to be monophyletic based on prior research from my lab. An intensive study of floral morphology of this group has shown that there are 11 distinct clusters that correspond to previously accepted species ( M. austromontana, M. citriodora, M. clinopodioides, M. fruticulosa, M. humilis, M. maritima, M. mexicana, M. pectinata, M. punctata, M. stanfieldii, and M. viridissima ) and five clusters that represent new species or combinations, illustrating that morphology can be useful in circumscribing taxa when highly variable DNA sequence regions have not been identified. This work constitutes the dissertation of former student Nathan Sammons. Our studies of the evolution of floral morphology received a major boost when we recieved a National Science Foundation grant to study how polyploidy affects floral morphology in three species of Phlox. We completed three field trips to collect material, collaborators in the project have used flow cytometry to determine the ploidy level of these samples, and we collected a massive amount of material so that we can document any differences in floral mophology among the ploidy levels. In the first analyses performed we have found a significant association between over half the examined characters (mostly floral) and ploidy in one of the three species, Phlox amabilis.

Publications

  • Type: Journal Articles Status: Awaiting Publication Year Published: 2014 Citation: Powers, K. E., L. A. Prather, J. A. Cook, J. Woolley, H. L. Bart, Jr., A. K. Monfils, and P. Sierwald. In Press. Revolutionizing the Use of Natural History Collections in Education. The Science Education Review 13(2).
  • Type: Theses/Dissertations Status: Published Year Published: 2009 Citation: Godden, Grant T. 2009. Phylogenetic relationships in Poliomintha and related genera in the Mentheae (Lamiaceae). Thesis (MS), Department of Plant Biology, Michigan State University.
  • Type: Theses/Dissertations Status: Published Year Published: 2011 Citation: Sammons, Nathan J. 2011. Morphometric analysis and monograph of Monarda subgenus Cheilyctis (Lamiaceae). Dissertation (PhD), Department of Plant Biology, Michigan State University.
  • Type: Conference Papers and Presentations Status: Published Year Published: 2013 Citation: Waselkov, K., B. Wright, L. A. Prather, S. D. Fehlberg and C. J. Ferguson. Polyploidy and diversity in the genus Phlox: development of polyploid assemblages as study systems for phylogenetic and phylogeographic work. 2013 Ecological Genomics Symposium, Kansas City, MO, 1-3 November 2013. (poster)
  • Type: Journal Articles Status: Published Year Published: 2010 Citation: Monfils, A.K. and L. A. Prather. 2010. Phylogeny of Cantua (Polemoniaceae): Evidence from Chloroplast and Nuclear DNA Sequence Data. Systematic Botany 35: 877-884.
  • Type: Book Chapters Status: Published Year Published: 2009 Citation: Prather, L. A. 2009. Cobaea. In: Flora Mesoamericana, Vol. 4. G. Davidse, M. Sousa S., S. Knapp, and F. Chiang, eds. Universidad Nacional Autonoma de Mexico, Missouri Botanical Garden, and The Natural History Museum (London), publishers.
  • Type: Other Status: Published Year Published: 2009 Citation: Bart, H. and L. A. Prather. 2009. CollectionsWeb, A Natural History Collections Community Building Project. In Society for the Preservation of Natural History Collections Newsletter. Quarterly. Newsletter for a professional organization of collections managers and preservationists.


Progress 01/01/13 to 09/30/13

Outputs
Target Audience: During this period my efforts were focused on the general public and serving other researchers. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? A graduate student, Matt Chansler, has joined the project to study Polyploidy in Phlox and we have employed 12 undergraduates in the MSU Herbarium that are studying aspects of museum curation and plant biology. How have the results been disseminated to communities of interest? As mentioned above we have added significantly to our public database of the flora of Michigan. We have also submitted to publications for review. What do you plan to do during the next reporting period to accomplish the goals? 1) We will continue our efforts to document floristic change on Baker Woodlot. 2) We will continue to add specimens to the database of the Flora of Michigan. 2) The study of the effect of polyploidy on floral morphology will continue with additional field trips, more collecting and generation of data.

Impacts
What was accomplished under these goals? During this reporting period a number of major accomplishments were achieved. First, our efforts to document the changes in the flora of Baker Woodlot continue and we made major progress this year. A number of rare species were relocated and we identified several potentially invasive species in the survey area. Many of these were intentionally planted, helping us identify a major source of threat to our native flora. We also are continuing our efforts to build a smartphone app to help identify Great Lakes Wildflowers. Our efforts at adding to our databse of the Michigan Flora increase 17-fold over last year, and these data are all available to the public. Second, our studies of the evolution of floral morphology received a major boost when we recieved a National Science Foundation grant to study how polyploidy affects floral morphology in three species of Phlox. We completed two field trips to collect materal, collaborators in the project have used flow cytometry to determine the ploidy level of these samples, and we collected a massive amount of material so that we can document any differences in floral mophology among the ploidy leves. these efforts are underway.

Publications

  • Type: Conference Papers and Presentations Status: Other Year Published: 2013 Citation: Waselkov, K., B. Wright, L. A. Prather, S. D. Fehlberg and C. J. Ferguson. Polyploidy and diversity in the genus Phlox: development of polyploid assemblages as study systems for phylogenetic and phylogeographic work. 2013 Ecological Genomics Symposium, Kansas City, MO, 1-3 November 2013. (poster)


