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OVERVIEW OF THE UNIVERSITY OF SOUTH HERBARIUM by Alan R. Franck, June 2018 (http://florida.plantatlas.usf.edu/documents/USF-Herbarium.pdf)

Figure 1. George Cooley with the USF Herbarium where it originated at Chinsegut Hill, ca. 1958–1959 (courtesy of The Tampa Tribune).

TABLE OF CONTENTS History of the USF Herbarium (1956–2018) ...... 1 George Ralph Cooley ...... 3 James Davis Ray, Jr...... 7 Robert William Long ...... 8 Olga Korhoven Lakela ...... 8 Maurice Mulvania ...... 10 Margaret Lois Gilbert ...... 10 Photographic Tour of the Herbarium ...... 13 Additional Historic USF Herbarium Photos...... 24 References ...... 30 Overview of the Collections ...... 32 Online Activity ...... 35 Filing System ...... 36 Personnel ...... 37 History & Roles of Herbaria (ca. 40,000 BC–present)...... 39 Specimen Protocols ...... 45 Collecting ...... 44 Mounting ...... 48 Modified Archer’s Glue Formula ...... 50 Digitization ...... 50 Morphology Photos (for Atlas) ...... 56 Plant/Fungal DNA Extraction & PCR ...... 57 Selected Publications Utilizing the USF Herbarium Collections...... 60 Appendices ...... 76

Table 1. Summary of University of South Florida Herbarium. ca. 300,000 No. of specimens

Taxa 93% vascular , 5% , 1% bryophytes, 1% fungi

Geography 42% Florida, 21% SE USA, 14% continental USA + Canada, 21% Latin America, 8%

Index herbariorum USF acronym

Mailing Address 4202 E. Fowler Avenue, ISA 2015, University of South Florida, Tampa, FL 33620, USA

Street Address 12010 USF Cherry Drive

Campus Building Code CMC 9–11 & Rooms

Website http://florida.plantatlas.usf.edu/

Phone 1(813)974-7602

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HISTORY of the USF HERBARIUM (1956–2016) The University of South Florida (USF) Herbarium was planned with the founding of USF in 1956, in cooperation with John S. Allen, the first president of USF. It originated at the Chinsegut Hill manor house in February 1958 under the patronage of George R. Cooley, an investment banker from New York and a personal friend of John Allen. The first specimen officially accessioned into the herbarium was a specimen of the Florida endemic Calydorea caelestina collected by John Allen in 1951. In 1959 USF hired the first director of the herbarium, James D. Ray, Jr., who was the first faculty member at USF and later became Dean of the College of Natural Sciences. The herbarium moved to “rooms 1029 and 1030” of the Administration Building (formerly ADM, now the John & Grace Allen Building [ALN]) at the Tampa campus in August 1960. At the time of its move to the Tampa campus, it contained 19,432 specimens, built up largely by the efforts of George Cooley. In the fall of 1960 during the opening convocation of USF, George Cooley was presented USF’s first degree, an Honorary Doctoral Degree.

Olga K. Lakela was hired as curator in 1960. Upon completion of the Life Sciences Building (LIF), the herbarium moved there in Sep 1961, eventually occupying rooms 170–179. Robert W. Long became curator in 1962 and was promoted to director of the herbarium in 1965. Clinton Dawes arrived to USF in 1964 and was nearly solely responsible for amassing the algal collection. The cultivated and Neotropical collections greatly benefited from the first director of the USF Botanical Gardens, Derek Burch, hired in 1969. Mosses of the Tampa Bay Area was published by Diane Wagner-Merner [Diane Te Strake] et al. in 1970. A Flora of Tropical Florida, a descriptive flora of Collier, Miami-Dade, and Monroe Counties written by Robert Long and Olga Lakela, was published in 1971 and shortly thereafter the herbarium accessioned its 100,000th specimen. Olga Lakela received an Honorary Doctoral Degree from USF upon her retirement in 1973.

Richard P. Wunderlin was hired as curator in 1973 and became director in 1976 after Robert Long’s death. In 1976, Ferns of Florida by Olga Lakela and Robert Long and Plants of the Tampa Bay Area by Olga Lakela, Robert Long, et al. were also published. Guide to the Vascular Plants of Central Florida, written by Richard Wunderlin, covered a 30-county region of peninsular Florida and was published in 1982. In the fall of 1989, Alice F. Tryon and Rolla M. Tryon began a several year long stay at USF, studying ferns and generously supporting departmental lectures.

The Atlas of Florida Vascular Plants was launched in 1995 by Richard Wunderlin, Bruce F. Hansen (curator from 1979–2014), and Edwin L. Bridges (of the FTG herbarium at the time) with technical assistance from Shawn Landry (a USF masters student at the time who later became part of the USF faculty). It was briefly distributed as a CD-ROM in 1996. The Atlas provided county- level distribution data and other details for all vascular plants native or naturalized in Florida. In 1996, a National Science Foundation Collections Improvement grant for nearly $138,000 was awarded. With matching funds from USF, this permitted the renovation and expansion of the herbarium, including the installation of compactors.

The first state-wide guide specific to the native and naturalized flora of Florida, Guide to the Vascular Plants of Florida, written by Richard Wunderlin, was published in 1998. Both the Guide

2 and the Atlas rely mainly upon specimen records from the four major Florida herbaria (FLAS, FSU, FTG, and USF). The Atlas interface is licensed by USF Patents and Licensing, with licenses having been sold to other institutions (e.g. to cover the floras of , Kentucky, Indiana, New York, and ).

In 2003, digitization of the herbarium’s collections was initiated as a component of the Atlas, providing images and searchable data of the specimens. Bruce Hansen, co-authored the second edition of the Guide to the Vascular Plants of Florida published in 2006 and the third edition published in 2011. The Seaweeds of Florida was released in 2008 by Clinton Dawes and Arthur Mathieson, covering nearly 700 of macroalgae. After residing in LIF for over 50 years and accruing about 276,000 specimens, the herbarium was moved to the College of Arts & Sciences Multidisciplinary Complex (CMC, formerly the Physics and Mathematics building [PHY]) in Nov 2014. In 2016 the online website was changed to the Atlas of Florida Plants upon inclusion of bryophytes in 2016.

Figure 2. Olga Lakela and Provost Carl Riggs holding the 1st specimen accessioned (Calydorea caelestina) into the herbarium, collected by USF President John Allen (taken in 1960s in LIF, photographer unknown).

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GEORGE RALPH COOLEY (1896–1986) As a long-time friend of USF’s first president John S. Allen, George R. Cooley had a central role in the initial development and continued support of the USF Herbarium. George collected many specimens himself for the herbarium and procured thousands of specimens from numerous institutions and individuals. He financed a large part of the expenses of the herbarium, including salary, equipment, and collecting expeditions throughout the 1950s–1980s. He also was responsible for the acquisition of countless important books in the herbarium library (the George R. Cooley Library), many rare and out-of-print.

Figure 4. Opening convocation for USF on 26 Sep 1960 with John Allen on the left and George Cooley front-right receiving USF’s first degree, an Honorary Doctoral Degree, in front of the ALN [formerly ADM] building (courtesy of The Tampa Tribune).

George and his wife Myra were very close friends with John and Grace Allen. The majority of George’s archives at the New York State Library consist of about 100 correspondences with John and Grace, from the 1940s–1980s. The earliest letters are dated in 1948, when John left New York to become vice president at the , along with J. Hillis Miller who became their

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new president. The final letters in the archives were from Grace just before John’s passing. John sought George’s advice and support extensively, especially concerning botany and other sciences at USF, as well as the solicitation of donations and scholarships for USF.

George had a passionate vision for botany at USF, writing “I entered into the planning with the suggestion Figure 5. The offices of George Cooley resided at 90 State Street (left) that the new State and 100 State Street (right) in Albany, NY. Photos by Alan Franck, 19 University [USF] acquire Dec 2014. all possible knowledge of the vegetation of the state of Florida to the point where it would be known throughout the world as the authority,” (letter to James Keenan, 21 June 1967). To then curator Olga Lakela, 18 September 1967, he further stated “It is still my hope that the University of South Florida will become the acknowledged depositary of all knowledge of the vegetation of that interesting state so that scientists, teachers, students and the public throughout the State, throughout the nation, and throughout the world will expect the University of South Florida to know everything about its vegetation and to be right about it in every detail.”

George was born in 1896 in Troy, NY. His father passed away when he was 8½ and George started a factory job three miles from town making $8 a week, five of which he gave his mother. In September 1917, with $52, he later went to Colgate University with the purpose of becoming “the greatest Baptist minister.” After six months he ran out of money and joined the army in March 1918, becoming First Lieutenant. After his time in the army, George Figure 6. The Chinsegut Hill manor house, where the USF entered the bond business. He Herbarium originated. It was kept on the E side, 1st floor, against the wall, facing the E windows of the main house here in front. worked for James Forrestal for Photo (facing N) by Alan Franck, 17 Jan 2016. six years as his best salesman. In 1924 he started his own

5 investment banking business, George R. Cooley & Co., Inc., in Albany, NY and made a small fortune. “My only claim to credit is that I’ve worked hard … I’ve had no love for money. I’ve enjoyed earning it, saving it.” George became very fond of Colgate University and was an important benefactor of the university, where he became acquainted with John Allen, an astronomer at the time.

George soon developed an interest in learning the local plants, stating “I thought, ‘what an interesting hobby for an old man.’” George and Myra bought land around Hickory Hill, Rennsselaerville where George would often do some botanical work. Some of his earliest herbarium specimens date from 1945 at Hickory Hill. While John Allen was at UF, George financed a plant collecting expedition to for Erdman West and Lillian Arnold in 1951– 1952 and also collected himself there with George R. Proctor around the same time. After retiring in 1953 and handing the business over to his son-in-law Daniel McNamee, George became d strong interest in contributing to a modern flora of the southeastern USA. George began collaborations with botanists at Harvard in 1954 as a research fellow. In 1955, George published “The vegetation of Sanibel Island, Lee County, Florida” in Rhodora based predominantly upon his own collections from the area during stays in the winter and spring of 1952–1954. In 1955, George made extensive collections in Alabama, Arkansas, Louisiana, and accompanied by L.J. Brass, D. Demaree, or A.S. Pease. That same year his interest in the southeastern flora likely brought him into contact with James D. Ray, Jr. at Mississippi State College. In January 1957, George collected plants in Sri Lanka. In November 1957, George and Myra began a stay at Chinsegut Hill, to serve “as a kind of headquarters for his southeastern project” to make progress on a successor to John K. Small’s Manual of the Southeastern Flora (1933). This stay marks the very beginnings of the USF Herbarium. Presumably the bulk of the “Cooley Herbarium” was delivered to Chinsegut Hill around February 1958, as this was the given date for the start of the USF Herbarium by Robert Long (1972). John and Grace Allen visited the Cooleys often at Chinsegut Hill during the winter and spring months of the late 1950s. The herbarium curator Olga Lakela and director James Ray also spent much time there. Chinsegut Hill was the type locality for the endemic Campanula robinsiae, collected by John K. Small and Raymond Robins, ca. 1924. In 1958, USF took over management of Chinsegut Hill after the University of Florida’s lease expired.

George was a trustee officer for the University of South Florida Foundation from 1958–1964. As part of USF’s opening convocation on September 26, 1960, John Allen presented the Doctor of Science to George Cooley, hooded by Dean Sidney French, recognizing his efforts to “enlarge our knowledge of native plants and to help establish a pattern for research at the university.” This was the first degree awarded by USF. In a letter to George, 20 June 1961, John Allen wrote “Through your vision and help, the University of South Florida has gotten off to a wonderful start in plant .” George’s diligent plant collecting continued as he traveled the world, including trips to the West Indies (Barbados, Dominica, St. Kitts, St. Lucia, St. Vincent, 1962), Africa (including Basutoland, Kenya [with Richard A. Howard], South Africa, 1963), Australia with R.A. Howard (1963), New Zealand (1963), Jamaica with George R. Proctor (1965), Guadeloupe (1967), and St. Vincent with R.A. Howard (1971). As a recognized benefactor of Colgate University, Colgate conferred an honorary degree to him in 1969.

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Figure 7. Collecting trip in the Everglades, Oct 1962, showing left to right: Olga Lakela, Frank Craighead, James Ray, Myra Cooley, and George Cooley (and Robert Long the likely photographer).

Figure 8. Collecting trip in the Everglades, Oct 1962, showing left to right: Frank Craighead, Richard Jefferson Eaton, Myra Cooley, George Cooley, and James Ray (and Robert Long the likely photographer).

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On 29 June 1970, John Allen made George Cooley and Olga Lakela honorary curators of the USF Herbarium, stating “Each of you have contributed so much to the sum total of our knowledge of the native plants of Florida [...] that you have made the University of South Florida Herbarium known throughout the botanical fraternity, and you have brought kudos to the University of South Florida.” His support of the herbarium remained crucial to its activities in the 1970s and 1980s and he continued to make visits when Richard Wunderlin was director of the herbarium.

On 16 September 1976, James Ray wrote to George, “My heart goes out to you and Mrs. Cooley for all that you’ve done – for me – Bob [Robert] Long, our families, so many others and the many in situations which through your generosity and feeling have been given a chance.” In 1981, George was given the Class of ’56 Award by USF. Andy Rooney, who had also once attended Colgate, wrote an open letter to George on 15 August 1985, “I don’t know anyone who has done more of everything after he was 65 than you. You’re a person who could give rich people a good name.” USF President John Lott Brown, 18 December 1985, wrote to him “we [the University] will never forget our old friends such as you.” From his daughter, Robin Krivanek stated “he loved people and he loved telling stories about them.”

George was a philanthropist and supporter of innumerable persons, institutions, and programs, including USF, Colgate University, the Nature Conservancy (e.g. Tiger Creek Preserve), Harvard University herbaria, Herbarium Plumier at the Muséum National d’Histoire Naturelle, Carnegie Museum, Dept. of Botany at the University of North Carolina, King’s Park and Botanic Garden, the Botanical Society of South Africa, Swarthmore College (George R. Cooley Curator of the Peace Collection), World Policy Institute, and the University of Montana. He was a member of several professional botanical and biological societies. He donated hundreds of thousands to millions of dollars to many causes throughout his life.

Three plant names have been dedicated to George Cooley for his outstanding work in botany: cooleyi Monach. & Leonard occurring in Florida, Thalictrum cooleyi H. E. Ahles known from the southeastern USA including Florida, and Thelypteris cooleyi Proctor found in the Lesser Antilles.

JAMES DAVIS RAY, JR. (1919–1990) A native of Mississippi, James Ray, Jr. received his bachelor’s degree in botany from Mississippi State University in 1939 and later his Ph.D. from the University of Illinois in 1951, presumably under the auspices of George Neville Jones. As a faculty member at Mississippi State University, Jim met George Cooley, who would later invite him to work at USF. Jim was the first faculty member to be hired at USF, in 1959, a year before the first classes. He later became the first director of the USF Herbarium in 1960, and was extremely active collecting 1000s of specimens for the herbarium. Soon Jim became the Dean of the College Figure 9. Undated photo of Natural Sciences. of James Ray from USF Oracle, 9 Mar 1990. In a letter to Cyrus Mark, 13 September 1965, George Cooley wrote “He [James D. Ray] was of such value to the president in other matters

8 that he is now wholly in administrative work.” Jim and George remained good friends and Jim visited him at Hickory Hill in 1981. James Ray was described by Carolyn Borders (then the business manager for the College of Natural Sciences) as a “teacher’s teacher; a scholar’s scholar; and above all, a gentleman’s gentleman. The university is a much better place because he was here.”

ROBERT WILLIAM LONG (1927–1976) Born in Kentucky, Robert Long received his bachelor’s degree from Wesleyan University in 1950. He was an early Ph.D. student of Charles Heiser at Indiana University, working on the Helianthus () for his dissertation which he completed in 1954. He became employed at Ohio Wesleyan University in the Department of Botany & Bacteriology from 1954–1962, before becoming the curator of the herbarium at USF (also in the Department of Botany & Bacteriology at the time) in 1962. Soon thereafter, Bob became director after James Figure 10. Photo of Robert Long Ray transitioned into administrative work. He was credited with ca. 1970 in the herbarium (from suggesting the present location of the gardens during talks with jacket cover of A Flora of John Allen in May 1965. Tropical Florida, courtesy of the University of Miami Press). Bob taught many classes at USF, including Plant Ecology and Systematic Botany, and was very supportive of his students, considering his main objective to be a teacher. It was in his Systematic Botany class that he became impressed by the plant knowledge of an undergraduate student and Tampa native, Frank Almeda (class of 1968), whom he quickly hired to work at the herbarium.

