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Systematics, taxonomy, and the new flora of the Bahamian Archipelago.

Introduction

As part of the celebration of the thirtieth anniversary of the publication of the Flora of the Bahama Archipelago (Correll & Correll, 1982), it is informative to look back at the history of the study of the flora of the region, (which includes both the Bahamas and the Turks and Caicos Islands), and to look forward to the next phase in the study of the plants of the archipelago. In this paper, we present a comprehensive history of botanical exploration of the archipelago, show how the flora, as presented by Correll & Correll (1982), was compiled, and describe the explorations that have occurred since that date. This should provide an understanding of the need for a new, up-to-date Flora for the archipelago.

Many components go into the making of a flora: field work, specimens, historical writings on the flora, and literature on the taxonomy, systematics, and phylogeny of the families, genera, and species in the region. We will discuss the history of botanical exploration in the Bahama Archipelago, followed by a discussion of the impact of other types of research on a floristic treatment.

Over the years, several writers have provided synopses of botanical explorations in the Bahama archipelago. Coker (1905), Britton & Millspaugh (1920), and Correll & Cornell (1982) gave historical overviews of such explorations, and these serve as the backbone of what is presented here. Although many other notes and papers were published in which Bahamian plants and agriculture are discussed, we will limit this history to those publications that are tied to specimens procured in the archipelago. We divide the history of exploration in the Bahamas into three major periods:

1. 1700 to 1920 and the publication of Britton & Millspaugh's flora;

2. 1920 to 1984 and the publication of up the Correll & Correll flora; and

3. 1984 to present, during which time our understanding of the flora has been greatly enhanced by numerous field studies, the recognition of many new taxa, and tremendous changes in our understanding of plant phylogeny.

This paper begins with a historical overview of botanical exploration first, followed by changes in nomenclature and relationships among the taxa in the region, followed by how a new flora will proceed in these modern and rapidly changing times.

A History of Botanical Exploration in the Bahama Archipelago

A 1700 to 1920 and the Publication of Britton & Millspaugh's Flora

The earliest known botanical study of the Bahama Archipelago was done by Captain Thomas Walker (ca. 1659-1722), a British planter who served as governor of the Bahamas in 1706. Walker sent his plant specimens to his father, who had a garden in Westminster. At least some of these specimens were seen by Leonard Plukenet (1642-1706), who used some for figures in his Phytographia (Plukenet, 1691-1696; Steams, 1970) and for descriptions in his Almagestum Botanicum (Plukenet, 1696). Walker also sent specimens to James Petiver (ca. 1655-1718), who referred to them in his Musei Petiveriani (Petiver, 1692-1703). Petiver's herbarium, which still contains Walker specimens, is housed in the Sloane Herbarium (BM) (herbarium acronyms from Thiers, continuously updated).

Another early explorer in the Bahamas was Mark Catesby (1682-1749), an English naturalist who came to the colonies as a collector of plant and animal specimens on behalf of a group of English patrons (Meyers & Pritchard, 1998). Catesby visited the islands of "Providence", "Ilathera", Andros, and "Abbacco" for a few months in 1725 (Steams, 1970), where he made notes, drawings, and paintings of plants and animals. During his time in the Bahamas, Catesby focused more on fish than plants. Mr. Catesby sent collections of plants and seeds to Hans Sloane. Two bound volumes of Catesby's specimens are housed in Sloan's herbarium, now in BM [source: www.nhm.ac.uk/ research-curation/research/projects/sloane-herbarium/peopleinsloaneslife.htm--24 Oct 2013]). In addition to the Sloane herbarium, material at BM may be found among the herbarium of Samuel Dale (see below), to whom Catesby presented specimens (W.T. Steam, in Linnaeus, 1957). Catesby also sent specimens to William Sherard and Charles DuBois, both of whose herbaria are now housed at OXF, and to John James Dillenius, Dutch botanist who was brought to Oxford as professor of Botany by Sherard (Allen, 1937). Thirty Catesby specimens labeled "Providence" are in Sherard's collection, and specimens derived from seeds collected by Catesby are also found there [source: www.herbaria.plants.ox.ac.uk/bohcatesby--24 Oct 2013]).