Progress 01/01/12 to 12/31/12

Outputs
OUTPUTS: WORKSHOPS AND SYMPOSIA (2): Participant and speaker ("Major Initiatives in Specimen Digitization"), Small Collections Herbarium Workshop, Hanes Foundation supported workshop, Central Michigan University, Mt. Pleasant, MI. Participant, Global Plants Initiative, Mellon Foundation supported workshop at the Real Jardin Botanico de Madrid, Spain. PRESENTATIONS FOR LAY AUDIENCES: Botanicals in Captivity: How Plants Get From the Wild to Our Gardens. Shipboard presentation to MSU Alumni on the MSU Alumni Expedition for Life Along the Waterways of Holland and Belgium; Herbaria, Plant Collecting, and Our Dynamic Flora, meeting of the Red Cedar Chapter of the Michigan Botanical Club, Lansing, MI. Two graduate and one undergraduate students have continued the project to record the plant species present at Baker Woodlot on the MSU Campus and compare species that grow there today to those that did 30 years ago. So far this project has generated about 700 new specimens that have been deposited in the MSU Herbarium. MSU HERBARIUM DATABASE: The Michigan State University Herbarium operates an online database of specimen data for specimens from the state of Michigan and elsewhere. We have added over 1,100 specimens (to date) in this calendar year. The primary means of dissemination of the Michigan Flora is the MSU Herbarium website, at herbarium.msu.edu. We continued the grant (Digitization of the MSU Herbarium Type Collection) from the Global Plants Initiative project funded by the Arthur J. Mellon Foundation to capture high-resolution images of all the type collections held in herbaria world-wide, and make these and the accompanying collection data available on-line. The MSU Herbarium has been participating in this project and has imaged all of its 1934 type collections (1656 Vascular Plants, 278 Lichens). To date, 613 of our type images and associated data are available online via the GPI JStor web site (http://plants.jstor.org/partner/MSC), with a further 489 currently being processed by JStor, and the remainder being prepared for submission in the near future. We are also involved in floristic work on lichens and bryophytes that are associated with climate change via the "Lichens, Bryophytes and Climate Change (LBCC)" Thematic Collections Networks project funded under the NSF's Advancing Digitization of Biological Collections (ADBC) program. The aim of this project is to computerize the label data from all North American lichen and bryophyte collections held in US herbaria. As part of this project we are imaging the label data of all our North American bryophytes (the label data of our lichens having already been digitized as part of a previous NSF Award) and uploading these to the LBCC server in Wisconsin to be digitized by optical character recognition. To date 6400 labels have been imaged, of which 1000 have been uploaded to the LBCC server and 100 are available on-line (http://bryophyteportal.org/portal/collections/misc/collprofiles.php collid=16). An undergraduate student prepared a treatment for the genus Pogogyne outlining the diversity of this plant genus for the Flora of North America (a series of books, see FNA.org) that will be submitted soon. PARTICIPANTS: Former student Nathan Sammons contributed to work on a manuscript submitted for publication describing a new species of the genus Monarda. This manuscript is in review. Former student Orlando Alvarez-Fuentes contributed to the monographic outcomes (Thelypteridaceae) and a manuscript in preparation. Former undergraduate student Elizabeth Straley co-wrote the Pogogyne treatment for the Flora of North America. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
The multi-year floristic study of Baker Woodlot, a 70+ acre parcel on the MSU campus, is ongoing. The primary goal of this study is to detect changes in the flora of this parcel and determine the extent of change during this 30 year interval. Over 70 species are recorded as new since the survey 30 years ago, many of them invasive species. This suggests that invasive species can move into protected areas very rapidly. About 30 species that were present at the time of the previous survey have not yet been found during the present survey. Many of these species have likely been extirpated from the site suggesting that species can easily be lost, even from protected areas, however in the last year we have found about 15 of these "missing" plant suggesting that more collecting work remains. This research, along with discovery of new species, additional data on species distributions, new phylogenetic work on plants, and elaboration of plants in North America add significantly to our understanding of the plant biodiversity of Earth.

Publications

  • No publications reported this period


Progress 01/01/11 to 12/31/11

Outputs
OUTPUTS: WORKSHOPS AND SYMPOSIA (5): Organizer and co-host, CollectionsWeb Workshop IV: The Digitization Challenge: Biology and Engineering. NSF-supported workshop in Norman, OK). Organizer and co-host, CollectionsWeb Workshop IV: New Approaches to Specimen-based Education. NSF-supported workshop in Radford, VA. Moderator, Plenary Session on Collections and Sustainability, Annual Meeting of the Society for the Preservation of Natural History Collections, San Francisco, CA. Participant, Scientific Software Institute in Biological Collections Digitization. NSF Cyberinfrastructure-supported workshop (BIOSYNC, Field Museum of Natural History, Chicago, IL). Participant, Integrated Digitized Biocollections (iDigBio) Summit, NSF supported workshop at the Florida Museum of Natural History, University of Florida, Gainesville, FL. STUDENTS (5) During 2011 I mentored (advised) one PhD student, as well as four undergraduate students. One PhD student completed his degree: Nathan Sammons, "Morphometric analysis and monograph of Monarda subgenus Cheilyctis (Lamiaceae)". PRESENTATIONS FOR LAY AUDIENCES: Plant Materials Identification, meeting of USDA Plant Materials Researchers (Rose Lake Research Station) at the MSU Herbarium. Two undergraduate students have continued the project to record the plant species present at Baker Woodlot on the MSU Campus and compare species that grow there today to those that did 30 years ago. So far this project has generated about 600 new specimens that have been deposited in the MSU Herbarium. MSU HERBARIUM DATABASE: The Michigan State University Herbarium operates an online database of specimen data for specimens from the state of Michigan and elsewhere. We have added 378 specimens (to date) in this calendar year. For the first part of the year we added to this database as part of a projected funded by a grant from the Kalamazoo, MI-based Hanes Foundation. In spring 2011, we acquired a grant from the Mellon Foundation (Digitization of the MSU Herbarium Type Collection) that will fund digitization (both imaging and text entry) of our substantial and important type collection. As part of this project we have digitized 1754 type specimens. DISSEMINATION: The primary means of dissemination for the symposia and workshop above is on the website, CollectionsWeb.org that is the primary site for the NSF-funded Research Coordination Network (Prather, PI) that has funded much of the work above. The primary means of dissemination of the Michigan Flora is the MSU Herbarium website, at herbarium.msu.edu. The type specimens imaged as part of the Mellon Project will be released on the JSTOR Plants Resource pages online, with the first sets to be released in 2012. Much dissemination also happened at the workshops reported above. PARTICIPANTS: Former student Nathan Sammons contributed to work on a manuscript submitted for publication describing a new species of the genus Monarda. Former student Orlando Alvarez-Fuentes contributed to the monographic outcomes (Thelypteridaceae) and a manuscript in preparation. Former student Anna Monfils contributed to the phylogenetic work (Cantua) that was published in very late 2010 and is first reported this year. Henry Bart is co-PI on the NSF RCN, CollectionsWeb. He and the rest of the steering committee of CollectionsWeb (James Woolley of Texas A & M University, Karen Francl of Radford University, Meredith Blackwell of Louisiana State University, Michael Mares of the University of Oklahoma, and Joe Cook of the University of New Mexico) helped organize the two CollectionsWeb workshops mentioned above. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
The multi-year floristic project continues on Baker Woodlot, a 70+ acre parcel on the Michigan State University campus. This area was first subject of a botanical survey about 30 years ago. The primary goal of this study is to detect changes in the flora of this parcel and determine the extent of change during this 30 year interval. So far over 65 species are recorded as new since the survey 30 years ago, many of them invasive species. This suggests that invasive species can move into protected areas very rapidly. Almost 45 species that were present at the time of the previous survey have not yet been found during the present survey. Many of these species have likely been extirpated from the site suggesting that species can easily be lost, even from protected areas. Phylogenetic and monographic studies of flowering plants continue. A report on the phylogeny of the genus Cantua (Polemoniaceae or the Phlox Family) was published (with former student Anna Monfils). A dissertation on the genus Monarda was completed by graduate student Nathan Sammons and a manuscript based in part on that dissertation was submitted.