Bob intensively studied at USF, especially the genus Ruellia. It is speculated that Ruellia ciliatiflora is abundantly common on the USF Tampa campus because of his research. Carmen Rose Broome often accompanied Bob on collecting expeditions and received her Master’s degree in 1969 under his supervision studying Ruellia (Acanthaceae). Martha Meagher was one of Bob’s Ph.D. students who studied Justicia (Acanthaceae) and herself contributed hundreds of collections to the herbarium from south Florida and the West Indies. Bob contributed 1000s of specimens to the herbarium including many from Belize, Honduras, and . Together with Olga, they published floristic works such as the Flora of Tropical Florida to succeed much of John Small’s work in the early 1900s.

With polycystic kidney disease, Bob’s kidneys eventually became critically weak in 1974. Bob regularly underwent dialysis, and despite this, remained dedicated and active in teaching and research until near his death.

OLGA KORHOVEN LAKELA (1890–1980) Olga Lakela was from Kestilä, Finland and moved to the northern USA at the age of 15 in 1906 without much knowledge of the English language. Some of her earliest collections are from North Dakota, starting in 1921, and deposited at Minot State University (MISU) and the University of Minnesota (DUL), herbaria she founded. Olga earned her Ph.D. studying the local flora and

9 conducting a monograph of Heuchera (Saxifragaceae) in the 1930s at the University of Minnesota- Duluth, teaching there from 1935–1958. Her name was inscribed on the Wall of Fame at the 1940 New York World’s Fair. Eventually she published A Flora of Northeastern Minnesota in 1965.

Olga was probably contacted by George Cooley and/or James Ray to solicit her help in developing the herbarium at USF, which quickly developed into her studying the local flora for the next 14 years. She began working at the herbarium in 1960 and for awhile lived in a small apartment complex near campus. Olga went on a collecting trip to the Pineola area in Citrus County with the Cooleys, Allens, and Lassiters. She compiled a list of species from Collier, Dade (now Miami-Dade), and Monroe Counties from her studies of specimens at FLAS, the University of Miami, and Figure 11. Photo of Olga Lakela ca. 1970 with help from Frank C. Craighead and George Avery. in the herbarium (from jacket cover of A Together with Robert Long, Olga co-authored the Flora of Tropical Florida, courtesy of the tome A Flora of Tropical Florida. University of Miami Press). Olga hired an undergraduate, David Nellis (class of 1964), as an assistant and guide for her work in the early 1960s; he carried much of her specimens in the field and navigated them back to their automobile after the collecting was finished. David described her as a dynamo and very enthusiastic whereby seemingly everyday and every plant held excitement. He considered her nearly fearless in the way she trudged straight on through the natural landscape. David recounted one night in a motel in Immokalee which they suspected of being a brothel with the constant noise and disruption. Finding nothing else available, they asked the sheriff if they could use the jail, and, for one night in Immokalee, David and Olga spent the night at the jail.

Frank Almeda served to drive Olga to many sites Figure 12. Photo of Frank Almeda and Olga in Florida for collecting plant specimens from Lakela in the summer of 1977 in Clearwater, 1966–1968, noting that Olga was particularly fond where Olga lived her remaining years. of stopping for coffee and ice cream at Howard Johnson’s. He recalled once getting lost together among cypress domes in southwest Florida as Olga had lost her compass. They made it back to the Illinois Motel to press the plants and take notes. Olga mentioned that she had been very scared during the incident, but selflessly explaining she was not so much for scared herself, as she had already lived a great life. On another trip, Frank recounted Olga reacting with disapproving surprise when Bob Long ordered a beer at a restaurant

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Glenn Fleming (who himself deposited 1000s of specimen at USF) described Olga as “independent, delightfully considerate and an inspiration. A dedicated caretaker, her life work was preserving the natural beauty of the Earth. She knew no other.” Olga has collected more specimens for the USF herbarium than almost anyone else, with about 10,000 specimens. Upon her death, she willed a substantial amount to the herbarium which established an endowment. The plant Lechea lakelae was named in her honor, to which she responded to Robert Wilbur to thank him for “his choice of a euphonic name, not for naming it for me.”

MAURICE MULVANIA (1874–1961) Born in Missouri, Maurice Mulvania earned an M.S. from the University of Tennessee in 1908 and worked there until 1923. He was the Dean of the Pre-Medical School for the University of Tennessee but was dismissed for controversial issues surrounding a course taught by Jesse W. Sprowls which sought use of a textbook containing ideas on evolution. Jesse had been prevented from using the text by others in the university. Maurice, Jesse, and others were dismissed after they had submitted a petition to the American Association of University for an investigation into the matter. Shortly thereafter the Butler Act was enacted in Tennessee in 1925, criminalizing the teaching of evolution in public schools, which led to the prosecution of a high school teacher, John T. Scopes.

Figure 13. Maurice Mulvania at Florida Southern College (Graves 1946). Maurice earned his Ph.D. under James Johnson from the University of Wisconsin in 1925. He then worked at Florida Southern College from 1926–1947 and is believed to be responsible for the collections in the Florida Southern College herbarium (FSCL) from the 1920s–1930s (the FSCL herbarium was donated to USF in 2015). Maurice was alleged to have a “herbarium of more that [than] 2,000 plants” at FSCL and had aspirations to compile “an illustrated book of the flora of Florida”. He reported on the first discovery of the naturalization of Chamaecrista rotundifolia and Solanum jamaicense in Florida. He was married to Eunice V. Mulvania and they had three children: Sarah H., Walton M., and Robin C. Mulvania.

MARGARET LOIS GILBERT (1928–2011) From Rhode Island, Margaret Gilbert earned her Ph.D. under John T. Curtis from the University of Wisconsin in 1953 studying the ecology of trees and understory herbs. Margaret taught at Northwestern State College [now University] of Louisiana in 1953–1954. She began her career at Florida Southern College (FSC) in 1954 and substantially helped the growth of the FSCL herbarium (donated to USF in 2015). She was a very well-respected mentor at FSC and was described as inspirational, supportive, and strict. From 1962–1964, Margaret and John B. Funderburg, Jr. were awarded several National Science Foundation (NSF) grants to conduct a

11 comprehensive study of plant and animal life of central Florida (the NSF grants under the Undergraduate Science Education Program were no. 6219766 ($14,640) in 1962, 6325496 ($10,950) in 1963, 64E4900 ($5,470), 64E4161 ($12,600), and 640U724 ($10,260)). Through these grants, they collected thousands of specimens from various places in Florida. The specimens from the FSCL herbarium (donated to USF in 2015) funded by these grants were labeled as “NSF” for the collector. This mainly refers to Carol Florence Caudle [now Carol C. Baskin], as she was in charge of the plant collecting as well as the herbarium work, but nearly always Margaret and/or Mary Sue Tisdale were involved. In the summer, they were often in groups with about eight FSC undergraduates along with Margaret and John. They camped, canoed (e.g. along the Peace River often), or sometimes stayed in cabins at Methodist youth camps (abbreviated MYC on many labels). In 1962, they had one truck and in 1963 they secured an additional truck which was once a weapons carrier. John was most interested in herpetology and collecting snakes. Margaret was influential in helping Carol get admitted to Vanderbilt University for her Ph.D. under Elsie Quarterman. Carol and her husband Jerry wrote the tome : Ecology, Biogeography, and Evolution of Dormancy and Germination.

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Figure 14. Article about FSC faculty and students studying Florida’s wildlife. On 1962 herbarium specimens from FSCL, “NSF” denotes this particular group (courtesy of the Lakeland Ledger).

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PHOTOGRAPHIC TOUR OF THE HERBARIUM AT USF

Figure 15. Chinsegut Hill manor house, the same room where the USF herbarium collections were first stored and photographed with George Cooley as seen in Figure 1. Photo (facing S) by Alan R. Franck with Carmel vanHoek and Colleen Werner, 17 Jan 2016.

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Figure 16. The Administration Building (formerly ADM, now ALN) in October 1960, when it contained the USF Herbarium. Photo (facing NW) by Joel Jackson.

Figure 17. Courtyard of the ADM building (now ALN) in November 1960, where the herbarium resided and contained nearly 20,000 specimens before its move to LIF. Photo (facing E) by Joel Jackson.

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Figure 18. The Chemistry Building (CHE) with LIF under construction in the background in October 1960. Photo (facing NW) by Joel Jackson.

Figure 19. The LIF building, ca. 1968–1970. Photo (facing SW) by unknown.

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Figure 20. On the left, the LIF building (where the herbarium resided) and the CHE building to the right in 1965. Photo (facing NE) by Joel Jackson.

Figure 21. Overview of USF Tampa campus ca. 1968–1970, with LIF in the left-center. Photo (facing NE) by unknown.

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Figure 22. The east wing of the LIF building where the herbarium once resided. Photo (facing NE) by Alan Franck, 9 Nov 2014.

Figure 23. The east wing (left) of the LIF building where the herbarium once resided along with the truck Alan was using to move the final bits of the herbarium to CMC. Photo (facing south) by Alan Franck, 9 Nov 2014.

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Figure 24. The entrance to the herbarium in LIF. Photo (facing E) by Alan Franck, ca. 2010.

Figure 25. Richard Wunderlin in the herbarium, with Bruce Hansen in far-right background. This room was once the lab of Robert Long and eventually held the compactor system for the “Sympetalae” families. Photo (facing SE) from USF Magazine, 1990.

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Figure 26. The reception area (left) which housed the cultivated specimens and a workspace (right) which was once Long’s lab in LIF. Photos (facing NW and NE) by Peter Simoes, 16 Oct 2014.

Figure 27. Herbarium work area (left) and mounting room (right) in LIF. The mounting room was once a classroom where courses such as Systematic Botany were taught by Robert Long. Photos (facing W) by Peter Simoes, 16 Oct 2014.

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Figure 28. Mounting room and Bruce Hansen’s office in LIF. Photos (facing E) by Peter Simoes, 16 Oct 2014.

Figure 29. The George Cooley Library in LIF packed in boxes ere its move to the CMC building. Photo (facing E) by Peter Simoes, 16 Oct 2014.

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Figure 30. Compactor system being disassembled (left) and after disassembly (right) in LIF. Photos (facing NE and north) by Alan Franck, 20 Oct and 23 Oct 2014.

Figure 31. Richard Wunderlin packaging cucurbit specimens in LIF ere being moved to CMC. Photo (facing NW) by Alan Franck, 28 Oct 2014.

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Figure 32. Herbarium cases being re-assembled onto compactor systems in CMC. Photos (facing W) by Alan Franck, 27 Oct 2014.

Figure 33. The last part of LIF to be torn down, the west wing, where most biology labs were once taught until the ISA building was erected. Photo (facing SW) by Alan Franck, ca. Mar 2015.

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Figure 34. Imaging of herbarium specimens by Peter Simoes (left) and mounting of specimens by Aline Sousa (right) in the CMC building. Photo (facing SE and S) by Alan Franck, 2015.

Figure 35. Main herbarium collections room in CMC. Photo (facing E) by Alan Franck, 2015.

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ADDITIONAL HISTORIC USF HERBARIUM PHOTOS

Figure 36. Frank C. Craighead in the Everglades, Jun 1962. Photographer unknown.

Figure 37. Likely Long’s Plant Taxonomy students at Fairchild Tropical Botanic Garden, Apr 1964. Photo likely by Robert Long.

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Figure 38. George N. Avery on a collecting expedition on Big Pine Key with Robert Long, J. De Boer, Rosalind Hebert, and G. Mills, Aug 1966. Photographer likely Robert Long.

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Figure 39. George N. Avery on a collecting expedition on Big Pine Key with Robert Long, J. De Boer, Rosalind Hebert, and G. Mills, Aug 1966. Photographer likely Robert Long.

Figure 40. Likely Long’s Plant Taxonomy students at Fairchild Tropical Botanic Garden, 1968. Photo likely by Robert Long.

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Figure 41. Likely Long’s students at unknown location in a football game, with Robert Long second from the left in the foreground, 1970. Photographer unknown.

Figure 42. Likely Long’s Plant Taxonomy students at Fairchild Tropical Botanic Garden, Jun 1971. Photo likely by Robert Long.

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Figure 43. Marvin E. Ragan with a specimen of Sobralia (USF accession 149659, 149663, or 151775) in Ecuador on a collecting expedition with Lawrence J. Davenport, Bruce Hansen, Harry E. Luther, Ruben P. Sauleda, John H. Wiersema, and Richard Wunderlin, Jul–Aug 1980. Photographer unknown.

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Figure 44. Marvin E. Ragan (right) in Ecuador on a collecting expedition with Lawrence J. Davenport, Bruce Hansen, Harry E. Luther, Ruben P. Sauleda, John H. Wiersema, and Richard Wunderlin, Jul–Aug 1980.

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REFERENCES: Almeda, Frank and Mary Beth Almeda. 2016. Personal interview [14 Jan 2016]. CMC 10, Herbarium, University of South Florida. [Anonymous]. 1923. University disturbed by row over faculty dismissals. New York Times, 22 Jul 1923. [Anonymous]. 1925. Freedom of education on trial. The Aspen Democrat-Times, 4 Jun 1925. [Anonymous]. 1933. Dr. Mulvania compiles illustrated flora book. The Southern, 6 May 1933. [Anonymous]. 1947. Dr. Mulvania retires after 21 years. The Southern, 16 May 1947. [Anonymous]. 1960. Classes start after ceremonies. The Tampa Times, 26 Sep 1960. [Anonymous]. 1961. Dr. Mulvania and wife die within 3 hours. Lakeland Ledger, 19 Oct 1961. [Anonymous]. 1962. National Science Foundation Grant Aids Wildlife Study. Lakeland Ledger, 23 Sep 1962. [Anonymous]. 1963. Science Foundation Gives College $10,950 Grant. Lakeland Ledger, 23 Jan 1963. [Anonymous]. 1970. Olga Lakela, Botanist –– Author. Tampa Tribune, Florida Accent Magazine, 3 May 1970. [Anonymous]. 1973. Florida herb named for USF botanist. Tampa Tribune, 1 Apr 1973. [Anonymous]. 1975. Personalia. Plant Science Bulletin 21: 41. [Anonymous]. 1980. USF curator Lakela dies. USF Oracle, 21 May 1980. [Anonymous]. 1986. George R. Cooley dies at 90; established professorships. The New York Times, 30 Sep 1986. [Anonymous]. 1986. Conservation Legacy. To the Editor, Opinion, The New York Times, 4 Nov 1986. [Anonymous]. 1987. In memoriam: George R. Cooley––USF friend and benefactor. CNS Developments [USF College of Natural Sciences] 3: Summer. [Anonymous]. 1987. The USF Herbarium: A garden of wonder. CNS Developments [USF College of Natural Sciences] 3: Summer. [Anonymous]. 1990. First hired at USF dies. USF Oracle, 9 Mar 1990. [Anonymous]. 2011. Dr. Margaret Lois Gilbert. Ledger, Obituaries, 13 Mar 2011. Archives of the University of South Florida. Special and Digital Collections, University of South Florida Library, Tampa, Florida. Arthurs, R. 1978. Island visitor spearheads Tiger Creek protection effort. Sanibel-Captiva Islander, 31 Jan 1978. Baskin, Carol C. [previously Carol Florence Caudle]. 2016. Phone interview [19 Feb 2016]. Herbarium, University of South Florida. Berardino, J. T. 1973. USF’s Herbarium preserves the past. USF Oracle, 3 Oct 1973. Coen, L.D. and E.C. Milbrandt. 2010. Brief history of the Sanibel-Captiva Conservation Foundation (SCCF)’s marine laboratory, Sanibel, Florida: not to be confused with Doc Ford’s Sanibel Biological Supply Company. Gulf of Mexico Science 2010: 200–213. Cooley, George R., Papers, 1941–1986. New York State Library, Albany, New York. 14 boxes [consisting mostly of personal letters and a taped interview ca. 1986 by an unknown archivist from Colgate University]. SC 18858. Donated by Barbara C. M. Dudley, processed by Thomas E. Lavery, June 1990. Fred, E.B., W.H. Peterson, and M. Mulvania. 1926. The effect of lactic acid bacteria on the acetone-butyl alcohol fermentation. J. Bacteriol. 11: 323–343.