Catesby's collections in the New World form the basis of A Natural History of Carolina, Florida, and the Bahama Islands (1730-1747), in which he described 71 plant species directly attributed to the Bahamas, 65 of which are depicted in plates. Catesby used polynomials for the species included in the work, and several attempts have been made to correlate these polynomials to modern binomials (Cates, 1996; Cates, 1998; Howard & Staples, 1983).

In addition to the Bahamian plants described in his Natural History, Catesby (1767) mentioned a Bahamian Juniperus in his Hortus europae-americanus, though he conflated it with the eastern North American species now known as Juniperus virginiana L.

Francis Dale, an English naturalist, collected specimens in the Bahamas from 1730 to 1732 or 1733 (Britton & Millspaugh, 1920; Howard, 1975). These he sent to his uncle Samuel Dale (Boulger, 1883) in England; some of these specimens are preserved in herbarium BM.

Carolus Linnaeus (1707-1778), considered by many to be the "Father of Plant Taxonomy", published Species Plantarum in May 1753. This publication serves as the starting point for all modern plant names because of the consistent use of binomials for the included species. While Linnaeus did not himself collect specimens in the Bahamas, he referred to previous publications, e.g., Catesby, Plukenet, and Petiver, as sources of species he included in the two volumes (Linnaeus, 1753). Some of Catesby's specimens cited by Linnaeus as from the Carolinas were actually from the Bahamas (W.T. Steam, in Linnaeus, 1957). Only eight species are attributed directly to the Bahamas, even though many species included in the publication occur in the archipelago. These species are enumerated here (with the volume and page number on which the name was originally published in Species Plantarum (Linnaeus, 1753), as well as the name we currently accept for the taxon): Cassia ligustrina (Virginia, Bahama; 1: 378) = Senna ligustrina (L.) H.S. Irwin & Bameby (Fabaceae); Catesbaea spinosa (Providentia; 1: 109) = Catesbaea spinosa L. (Rubiaceae); Convolvulus alsinoides (Malabaria, Zeylonia, Bisnagaria, Bahama; 1: 157) = Evolvulus alsinoides (L.) L. (Convolvulaceae); Passiflora cupraea (Providentia, Bahama; 2: 955) = Passiflora cupraea L. (Passifloraceae); Schoenus coloratus (Jamaica, Bahama; 1: 43) = Rhynchospora colorata (L.) H. Pfeiff. (Cyperaceae); Sida crispa (Carolina, Providentis, Bahama; 2: 685) = Herissantia crispa (L.) Brizicky (Malvaceae); Solanum bahamense (in Americae insula Providentia; 1: 188) = Solanum bahamense L. (Solanaceae); Waltheria americana (Bahamas, Barbiches, Surinimao; 2: 673) = Waltheria americana L. (Malvaceae).

No collections are known to have been made between the early 1730's and the 1780's, at which point two expeditions were undertaken that included the Bahamas. In 1783-1784, Franz Joseph Matter (1753-1827), botanist to Emperor Joseph II of Austria, along with Franz Boos (1753-1832), gardener at Schonbrunn Palace in Vienna, and Johann D. Schopf (1752-1800), a geologist and natural historian, visited the Bahamas and collected specimens and live plants that were sent back to Vienna (Carozzi & Carozzi, 1994; Coker, 1905; Kronfeld, 1892; King, 2011). Marter sent a letter back to von Born, editor of Physikalische Arbeiten, in which he described the flora of the islands and the plants species he found there (Marter, 1786). Specimens from this voyage are housed in herbaria in Vienna (W) and Buxelles (BR).