Publications

  • Monfils, A.K. and L. A. Prather. 2010. Phylogeny of Cantua (Polemoniaceae): Evidence from Chloroplast and Nuclear DNA Sequence Data. Systematic Botany 35: 877-884.


Progress 01/01/10 to 12/31/10

Outputs
OUTPUTS: WORKSHOPS (5): Participant, Community Round-Table Meeting on Building Community Support for an NSF-funded HUB for Advancing Digitization of Biological Collections (University of Colorado, Boulder, CO). Participant, Biological Collections Digitization Focus Meeting #2, NSF-supported workshop at the National Evolutionary Synthesis Center (Durham, NC). Co-organizer, CollectionsWeb Workshop III: New Research Opportunities Emerging from Integrating Data Across Different Taxonomic Collections. NSF-funded workshop (New Orleans, LA). Participant, United States Virtual Herbarium Workshop. NSF-Supported workshop (St. Louis, MO). Participant, SciColl: International Coordination of an Interdisciplinary Global Research Infrastructure, Organized under the auspices of the OECD Global Science Forum and hosted by the Royal Belgian Institute of Natural Sciences, Brussels. STUDENTS (6) During 2010 I mentored (advised) two PhD students, as well as four undergraduate students. One PhD student completed his degree. PRESENTATIONS FOR LAY AUDIENCES (2): "Top ten unexpected things you can do with herbarium specimens: the role of the Herbarium specimens in research at MSU." September 2010 Meeting of the Red Cedar Chapter of the Michigan Botanical Club. "A Field Trip in Winter at the MSU Herbarium." February 2010 Meeting of the East Lansing Chapter of The Wild Ones", East Lansing, MI. One graduate student and one undergraduate have continued the project to record the plant species present at Baker Woodlot on the MSU Campus and compare species that grow there today to those that did 30 years ago. So far this project has generated about 500 new specimens that have been deposited in the MSU Herbarium. MSU HERBARIUM DATABASE: Under my direction, the Michigan State University Herbarium has released a database of specimen data for specimens from the state of Michigan. We have added 5403 (to date) specimens in this calendar year. We are actively adding to this database by paying student employees that are funded by a grant from the Kalamazoo, MI-based Hanes Foundation. DISSEMINATION: The primary means of dissemination for the symposia and workshop above is on the website, CollectionsWeb.org that is the primary site for the NSF-funded Research Coordination Network (Prather, PI) that has funded much of the work above. The primary means of dissemination of the Michigan Flora is the MSU Herbarium website, at herbarium.msu.edu. Much dissemation also happened at the five workshops reported above and the two presentations to lay audiences. PARTICIPANTS: Former student Orlando Alvarez Fuentes contributed to the monographic outcomes (Thelypteridaceae). Former student Anna Monfils contributed to the phylogenetic work (Cantua) that will be published in very late 2010. Henry Bart is PI on the NSF funded RCN, CollectionsWeb. He and the rest of the steering committee of CollectionsWeb (James Woolley, Karen Francl, Meredith Blackwell, and Michael Schauff) helped organize CollectionsWeb Workshop III. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
The multi-year floristic project continues on Baker Woodlot, a 70+ acre parcel on the Michigan State University campus. This area was first subject of a botanical survey about 30 years ago. The primary goal of this study is to detect changes in the flora of this parcel and determine the extent of change during this 30 year interval. So far over 45 species are recorded as new since the survey 30 years ago, most of them invasive species. This suggests that invasive species can move into protected areas very rapidly. Almost 75 species that were present at the time of the previous survey have not yet been found during the present survey. Many of these species have likely been extirpated from the site suggesting that species can easily be lost, even from protected areas. Phylogenetic and monographic studies of flowering plants continue. A manuscript on the phylogeny of the genus Cantua (Polemoniaceae or the Phlox Family) will be published (with former student Anna Monfils) in the last issue of Systematic Botany in (very) late 2010. A thesis on a members of the genus Thelypteridaceae by Dr. Orlando Alvarez-Fuentes was submitted to MSU. This dissertation found that the widespread genus Thelypteris is non-monophyletic and should be split into at least 5 genera. Several new species were also described in the dissertation.