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Funderburg, J.B., D.S. Lee, and M.L. Gilbert. 1969. An unusual salamander from the Ocala National . Quart. J. Florida Acad. Sci. 32: 230–232. Gilbert, M.L. and J.T. Curtis. 1953. Relation of the understory to the upland in the prairie-forest border region of Wisconsin. Trans. Wisc. Acad. Sci. Art. Lett. 42:183–195. Graves, R. (ed.). 1946. The Interlachen. Students of Florida Southern College, Lakeland, Florida. Harper, Francis, Collection, MS 245. Department of Special Collections, Kenneth Spencer Research Library, University of Kansas Libraries. Howard, R.A. 1987. George Ralph Cooley (1896–1986). Taxon 36: 696–698. KQDS Staff. Northland uncovered: Olga Lakela herbarium. Fox 21, Duluth, MN, 4 Apr 2017. 5Kosharsky, R. [1980?]. Olga Lakela, was curator of herbarium at USF. St. Petersburg Times, Obituaries, [1980?]. Krivanek, Robin C. 2015. Email correspondence [28 Feb 2015]. Lakela, O. 1939. A floristic study of a developing plant community on Minnesota Point, Minnesota. Ecology 20: 544–552. Liller, David. 1990. A Growing Reality . . . Some of Florida’s plants, irreplacable [irreplaceable] food and medicinal drug sources, may in fact be facing their last chance. USF Magazine 32(7): 6–7. Long, R.W. 1972 [no date given but the accession 100,000 appears to have happened in 1972]. The University Herbarium 1971. A report and summary of activities for 1960–1971 of the herbarium. University of South Florida, Tampa. 20 pp. McNamee, B.C. 1977. Hickory Hill Zucchinis, 2nd ed. Press of the First Albany Corporation, Albany, NY. 81 pp. Mulvania, M. 1915. Observations of Azotobacter. Science 42: 463–465. Mulvania, M. 1919. A comparison of Azotobacter with yeasts. Agricultural Experiment Station of the University of Tennessee 122: 1–5. Mulvania, M. 1925. Cultivation of the virus of tobacco mosaic by the method of Olitsky. Science 62: 37. Mulvania, M. 1930. New types of plants in Florida. Science 72: 603–604. Mulvania, M. 1931. Ecological survey of a . Ecology 12: 528–540. Nellis, David. 2015. Phone interview [2 Nov 2015]. Herbarium, University of South Florida. Otto, S. 4 graphs too few to chronicle a life [George Cooley] worth volumes. The Tampa Tribune, 15 Oct 1986. Palmer, Tom. 2005. Botanical explorer’s old records prove a Florida treasure trove. Sarasota Herald-Tribune, 7 Sep 2005. Ray, Jr., J.D. 1956. The genus Lysimachia in the New World. Illinois Biological Monographs 24:1–160. Rollins, R.C. and R.A. Howard. 1986. A tribute to George R. Cooley. Syst. Bot. 12: 444–448. Rooney, A. 1985. One zucchini is one zu many. St. Louis Globe-Democrat: 4C. 15 Aug 1985. Thomas, G. 1970. William Henry Browne, III. Obituaries, Florida Times-Union, Jacksonville, Florida, 16 Dec 1970. Wargelin, M. [without date]. Review of the Finnish American Culture. Kauppisten Sukuseura. Winter, F. 1969. Conservancy buys large Collier tract. Fort Myers News Press, 12 Dec 1969. Wunderlin, R.P. 1977. Robert William Long. Bull. Torrey Bot. Club 104: 66. Wunderlin, R.P. 1978. A brief history of pre-Darwinian systematic botany. Ex Libris, Journal of the USF Library Associates, Winter 1978: 5–12.

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OVERVIEW OF THE USF HERBARIUM COLLECTIONS As of June 2018, there are approximately 300,000 accessioned specimens at the USF Herbarium, consisting of about 280,000 specimens of vascular plants, 14,000 algae, 2,800 bryophytes, 1,300 lichens, and 1,000 macrofungi. The collection contains approximately 520 type specimens (predominantly vascular plants with a few algae, lichens, and macrofungi). The USF Herbarium is richest in specimens from Florida (~40%), with additional holdings from north of

Figure 45. Map showing which herbarium has the largest collection for each county in Florida, based on data and estimates current to 2016.

Mexico (~35%), Latin America and the West Indies (~15%), and the Eastern Hemisphere (~10%). Specific groups well represented in the herbarium include Acanthaceae, , Begoniaceae, , , , and pteridophytes (ferns), due to the research

33 interests of previous staff. There is a unique set from Mexico covering indigenous ethnobotanical specimens (of the Amuzgo, Cora, Mixtec, Otomí, Triqui or Trique, and Zapotec) that were acquired by Gerald G. Robinson (1933–2008), who started at USF in 1960; in addition to Robinson, other collectors for this research included Cornelio Basilio G., Barbara Hollenbach, Albertha Kuiper, Livorio, Barcimeo Lopez C., Martías (or Matías), Enrique Romero L., Jesús Santiago Martínez, and Sebastián. Other significant collections include those of James Keenan from Myanmar (Burma) and M.L. Banerji specimens from Nepal.

Figure 46. Global distribution of specimens at the USF Herbarium.

Since its beginning, the USF Herbarium has sent out 500+ loans totaling 40,000+ specimens to 100+ institutions. Researchers based at USF have studied 50,000+ specimens from 650+ loans from 60+ institutions. With the advent of online specimen databases (e.g. Atlas of Florida Plants), herbarium specimens are now more widely viewed and used than ever, and conversely loans requesting physical access to specimens have understandably decreased. Digitization of the collections began in November 2003 with transcription of vascular plant label data. Imaging of the vascular plants began in February 2006. Initial digitization of bryophytes and lichens (~2,500 specimens) was accomplished by FLAS in 2013. NHA digitized the initial ~13,800 algae specimens in 2015. Nearly all of the cryptogamic specimens (algae, bryophytes, lichens, and macrofungi) at USF are digitized.

To enrich a herbarium’s collections, herbaria typically exchange specimens with each other. Over the span of its history, USF has exchanged specimens with 80+ herbaria worldwide. Over 50,000

34 specimens have been sent out from USF and a nearly equal number of specimens have been received, averaging about 900–1000 specimens sent out and an equal number received annually. The most active specimen exchange programs with USF have been with the herbaria of the Fairchild Tropical Botanic Garden (FTG), Harvard University (A, FH, GH), Marie Selby Botanical Gardens (SEL), Missouri Botanical Garden (MO), New York Botanical Garden (NY), University of Florida (FLAS), and University of Wisconsin (WIS).

Figure 47. Loan and exchange activity from 1962–2015. Bars in black are exchange specimens sent from USF, bars in white are numbers of exchange specimens received to USF, bars in dark gray are loans sent from USF, and bars in light gray are loans received to USF.

The herbarium occasionally becomes the recipient of large collections from other institutions or individuals. Specimens have been received from the Florida Southern College herbarium (FSCL, ~7,000, through the help of Eric Kjellmark), Florida Marine Research Institute in St. Petersburg (~1,000), University of Tampa (~1,000), bryophyte herbarium of Rita Lassiter (~1,500), algae herbarium of Harold Humm (~1,000), algae herbarium of John Schwede (~500), lichen herbarium of Julia A. Schulten [previously Julia A. Schutte] (~1,000), and Eckerd College herbarium (~200). The main collectors within the FSCL collections were M. H. Berry, C.C. Carter, Carol F. Caudle, Joe E. Crumpton, D.E. Dimmick, M. A. Ford, Margaret L. Gilbert, Maurice Mulvania, Paul O. Schallert, and Victor W. Yingst. From the Eckerd collections, the main collectors were D.E. Anderson, Michael R. Blake, Stephan Imhof, and A.M. Wooten.

Specimens collected by 1000s of different individuals are deposited at the USF Herbarium. Some of the most active collectors with specimens at USF include G.R. Cooley (~6,000 specimens), R. K. Godfrey (~3,000), J.D. Ray, Jr. (~4,000), R. Kral (~2,500), O.K. Lakela (~10,000), R.B. & J.A. Lassiter (~3,000), R.W. Long (~4,000), C.J. Dawes (~9,000), Margaret L. Gilbert (~3,000), D. Burch (~4,000), R.P. Wunderlin (~8,000), B.F. Hansen (~10,000), R.P. Sauleda (~5,000), A.G. Shuey (~3,000), S.L. Orzell and E.L. Bridges (~7,000), C. vanHoek (~3,000), J.M. Kunzer (~2,500), and A.R. Franck (~4,000).

Some of the oldest specimens at the USF Herbarium include those collected by Ferdinand Rugel

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(1830s–1840s, ~200 specimens), Jacob Wolle (1840s, ~500), Alvin W. Chapman (1840s–1860s, ~10), William G. Farlow (1880–1890s, ~200), C.F. Batchelder (1880s–1930s, ~300), Allen H. Curtiss (1890s–1900s, ~200), William H. Harris (1890s–1900s, ~10), John K. Small (1890s– 1920s, ~500), Maurice Mulvania (1920s–1930s, ~350), H. Harold Hume (1930s, ~30), and W.M. Buswell (1930s–1950s, ~200).

In the 1970s, there were a few hundred specimens of fungi at USF. A letter from Robert Long to George Cooley (10 July 1975) speaks of the “addition of a very large mycological collection from the University of Michigan” and was mentioned in the context of the 2nd International Mycological Congress that later occurred at USF in 1977 (Simmons 1978). They were held in a teaching lab in the west wing ground floor of LIF. Apparently, a student’s plant press slipped towards the light of a dryer, causing a fire that ruined the fungi specimens in the late 1970s (Wunderlin, pers. comm.). The fungi collections have since been re-established, primarily with new collections since the 2000s.

Figure 48. Number of specimens accessioned and digitized in the USF herbarium. Nearly all cryptogamic specimens (algae, bryophytes, fungi, & lichens) are digitized and about 200,000 vascular plant specimens are digitized, as of June 2018.

ONLINE ACTIVITY Over half of the vascular plant specimens are digitized and available on the Atlas of Florida Plants. Nearly all Florida and West Indies specimens of vascular plants are digitized. The rest of the Americas and the eastern hemisphere are actively being digitized (as of June 2018). Nearly all of the cryptogamic specimens (algae, bryophytes, lichens, and macrofungi) at USF are digitized and available through their respective online herbaria consortiums.

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Digitized data are shared with several other data aggregators: Global Biodiversity Information Facility (GBIF), iDigBio, Consortium of North American Bryophyte Herbaria, Consortium of North American Lichen Herbaria, Macroalgal Herbarium Portal, Mycology Collections Portal, and Southeast Regional Network of Expertise and Collections (SERNEC). These data are used for multitudes of applications.

The Atlas of Florida Plants now (June 2018) treats 4,700+ species which amounts to 18,000+ photos of plant morphology, 75,000+ mapped county data points, and 50,000+ taxonomic names (including synonyms). Data are also provided for traits of each species such as native, endemic, status, and threatened/endangered status. Over 3,000 names (including synonyms) excluded from the Florida flora are listed along with the explanation.

On average per year, Google Analytics records about 130,000 users (unique devices), 240,000 sessions (average of 5.5 minutes and 8 pages per session), and 2.0 million page views (49.65% bounce rate) for the Atlas of Florida Plants, which has remained fairly consistent since data collection began for the Atlas in Apr 2009. The majority of uses are from the USA (84%), followed by India (2.1%), Philippines (0.8%), Canada, Brazil, Mexico, United Kingdom, Indonesia, and about 200 other countries. The USF Water Institute now handles data, patents, and licensing for other states (i.e. Alabama, Kentucky, Indiana, New York, and Tennessee) that utilize the Atlas technologies.

Figure 49. Percent usage from Apr 2009–Apr 2016 for the Atlas of Florida Plants based on Google Analytics (a blue dot at this scale represents at least several thousand sessions).

FILING SYSTEM Vascular plant specimens are arranged in the USF Herbarium by a modified Dalla Torre & Harms (1908) system, whereby each family is given a number, and the sequence generally follows evolutionary relatedness based on their understanding in 1908. The order of USF cases is ferns, gymnosperms, monocots, and dicots. The ferns are given parenthetical family numbers, i.e. (1)– (32). The gymnosperms comprise family numbers 1–7, the monocots 8–50, and the dicots 51–280.

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Of course, many families have been recircumscribed since this system was founded and slight modifications continue to be enacted to accommodate this. However, many families remain out of place with respect to their nearest relatives (e.g. the Salicaceae and Flacourtiaceae, though now one family, are very distant in the filing system).

Within each family, genera are arranged alphabetically. Within each genus, specimens are arranged geographically. All Florida specimens are found first within manila folders. Next are red folders containing taxa from the rest of the southeastern USA (Alabama, Arkansas, Georgia, Louisiana, Mississippi, North Carolina, , Tennessee). Next are yellow folders containing the rest of the continental USA and Canada. Then are filed gray folders for the West Indies. Green folders then contain the rest of the Neotropics. Blue folders comprise the eastern hemisphere.

Within each geographic section of each genus, species are arranged alphabetically. Each taxon has its own folder for Florida. For the rest, a taxon is given its own folder only if there are 3 or more specimens per geographic area. Otherwise, taxa are alphabetized within a genus folder at the front of the geographic section. Any unidentified specimens are placed at the front of their respective lowest known taxonomic level. Vascular plant specimens without known provenance, i.e. known only from cultivation, are in separate filing cabinets from wild-collected material (although occasional cultivated specimens are placed with wild-collected material).

The general order of the algae collections are cyanobacteria, red algae, green algae, brown algae, and miscellaneous eukaryotic algae. The mosses are are alphabetized by order, family, genus, then species. The liverworts and fungi are alphabetized by genus. The lichens are are alphabetized by class, order, family, genus, then species.

HERBARIUM PERSONNEL

Directors: James Ray (1960–1964), Robert Long (1964–1976), Richard Wunderlin (1976–2009), Alan R. Franck (2016–2018), Diane Te Strake (2009–2016, 2018)

Curators: George R. Cooley (1958), James Ray (1959–1962), Olga Lakela (1960–1973), Robert Long (1962–1964), Richard Wunderlin (1973–1976), David Crewz (1976–1979), Bruce Hansen (1979–2014), Alan R. Franck (2014–2016), Arian Farid (2016–2018)

Assistant Curators: Martha Gillon (1960s), Thelma Fricks (1965–1968), Frank Almeda (1966– 1968), unknown (1970s, 1980s, 1990s), John Kunzer (2003–2009), Alan Franck (2009–2014), Bob Upcavage (2009–2011), Ronald Chicone (2011–2014), Arian Farid (2014–2016), Peter J. Simoes (2014–2015), Rose Odonovan (2014–2015), Sarah R. Funaro (2015–2016), Samra Kazim (2016–2018), Leenil Noel (2016–2018), Joshua Campbell (2016–2018), Allison Blakely (2017), Qiara Perez (2017), Daley Drucker (2017–2018)

Director Emeritus: Richard P. Wunderlin; Curator Emeritus: Bruce F. Hansen; Associate Staff: Kathleen Burt-Utley, Clinton J. Dawes, Frederick B. Essig, Diane Te Strake, John F. Utley Information Technology: USF Water Institute (Keith Bornhorst, Shawn Landry, Jason Scolaro)

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Volunteers (data deficient for 1960s–2000s). Importantly, the volunteers have performed a significant part of the digitizing, mounting, filing, and other curatorial duties over the years. 1970s–1980s: Glenn Fleming & Pierre Genelle. 1980s–1990s: Marie van Etten (volunteer librarian & fern specialist). 2010–2014: Aarzoo Bukhari, Ryan Coddington, Chuck Courtney, Ambang Ebie, Alexandra Ferrero, Emma Fontaine, Sarah Funaro, Michael Inman, Jabari Lee, Mike Meads, Thomas Richards, Austin Ritenour, Janice Rivera, Bob Upcavage, Brittany Wert, Marilyn Whetzel, David H. Williams, Gang Xian. 2015: Zein Barakat, Joshua Campbell, Ryan Coddington, Linda Dyet, Natalia Ferrer, Alexandra Ferrero, Nicholas Fethiere, Sarah Funaro, Lauren Grant, Samra Kazim, Hyo Kim, Vivian Lee, Leenil Noel, Jacob Norman, Charles Nguyen, Julie Nguyen, Thomas Nguyen, Thuy Nguyen, Nicole Perez, Victor Perez, Advaitaa Ravipati, Thomas Richards, Michael Simoes, Aline Sousa, Sean Suba, Briana Williams. 2016: Jahnii Bell, Nitesh Coomar, Natalia Ferrer, Nicholas Fethiere, Samantha Kane, Vivian Lee, Christian Marr, Catherine McInnes, Nicole Perez, Thomas Richards, Deborah Robison, Sarah Robison, Gloria Russell, Misha Sholeh, Amanda Shuttleworth, Ling-Yu Tsai, Crystal Williams, Titilola Wright. 2017: Anika Alam, Sabrina Burkholder, Felipe Crespo, Ashley Denslow, Shrhidi Desai, Thi Doan, Daley Drucker, Medina Dzaferi, Yasmin Espinoza, Ami Faust, Victoria Forbrich, Robert Frank, Justin Jimenez, Jessica Jordahl, Chynna Liu, Charles Nguyen, Victoria Okeke, Chaimae Oualid, Sofia Perez, Danielle Pichardo, Matthew Polner, Caitlin Powell, David Rongo, Alexis Smith, Misha Sholeh, Shivani Sookchand, Abram Suneson, Nader Tabsch, Latina Tang, Nguyet Tran, Ling-Yu Tsai, Dino Tsilionis, Akhil Tumpudi, Marion Ugochukwu, Rampses Viracacha. 2018: Timothy Chafin, Deanna Chan, Rachel Fanning, Victoria Forbrich, Francisco Gonzalez, Jad Halawani, Madison Hatch, Morgan Hendrie, Gilbert Immanuel, Frances Laiuppa, Joshua Garcia, Meaghan Kane, Carly Mott, Chaimae Oualid, Dylan Passero, Sofia Perez, Phuc Phan, Carina Pinkston, Saranda Rossin, Sabrina Sarun, Nguyet Tran, Dino Tsilionis, Akhil Tumpudi, Leticia Vasquez, Caroline Wagstaff, Michelle Woronka, Mery Yanez Yuncosa, Jingjie Yu

Donors The USF herbarium has benefited from generous donations from multiple individuals and institutions since its beginning. Past individual donors included George R. Cooley, Olga Lakela, Glenn Fleming and Pierre Genelle, Marie Van Etten, and Alice F. Tryon and Rolla M. Tryon. Donors of 2015–2018 include Amec; Aquatic Plants, Inc.; Breedlove, Dennis & Associates, Inc.; CH2M HILL; Donald Richardson; EarthBalance; Flatwoods Consulting Group; Florida Wildflower Foundation; Heartland Chapter of the Florida Native Plant Society; HDR, Inc.; Kleinfelder; Onsite Environmental Consulting, LLC; Quest Ecology; Richard Gorsira; Jeffrey Norcini; Richard Pragluski; Scheda Ecological Associates; Southeast Environmental Solutions, Inc.; W. Dexter Bender & Associates, Inc.; and Wood Environment & Infrastructure Solutions, Inc.