Andre Michaux (1746-1802; Savage & Savage, 1986), French botanist working under the auspices of King Louis XVI, explored and collected in the United States and the Caribbean. In 1789, he visited the Bahamas, arriving on February 25 in New Providence, and departing in March (Britton & Millspaugh, 1920). While in the Bahamas, he collected specimens and seeds of many plants, and dug up many trees that were sent back to a nursery in South Carolina (Michaux & Sargent, 1888). His herbarium specimens are in the Museum national d'Histoire naturelle (P) in Paris, France. Michaux named two species from the Bahamas collections (Michaux 1803): Laurus catesbyana Michx. (=Nectandra coriacea L., Lauraceae), and Limodorum trifidum Michx. (=Bletia purpurea (Lam.) DC., Orchidaceae).

Early in the 19th century, only a few botanists collected specimens in the archipelago. John Fraser made collections during a visit to the Bahamas in 1802; these specimens are probably in herbarium LINN (Britton & Millspaugh, 1920), though this has not been confirmed. Johann Wilhelm Crudy (1753-1810) (Britton & Millspaugh, 1920; Solereder, 1912) made collections in the Bahamas sometime prior to 1810; his specimens are in herbaria BM, GH. M. Jean Baptiste Ricord (or Ricord-Madiana) (1777-1837), born in France and trained in New York as a physician (Wilson & Fiske, 1900), lived on the island of Guadaloupe, and gathered specimens in the Bahamas in about 1810; some of his specimens are in GH (Britton & Millspaugh, 1920).

After 1810, no new botanical materials were gathered from the Bahamas, as far as is known, until 1830. Gulielmus (William) Hamilton (1783-1856) wrote an early synopsis of the flora of the West Indies (Hamilton, 1825), but only mentioned 3 species for Bahamas: Catesbaea spinosa L., Croton bahamensis Desvaux ex Hamilton, Phymosia abutiloides (L.) Ham. Hamilton's specimens appear to have been placed in Desvaux's herbarium (now at P), though it is unclear if he ever collected in the Bahamas himself. Howard et al. (1981) discussed the difficulties in determining the meanings of Hamilton's names, and the typification and current applications of many of them. William Swainson (1789-1855), collected in the Bahamas from 1830 to 1842 (Coker, 1905; Urban, 1898-1928). His collections were studied by Grisebach for his Flora of the West Indian Islands (Grisebach, 1864), and are housed in the herbarium of the Royal Botanic Gardens, Kew (K); some Swainson material is also in CGE and LIV (Vegter, 1986).

In the 1850's, several naturalists visited the archipelago and gathered plant specimens. William Freeman Daniell (1817-1865), a surgeon in the British army, was stationed in the Bahamas from 1857 to 1858 (Nickalls, 2012). During his stay, he collected specimens that were sent back to the British Museum (BM) (Urban, 1898-1928). He also gathered data on the medicinal properties of Croton species (cascarilla), about which he published a medical monograph (Daniell, 1862). At about the same time as Daniell was collecting in the Bahamas, Justus Adelric Hjalmarson, a Swedish naturalist, visited Grand Turk Island for 14 days in May, 1858 (Coker, 1905; Howe and Wilson, 1908). Hjalmarson's specimens were studied by Grisebach, and are to be found in herbaria B, GOET, and K. In 1859, the American botanist William Cooper collected about 100 specimens on New Providence for John Torrey (Britton, 1890); his specimens are at NY (Coker, 1905). Henrick Johannes Krebs (1821-1907), a Dane living on St. Thomas, collected in about 1866 (Britton & Millspaugh, 1920) on New Providence. He returned to Copenhagen in 1870, and his collections are in herbarium C (Coker, 1905; Rydberg, 1907).

In 1864, A.H.R. Grisebach (1814-1879) published Flora of the British West Indian Islands (1864) at the behest of William Jackson Hooker, Kew. Grisebach was given a grant of 300 [pounds sterling] by the British government to cover expenses for the project (Grisebach, 1864). While Grisebach did no field work in the Bahamas himself, he had access to the important collections at the Royal Botanical Garden (K), including the Hookers' own herbarium, the British Museum (BM), various other herbaria in Europe, and his own at Gottingen (Goet). Grisebach's flora is the first substantive manual that included the Bahama flora; it served as a foundation for subsequent floras of the region. Of particular interest is that Grisebach's flora is the first to mention the economically and floristically important Bahamian pine. His description of Pinus bahamensis Griseb. (p.503; = P. caribaea var. bahamensis (Griseb.) W.H. Barrett) is based only on a cone collection.