Publications

  • Prather, L. A. 2009. Cobaea. In: Flora Mesoamericana, Vol. 4. G. Davidse, M. Sousa S., S. Knapp, and F. Chiang, eds. Universidad Nacional Autonoma de Mexico, Missouri Botanical Garden, and The Natural History Museum (London), publishers.


Progress 01/01/09 to 12/31/09

Outputs
OUTPUTS: SYMPOSIA (2) Symposium co-organizer, Biodiversity Informatics, Zach Murrell and L. A. Prather. Annual Meeting of the Association of Southeastern Biologists (Birmingham, AL). Symposium speaker, CollectionsWeb: Building a Community of Natural History Collections. L. A. Prather. Annual Meeting of the Association of Southeastern Biologists (Birmingham, AL). WORKSHOP (1) Workshop host and co-organizer, CollectionsWeb Workshop II: Enhancing the Interface between Collections and Systematists. J. Woolley, L. Page, and L. Alan Prather. NSF-supported workshop (Fairbanks, AK). STUDENTS (4) During 2009 I mentored (advised) two PhD students and two masters students, as well as three undergraduate researchers. My two masters students both received their degrees in 2009. PRESENTATIONS FOR LAY AUDIENCES (2) "The Changing Face of Natural History Collections" and "Flower Pollination in the Phlox Family." Shipboard presentations to MSU Alumni on the Alumni Association Expedition to the Amazon River, Peru. PLANT COLLECTIONS: One graduate student and one undergraduate student began a project to record the plant species present at Baker Woodlot on the MSU Campus, and compare species grow there today compared to when the most recent census was taken, almost 30 years ago. So far this project has generated several hundred new specimens that have been deposited in the MSU Herbarium. MSU HERBARIUM DATABASE: Under my direction, the Michigan State University Herbarium has released a database of specimen data for specimens from the state of Michigan. We estimate that we have 130,000 specimens of plants from Michigan, and the database has 30,391 individual records so far. We are actively adding to this database by paying student employees that are funded by a grant from the Kalamazoo, MI-based Hanes Foundation. DISSEMINATION: The primary means of dissemination for the symposia and workshop above is on the website, CollectionsWeb.org that is the primary site for the NSF-funded Research Coordination Network (Prather, PI) that has funded much of the work above. The primary means of dissemination of the Michigan Flora is the MSU Herbarium website, at herbarium.msu.edu. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
A multi-year floristic project was begun on Baker Woodlot, a 70+ acre parcel on the Michigan State University campus. This area was first subject of a botanical survey about 30 years ago. The primary goal of this study is to detect changes in the flora of this parcel and determine the extent of change during this 30 year interval. Based on the results so far, it clear that some potentially invasive species have established populations on the site. Some species that were present 30 years ago have not been relocated, suggesting that they may have been locally extirpated. This work is a part of a broader research focus on the flora of Michigan and how it is changing. My research team is also continuing to help establish baseline data for the flora for the entire state and have entered thousands of new records into the MSU Herbarium database. Phylogenetic and monographic studies of flowering plants continue. A manuscript on the phylogeny of the genus Cantua (Polemoniaceae or the Phlox Family) has been accepted for publication and will be published in 2010. A thesis on the tribe Mentheae in the Lamiaceae (or Mentheae) by Mr. Grant Godden of my lab was submitted to MSU. This thesis demonstrated that many genera in the Lamiaceae (Clinopodium, Hedeoma, Hesperozygis, Poliomintha, etc.) are non-monophyletic and that much more phylogenetic and taxonomic work in the Lamiaceae needs to be done.

Publications

  • Bart, H. and L. A. Prather. 2009. CollectionsWeb, A Natural History Collections Community Building Project. In Society for the Preservation of Natural History Collections Newsletter. Quarterly. Newsletter for a professional organization of collections managers and preservationists.


Progress 01/01/08 to 12/31/08

Outputs
OUTPUTS: During 2008 I made several oral reports on my research, including being the moderator and organizer for a policy breakout session at the National Conference on Science and the Environment conference (2008 theme: Biodiversity in a Rapidly Changing World). During this breakout session on biodiversity and museums we enumerated eight policy recommendations that became part of a document that was discussed and passed onto members of the Obama transition team. I also hosted a workshop at Michigan State University that developed a strategy for small biodiversity collections to play a larger role in conservation, science and education endeavors. I attended a workshop at the National Museum of Natural History (Smithsonian Institution) to help guide the National Ecological Observatory Network staff in developing a plan to achieve their biological collections-related activities. I hosted a symposium at the Society for the Preservation of Natural History Collections and Natural Science Collections Alliance joint meeting on CollectionsWeb (see below). Additionally, I made several presentations at scientific meetings about my research. I mentored four graduate students during this time (two Masters and one PhD). I remain the PI (Hank Bart, Tulane, co-PI) on a new NSF funded research coordination network, and as such I have primary responsibility for the associated website (www.collectionsweb.org). This project will coordinate a community effort to enhance collections-related research in all areas of biology. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Phylogenetic and monographic studies of flowering plants continue. Two manuscripts were published based on this work. A manuscript on the taxonomy and floral biology of a new species of Pedicularis was published is now in press and will be published in 2008, in collaboration with a former PhD student, Anna Monfils. An extensive examination was undertaken to detect and explain patterns of infraspecific variation among populations of Pedicularis densiflora, including populations formerly recognized as P. densiflora subsp. aurantiaca. Several statistical analyses (UPGMA, ANOVA, and PCA) were conducted on floral traits measured in the field. All three statistical analyses indicated consistent differences in floral morphology among populations. These differences were confirmed by a review of over 1000 herbarium specimens. Two series of populations could easily be separated, and two species are now recognized, P. densiflora and P. aurantiaca. Pedicularis aurantiaca had large calyces with floral tubes included at anthesis, reduced lower labia, and enlarged galea openings. Pedicularis densiflora had short calyces with fully exserted floral tubes, enlarged lower labia, and smaller galea openings. The taxonomy, phenology and floral differences are discussed in the context of previously published data on pollinator visitation. This study helps develop a system that can be used to study the potential role of pollination biology in speciation. The second publication, in collaboration with J. Mark Porter and Rancho Santa Ana Botanic Garden describing morphological variation in the genus Cantua, describing a new species, and providing correct taxonomic names for two other species was published in the journal Aliso. This paper focuses on how floral morphology varies among these rare plants. A manuscript on the phylogeny of the genus Cantua (Polemoniaceae) is in manuscript. This manuscript will be submitted in 2009 and will present information from three chloroplast and one nuclear DNA analysis to show how the species of this florally diverse genus are related. This manuscript is also from work completed in collaboration with Dr. Monfils.