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Figure 50. Driving directions to the USF Herbarium

HISTORY & ROLES OF HERBARIA (ca. 40,000 BC–present) In order to understand the purpose and relevance of a herbarium in society, it is necessary to trace the initial investigations into plants by humans.

History of Plant Depictions & Writings (ca. 40,000 BC–1500 CE) For at least the past 40,000 years (Thackeray 1983; Pike et al. 2012; David et al. 2013; Aubert et al. 2014; Middleton et al. 2014), humans (and maybe other Homo spp.) have created indelible art extant on such media as rock, metal, parchment, and paper. Rock art bears illustrations of environmental animals from as early as 30,000–35,000 years ago (Aubert et al. 2014; Hoffman et al. 2016) yet are noticeably bereft of plant depictions (except for tools made from plants such as weapons), despite plants being vital to their livelihood (e.g. fire, tools, and nutrition; Hardy et al. 2012; Hardy et al. 2015; Hardy et al. 2017). The depiction of somewhat identifiable plant species (e.g. cereal grains, date palm, and grapes) during the Bronze Age appears to follow the importance of plant domestication and agriculture.

Examples of some of the earliest plant illustrations include a Chogha Cheshmeh pottery sherd (ca. 4,000 BC, Iran; A 40720, 3668, The Oriental Institute of The University of Chicago), a date palm on the Battlefield Palette (ca. 3,100 BC, Egypt; in the British Museum; Gadd & Edwards 1953), and cereal florets from an Abydos tomb (ca. 3,000 BC, Egypt; Singer 1958). In the tomb of

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Khnumhotep II (ca. 1,900 BC) appear to be depictions of water lilies and acacias (David 2014). Excellent illustrations of viticulture are found in the tombs of Nakht, Sennefer, and Userhet (ca. 1,400 BC, Egypt). Other ancient representations may include cereal carvings on Oriental tombs and various plant species on Greek coins (Garrison 1931). A branched dicot carved on an antler might be dated ca. 30,000 BC if relatable to the dates of Grotte du Renne artefacts, Arcy-sur-Cure, France (Singer 1958; Highams et al. 2010),

The oldest herbals and botanical texts reveal increased attention paid toward specific plant species with regard to their agriculture, nutrition, and medicine. Some of the earliest known examples include Assyrian and Babylonian cuneiform (ca. 2500 BC; Thompson 1924; Borchardt 2002), Ebers Papyrus (ca. 1500 BC, Egypt), The Hippocratic Corpus (ca. 450 BC, probably authored by many individuals), On Plants (ca. 350 BC, by Aristotle, not extant or unclear if ever written), Enquiry into Plants (ca. 300 BC, by Theophrastus), texts by Diocles of Carystos (ca. 300 BC), On Agriculture (ca. 160 BC by Marcus Cato), Alexipharmaca (ca. 150 BC, by Nicander of Colophon; Scarborough 1977), Rhizotomikon (ca. 100 BC, by Crateuas, possibly the first known illustrated herbal), On Plants (ca. 30 BC by Nicolaus of Damascus), Natural History (ca. 70 CE by Pliny the Elder), and Materia Medica (ca. 80 CE by Dioscorides). Many of these dates are very rough estimates which often rely upon the works of countless others who have copied and modified these texts over hundreds of years.

The origins of the equally important ancient Ayurvedic and Chinese texts are difficult to date and are variously ascribed from ca. 300 BC (or earlier) to 100 CE (e.g. Bhishagratna 1907; Atal et al. 1981; Harper 1998). These ancient herbal texts include Charaka Saṃhitā, Sushruta Saṃhitā, Shen Nong Ben Cao Jing, and Wushi’er Bingfang. Numerous other herbals and botanical texts continued to appear during the common era (CE) (Arber 1912; Singer 1958), such as Galen’s works (ca. 200 CE), Herbarius (ca. 300 CE, by Pseudo-Apuleius), Ex Herbis Femininis (ca. 500 CE; Riddle 1981), The Canon of Medicine (ca. 1025 CE, by Avicenna), and De Proprietatibus Rerum (ca. 1240 CE, by Bartholomeus Anglicus).

Historical documentation of plants in many other cultures during this time period (Ritter & Ritter 1976; McKissack & McKissack 1994; Singleton 2004; Lozoya 2006) cannot be discounted and requires synthesis in the timeline of human evolution and plant knowledge. Certainly much has been lost due to plundering, looting, and destructive invasions which have occurred throughout human history, the significance of which can oft only be speculated. Recent demonstrative examples include the deliberate catastrophic losses of Mayan texts in the 1500s (Arbagi 2011), the Benin Punitive Expedition in 1897 (Ratté 1972), and the looting of the Iraq Museum in 2003 (Bogdanos 2005).

The Herbarium (ca. 1500–present) The art of preserving physical plants as herbarium specimens for study appears to trace its origins to present-day Italy during the 1500s when paper became an accessible commodity (Meyer 1863; Anonymous 1885; Druce 1889–1890; Lane 1996). Some of the earliest known extant herbarium specimens are of Gherhardo Cibo dated to 1532 and held in the Angelica Library in Rome, specimens of Michele Merini dated to the 1500s and held at the University of Florence Herbarium (FI), and specimens of Andrea Cesalpino dated to 1563 and also at FI (Moggi 2009). Luca Ghini

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(Smith 2017) is credited with directly catalyzing the preservation of specimens undertaken by Cibo, Merini, and Cesalpino. The En Tibi specimens and Leonhart Rauwolf specimens also date to the 1500s and reside in the National Herbarium of the Netherlands (L) in Leiden (Van Steenis- Kruseman 1979; van Andel 2017).

There are now over 2,500 herbaria across the globe altogether containing hundreds of millions of specimens of plants, algae, and fungi (Thiers 2016). The majority of herbaria were founded between 1951–1975 (Thiers 2016).

Purpose As humans thrive from information, a herbarium preserves information about plants, algae, and fungi on Earth. We depend on plants for food, medicine, water and air quality, soil health, paper, fabrics, cosmetics, building materials, and beautification. Herbarium plant specimens contain vital information that help us ensure the health and longevity of our species and the ecosystem. The most basic function of a herbarium is in identification, which has wide ranging applications. In nearly any scientific study, the documentation of plants is preferentially done by depositing a specimen in a herbarium to corroborate species identification and allow others to reanalyze data from these studies. The utility of herbarium specimens to answer inquiries into several disciplines is exemplified below (see also Funk 2004; Schilthuizen et al. 2015; Heberling & Isaac 2017).

For example, biodiversity measures rely on herbarium specimens to enumerate species lists. Climatic inferences are made from herbarium specimens by examining the change in phenology (e.g. flowering time) vs. temperature or stomatal density vs. carbon dioxide levels over time. DNA can be extracted from specimens to address evolutionary relationships and other questions concerning genetics, though DNA notably fractures and crosslinks as it ages in specimens. Ecological change in species compositions and associations can be determined from specimens. Endophytes (bacteria and fungi) can be studied within preserved plant tissues. Ethnobotanical or medicinal knowledge can be extracted from notes given on specimens. Extinct species are sometimes represented by extant herbarium specimens. Fecal samples of animals can be compared with specimens to determine their dietary preferences. Forensics can utilize specimens to establish crime scene localities and identify suspects based on the geographic localizations of certain species. Geographic ranges of species are typically based on specimens. Insects occurring directly on specimens can be studied (e.g. miners). Metal concentrations in specimens can be analyzed to gain insight into ecology, soil characteristics, and pollution. Microscopic and morphological characters can be studied from specimens to discover synapomorphies and yield insights into evolution. Nativity of a species can be partly determined based on the earliest specimen known from an area. Non-native and can be mapped over time to retrace their time of introduction and spread. Restoration ecology may select reintroduction of particular species based on historic or local occurrences as documented in specimens. Soil adhering to can be analyzed (e.g. for bacteria or mycorrhizae). Species are continually described from herbarium specimens, some collected several decades ago before being recognized as a new species. Species delimitation and infraspecific variation is understood through collection and study of herbarium specimens. The herbarium is also the cornerstone of nomenclature and taxonomy, by securing a name to a plant specimen thus stabilizing the identity and communication of a particular species.

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Nomenclatorial role of the herbarium Just as a word in the dictionary has a definition, organismal scientific names also have definitions. Scientific names are defined by their type or type specimen. Type specimens can be either a preserved physical specimen or a printed illustration. Most scientific names of plants, algae, and fungi are defined by a physical type specimen preserved at a herbarium. Otherwise the illustrations serving as type specimens usually reside in historic books and journals.

The International Code of Nomenclature for Algae, Fungi, and Plants governs rules for utilizing the proper scientific name for plants, fungi, and algae as based on type specimens. In this capacity, it offers a way to communicate internationally with an acceptable, and, hopefully, agreeable name, depending on taxonomic opinions.

Taxonomy at the herbarium Taxonomy is an arbitrary art of applying one name to a group of organisms, a group which is typically thought to be a coherent evolutionary unit. The rules of nomenclature determine what names are available for use in naming a group of organisms. Taxonomy is a practice of deciding upon or creating an available name to use. If a herbarium contains specimens representative of the extent of diversity inherent in the natural world, these specimens can be used to help define the limitations of a taxonomic name such as the delimitation of a species.

A herbarium is organized by taxonomy. Usually all organisms considered to be in the same taxonomic family are shelved into the same or adjacent cases. All species within a genus, typically reside adjacent to one another. All specimens considered the same species and from the same geographic region are usually filed together in a folder.

Species identification Unknown specimens are compared with confidently identified specimens and type specimens in the herbarium to find the best match. Frequently, to corroborate the identification, microscopic characters must be viewed. If adequate specimens do not exist in the herbarium, identification must rely upon written descriptions or illustrations of specimens.

Epilogue Though we continue to use a wide variety of plants and other organisms in our daily lives, we may be unaware of the living state of the organisms that we use, directly or indirectly. In particular, urbanization, commerce, and trade enable greater ignorance of ecosystems and species. Through specimens contained at a herbarium, we learn about the diversity of plants and associated organisms and are able to maintain this knowledge.

References Alizadeh, A. 2008. Chogha Mish II: The development of a prehistoric regional center in lowland Susiana, southwestern Iran. Final report on the last six seasons of excavations, 1972–1978. Oriental Institute Publications 130: 1–395. Anonymous. 1885. Communications, Séance du 5 Mai 1885. Bull. Soc. Bot. Lyon Compt. Rend. Séances 3: 61–64 [English version as “The Origin of Herbaria” in Bull. Torrey Bot. Club 12: 129–131].

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Arbagi, M. 2011. The Catholic Church and the preservation of Mesoamerican archives: an assessment. Archival Issues 33: 112–120. Arber, A. R. 1912. Herbals: Their Origin and Evolution: A Chapter in the History of Botany (1470– 1670). University Press, Cambridge. Atal, C.K., U. Zutshi, and P.G. Rao. 1981. Scientific evidence on the role of Ayurvedic herbals on bioavailability of drugs. J. Ethnopharm. 4: 229–232. Aubert, M., A. Brumm, M. Ramli, T. Sutikna, E.W. Saptomo, B. Hakim, M.J. Morwood, G.D. van den Bergh, L. Kinsley, and A. Dosseto. 2014. Pleistocene cave art from Sulawesi, Indonesia. Nature 514: 223–227. Bhishagratna, K.K.L. (ed.). 1907. An English translation of the Sushruta Samhita. Vol. I – Sutrasthanam. No. 10, Kashi Ghose’s Lane, Calcutta. Borchardt, J.K. 2002. The beginnings of drug therapy: Ancient Mesopotamian medicine. Drugs New Persp. 15: 187–192. Bogdanos, M. 2005. The casualties of war: The truth about the Iraq Museum. Amer. J. of Arch. 109: 477–526. Bortolus, A. 2008. Error cascades in the biological sciences: the unwanted consequences of using bad taxonomy in ecology. Bradley, R.D., L.C. Bradley, H.J. Garner, and R.J. Baker. 2014. Assessing the Value of Natural History Collections and Addressing Issues Regarding Long-Term Growth and Care. BioScience 64: 1150–1158. Buenz, E.J., D.J. Schnepple, B.A. Bauer, P.L. Elkin, J.M. Riddle, and T.J. Motley. Techniques: Bioprospecting historical herbal texts by hunting for new leads in old tomes. Trends in Pharm. Sci. 25: 494–498. Culley, T.M. 2013. Why vouchers matter in botanical research. Applications in Plant Sciences 1: 1300076. David, A. 2014. Hoopoes and acacias: Decoding an ancient Egyptian funerary scene. Journal of Near Eastern Studies 73: 235–252. David, B., J.-M. Geneste, F. Petchey, J.-J. Delannoy, B. Barker, and M. Eccleston. 2013. How old are Australia’s pictographs? A review of rock art dating. J. Arch. Sci. 40: 3–10. Druce, G.C. 1889–1890. History of Botany.—(Herbaria). Pharm. J. Trans. [ser. 3] 20: 511–513, 536–539. Funk, V. 2004. 100 uses for an herbarium. US National Herbarium, Smithsonian Institution. http://botany.si.edu/documents/2003_Funk_100Uses.pdf Gadd, C.J. and I.E.S. Edwards. 1953. The growth of the Egyptian and Assyrian Collections. The British Museum Quarterly 18: 81–83. Garrison, F.H. 1931. Herbals and bestiaries. Bull. New York Acad. Med. 7: 891–904. Gaur, A. 1992. A History of Writing. Cross River Press, New York. Hardy, K., J. Brand-Miller, K.D. Brown, M.G. Thomas, and L. Copeland. 2015. The importance of dietary carbohydrate in human evolution. Quart. Rev. of Biol. 90: 251–268. Hardy, K., Buckley, S., Collins, M.J., Estalrrich, A., Brothwell, D., Copeland, L., GarcíaTabernero, A., García-Vargas, S., de la Rasilla, M., Lalueza-Fox, C., Huguet, R., Bastir, M., Santamaría, D., Madella, M., Wilson, J., Cortes, A.F., Rosas, A., 2012. Neanderthal medics? Evidence for food, cooking, and medicinal plants entrapped in dental calculus. Die Naturwissenschaften 99, 617e626. Hardy, K., et al. 2017. Diet and environment 1.2 million years ago revealed through analysis of