Rather few collections are known from the archipelago from between 1860 and 1875. Rawson William Rawson (1812-1899), governor of the Bahamas from 1864 to 1869, appears to have collected plant specimens between 1867 and 1868 (Britton & Millspaugh, 1920); his specimen may be found in BM, K, and perhaps B (if extant). H.E.W. Robinson (1836-1912), governor of the Bahamas from 1874 to 1880, collected 192 specimens in 1877, which are in herbarium K (Britton & Millspaugh, 1920). He was very interested in finding economic uses for plants of the islands, and suggested using stem sections of Agave species from the Bahamas as "razor strops" (Hooker, 1880).

Lewis Jones Knight Brace (1852-1938; Burkill, 1941), was born in Nassau, Bahamas, and educated in London, from which he returned to Nassau and studied the flora using Grisebach's book. His interest came to the attention of Governor Robinson, who suggested that Brace send a duplicate set of his specimens to the Royal Botanic Gardens, Kew, for identification. As a result, Brace sent 525 specimens to herbarium K between 1875 and 1887 (Britton, 1890; Britton & Millspaugh, 1920). Brace was first cousin of Elizabeth Britton, wife of Nathaniel Lord Britton, director of the New York Botanical Garden, and because of this connection, he sent about 4000 specimens to NY between 1904 and 1919 (Anonymous, 1905; Burkill, 1941).

Brace wrote an unpublished list of the plants of New Providence, a copy of which was given to John Gardiner (1863-1900), scientific advisor to the Agriculture Board of the Bahamas (Gardiner et al., 1889), and, later, professor of biology, University of Colorado (Anonymous, 1900, p. 934). Gardiner made many additions to Brace's list, but unfortunately did not keep specimens because of the difficulties of doing so while aboard-ship (Gardiner et al., 1889). A copy of the provisional list was given to Charles Sumner Dolley, who edited it and added to the list from his own collections and those of Francis Hobart Herrick, and in 1889 published A provisional list of the plants of the Bahamas Islands (Gardiner et al., 1889).

In 1885, Anna H. Searing (1830-1912), a physician from Rochester, New York, and early supporter of evolutionary theory (McKelvey, 1946), made a collection of plants from the Bahamas (Dolley, 1889). The whereabouts of her specimens is unknown. In June of 1886, Francis Hobart Herrick of Adlebert College, Cleveland, Ohio, collected specimens on Abaco Island and adjoining cays (Herrick, 1886). His specimens were identified by D.C. Eaton and W.A. Setchell, Yale University (Eaton & Setchell, 1886), and specimens are deposited in herbarium YU, with additional material reported as in the herbarium of Adelbert College (Britton, 1890; Dolley, 1889). The fate of Herrick's Bahamian specimens from Adelbert College is unknown, though that herbarium was likely combined with the herbarium of Western Reserve College (later Case Western Reserve University); the Case Western herbarium eventually went to the Holden Arboretum, Kirtland, Ohio (Cusick & Snider, 1984), and then to CLM (James Bissell, pers. comm.). In 1887-1888, Baron F.H.A. von Eggers collected on Grand Turk, Acklins, Fortune, Long, and Hog Islands under the auspices of the British Association for the Advancement of Science, from which he received a grant of 100 [pounds sterling] (Eggers, 1892; Thiselton Dyer, 1888). Carruthers et al. (1889) reported that 357 specimens were collected, and sent to herbaria BM and K, with duplicates at B. Eggers' collections were studied by Urban (1898-1928), who named several new species. In 1888, Charles Sumner Dolley (1856-1948), a biologist and microbiologist, collected plant specimens in the Bahamas, which were deposited in herbarium PENN (now in PH) (Dolley, 1889). Alexander Keith, a Scotsman on a sisal plantation on Andros Island, made a few collections from 1889 to 1891 that he gave to Alice Northrop (Northrop, 1902, 1910; Britton & Millspaugh, 1920); his specimens are at NY.