Publications

  • Porter, J. M and Prather, L. A.. (2008). Cantua dendritica (Polemoniaceae), a new species from Peru and two new Cantua names, Aliso 21: 31-35. Monfils, A. K. and Prather, L. A. (2008). Pedicularis aurantiaca and Pedicularis densiflora (Orobanchaceae): Taxonomy, Phenology, and Floral Morphological Variation, Madrono 54: 306-321.


Progress 01/01/07 to 12/31/07

Outputs
During 2007 I made several oral reports on my research, including a keynote address ("The Future of Specimen-based Research: Anticipation, Preparation and Participation") at the Fourth Annual Great Plains Plant Systematics Symposium in Manhattan, KS, a plenary address ("Building a Community of Natural History Collections") at the Natural Science Collections Alliance meeting in Washington, DC, and two oral papers ("Building a Community of Natural History Collections" at the American Society of Plant Taxonomists curators meeting in Chicago, IL and a presentation by the same title at the Society for the Preservation of Natural History Collections meeting in St. Paul, MN). Four graduate students (three PhD and one MS) students and one undergraduate were also mentored during this time and three of them are expected to graduate in 2008). I am the PI (Hank Bart, Tulane, co-PI) on a new NSF funded research coordination network, and as such I have primary responsibility for the associated website (www.collectionsweb.org). This project will coordinate a community effort to enhance collections-related research in all areas of biology.

Impacts
Phylogenetic and monographic studies of flowering plants continue. A manuscript on the taxonomy and floral biology of a new species of Pedicularis is now in press and will be published in 2008, in collaboration with a former PhD student, Anna Monfils. An extensive examination was undertaken to detect and explain patterns of infraspecific variation among populations of Pedicularis densiflora, including populations formerly recognized as P. densiflora subsp. aurantiaca. Several statistical analyses (UPGMA, ANOVA, and PCA) were conducted on floral traits measured in the field. All three statistical analyses indicated consistent differences in floral morphology among populations. These differences were confirmed by a review of over 1000 herbarium specimens. Two series of populations could easily be separated, and two species are now recognized, P. densiflora and P. aurantiaca. Pedicularis aurantiaca had large calyces with floral tubes included at anthesis, reduced lower labia, and enlarged galea openings. Pedicularis densiflora had short calyces with fully exserted floral tubes, enlarged lower labia, and smaller galea openings. The taxonomy, phenology and floral differences are discussed in the context of previously published data on pollinator visitation. This study helps develop a system that can be used to study the potential role of pollination biology in speciation. A manuscript on the phylogeny of the genus Cantua (Polemoniaceae) is in manuscript. This manuscript will be submitted in 2008 and will present information from three chloroplast and one nuclear DNA analysis to show how the species of this florally diverse genus are related. This manuscript is also from work completed in collaboration with Dr. Monfils.

Publications

  • No publications reported this period


Progress 01/01/06 to 12/31/06

Outputs
Collaborative work with Jeffrey Conner (Kellogg Biological Station) and others on genetic constraints in floral evolution continues. Genetic correlations are perhaps the most commonly studied of all potential constraints on adaptive evolution and tests are needed to understand whether genetic correlations, in fact, cause evolutionary constraints. Our data from wild radish shows that correlations do not necessarily constrain evolution. The genetic correlation between the filament and corolla tube in wild radish flowers is one of the highest ever reported in nature, is caused by pleiotropy, and is stable across environments, populations, and related species. Thus, this correlation should cause a constraint. On the contrary, artificial selection produced rapid independent evolution of these traits. This study integrates artificial selection studies (by Conner and colleagues) with phylogenetic analyses of the evolutionary pattern in character correlation. My lab contributed a phylogenetic analysis of a number of species of the Mustard Family based on the chloroplast gene ndhF. This well-resolved phylogeny illustrated that the high genetic correlation between filament length and corolla length is evolutionarily very stable. We plan to submit this work for publication in early 2007. Our work on invasive species in Michigan has shown that species are more likely to enter Michigan in the southeast part of the state than any other, and that rates of spread vary considerably across taxa. These data were presented by an undergraduate in my lab at an invasive species symposium held at Michigan State University in Spring 2006. Phylogenetic and monographic studies of the tropical members of the Polemoniaceae are continuing.

Impacts
The collaborative research on the genetic basis of floral correlations will provide the foundation needed to understand how the species can quickly evolve in new habitats and maintain high levels of reproduction which contributes to its invasive nature. The herbarium-based studies of plant invasions may help us predict the rate and direction of spread of future invasions.