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dental calculus from ’s oldest hominin at Sima del Elefante, Spain. The Science of Nature 104. doi:10.1007/s00114-016-1420-x Harper, D. 1998. Early Chinese Medical Literature: The Mawangdui Medical Manuscripts. Routledge, UK. Heberling, J.M. & B.L. Isaac. 2017. Herbarium specimens as exaptations: New uses for old collections. American Journal of Botany 104: 963–965. Higham, T., R. Jacobi, M. Julien, F. David, L. Basell, R. Wood, W. Davies, and C.B. Ramsey. 2010. Chronology of the Grotte du Renne (France) and implications fro the context of ornaments and human remains within the Châtelperronian. PNAS 107: 20234–20239. Hoffman, D.L., P. Utrilla, M. Bea, A.W.G. Pike, M. García-Diez, J. Zilhão, and Rafael Domingo. 2016. U-series dating of Paleaolithic rock art at Fuente del Trucho (Aragón, Spain). Quaternary International: http://dx.doi.org/10.1016/j.quaint.2015.11.111 Lane, M.A. 1996. Roles of natural history collections. Ann. Missouri Bot. Gard. 83: 536–545. Lozoya, X. 2006. Natural History and Herbal Medicine in Sixteenth-Century America. In: Saldaña, J.J. (ed.) and B. Madrigal (translator). Science in Latina America. University of Press. pp. 29–50. McKissack, P. and F. McKissak. 1994. The Royal Kingdoms of Ghana, Mali, and Songhay: Life in Medieval Africa. Henry Holt & Co., New York. Meyer, E.H.T. 1863. On the origin of herbaria. J. Bot. 1: 297–302. Middleton, E.S., G.M. Smith, W.J. Cannon, and M.F. Ricks. 2014. Paleoindian rock art: establishing the antiquity of Great Basin Carved Abstract petroglyphs in the northern Great Basin. J. Arch. Sci. 43: 21–30. Moggi, G. 2009. L’Erbario di Andrea Cesalpino. In: Raffaelli, M. (ed.). Il Museo Di Storia Naturale dell’Università degli Studi di Firenze. pp. 65–71. Nesbitt, M. 2014. Use of herbarium specimens in ethnobotany. In: Curating Biocultural Collections. Eds. Salick, J., K. Konchar, M. Nesbitt. Ch. 22, pgs. 313–328. Newman, D.J., G.M. Cragg, and K.M. Snader. 2000. The influence of natural products upon drug discovery. Nat. Prod. Rep. 17: 215–234. Pike, A.W.G., D.L. Hoffman, M. García-Diez, P.B. Pettitt, J. Alcolea, R. De Balbín, C. González- Sainz, C. de las Heras, J.A. Lashers, R. Montes, and J. Zilhão. 2012. U-series dating of Paleolithic art in 11 caves in Spain. Science 336: 1409–1413. Ratté, M.L. 1972. Imperial looting and the case of Benin. M.A., University of Massachusetts. Riddle, J.M. 1981. Pseudo-Dioscorides’ Ex herbis femininis and Early Medieval Medical Botany. 14: 43–81. Ritter, D.W. and E.W. Ritter. 1976. Prehistoric pictography in North America of medical significance. In: Grollig, F.X. and H.B. Haley (ed.). Medical Anthropology. Mouton Publishers, The Hague, Paris. pp. 137–228. Scarborough, J. 1977. Nicander’s toxicology I: Snakes. Pharmacy in History 19: 3–23. Schilthuizen, M., C.S. Vairappan, E.M. Slade, D.J. Mann, & J.A. Miller. 2015. Specimens as primary data: Museums and “open science”. Trends in Ecology & Evolution 30: 237–238 . Shetler, S. G. 1969. The Herbarium: Past, Present, and Future. Proc. Biol. Soc. Wash. 82: 687– 749. Simmons, E.G. 1978. Proceedings of the Second International Mycological Congress, University of South Florida, Tampa, Florida, USA 27 August – 3 September 1977. Mycologia 70:

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253–265. Singleton, B.D. 2004. African bibliophiles: Books and libraries in medieval Timbuktu. Libraries & Culture 39: 1–12. Smith, J.P. 2017. The Herbarium. Humboldt State University, California. Thackeray, A.I. 1983. Dating the rock art of southern Africa. The South African Archaeological Bulletin Goodwin Series 4: 21–26. Thiers, B. (ed.). 2016 [access date, but continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg.org/ih/ Thompson, R. C. 1924. The Assyrian Herbal. Luzac and Co., London. van Andel, T. 2017. Open the treasure room and decolonize the museum. Universiteit Leiden. [https://www.universiteitleiden.nl/binaries/content/assets/science/ibl/oratie-van- andel.pdf] Van Steenis-Kruseman, M.J. 1979. The collections of the Rijksherbarium. Blumea 25: 29–55.

SPECIMEN PROTOCOLS

COLLECTING

The plant data on the Atlas of Florida Plants (http://www.florida.plantatlas.usf.edu/) rely on well- made herbarium specimens for physical documentation of each species in each county. Generally photographs are not used because only physical specimens can be further studied and verified (e.g. microscopic characters, DNA, etc.).

The knowledge base of the Atlas of Florida Plants has relied upon the discoveries of 1000s of curious naturalists. Specimens are typically sent pressed and dried and placed in a sheet of folded newspaper. If preferring to send a fresh specimen, they generally have to be placed in a bag (to retain moisture) and shipped overnight. Then the herbarium can dry the specimen.

The specimen should be as informative as possible and include notes on the location (e.g. latitude & longitude), habitat, name of the collector(s), and the date of collection. The specimen(s) altogether should be about 11 × 17 inches.

Below is a concise outline of things to consider when collecting plant specimens. For examples, browse the USF herbarium specimen database: http://florida.plantatlas.usf.edu/Specimen.aspx. For additional guidelines, please refer to: http://www.flmnh.ufl.edu/herbarium/voucher.htm.

Lastly, collecting activities should not jeopardize a species or its diversity, such that it could cause local extinction. Generally, less than 5% of a population should ever be collected in one location for scientific research.

“Whatever is worth doing at all is certainly worth doing well. Neatness is the secret of success in all botanical work. Every step in the several processes of collecting, preserving, and mounting and plants should be most carefully considered, since lack of attention at a single point

46 will often destroy the entire value of a season’s work.” –Edward Thompson Nelson in Herbarium and Plant Descriptions (1895)

1. PRIOR TO COLLECTING a. Permission: from landowner/land manager/supervisor b. Prohibited plants: noxious weeds (5B-57.007), endangered & commercially- exploited plants (5B-40.0055), prohibited aquatic plants (5B-64.011); all require state permit c. Reasons to collect: contribute knowledge to society, as hobby, to know your plants d. Collecting philosophy: minimize disturbance, collection not detrimental to population e. Tools: plant press, spacers (cardboard), appropriately sized newspaper (~40 x 29 cm), pruners, digging utensil f. Precautions: companion, cell phone, water, food, protective clothing, inform others of your location or plans

2. WHERE TO COLLECT a. Paths/Edges/Roadsides: often have great diversity (natural habitat merging with disturbed habitat) b. Interior pristine/natural habitat: often has rare or unique taxa

3. WHAT SPECIMENS TO COLLECT a. Uniqueness: consider the value of the specimen–what is unique/valuable about this specimen?, range extension?, new to area?, under-collected area?, rare/unknown species?, characters rare/unknown on most specimens from species? (e.g. rarely collected for many species), inventory of an area? b. Duplicates: if especially valuable or unique, making multiple specimens for distribution to multiple herbaria is especially desirable. Herbaria almost always welcome well-made collections.

4. COLLECTING THE SPECIMEN a. Selection: specimen should exemplify maximum amount of information while being as compact as possible for storage and study (e.g. , fruits, buds, branching, roots, ) b. Notes: note characters unable to display on specimen (color, odor, bark, branching, height, orientation), photo often useful & can be attached with specimen c. Habitat: note the habitat: i.e. why is the plant here?, ruderal-disturbed area?, habitat edge?, roadside/path?, natural vegetation? hydric, xeric? cultivated?; utilize the Florida Natural Areas Inventory guide.

5. SPECIMEN COLLECTION – SPECIAL HANDLING a. Dissection: some plant parts should be dissected to expose internal characters before drying (e.g. flowers to show /pistils, fruits to show seeds or carpels) b. Delicate Aquatics: often should be floated in water directly onto archival paper

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to avoid clumping (e.g. confervoides, various algae) c. Delicate Plant Parts: can be dried directly onto archival paper (e.g. flowers of Convolvulaceae, , plants of Lentibulariaceae) d. Tall/long Plants: fold them to fit onto standard herbarium sheet e. Succulent Parts (succulents/fruits): dissect to expose interior and hasten drying; if succulence is copious, remove majority of succulent tissue and leave exterior (e.g. , Aloe, various cacti, fleshy fruits) f. Bulky Plants: if plant parts are larger than standard herbarium sheet, make multiple sheets from one collection (e.g. various palms) g. Bulky Plant Parts: if plant parts have considerable 3-dimensional shape which is to preserved, use a foam sheet on top of the newspaper holding the specimen; the foam will expand to press and retain the leaf shape but contract around the 3-dimensional object (e.g. acorns, hickory nuts, various fruits) h. Roots: brush or wash roots to remove soil (soil may be packeted for later study, too) i. Parasitic Plant: include host plant with collection j. Individuals: herbaceous habitats often have many plants growing in tight proximity to each other and individuals should be teased apart to reveal growth form k. Photos: photos can be included with the specimen to show additional features (e.g. the growth form of a palm)

6. POST-COLLECTION HANDLING a. Press: immediately press specimen, avoid wilting or shriveling; arrange specimen neatly for ease of study (specimen can be placed in a container to retain moisture until pressing) b. Dry: maximize air-flow, minimize humidity, minimize microbial decay, low heat often used; do not microwave, do not use excessive heat, or do not use chemical preservatives (they can degrade molecules such as DNA) c. Mounting: use archival acid-free ink & paper, archival glue, Archer’s glue, Elmer’s Glue, or straps & strings; if sending to herbarium do not mount, the herbarium will mount based on their best practices

7. SPECIMEN LABEL (complete ASAP before the specimen is neglected) a. Concise: be as brief as possible yet avoid ambiguity; provide details but avoid being too verbose b. Location: country, state, county, city or nearby city, water bodies, mountain range, roads (consider near-permanent reference points), latitude & longitude c. Identification: genus species if known, or family, or leave blank (can be identified years later) d. Plant Habit: characters not evident on specimen (e.g. color, odor, bark, branching, height, orientation, submergent/emergent) e. Habitat: ask ‘Why is this plant growing here?’ I.e. is the habitat hydric, mesic, or xeric? gravel, sand, or clay soil? forest, shrubland, grassland, disturbed? burned? cultivated? f. Your Name (and additional persons): Full name with middle initial is standard g. Collection Number: typically starts with 1 and proceeds sequentially

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h. Date: day, month (spelled out, not as a number), year i. Timeliness: make the label ASAP. Thousands of specimens in herbaria are thrown away because there is no label (only a dried plant without any written information).

MOUNTING Plants are glued to archival paper with archival labels. The glue prevents plant parts from moving around and being broken off and also allows easy study. For frugality, USF has often used Elmer’s glue, a polyvinyl acetate (PVA) glue, although it is thought to be slightly acidic. For strictly archival standards, other PVA-based glues with a neutral pH and more flexibility once dry may be preferable.

The archival paper is 100% cotton and acid-free to prevent degradation over time. It is 11.5″ × 16.5″ and 0.015 caliper. For recent orders, paper from Herbarium Supply Co. (Bozeman, MT) has tended to be thicker and more durable than paper from St. Louis Paper & Box Co. (St. Louis, MO).

BASIC OVERVIEW at USF 1. Dried plant specimens are found in newspaper and consist of two items: a) the dried plant pieces b) the label

2. Elmer’s glue is used to adhere the plant to the mounting paper. The glue gun is used for gluing the label and a fragment packet.

3. Then the glued specimen is brought to the mounting stacks, which consists of five items for each specimen, listed in order from bottom to top: a) the cardboard b) the mounted specimen c) wax paper d) metals weights e) wooden blocks

MOUNTING PROCEDURE 1. Place the mounting paper on the right side of the newspaper, exactly in the fold of the newspaper and do not allow it to move.

2. Stage the plant parts on the mounting paper with the label flush with the lower right corner. Display the plant on the paper in a way that maximizes the amount of accessible information. Generally, a) roots are situated near the bottom b) plant parts are placed so that some face up, some face down c) don’t hide flowers, fruits, or terminal buds under leaves d) individual plants or separate parts that are not attached to each other should be spaced apart on the paper (not glued in a messy, overlapping pile) e) plant parts should be as flat as possible on the paper (parts that are protruding off the

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paper will get broken off and damaged) f) loose or delicate material such as seeds, flowers for dissection, or extra leaves (e.g. for DNA extraction) are placed in a fragment packet

Check to make sure: a) plant parts are not overlapping with the label b) plant parts are not hanging off the edge of the mounting paper c) there is ample space below the USF logo in the upper right to stamp the accession number

If there any problems with the above, DO NOT MOUNT. Place the specimen aside and work on the next one. If there are no problems, continue to step 3:

3. Place the label aside. 4. Fold the newspaper onto the plant. 5. Grab the newspaper, plant, and mounting paper so that they all stay in position. 6. Flip them over so that the plant is now on the left side of the newspaper and turn the mounting paper over so it is on the right side of the newspaper and in the fold (exactly where it was at step #1). 7. Apply moderate glue (Elmer’s) to the back side of the plant parts. Be sure to apply to most of the plant, especially the ends of the plant parts. Large, bulky pieces require more glue. Thin, delicate parts require very little wisps of glue. 8. Flip the mounting paper and newspaper to the left and onto the glue and plant parts. 9. Press and smooth to distribute the glue evenly. 10. Grab the newspaper, plant, and mounting paper so that they all stay in position and flip them back over to the right. 11. If there is excess glue, dab it up with the newspaper. 12. Any loose pieces belong in a fragment packet. 13. Glue the fragment packet onto the mounting paper with the glue gun. 14. Glue the label in the bottom right with the glue gun. a) apply glue to the upper half of the label b) then place the upper half of the label in the bottom right c) slide it upward to smear the glue across the whole label d) you only have a few seconds to reposition the label as the glues dries fast e) do not continue tinkering with the label as your skin oils and the glue will combine to smudge the label ink

MOUNTING STACKS Be sure to know which stacks are dry and which are wet. It takes about a full 12–24 hours for all of the glue to have time to dry completely. Up to 3 stacks of specimens can be mounted in a day. A new stack (at the far left or right end, not in the middle) should be started for the first specimen of the day. 1. Place the specimen with wet glue on a piece of cardboard. 2. Place wax paper on the specimen. 3. Fold the corner of the wax paper so it does not touch the label (could smear the ink).

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4. Obtain metal weights from the adjacent stack, from a dry specimen. 5. Place metal weights as densely as possible on all plant parts and fragment packet. 6. Place 6 wooden blocks. Four set neatly at the corners. Two set neatly at the middle left and right edges. The smooth sides of the blocks face up or down. The rough edges are on the sides. 7. Place cardboard on top of the 6 blocks. 8. Make sure the stacks are straight and the blocks are uniform in their placement in the stacks. 9. You are now ready to glue the next specimen.

MODIFIED ARCHER’S GLUE FORMULA We use of a variation of the Gray Herbarium modified Archer’s formula glue (Archer 1950; Rollins 1955) for adhering the label and making a kind of plastic for “strapping” bulky plant parts. To make this glue: we mix 880 mL toluene, 220 mL methanol, 75 mL Dow Resin 276 V-2 (alpha- methylstyrene polymer [with methyl abietate?]), and usually 1000 mL (~250 g) of ethyl cellulose (viscosity ~10 cps). Oddly, ethyl cellulose seems to vary with density and color, being clear to brown in solution; so 1000 mL may not always be the proper amount. Toluene and methanol function as the solvent to fluidize the mixture for application, which evaporate after application. The Dow Resin is a plasticizer that helps the ethyl cellulose to fuse together as a supple continuous film. The ethyl cellulose is latex like, but somewhat brittle without the plasticizer. This should all be mixed in a fume hood to avoid the volatile exposure to the solvents and avoid dust-like particulates of the ethyl cellulose. We typically make twice this amount at a time. This glue is dispensed in an oiler or oil can pump to facilitate easy application and minimize volatile exposure.

References Archer, W.A. 1950. New plastic aid in mounting herbarium specimens. Rhodora 52:298–299. Hall, H.S., K.D. Lillie, & R.E. Pondell. s.d. Comparison – Aqueous vs. Solvent based Ethylcellulose films. 80-2: 1–8. http://www.coatingplace.com/images/stories/pdf/80- 2%20Comparison%20Aqueous%20vs.pdf Rollins, R.C. 1955. The Archer method for mounting herbarium specimens. Rhodora 57:294– 299.

DIGITIZATION

SPECIMEN PREP: 1. Assess the need for making additional folders. 2. Confirm correct name is shown, as reflected on the Atlas website (or Tropicos, , IPNI). 3. Confirm accession number is legible alongside words “University of South Florida Herbarium”. 4. Dot with archival red ink the county/parish/province/state location on map, if present on label. 5. Repair any loose material (e.g. re-glue).

DATABASING (identify, parse, and enter the following in order): 1. USF accession number. 2. Primary collector (recognition field by surname).

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3. Additional collectors (initials and surname, e.g. “J.R. Clark, G.R. Jerry”) 4. Collection number (“s.n.” if no number is given) 5. Date by day, 3-letter month, & year; e.g. “21 Jun 2002” or “Sep 1893”; (“s.d.” if no date given) 6. Plant description (abundance, habit, characters; e.g. “Frequent to 1.5 m tall; flowers white; leaves chartaceous.”). 7. Habitat (e.g. “Sand pine [] - turkey oak [Quercus laevis] scrub along river; with Polygala, Thysanella [Polygonella], Crinum.”). 8. Reproductive state (recognition field by “flower”, “bud”, “”, “immature fruit”, or simply enter “fertile” if difficult to distinguish by eye). 9. Location, see Appendix 2 (recognition field by Country, State, County, Named Place [only used for long names such as “Hillsborough River State Park”]) 10. Detail location (e.g. “Ca. 3.2 mi. N of C-734, 0.5 km W of FL 21, just S of I-4, near US 27 and Christmas; Alafia River; behind railroad tracks next to fence. T32S, R31E, Sec. 24, NW¼ of SW¼ of NE¼ of NE¼.”). 11. Lat/long (only use the upper row, ignore bottom row) 12. Elevation

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Figure 51. Database program for transcribing label data.

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Figure 52. Database program for transcribing determinations/identifications.

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FORM FOR DATABASE m, km meters, kilometers mi., ft., in., yds. miles, feet, inches, yards 0.25, 0.33 always use numbers for distances (not 1/4 or 1/3) ¼, ½ Only for Township, Range, and Section (Alt + 0188 or 0189) ca. For circa, about, or approximately. C-733 For county roads, use C- PR-605 For parish roads, use PR- I-75 For interstate highways, use I- FL 70 For state highways, use 2-letter state abbreviation, such as FL US 41 For federal highways, use US St. Saint NW, N, SSW directions (e.g. northwest, north, south-southwest)

Spell out all other abbreviations found on labels in locality field. Street St. Road Rd. Avenue Ave.

DETERMINATION PAGE (almost always entered chronologically, however must enter most recent currently accepted determined name last if not most recent) 1. Family (recognition field) 2. Genus (recognition field) 3. Species 4. First Author (recognition field by surname) 5. Second Author (recognition field) 6. Infraspecies (recognition field; “subsp.”, “var.”, “fma.”) 7. Infraspecies first author (recognition field) 8. Infraspecies second author (recognition field) 9. Determiner/identified by (recognition field) 10. Date

Notes: “Adv. Filter” button on Determination tab allows one to filter for specimens by taxonomy.

Right-click on the “Barcode number” field to select a particular specimen by accession number.

The program will prompt one to drop changes to any fields (excluding determinations) made (if the changes are made after the day the specimen entry was made). After the prompt, then changes can be made and saved (by pressing the button to proceed to the next specimen).

If one uses the simple arrow (not the arrow with asterisk) to make a new entry, then the previous determination will be copied into the new entry (however, one must edit the entry to save it, at least by putting the cursor into a blank author field and pressing the “delete” button).

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PHOTOGRAPHY 1. Turn on both flashes and camera 2. Open Digital Photo Professional, open EOS Capture 3. Take photo (press the spacebar) with ruler on specimen (confirm focus and framing on first image) 4. Mark a circle with x in the upper left corner

BATCH PROCESS IMAGES: 1. Open FastStone 2. Tools, Batch Convert Selected Images (F3) 3. Add All 4. Settings (JPEG quality 75) 5. Check Advanced Options, Rotate 90 6. Open image in desktop folder in FastStone 7. Double-click for full-screen and Ctrl+s for save-as the accession number in a subfolder

CAMERA SETUP: We use a Canon 5D mounted on a stand in a fixed position (ca. 72 cm above the specimen), so that we can set up the manual focus once and not worry about it (set near 0.72 m on the focus lens). The camera is pointed down. The height (or zoom) of the camera is set so that the width of the specimen fits perfectly in the photo, which usually yields about an inch or two of extra space at the top and bottom of the photo. The specimens are photographed on black cloth. We don't crop out the black cloth that shows in the background in the extra space of the photo.

We have two strobelights (JTL Versalight D-301 or D-501) to provide a flash on the top and bottom half of the specimen. One is hooked up to the camera (set to 26–30% flash) and the other operates as a slave (set to 24–30%). The camera is operated through Digital Photo Professional software using its EOS Capture. We use a high f-stop (f14) for better depth of field to keep things like large pine cones in focus. A higher F-stop (lower aperture) increases depth of field (that which is in focus), but also allows less light in, making a darker picture. A lower F-stop does the opposite, of course. The exposure time is kept at a short exposure (1/100th of a second). The iso is 100. AWB is auto balance on the EOS settings. Custom white balance results in more blue or more red; auto is best it seems. White balance is achieved through settings within the camera itself. Select the whitest paper one can find and photograph through the settings within the camera. Resolution was once set to S for small file in the EOS window (for the first 50,000 or so images). The resolution is now set to L for larger file sizes with higher zoom/resolution.

We put some sort of guiding paper down so we know where to line up the specimen with the camera's field of view. Then we snap the photo and save the jpg. Our method is to just snap the photo, rotate it, and resave it with our accession number. No cropping or realigning is done to save time. The resolution is lower than tiff but we feel that the jpg resolution and file size is more efficient for these purposes. The need for more resolution usually means the specimen requires examination under a dissecting scope and may need to be loaned to another herbaria for further study. Once it's all setup, the photographing is a quick process for us.

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The camera has been repaired twice since 2006. The mirror lens had to be re-glued, a very cheap repair. The ports had to be replaced, also not very expensive. Both D-301 strobelights have been replaced once over the course of ~120,000 photos.

MORPHOLOGICAL PLANT PHOTOS (for the Atlas) 1. Open in GIMP 2. Windows, Recently Closed Docks (to recover toolbox, layers) 3. Tools, Transform Tools, Crop: crop photo to frame plant or parts of interest 4. Image, Scale Image, pixel resolution=72, pixel width: a. 850-650 for Landscape b. 750-550 for Square c. 700-450 for Portrait d. 500-300 for Tall, slender 5. Layer, New Layer (up to 3 layers for Taxon Name, Photographer, Black Box) 6. Tools, Text, color pure yellow (R=255, Green=255, Blue=0) a. Taxon name (Genus species, ex. “Bidens alba”) is in Times new roman, bold italics, 24–26 pt. b. Photographer name is 14–16 pt, (Ex. “Photo by Keith Bradley”) or drag image onto background (see below) 7. (optional). Tools, Selection Tools, Rectangle Select. Edit, Fill with Foreground Color (black). Use only if names are illegilble because they blend in with photo. 8. Tools, Transform Tools, Move: to move layers around 8. Image, Merge Visible Layers 9. File, Export as “Genus_species.jpg”, “Genus_species2.jpg”, etc. 10. Quality 100.

SAVING LAYER AS IMAGE FOR REUSE(an example) 1. Using Photoshop, load background image at appropriate size (step 5 above) 2. Create text layer (step 6b above) (Ex. “Photo by Keith Bradley”) 3. Right-click layer and select “Duplicate layer” then for “Document” select “New” 4. “Image, Trim”, “Transparent Pixels” for new document 5. “File, Save for Web” 6. “PNG-24, Transparency, Interlaced, Embed Color Profile”, “Metadata: None”, uncheck “Convert to sRGB” 7. Now this image can be dragged & dropped onto background plant image at appropriate size (steps 5-6b above)

DUPLICATING LAYER FOR SAME PLANT SPECIES 1. Using Photoshop, load all images of same plant species 2. Edit all to make appropriate sized image (steps 2-5 above) 3. Create layers from steps 6a, 6b (7 optional) on the first image 4. Select all layers needed for duplication for additional images 5. Right click layers and select “Duplicate layer” then for “Document” select the desired next image

IMAGE CONSIDERATIONS

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1. CROPPING: Maximize the information in the frame of the photo a. Maximize frame size to include plant parts and include peripheral objects to give some indication of size and scale, and habitat context b. Minimize frame to exclude other plants that could be confused as the focal plant or to exclude other uninformative objects c. Minimize frame to enlarge details (like seeing fine hairs on stems) 2. ADJUSTMENTS: Only use “Shadows/Highlights” a. Maybe one in five typically are improved with up to a 7% change (rarely, up to 25– 50% change can be applied to severely bright/dark images)

PLANT/FUNGAL DNA EXTRACTION & PCR

1. Make a stock buffer of 100 mM Tris-base (not Tris-HCl), 1.4 M NaCl, and 20 mM EDTA. Per 100 mL is 8.18 g NaCl, 0.76 g EDTA, and 1.21 g Tris-base. Add HCl to obtain a pH of 8.0. Approximately 7 mL of 9:1 diluted HCl (9 mL water to 1 mL of HCl with a stock concentration of 35–38%). This stock buffer can be stored long-term.

2. Add the following ingredients fresh before use. Per 10 mL of stock buffer, add 0.2 g CTAB, 0.4 g PVP (avg. mol. wt. 40,000), and 14 mg N-phenacylthiazolium bromide (PTB). Let dissolve for 1–2 hours or use low heat to dissolve more quickly. Then add 50 µL 2-mercaptoethanol, 25 µL proteinase K (20 mg/mL), and 2 µL RNase A (100 mg/mL). Final concentration of proteinase K should be 50 µg/mL and RNase A 20 µg/mL. 10 mL of CTAB buffer is enough for 20 extractions.

3. The final concentrations of the CTAB buffer are 100 mM Tris-base, 1.4 M NaCl, 20 mM EDTA, 2% (m/v) CTAB, 4% (m/v) PVP, 0.5% (v/v) 2-mercaptoethanol, 5 mM PTB, 0.0005% (m/v) proteinase K, and 0.00000002% RNase.

4. Weigh out 10–20 mg of dried plant tissue (preferably crushed with liquid nitrogen) and put into a 1.5 mL tube. Exceeding 20 mg may cause overabundance of polysaccharides and downstream difficulties. PTB may be included for herbarium specimens (to free DNA cross-linked to proteins).

5. Add 500 µL CTAB buffer and incubate for 3 hours in a 55°C water bath. Ensure whole sample is exposed to buffer. Sample may be vortexed. Shorter or longer incubations may be used.

7. Add 500 µL 24:1 chloroform:isoamyl alochol and vortex. (24:1 chloroform:octanol should also work.)

8. Centrifuge for 7 minutes. Carefully, slowly pipette top aqueous phase into a new 1.5 mL tube (avoid drawing in mucousy strands of polysaccharides), discard the bottom two layers (polysaccharides, denatured proteins, chloroform and dissolved lipids).

9. Repeat step 7 and 8 to further remove and precipitate polysaccharides. (Can be repeated again if necessary.)

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10. Add 0.08 volumes of 7.5 M ammonium acetate (usually about 30 µL) and 0.60 volumes of isopropanol (usually 300 µL). Mix. (Longer, colder incubation may increase co-precipitation of polysaccharides.)

11. Centrifuge for 5 minutes and discard (decanting is fine) supernatant. Precipitated pellet is DNA. Pellet may be somewhat translucent.

12. Add 700 µL of 70% ethanol and mix to dislodge pellet. Centrifuge for 5 minutes and discard supernatant. Salts are removed here. Pellet should be more visible here.

13. Add 700 µL of 100% ethanol and mix to dislodge pellet. Centrifuge for 5 minutes and discard supernatant.

14. Dry the pellet in a vacuum centrifuge under medium heat for 2–3 minutes. Samples can also be air dried for 30–60 minutes.

15. Dissolve pellet in 65 µL of TE buffer.

16. Aliquot 60 µL for storage and subsequent use in 4°C. Samples stored in 4°C should work months later. Samples can be pipetted up and down to ensure DNA is mixed in solution for the aliquot. Store the rest in -20°C for later use.

Notes: There a couple untested things which may not matter. For example, longer incubation with CTAB buffer may yield more DNA. Shorter times can be used; some protocols do it for an hour only.

TE Buffer 10 mM Tris, 0.1 mM EDTA, pH 8.0

PCR 20 µL volume 1 unit IDProof DNA polymerase (ID Labs, London, Ontario, Canada) ($150) 2 µL 10X IDPol buffer 3mM MgCl2 250 µM dNTPs ($28) 120 ng of each primer ~40 ng DNA

20uL volume Water=14.72 µL MgCl=1.2 µL (of 50mM MgCl2) buffer=2 µL (of 10X buffer) dNTP=0.4 µL (of 250mM dNTPs) primer=0.24µL (of 0.5ug/uL primer stock [e.g. 0.38 mg=380 µg primer in 760uL=0.5 µg/µL]) Taq=0.2 µL (of 5U/µL)

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1 µL DNA (~40ng)

Make master mix of all reagents except DNA. Mix (vortex) and aliquot to PCR tubes (0.2mL tubes). Lastly, add 1uL DNA to each labelled tube.

THERMOCYCLER

All PCR cycles followed these cycle parameters with two variables, annealing temperature (x) and extension time (y). The lid temperature is 103°C and not preheated. Initial denaturation is typically 3 minutes at 94°C. Then 40 cycles of denaturation at 94°C for 45 seconds, annealing at x°C for 45 seconds, followed by extension at 72°C for y seconds. A final extension step of 2 minutes at 72°C completes the PCR. Samples can be held at 10°C in the thermocycler or in a refrigerator at 4°C until analysis by gel electrophoresis. The extension times could probably be lessened.

GEL ELECTROPHORESIS 0.2 g Bromophenol Blue 6 ml 50% glycerol 4 ml milipore water

TAE Buffer 50X Modified TAE Stock Solution For each litre of solution: 242 g Tris Base (MW=121.1) 57.1 mL Glacial Acetic Acid 10 mL 0.5 M EDTA mix Tris with stir bar to dissolve in about 600 mL of ddH2O. add the EDTA and Acetic Acid, pH to 8.0. bring final volume to 1 L with ddH2O. store at room temperature. Note: Final (1x) working concentration : 0.04 M Tris - Acetate 0.0001 M EDTA

AUTOCLAVING

Put liquids in a autoclavable bin. Add water to bin, up to about ¼ of the height of containers in bin. Also, for 100mL, for example, not a bad idea to add 5-10 mL extra to jar (as 5-10mL evaporates during autoclaving).

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SELECTED WORKS UTILIZING USF HERBARIUM RESOURCES (including research based on USF herbarium specimens, Atlas of Florida Plants citations, & USDA citations for Florida records based on the Atlas)

[list not comprehensive, especially incomplete for works 1965–1989]

1970 Wagner-Merner, D. Te Strake, G. Griepenburg, and K. Tyson. Mosses of the Tampa Bay Area. University of South Florida. 56 pp.

1973 Andorfer, J. Autoecological and synecological study of Swietenia mahagoni (Meliaceae). Ph.D. dissertation, University of South Florida, Tampa.

1974 Carlton, J.M. An ecological survey of selected communities in Florida. M.S. thesis, University of South Florida, Tampa. Wunderlin, R.P. Woody plants of the Tampa Bay area. University of South Florida, Tampa. 1–49.

1975 Cheney, D.P. A biosystematic investigation of the red algal genus Eucheuma (Solieriaceae). Ph.D. dissertation, University of South Florida, Tampa. Cole, D.P. Vegetational analysis of selected coastal Indian shell mounds in Florida. M.S. thesis, University of South Florida, Tampa.

1976 Poppleton, J.E. A Chromatographic and morphometric investigation of variation within peninsular Florida scrub oaks (with special emphasis on Q. chapmanii and Q. virginiana var. maritima). M.S. thesis, University of South Florida, Tampa. Small, E. 1976. A numerical taxonomic analysis of cannabis with special reference to species delimitation. Syst. Bot. 1:67–84. Sreemadavan, C.P. Leaf architecture and systematics of Acanthaceae and related families. Ph.D. dissertation, University of South Florida, Tampa.

1978 Cooper, S.T. Floristic assessment of Mound Key, Lee County, Florida. M.S. thesis, University of South Florida, Tampa. Hilsenbeck, R.A. A taxonomic and biosystematic investigation of Dyschoriste Nees (Acanthaceae) in Florida. M.S. thesis, University of South Florida, Tampa.

1979 Arcuri, A. A floristic survey of the vascular plants of Hillsborough River State Park. M.S. thesis, University of South Florida, Tampa.

1980

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Wagner-Merner, D. Te Strake. Bryophytes of Hillsborough River State Park, Florida. I. Mosses. Castanea 45: 97-100. Eilers, F.I., D. Te Strake Wagner-Merner and J.A. Kimbrough. Rare occurrences of large mushrooms {Tricholoma)sp. Florida Scientist 43: 50-54.

1982 Ragan, M.E. and R.P. Sauleda. A new natural hybrid of Oncidium Sw. from . American Orchid Society Bulletin 51: 388–390. Young, B. The palm genus Nengella Becc. M.S. thesis, University of South Florida, Tampa.