Publications

  • No publications reported this period


Progress 01/01/05 to 12/31/05

Outputs
In collaboration with Jeffrey Conner (Kellogg Biological Station) and James Hancock (Horticulture), we are studying the rapid evolutionary divergence in anther exsertion in wild radish, a weed, using QTL mapping on two crosses between artificial selection lines. Three sets of crosses were performed between individuals from the high and low exsertion lines. We screened over 400 microsatellite primer pairs developed for Brassica and found about 50 that amplify well and were polymorphic in our population of wild radish. We have scored 12 of these in 516 F2 from one of the two outbred crosses. We are also developing additional microsatellite loci using the Refseth et al. method (1997); in our first set of 12 AG enriched clones, we identified and sequenced 11 new microsatellite loci, most or all of which are variable in our population. We have also screened our four parents from one of the outbred crosses for AFLP loci using two pairs of selective amplification primers. We found 5 and 10 highly repeatable and fully informative loci from these two primer pairs respectively. Therefore, by screening additional selective primer pairs we will be able to find ample numbers of informative AFLP for our future work. My lab has begun to explore the history of invasive plant species in Michigan using herbarium records. We are in the process of acquiring data from thirteen of the most invasive plant species in Michigan habitats to try to learn if there are any commonalities in plant invasions in the state. So far we have learned that more early records of invasive plants originated from the southeastern portion of the state. We are in the process of determining if this is a statistically significant result if reporting bias is taken into account. We may also be able to learn if there are commonalities in the pattern of spread. Much progress has been made toward my studies of the evolution of pollination biology and floral morphology. A monographic treatment of Cantua is nearly complete. My own South American collections of tropical Polemoniaceae collected as a part of this study have added signficantly to the specimens available for study. These studies have uncovered four previously undescribed taxa in Cantua. My lab has generated both nuclear and chloroplast DNA sequence data to resolve species level relationships in the Cobaeoideae and to resolve the phylogenetic position of Acanthogilia according to the following breakdown: 51 partial ndhF sequences, 32 cp trnT-trnL spacer sequences, 48 trnL-trnF sequences, and 50 nrITS sequences. There is some evidence for hybridization in Cantua based on incongruence between the chloroplast and nuclear phylogenies. This evidence is in agreement with morphological evidence generated by the monographic studies. Phylogenetic analyses of the cpDNA sequences confirm paralellism in extreme floral morphologies that the ITS phylogeny had suggested. Scanning electron micrographs of the pollen grains of all species of the Cobaeoideae have been taken at the MSU Center for Electron Optics.

Impacts
The collaborative research on the genetic basis of floral correlations will provide the foundation needed to understand how the species can quickly evolve in new habitats and maintain high levels of reproduction which contributes to its invasive nature. The herbarium-based studies of plant invasions may help us predict the rate and direction of spread of future invasions.

Publications

  • No publications reported this period


Progress 01/01/04 to 12/31/04

Outputs
In collaboration with Jeffrey Conner (Kellogg Biological Station) and James Hancock (Horticulture), I am studying the rapid evolutionary divergence in anther exsertion in wild radish, a weed, using QTL mapping on two crosses between artificial selection lines. This research on the genetic mechanisms of adaptation and integration among floral traits in a weed should provide a better understanding of the genetic mechanisms underlying rapid evolution and the generalized pollination phenotype in weedy plants. We have screened all 64 primer combinations on the parent tissues in an effort to find markers suitable for genetic mapping. Of these 64 combinations, 36 amplified well. From these 36 primer combinations that amplified well, we identified 25 bands that are present in the correct patterns, which translates to 69% of the primer combinations yielding useable bands. My lab is continuing research on floral biology of rare plants in Michigan. In 2004, we published two papers that question the baseline data needed to understand rarity across the entire contiguous United States. These papers focused on understanding the rate of scientific collecting, which obviously and directly affects our ability to determine which plants are rare. We found that across the United States, and here in Michigan, plant collecting is in decline. In order to have a full understanding of rarity, especially patterns through time, and to understand invasiveness of plants, we need a well-documented flora, which is not currently being achieved. This has broad implications in land management, management of endangered species, invasive species biology, and other aspects of biology such as climate change. Research on the phylogenetics of the tropical Polemoniaceae is continuing. We have now published a study of pollen morphology that shows that rates of pollen morphological evolution are not consistent, even among close relatives. We expect that in the coming year we will submit for publication a molecular phylogeny of Cantua based on nuclear and chloroplast genetic sequence data.

Impacts
Three of the eighteen known populations of the endangered plant, Michigan monkeyflower (Mimulus michiganensis) have been extirpated. Most others are in danger of extirpation. Our work has shown that most plants are completely sterile and that reproduction is therefore a critical factor in the rarity of the species. This research will direct future research on recovering fertility in the species, which may ultimately lead to restoration efforts. The collaborative research on the genetic basis of floral correlations will provide the foundation needed to understand how the species can quickly evolve in new habitats and maintain high levels of reproduction which contributes to its invasive nature.

Publications

  • Prather,L.A., Alvarez-Fuentes,O., Mayfield,M.H., Ferguson,C.J. 2004. The decline of plant collecting in the United States: A threat to the infrastructure of biodiversity studies. Systematic Botany 29:15-28.
  • Prather,L.A., Alvarez-Fuentes,O., Mayfield,M.H., Ferguson,C.J. 2004. Commentary: Implications of the decline in plant collecting for systematic and floristic research. Systematic Botany 29:216-220.
  • Monfils,A.K., Prather,L.A. 2004. The conserved nature and taxonomic utility of pollen grain morphology in Cantua. Grana 43:249-256.