1984 Sauleda, R.P. A monographic study of the Caribbean genus Psychilis Raf. (Orchidaceae). Ph.D. dissertation, University of South Florida, Tampa. Te Strake, D. and H. Aldrich. Ultrastructure of two associations involving marine fungi and green algae. Botanica Marina 26: 515-519.

1985 Hansen, B.F. A revision of Forsteronia (Apocynaceae). Ph.D. dissertation, University of South Florida, Tampa. Richardson, D.R. Allelopathic efects of species in the sand pine scrub of Florida. Ph.D. dissertation, University of South Florida, Tampa. Robinson, G. A revision of Fevillea (). M.S. thesis, University of South Florida, Tampa. Te Strake, D., R.B. Lassiter, I.A. Lassiter and D.A. Breil. Bryophytes from and adjacent shoreline plant communities of Tampa Bay, Florida. Florida Scientist 49: 31-39. Te Strake, D. Fungi, their occurrence and potential roles in the Tampa Bay Ecosystem. p. 305-327. Tampa Bay Area Scientific Information Symposium. Simon, J. and E. Esteves (eds.).

1986 MacDonald, L.A. The diet of the gopher tortoise Gopherus polyphemus in a sandhill habitat in central Florida. M.S. thesis, University of South Florida, Tampa. 1987 Botts, P.S. Abundance and distribution of herbaceous angiosperms in grass-sedge marshes of west- central Florida: the effect of seasonal water-level fluctuation. M.S. thesis, University of South Florida, Tampa. 1988 Kaczor, S. Effects of gopher tortoise (Gopherus polyphemus) disturbances on the microenvironment and vegetation of a Florida sandhill. M.S. thesis, University of South Florida, Tampa.

1990 Brenner, M., B.W. Leyden, and M.W. Binford. Recent sedimentary histories of shallow lakes in the Guatemala savannas. J. Paleolimnology 4:239-252. Clancy, K. E. and M.J. Sullivan. Distribution of the needle palm, Rhapidophyllum . Castanea 55:31-39.

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Dunn, C.P. and R.R. Sharitz. The history of Murdannia keisak (Commelinaceae) in the southeastern . Castanea 55:122-129. Essig, F.B. The Clematis virginiana complex in the Southeastern United States. Sida 14:49-68. Herwitz, S.R. and R.P. Wunderlin. Vascular plant species diversity on two barrier islands in southwest Florida. J. Coastal Research 6:311-322. Luteyn, J.L. The Plumbaginaceae in the flora of the southeastern United States. Sida 14:169-178. Sauleda, R.P., and R.A. Adams. The Orchidaceae of the Bahama Archipelago: additions, distributional extensions, and nomenclatural change. Brittonia 42:286-291. Smith, G.L. and W.S. Flory. Studies on henryae (). Brittonia 42:212- 220. Terrell, E.E. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 68:125-133. Ward, D.B. and F.C. Craighead. Deletions and restorations in the flora of southern Florida. Sida 14:287-304.

1991 Anderson, L.C. Paronychia chartacea ssp. minima (): a new subspecies of a rare Florida endemic. Sida 14:435-441. Bornstein, A.J. The Piperaceae in the southeastern United States. J. Arnold Arbor., Suppl. Ser. 1:349-366. Eilers, R. Revision of subgenus Bauhinia Section Schnella (Cercideae: Caesalpinioideae: Fabaceae). M.S. thesis, University of South Florida, Tampa. Medley, M.E. Notes on Cleistes and Triphora (Orchidaceae). Selbyana 12:102-103. Moore, M.O. Classification and systematics of eastern North American Vitis L. (Vitaceae). Sida 14:339-367. Wise, E. A taxonomic study of the Rosoideae (Rosaceae) in Florida. M.S. thesis, University of South Florida, Tampa.

1992 Austin, D.F. and J.N. Burch. Status of Stylisma abdita (Convolvulaceae) in southwestern Florida. Florida Sci. 55:99-102. Burckhalter, R.E. The genus Nyssa (Cornaceae) in North America: a revision. Sida 15:323-342. Essig, F.B. A new species of (Palmae) from New Britain. Principes 36: 4-6. Herndon, A. The genus Hypoxis (Hypoxidaceae) in Florida. Florida Sci. 55:45-55. Matthews, J.F., D.W. Ketron, and S.F. Zane. The reevaluation of Portulaca pilosa and P. mundula (Portulacaceae). Sida 15:71-89. Taylor, W.K. and Z.A. Prusak. Distributions of Florida sneezeweeds (Helenium), with a range extension for Helenium autumnale L. Florida Sci. 55:244-245.

1993 Bridges, E.L., S.L. Orzell, and J.R. Burkhalter. Cladium mariscoides () in the western Florida panhandle and its phytogeographic significance. Phytologia 74:35-42. Essig, F.B. Clematis. In J.D. Hickman. The Jepson Manual: Higher Plants of California. p. 914. Berkley, Los Angeles, and London: University of California Press. Herndon, A. A revision of the Chamaesyce deltoidea (Euphorbiaceae) complex of southern Florida. Rhodora 95:38-51.

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Jones, S.D. and J.K. Wipff. Eustachys retusa (Poaceae), the first report in Florida and a key to Eustachys in Florida. Phytologia 73:274-276. Lambert, B. The effects of light, soil, disturbance, and presence of organic litter on the field germination and survival of the Florida goldenaster ( floridana Small). M.S. thesis, University of South Florida, Tampa. Leyden, B.W., M. Brenner, D.A. Hodell, and J.A. Curtis. Lake Pleistocene climate in the central American lowlands. Pp. 165-178. In P.K. Swart, K.C. Lohmann, J. McKenzie, and S. Savin, eds. Climate Change in Continental Records. Washington, DC: American Geophysical Union. Matthews, J.F., D.W. Ketron, and S.F. Zane. The biology and taxonomy of the Portulaca oleracea L. (Portulacaceae) complex in North America. Rhodora 95:166-183. Miller, H.A. A new species of Dicerandra (Lamiaceae) from Florida. Phytologia 75:185-189. Orzell, S.L. and E. L. Bridges. Eriocaulon nigrobracteatum (Eriocaulaceae), a new species from the Florida panhandle, with a characterization of its poor fen habitat. Phytologia 74:104- 124. Wunderlin, R.P. Callitris (Cupressaceae). In Flora of North America Editorial Committee. Flora of North America North of Mexico. 2:421-422. New York and Oxford: Oxford University Press. Wunderlin, R.P., B.F. Hansen, K.R. DeLaney, M. Nee, and J.J. Mulahey. Solanum viarum and S. tampicense (Solanaceae): two weedy species new to Florida and the United States. Sida 15:605-611.

1994 Leyden, B.W., M. Brenner, D.A. Hodell, and J.H. Curtis. Orbital internal forcing of climate on the Yucatan Peninsula for the past ca. 36 ka. Palaeogeography, Palaeoclimatology, Palaeoecology 109:193-210.

1995 Clemants, S.E. Bejaria (Ericaceae subfamily Rhodendroideae). In J.E. Luteyn. Ericaceae. Flora Neotropica Monogr. 66:54-106. Essig, F.B. A new species of Gronophyllum from the Bismarck Archipelago. Principes 39:100- 101. Hansen, B. F. Forsteronia. In J.A. Steyermark, P.E. Berry, B.K. Holst, and K. Yatskievch. Flora of the Venezuelan Guayana. 4:496-498. St. Louis: Missouri Botanical Garden Press. Hansen, B. F. Odontadenia. In J.A. Steyermark, P.E. Berry, B.K. Holst, and K. Yatskievch. Flora of the Venezuelan Guayana. 4:537-541. St. Louis: Missouri Botanical Garden Press. Manka, T. A survey and study of the fruit anatomy of the Iguanura alliance (). M.S. thesis, University of South Florida, Tampa. Taylor, Y. Comparative study of the distribution patterns and leaf surface anatomy of silver and green forms of saw palmetto, repens (Bart.) Small. M.S. thesis, University of South Florida, Tampa. Wunderlin, R. P., B. F. Hansen, and E. L. Bridges. [1995-2000]. Atlas of the Flora of Florida. University of South Florida, Tampa. (www.usf.edu.edu~isb)

1996

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Campbell, K.N. An analysis of epiphyte populations in naturally occurring forest patches in Myakka River State Park. M.S. thesis, University of South Florida, Tampa. Haddock, A.L. Fungi associated with frutescens (L.) DC. M.S. thesis, University of South Florida, Tampa. Herwitz, S.R., R.P. Wunderlin, and B.F. Hansen. Species turnover on a protected subtropical : a long term study. J. Biogeography 23:705-715. Landry, S. The importance of availability and microsite factors relating to the distribution of vascular epiphytes in west central Florida. M.S. thesis, University of South Florida, Tampa. Leyden, B.W., M. Brenner, T. Whitmore, J.H. Curtis, D.R. Piperno, and B.H. Dahlin. A record of long and short-term climatic variation from northwest Yucatan: Cenote San Jose Chulchaca. Pp. 30-50. In S. Fedick, ed. Maya Subsistence Resource Management. Salt Lake City: University of Utah Press. Sauleda, R.P. and M.E. Ragan. A new species of Tolumnia Raf. from Puerto Rico. Orchid Digest 60:183–187. Smith, G.L. and M.A. Garland. Taxonomic status of Hymenocallis choctawensis and Hymenocallis puntagordensis (Amaryllidaceae). Sida 17:305-319. Whitmore, T.J., M. Brenner, J.H. Curtis, B.H. Dahlin, and B.W. Leyden. Holocene climatic and human influences on lakes of Yucatan Peninsula, Mexico: an interdisciplinary, palaeolimnological approach. The Holocene 6:273-287. Wunderlin, R. P., B. F. Hansen, and E. L. Bridges. Atlas of the Flora of Florida: CD-ROM. Institute for Systematic Botany, University of South Florida, Tampa, FL.

1997 Moreno, N. and F.B. Essig. Clematis subgenus Clematis (Ranunculaceae). In Flora of North America Editorial Committee. Flora of North America North of Mexico. 3:159-164. New York and Oxford: Oxford University Press. Wunderlin, R.P. Central Highlands of Florida, U.S.A. In S.D. Davis, O. Herrera-MacBryde, J. Villa-Lobos, and A.C. Hamilton. Centres of Plant Diversity: A Guide and Strategy for their Conservation. 3:87-90. Cambridge, England: World Wide Fund for Nature & International Union for Conservation of Nature. Wunderlin, R.P. Moraceae. In Flora of North America Editorial Committee. Flora of North American North of Mexico. 3:388-399. New York and Oxford: Oxford University Press.

1998 Cochrane, C.B. Antibacterial and antifungal screening of Florida's exotic invasive plant species. M.S. thesis, University of South Florida, Tampa. Dawes, C.J. Marine Botany. Second Edition. New York: John Wiley and Sons Publ. 480pp. Hill, S.R. and A. James. 1998. New plant records for Dominica, Lesser Antilles. Sida 18:297–305. Lai, M., J. Sceppa, J.A. Ballenger, J.J. Doyle, and R.P. Wunderlin. Polymorphism for the presence of the rpl2 intron in chloroplast genomes of Bauhinia (Leguminosae). Syst. Bot. 22:519– 528. Leyden, B.W., M. Brenner, and B.H. Dahlin. Cultural and climatic history of Coba, a lowland Maya city in Quintana Roo, Mexico. Quarternary Research 49:111–122. Markham, L. The molecular ecology of the Florida goldenaster, Chrysopsis floridana (Asteraceae). M.S. thesis, University of South Florida, Tampa.

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Teasdale, B. The ultrastructure of the outer wall in three species of the red macroalga Gracilaria and attachment by the green epiphyte Ulva lactuca. M.S. thesis, University of South Florida, Tampa. Wunderlin, R.P. Bauhinia. In Steyermark, J. A., P. E. Berry, and B. K. Holst, K. Yatskievych. Flora of the Venezuelan Guayana. 4:5-13. St. Louis: Missouri Botanical Garden Press. Wunderlin, R.P. Guide to the Vascular Plants of Florida. Gainesville: University Press of Florida. 806 pp.

1999 Delaney, K.R., N. Bissett, and J. D. Weidenhamer. A new species of Carphephorus (Asteraceae: Eupatorieae) from peninsular Florida. Bot. Expl. (Florida) 1:1-15. Kearns, D. and R.P. Wunderlin. Fabaceae. In P.M. Jørgensen and S. León-Yánez, eds. Catalogue of the Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:414-418. Essig, F.B., Y.R. Taylor, and D. Te Strake. Florida’s wax palm: the silver form of Serenoa repens (Arecaceae). Florida Sci. 63: 13-16. Neill, D.A., B.B. Klitgaard, and G.P. Lewis. Caesalpiniaceae. In P.M. Jørgensen and S. León- Yánez, eds. Catalogue of the Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:369-374. Schmidt, T.J. 1999. Novel seco-prezizaane sesquiterpenes from North American Illicium species. J. Nat. Prod. 62: 684–687.

2000 Bridges, E.L. and S.L. Orzell. Rhynchospora megaplumosa (Cyperaceae), a new species from central Florida, with supplemental notes and a key to Rhynchospora series Plumosae. Lundellia 3:19-25. Erickson, A. Associational resistence from and susceptibility to herbivory by the mangrove tree crab, Atatus pisoni, on the red mangrove, Rhizophora mangle, when in the presence of the black mangrove, Avicennia germinans, and the white mangrove, Laguncularia racemosa. Ph.D. dissertation, University of South Florida, Tampa. Hansen, B.F. and R.P. Wunderlin. Two new combinations in Florida selaginellas. Phytologia 85:300-302. Myers, J. A floristic inventory of Little Manatee River State Park, Hillsborough County, Florida. M.S. thesis, University of South Florida, Tampa. Norman, E. M. Buddlejaceae. Flora Neotropica Monogr. 18:1-225. Wunderlin, R. P. and B. F. Hansen. Flora of Florida. 1:1-365. Gainesville: University Press of Florida. Wunderlin, R. P., and B. F. Hansen. [2000-2005]. Atlas of Florida Vascular Plants. [S. M. Landry and K. N. Campbell (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa. (www.plantatlas.usf.edu).

2001 Brown, M. Drift algae of seagrass beds in Sunset , Cockroach Bay, and Tarpon Bay on west coast of Florida. M.S. thesis, University of South Florida, Tampa. Huck, R.B. Two infraspecific taxa in Florida Dicerandra (Labiatae). Novon 11:417-420.

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Siedo, S.J. A new species of Sida sect. Elliptifoliae (Malvaceae). Lundellia 4:69-75. Smith, G.L., L.C. Anderson, and W.S. Flory. A new species of Hymenocallis (Amaryllidaceae) in the lower central Florida panhandle. Novon 11:233-240. Wunderlin, R.P. Bauhinia (Caesalpiniaceae). In W.D. Stevens, C.U. Ulloa, A. Poole, and O.M. Montiel, eds. Flora of Nicaragua.1:522-526. St. Louis: Missouri Botanical Garden Press. Wunderlin, R.P. and B.F. Hansen. Seven new combinations in the Florida flora. Novon 11:366- 369.

2002 Bridges, E.L. and S.L. Orzell. Euphorbia (Euphorbiaceae) section Tithymalus subsection Inundatae in the southeastern United States. Lundellia 5:59-78. DeLaney, K. R. and R. P. Wunderlin. A new species of Chrysopsis (Asteraceae, Astereae) from central Florida. Bot. Explor. (Florida) 2:1-20. Naczi, R.F.C., C.T. Bryson, and T.S. Cochrane. Seven new species and one new combination in Carex (Cyperaceae) from North America. Novon 12:508-532. Orzell, S.L. and E.L. Bridges. Notes on Carphephorus odoratissimus (Asteraceae) in peninsular Florida, U.S.A. Sida 20:559-569 Terrell, E.E., and R.P. Wunderlin. Seed and fruit characters in selected Spermacoceae and comparison with Hedyotideae (Rubiaceae). Sida 20:549-557. Wunderlin, R.P., B.F. Hansen, and L.C. Anderson. Plants new to the United States and Florida. Sida 20:813-817.