Progress 01/01/03 to 12/31/03

Outputs
In collaboration with Jeffrey Conner (Kellogg Biological Station) and James Hancock (Horticulture), I am studying the rapid evolutionary divergence in anther exsertion in wild radish, a weed, using QTL mapping on two crosses between artificial selection lines. This research on the genetic mechanisms of adaptation and integration among floral traits in a weed should provide a better understanding of the genetic mechanisms underlying rapid evolution and the generalized pollination phenotype in weedy plants. During the past year we have made the first round of crosses and begun generating the f2 crosses. Screening microsatellites and AFLP makers to generate the genetic map is continuing. My lab is continuing research on Michigan monkeyflower, the only endangered plant that is found only within the state of Michigan. We have found that most plants are female sterile, which correlated with male sterility. Research on the phylogenetic of the tropical Polemoniaceae is continuing and we have now sequenced both nuclear and chloroplast DNA regions from the twelve species of Cantua. We have also found that the pollen morphology of Cantua is relatively conservative, unlike other floral characters which are highly labile. This work will be published soon.

Impacts
Three of the eighteen known populations of the endangered plant, Michigan monkeyflower (Mimulus michiganensis) have been extirpated. Most others are in danger of extirpation. Our work has shown that most plants are completely sterile and that reproduction is therefore a critical factor in the rarity of the species. This research will direct future research on recovering fertility in the species, which may ultimately lead to restoration efforts. The collaborative research on the genetic basis of floral correlations will provide the foundation needed to understand how the species can quickly evolve in new habitats and maintain high levels of reproduction which contributes to its invasive nature.

Publications

  • No publications reported this period


Progress 01/01/02 to 12/31/02

Outputs
In collaboration with Jeffrey Conner (Kellogg Biological Station) and James Hancock (Horticulture), I am studying the rapid evolutionary divergence in anther exsertion in wild radish, a weed, using QTL mapping on two crosses between artificial selection lines. This research on the genetic mechanisms of adaptation and integration among floral traits in a weed should provide a better understanding of the genetic mechanisms underlying rapid evolution and the generalized pollination phenotype in weedy plants. During the past year, we have characterized microsatellites and AFLP markers that will be used to construct a genetic map of wild radish, which will provide the foundation for the QTL study. This basic knowledge should ultimately be valuable in designing more effective weed control strategies and will likely provide an example of how basic research can contribute to applied research. My lab is continuing research on Michigan monkeyflower, the only endangered plant that is found only within the state of Michigan. We have completed a study of genetic diversity among populations using AFLP markers. A surprising finding is that the population with the greatest genetic diversity is on Beaver Island. Our study will provide much useful information regarding management of the remaining populations of this species. The phylogenetic analysis of Monarda was published in 2002, and illustrates the floral morphology is evolutionarily labile. Phylogenetic and taxonomic work in Cobaea continues. During the past year we have sequenced both nuclear and chloroplast markers from all species. This work is yielding valuable data regarding the evolution of floral morphology and pollination systems.

Impacts
Three of the eighteen known populations of the endangered plant, Michigan monkeyflower (Mimulus michiganensis) have been extirpated. Most others are in danger of extirpation. Our work on the genetic diversity among populations suggest that each populations harbors genetic diversity, and many of the populations harbor unique genetic markers. Our new understanding of the distribution of genetic diversity will help land stewards prioritize conservation efforts and may make possible reintroductions, as specified in the recovery plan for this species.

Publications

  • Prather,L.A., Keith,J.A. 2003. Monarda humilis (Lamiaceae), a new combination for a species from New Mexico, and a key to the species of section Cheilyctis. Novon 13:104-109 .
  • Posto,A.L., Prather,L.A. 2003. The evolutionary and taxonomic implications of RAPD data on the genetic relationships of Mimulus michiganensis (comb. et stat. nov.: Scrophulariaceae). Systematic Botany 28:xx-xx.


Progress 01/01/01 to 12/31/01

Outputs
A new phase of my project was begun this year with the initiation of research on the genetic mechanisms of adaptation and integration among floral traits in a weed. In collaboration with Jeffrey Conner (Kellogg Biological Station) and James Hancock (Horticulture), I am studying the rapid evolutionary divergence in anther exsertion in wild radish, a weed, using QTL mapping on two crosses between artificial selection lines. The results should provide a better understanding of the genetic mechanisms underlying rapid evolution and the generalized pollination phenotype in weeds. This basic knowledge should ultimately be valuable in designing more effective weed control strategies and will likely provide an example of how basic research can contribute to applied research. We also made two major advances in our studies of Michigan biodiversity. First, we completed a checklist of all the lichens in Michigan, which is the first such list every published for the state. Second, we completed data that provides support for the elevation of Michigan monkeyflower to species from variety. This has major implications in conservation of this endangered plant. Michigan monkeyflower is the only endangered plant that is found only within the state of Michigan. We studied genetic relationships of Michigan monkeyflower and two related species, James' monkeyflower and common monkeyflower, that have been implicated in its origin. We used random amplified polymorphic DNA (RAPD) markers to calculate genetic similarities and to construct a UPGMA phenogram. All samples of Michigan monkeyflower were highly similar to each other and formed a distinct cluster in the phenogram. This cluster was nested among the samples of James' monkeyflower, and the resulting cluster was, in turn, nested within the samples of common monkeyflower. This year my lab also began a major study of the phylogenetics and evolution of floral morphology in the Phlox family, and work on the Mint family continues. These studies will yield valuable data on how and why flower morphology changes among taxa.

Impacts
The elevation of Michigan monkeyflower from variety to species will have a major impact on the levels of protection awarded to this endangered species, and will clarify our understanding of biodiversity in Michigan species of monkeyflower. Michigan monkeyflower is the only plant species that occurs nowhere outside of the state of Michigan.

Publications

  • Fryday, A. M. Fair, J. B. Googe, M. S. Johnson, A. J. Bunting, E. A. Prather, L. A. 2001. Checklist of lichen-forming, lichenicolous, and allied fungi recorded from Michigan. Contributions of the University of Michigan Herbarium 23:145-223.
  • Sahagun-Godinez, E. Lomeli-Sencion, J. A. Prather, L. A. 2001. Habitat and range extension of Cobaea lutea (Polemoniaceae) in western Mexico. Acta Botanica Mexicana 54:29-37.
  • Prather, L. A. Monfils, A. K. Posto, A. L. Williams R. A. 2002. Monophyly and phylogeny of Monarda (Lamiaceae): Implications of sequence data from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA. Systematic Botany (In Press).


Progress 01/01/00 to 12/31/00

Outputs
In the past year, I completed the phylogenetic study of all 16 spp. of Monarda based on sequences of the internal transcribed spacer regions of nuclear ribosomal DNA. Thymus and Mentha were used as outgroups, and Blephilia, Clinopodium, Conradina, Hesperozygis, Monardella, Pycnanthemum, and Ziziphora were included in the ingroup to test the monophyly of Monarda. Three parsimony searches were performed: one on the complete data set with insertion/deletion events treated as binary characters, and two using different insertion/deletion events on a data set with redundant sequences removed. All searches yielded congruent results. There was strong support for the monophyly of Monarda and a close relationship was found between Monarda, Blephilia, and Pycnanthemum, with weak support for the latter as sister group to Monarda. The molecular phylogeny was completely congruent with the infrageneric classification of the genus. Our results were consistent with hypotheses of hybridization between M. fistulosa and M. lindheimeri in Texas. Despite considerable morphological variation among many species, especially in floral characters, little molecular diversification was found in those same species groups. Intraspecific polymorphism in ITS sequence was found in over half the species examined. These data and results have been submitted for publication. We have also acquired sequence data from several related genera in the Clinopodioid complex, which includes several important ornamental and herb plants, in a study of the relationships of this economically important group. We have further initiated phylogenetic studies of these taxa using low-copy nuclear genes, which may provide more resolution of relationships. In our study of Michigan monkey-flower, we investigated the genetic structure of 51 individuals from 16 populations and found genetic variation within 14 of the 16 populations studied and found 45 different multilocus genotypes in 51 individuals. We tested three hypotheses for the taxonomic origin and our evidence supports an ancient hybrid origin of Michigan monkey-flower from James' monkey-flower and Common monkey-flower. The Reese's Swamp population, which had never before been studied, has 0% viable pollen, similar to all other populations but one. There is considerable variation (27-52%) in pollen viability levels between individuals of the Maple River population, which has implications for mating, selection, and gene flow within the Maple River population. Among four seed germination regimes tested, we found best results at 25C in light. Michigan monkey-flower is a self-compatible plant taxon. Plants from the Maple River population are capable of self-pollination, and regularly set self fruits in the greenhouse. These studies are continuing.

Impacts
In the last year my lab has contributed substantially to our knowledge of the relationships of the Lamiaceae, especially the of the Clinopodioid group which includes many cultivated plants such as sage, salvia, bee-balm, Oswego tea, and pennyroyal. We have also learned that there is a substantial amount of genetic variation in the rare, endangered plant, Michigan monkeyflower. This is contrary to expectations because most populations of the plant are thought to be asexual.

Publications

  • Prather, L.A. Ferguson C.J. Jansen, R.K. 2000. Polemoniaceae phylogeny and classification: Implications of sequence data from the chloroplast gene ndhF. American Journal of Botany 87:1300-1308.


Progress 01/01/99 to 12/31/99

Outputs
My focus in the last year has been on phylogenetic aspects of this project. The evolution of floral morphology and pollination systems is a major component of this project. There are several components spelled out in the project description and I will address some below. Phylogeny of the genus Monarda as a framework for studying morphological evolution and the evolution of pollination systems: In the past year my lab has sequenced the ITS region of nuclear ribosomal DNA repeat regions to determine the relationships among nearly all species of Monarda (bee-balm and horsemint) as well as several related genera. This work is progressing well. We now know that the 'fistulosa' group is monophyletic and is the sister group to the rest of the genus. We are working on completing the phylogeny and studying character evolution in floral traits of the species. Use of Cobaea as a model system to understand the evolutionary lability of floral morphology: I published an 81 pp. monograph in 1999 that carefully describes the floral forms of all the species of Cobaea (cup-and-saucer vine) and summarized what we know about floral diversity in the group. Based on morphological data, I published a second paper in 1999 that shows that floral morphology is not more evolutionarily labile that vegetative morphology, even though there are numerous parallelisms in flower structure (see next). This paper develops a new technique for comparing levels of evolutionary lability between different subsets of data. Furthermore, a former my lab has accumulated new phylogenetic data from sequences of three chloroplast regions (two spacers and one gene) of all the species of Cobaea. This evidence suggests that very remarkable floral forms in Cobaea have arisen more than one time, i. e. are evolutionary parallelisms. This is a substantial finding, and we hope to publish these results in 2000. Investigation of three hypotheses proposed for the origin of Michigan monkey-flower with genetic analysis of its putative parental species: In collaboration with one of my graduate students, we have obtained preliminary evidence that supports one of the three models over the remaining two. Michigan monkeyflower has genetic markers in common with M. glabratus var. jamesii, suggesting it shares a common ancestor with that subspecies. This data suggests that the Common monkeyflower, M. guttatus, is not a potential parent species as has been suggested by other workers. We also now know that there is more than one genetic individual in some populations of the Michigan monkeyflower, which is a rare clonal species.

Impacts
Information provided by phylogenetic studies of mints, including several ornamentals and crops (e.g. bee-balm, sage, thyme), can help direct breeding programs and increases predictability when searching among species for compounds of interest. The work on the federally endangered Michigan monkeyflower, the only plant whose distribution is limited to Michigan, is of critical conservation value.

Publications

  • Prather, L. A. 1999. The relative lability of floral vs non-floral characters and a morphological phylogenetic analysis of Cobaea (Polemoniaceae). Botanical Journal of the Linnean Society. 131: 433-450.
  • Prather, L. A. 1999. Systematics of Cobaea (Polemoniaceae). Systematic Botany Monographs 57: 1-81.