2003 Bridges, E.L. and S.L. Orzell. Two new species and a new combination in southwestern United States Xyris (Xyridaceae) from Florida. Novon 13:16-25. Burks, K.C., V.V. Vandiver, D.W. Hall, and C.C. Jacono. Rotala rotundifolia (Lythraceae) new to Florida. Sida 20:1765-1769. DeLaney, K.R., R.P. Wunderlin, and J.C. Semple. Chrysopsis delaneyi (Asteraceae, Astereae), another new species from peninsular Florida. Bot. Explor. (Florida) 3:1-37. Hansen, B. F. Forsteronia. In: S.A. Mori, G. Cremers, C.A. Gracie. J-J. de Granville, S.V. Heald, M. Hoff, and J.D. Mitchell. Guide to the Vascular Plants of Central French Guiana. Mem. New York Bot. Gard. 76(2):76-77. Kral, R. and G.L. Nesom. Two new species of Liatris series Graminifoliae (Asteraceae: Eupatorieae) from the southeastern United States. Sida 20:1573-1583. Myers, J.H. and R.P. Wunderlin. Vascular flora of Little Manatee River State Park, Hillsborough County, Florida. Castanea 68:56-74. Slomba, J. The phylogenetic potential of the actin I intron in Clematis (Ranunculaceae). M.S. thesis, University of South Florida, Tampa. Wunderlin, R. P. and B. F. Hansen. Guide to the Vascular Plants of Florida. Second Edition. Gainesville: University Press of Florida. 787 pp.

2004 Ferguson, E. Vascular plant inventory of Jay B. Starkey Wilderness Park, Pasco County, Florida. M.S. thesis, University of South Florida, Tampa. Nesom, G.L. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United

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McGill, B., Morueta-Holme, N., Peet, R. K., Violle, C., Wiser, S. and Svenning, J.-C. Patterns and drivers of plant functional group dominance across the Western Hemisphere: a macroecological re-assessment based on a massive botanical dataset. Bot. J. Linn. Soc. 180: 141–160. doi:10.1111/boj.12362 Galasso G., Domina G, Adorni M, Ardenghi NMG, Banfi E, Bedini G, Bertolli A, Brundu G, Calbi M, Cecchi L, Cibei C, D’Antraccoli M, De Bastiani A, Faggi G, Ghillani L, Iberite M, Latini M, Lazzeri V, Liguori P, Marhold K, Masin R, Mauri S, Mereu G, Nicolella G, Olivieri N, Peccenini S, Perrino EV, Peruzzi L, Petraglia A, Pierini B, Prosser F, Roma- Marzio F, Romani R, Sammartino F, Selvaggi A, Signorile G, Stinca A, Verloove F, Nepi C. Notulae to the Italian alien vascular flora: 1. Italian Botanist 1: 17-37. doi: 10.3897/italianbotanist.1.8777 Hammer, R.L. Central Florida Wildflowers: A Field Guide to Wildflowers of the Lake Wales Ocala National Forest, Disney Wilderness Preserve, and More than 60 State Parks and Preserves. Falcon Guides. Hammer, R.L. Everglades Wildflowers: A Field Guide to Wildflowers of the Historic [2016 ed.]. Falcon Guides. Overholt, W.A., M. Rayamajhi, E. Rohrig, S. Hight, F. Allen Dray, E. Lake, M. Smith, K. Hibbard, G.P. Bhattarai , K. Bowers, R. Poffenberger, M. Clark, B. Curry , B. Stange, E. Calise, T. Wasylik, C. Martinez, J. Leidi. Release and distribution of Lilioceris cheni (Coleoptera: Chrysomelidae), a biological control agent of air potato (: Dioscoreaceae), in Florida. Biocontrol Science and Technology Ross, M. S., J.P. Sah, P.L. Ruiz, A.A. Spitzig, A. A. & S.C. Subedi.Inferring implications of climate change in south Florida hardwood hammocks through analysis of metacommunity structure. Diversity Distrib. doi:10.1111/ddi.12442 Salvatore Pasta, S., E. Badalamenti , G. Sala , T. La Mantia. Nicodemia madagascariensis (Lam.) R. Parker (family Scrophulariaceae), a casual alien plant new to Italy. Webbia 71. Schmalzer, P.A. & T.E. Foster. Flora and Threatened and Endangered Plants of Canaveral National Seashore, Florida. Castanea Semple, J.C. & R. Lopez Laphitz. On Solidago gypsophila and S. pringlei (Asteraceae: Astereae), rare and not so rare Mexican endemics: A multivariate study of the Tortifolia group of subsect. Triplinerviae. Phytoneuron 2016-29: 1–20. Weremijewicz, J., Almonte, J. I., Hilaire, V. S., Lopez, F. D., Lu, S. H., Marrero, S. M., et al. Downing, J. L. Microsatellite primers for two threatened orchids in Florida: Encyclia tampensis and Cyrtopodium punctatum (Orchidaceae). Applications in Plant Sciences, 4 apps.1500095. http://doi.org/10.3732/apps.1500095 Wunderlin, R.P. & B.F. Hansen. Flora of Florida. Vol. III. Dicotyledons, Vitaceae through Urticaceae. University Press of Florida, Gainesville. Wunderlin, R. P., B. F. Hansen, A.R. Franck, and F.B. Essig. [2016]. Atlas of Florida Vascular Plants. [S. M. Landry and K. N. Campbell (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa. (www.florida.plantatlas.usf.edu).

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APPENDICES

Appendix 1. Standardized entries for geographic entities and administrative divisions in the USF Herbarium database. The database field ‘continental regions’ are specialized for emphasizing certain constituent collections within the USF Herbarium. Some places are given country status for convenience (e.g. Guam) since they are geographically distant from their associated country.

Continental Region Country - Autonomous 1st level (e.g. State) 2nd level (e.g. County) Region Africa Algeria Province (Prov.) District (Distr.) Africa Angola Province (Prov.) Municipality (Mun.) Africa Benin Department (Dept.) Commune Africa Botswana District (Distr.) Sub-district Africa Burkina Faso Region Province (Prov.) Africa Burundi Province (Prov.) Commune Africa Cameroon Region Department (Dept.) Africa Cape Verde Municipality (Mun.) Parish (Par.) Africa Central African Republic Prefecture Sub-prefecture Africa Chad Region Department (Dept.) Africa Comoros Island Municipality (Mun.) Africa Congo Province (Prov.) District (Distr.) Africa Djibouti Region District (Distr.) Africa Egypt Governorate Region Africa Equatorial Guinea Province (Prov.) District (Distr.) Africa Eritrea Region District (Distr.) Africa Ethiopia Region Zone Africa Gabon Province (Prov.) Department (Dept.) Africa Gambia Local government area District (Distr.) Africa Ghana Region District (Distr.) Africa Guinea Region Prefecture Africa Guinea-Bissau Region Sector Africa Ivory Coast District (Distr.) Region Africa Kenya County (Co.) Sub-county Africa Lesotho District (Distr.) Constituency Africa Liberia County (Co.) District (Distr.) Africa Libya District (Distr.) N/A Africa Madagascar Region District (Distr.) Africa Malawi Region District (Distr.) Africa Mali Region Cercle Africa Mauritania Region Department (Dept.) Africa Mauritius Island District (Distr.)

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Africa Morocco Region Prefecture and Province Africa Mozambique Province (Prov.) District (Distr.) Africa Namibia Region Constituency Africa Niger Region Department and Commune Africa Nigeria State Local government area Africa Réunion N/A N/A Africa Rwanda Province (Prov.) District (Distr.) Africa Sao Tome & Principe Province (Prov.) District (Distr.) Africa Senegal Region Department (Dept.) Africa Seychelles District (Distr.) Region Africa Sierra Leone Province (Prov.) District (Distr.) Africa Somalia Region District (Distr.) Africa South Africa Province (Prov.) District (Distr.) Africa South Sudan State County (Co.) Africa Sudan State District (Distr.) Africa Swaziland Region Inkhundla Africa Tanzania Region District (Distr.) Africa Togo Region Prefecture Africa Tunisia Governorate District (Distr.) Africa Uganda Region District (Distr.) Africa Zambia Province (Prov.) District (Distr.) Africa Zimbabwe Province (Prov.) District (Distr.) Antarctica N/A N/A N/A Afghanistan Province (Prov.) District (Distr.) Asia Bahrain Governorate N/A Asia Bangladesh Division District (Distr.) Asia Bhutan District (Distr.) Block Asia Brunei District (Distr.) Sub-district Asia Myanmar [Burma] State District (Distr.) Asia Cambodia Province (Prov.) District (Distr.) Asia Province (Prov.) Prefecture Asia East Timor District (Distr.) Sub-district Asia India State District (Distr.) Asia Indonesia Province (Prov.) Regency and city Asia Iran Province (Prov.) County (Co.) Asia Iraq Governorate District (Distr.) Asia Israel District (Distr.) Sub-district Asia Japan Prefecture Sub-prefecture Asia Jordan Governorate Nahia Asia Kazakhstan Region District (Distr.)

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Asia Kuwait Governorate Area Asia Kyrgyzstan Region District (Distr.) Asia Laos Province (Prov.) District (Distr.) Asia Lebanon Governorate District (Distr.) Asia Malaysia State District (Distr.) Asia Maldives Province (Prov.) Atoll Asia Mongolia Province (Prov.) Sum Asia Nepal Province (Prov.) District (Distr.) Asia North Korea Province (Prov.) County (Co.) Asia Oman Governorate Province (Prov.) Asia Pakistan Province (Prov.) Division Asia Philippines Region Province (Prov.) Asia Qatar Municipality (Mun.) N/A Asia Russia Subject District (Distr.) Asia Saudi Arabia Region Governorate Asia Singapore District (Distr.) Constituency Asia South Korea Province (Prov.) County (Co.) Asia Sri Lanka Province (Prov.) District (Distr.) Asia Syria Governorate District (Distr.) Asia Taiwan Province (Prov.) County (Co.) Asia Tajikistan Province (Prov.) District (Distr.) Asia Thailand Province (Prov.) District (Distr.) Asia Turkey Province (Prov.) District (Distr.) Asia Turkmenistan Province (Prov.) District (Distr.) Asia United Arab Emirates Emirate Municipality (Mun.) Asia Uzbekistan Province (Prov.) District (Distr.) Asia Province (Prov.) District (Distr.) Asia Yemen Governorate District (Distr.) Central America Belize District (Distr.) Constituency Central America Costa Rica Province (Prov.) Canton Central America El Salvador Department (Dept.) Municipality (Mun.) Central America Guatemala Department (Dept.) Municipality (Mun.) Central America Honduras Department (Dept.) Municipality (Mun.) Central America Nicaragua Department (Dept.) Municipality (Mun.) Central America Panama Province (Prov.) District (Distr.) Europe County (Co.) Municipality (Mun.) Europe Andorra Parish (Par.) Quarter Europe Armenia Province (Prov.) Municipality (Mun.) Europe Austria State Local government area Europe Azerbaijan N/A District (Distr.) Europe Belarus Region District (Distr.)

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Europe Belgium Region Province (Prov.) Europe Bosnia & Herzegovina Entity Canton Europe Bulgaria Province (Prov.) Municipality (Mun.) Europe County (Co.) Municipality (Mun.) Europe Cyprus District (Distr.) Municipality (Mun.) Europe Czech Republic Region District (Distr.) Europe Denmark Region Municipality (Mun.) Europe England Region County (Co.) Europe Estonia County (Co.) Municipality (Mun.) Europe Finland Region Sub-region Europe France Region Department (Dept.) Europe Georgia Region District (Distr.) Europe Germany State County (Co.) Europe Greece Municipality (Mun.) Region Europe Guernsey Jurisdiction N/A Europe Hungary County (Co.) District (Distr.) Europe Iceland Municipality (Mun.) N/A Europe Ireland County (Co.) District (Distr.) Europe Italy Region Province (Prov.) Europe Latvia Municipality (Mun.) City and parish Europe Liechtenstein Municipality (Mun.) N/A Europe Lithuania County (Co.) Municipality (Mun.) Europe Luxembourg District (Distr.) Canton Europe Macedonia Region Municipality (Mun.) Europe Malta Region Municipality (Mun.) Europe Moldova District (Distr.) Municipality (Mun.) Europe Monaco Municipality (Mun.) Quarter Europe Municipality (Mun.) Settlement Europe Netherlands Province (Prov.) Municipality (Mun.) Europe Northern Ireland District and County N/A Europe Norway County (Co.) Municipality (Mun.) Europe Poland Province (Prov.) County (Co.) Europe Portugal District (Distr.) Municipality (Mun.) Europe Romania County (Co.) Municipality (Mun.) Europe San Marino Municipality (Mun.) N/A Europe Scotland Council area N/A Europe Region District (Distr.) Europe Slovakia Region District (Distr.) Europe Slovenia Region Municipality (Mun.) Europe Spain Autonomous Province (Prov.) community

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Europe Sweden County (Co.) Municipality (Mun.) Europe Switzerland Canton District (Distr.) Europe Ukraine Region District (Distr.) Europe Vatican City N/A N/A Europe Wales County (Co.) N/A North America Canada Province (Prov.) Municipality (Mun.) North America Greenland Municipality (Mun.) N/A North America Mexico State Municipality (Mun.) North America USA State County (Co.) Oceania American Samoa District (Distr.) County (Co.) Oceania Australia State Local government area Oceania Fiji Division Province (Prov.) Oceania Easter Island N/A N/A Oceania French Polynesia Island Commune Oceania Kiribati Island N/A Oceania Guam Village N/A Oceania Marshall Islands Municipality (Mun.) N/A Oceania Micronesia Island N/A Oceania Nauru District (Distr.) N/A Oceania New Caledonia Province (Prov.) Commune Oceania New Zealand Region District (Distr.) Oceania Northern Mariana Municipality (Mun.) N/A Islands Oceania Palau State N/A Oceania Papau New Guinea Region Province (Prov.) Oceania Pitcairn Islands N/A N/A Oceania Samoa District (Distr.) N/A Oceania Solomon Islands Province (Prov.) N/A Oceania Tonga Division District (Distr.) Oceania Tuvalu Island N/A Oceania USA Hawaii Island Oceania Vanuatu Province (Prov.) Municipality (Mun.) South America Argentina Province (Prov.) Department (Dept.) South America Bolivia Department (Dept.) Province (Prov.) South America Brazil State Municipality (Mun.) South America Chile Region (or Island) Province (Prov.) South America Colombia Department (Dept.) Municipality (Mun.) South America Ecuador Province (Prov.) Canton South America French Guiana N/A Department (Dept.) South America Guyana Region Council South America Department (Dept.) District (Distr.)

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South America Peru Region Province (Prov.) South America Suriname District (Distr.) Resort South America Department (Dept.) Municipality (Mun.) South America State Municipality (Mun.) West Indies Anguilla Island District (Distr.) West Indies Antigua & Barbuda Island Parish (Par.) West Indies Aruba N/A N/A West Indies Bahamas Island District (Distr.) West Indies Barbados Parish (Par.) N/A West Indies Parish (Par.) N/A West Indies British Virgin Islands Island District (Distr.) West Indies Caribbean Netherlands Island N/A West Indies Cayman Islands Island District (Distr.) West Indies Province (Prov.) Municipality (Mun.) West Indies Curaçao N/A N/A West Indies Dominica Parish (Par.) N/A West Indies Dominican Republic Province (Prov.) Municipality (Mun.) West Indies Grenada Parish (Par.) N/A West Indies Guadeloupe N/A Department (Dept.) West Indies Haiti Department (Dept.) Arrondissement West Indies Jamaica County (Co.) Parish (Par.) West Indies Martinique N/A Department (Dept.) West Indies Montserrat N/A Parish (Par.) West Indies Puerto Rico N/A Municipality (Mun.) West Indies St. Barthélemy Department (Dept.) N/A West Indies St. Kitts & Nevis Island Parish (Par.) West Indies St. Lucia Quarter N/A West Indies St. Maarten N/A N/A West Indies St. Martin N/A N/A West Indies St. Vincent & the Island Parish (Par.) Grenadines West Indies Trinidad & Tobago Island Region West Indies Turks & Caicos Islands Island District (Distr.) West Indies U.S. Virgin Islands Island District (Distr.)

Appendix 2. County codes for maps in the Atlas of Florida Plants. County County Code Alachua ALA Baker BAK Bay BAY

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Bradford BRA Brevard BRE Broward BRO Calhoun CAL Charlotte CHA Citrus CIT Clay CLA Collier CLL Columbia CLU DeSoto DES Dixie DIX Duval DUV Escambia ESC Flagler FLA Franklin FRA Gadsden GAD Gilchrist GIL Glades GLA Gulf GUL Hamilton HAM Hardee HAR Hendry HEN Hernando HER Highlands HIG Hillsborough HIL Holmes HOL Indian River IND Jackson JAC Jefferson JEF Lafayette LAF Lake LAK Lee LEE Leon LEO Levy LEV Liberty LIB Madison MAD Manatee MAN Marion MAR Miami-Dade DAD Monroe Keys MOK Monroe Mainland MOM

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Martin MRT Nassau NAS Okaloosa OKA Okeechobee OKE Orange ORA Osceola OSC Palm Beach PAL Pasco PAS Pinellas PIN Polk POL Putnam PUT Santa Rosa SAN Sarasota SAR Seminole SEM St. Johns STJ St. Lucie STL Sumter SUM Suwannee SUW Taylor TAY Union UNI Volusia VOL Wakulla WAK Walton WAL Washington WAS