The 1890's saw several expeditions to the Bahamas by botanists. John T. Rothrock (1839-1922), American geologist, botanist, and forester (Swanger, 2010), collected in the Bahamas and Jamaica in 1890-1891, under the auspices of American Philosophical Society of Philadelphia, from which he had a grant of $300 to perform field work in the West Indies (Rothrock, 1891). Rothrock reported that he visited "New Providence, Eleuthera, San Salvador, Watling and Inagua" during his brief visit to the Bahamas. Rothrock's specimens from this expedition are mainly in herbaria F and PH. At the same time, and on the same expedition, though funded by the Missouri Botanical Garden (Hitchcock, 1893), Albert S. Hitchcock (1865-1935), American plant taxonomist (Chase & Rendle, 1936), collected 953 species, many of which were from the Bahamas (Hitchcock, 1891; Hitchcock, 1893). His specimens are housed in herbarium MO. The last expedition of note in the 19th century was that of John and Alice Northrop, who visited the Bahamas from January to July in 1890, after a letter from Sir William Thistleton Dyer, urging continued exploration of the Bahamas, was received at the Columbia University herbarium (Northrop, 1902). John Northrop (1861-1891), American zoologist (Osborn, 1910), and his wife, botanist Alice R. Northrop (1864-1922; ultimately professor of botany at Hunter College, New York) collected plant specimens and animal specimens; the bulk of the botanical work was conducted on Andros Island. The Northrops collected ca. 200 plant specimens on New Providence and cays in its vicinity, and then spent 4 months on Andros, and explored it thoroughly; about 550 species were collected during their stay in the Bahamas (Northrop, 1902), among which were 29 plant species described as new to science by Alice Northrop and others. Sets of the specimens were sent to the following herbaria: B, F, GH, K, and Columbia University (now at NY); some specimens were also sent to experts at other institutions for determination (Northrop, 1902, 1910). In her paper, Northrop states that the type specimens of her new species are in the herbarium of Columbia University (now at NY). The status of her specimens was discussed in great detail by Kass (2005), who concluded that specimens at NY should be regarded as holotypes where possible, and that other species for which no specimen was extant at NY should be lectotypified by specimens at F or other herbaria.

In addition to the collections described above, Bertha Wilson, State University of Iowa, collected in the Bahamas in 1893 (Nutting, 1895), but the specimens appear to have been discarded (Britton & Millspaugh, 1920). Boston pteridologist George Franklin Curtiss (1861-1895; Anonymous, 1896) collected in 1893-4 on Andros Island (Britton & Brown, 1920); these specimens are in herbarium F.

At the start of the new century, a small collection was made by a visitor to the Bahamas. Emilia Royce Anthony (1834-1904), of Gouvemeur, New York, in 1900-1902 (Anthony, 1902; Britton & Millspaugh, 1920). Her personal herbarium was destroyed in a fire, but duplicates were in the personal herbaria of B.D. Gilbert (formerly in the Hamilton College herbarium (Kittredge, 1939), now at GH), and Willard N. Clute (now in BUT). John W. Harshberger (1869-1929; Nichols, 1930), botanist at the University of Pennsylvania, collected in the Bahamas in 1901 (Britton & Millspaugh, 1920); these specimens, originally housed at PENN, are now in PH and US.

William C. Coker (1872-1953), a botanist at the University of North Carolina in Chapel Hill, conducted the first major expedition to the Bahamas in the new century. With funds provided by the Geographical Society of Baltimore, Maryland, Coker and others undertook a survey of the islands in 1903, studying their physiography, geology, fossils, tides, magnetism, climate, soils, plants, animals, diseases, and history (Shattuck, 1905). Coker's study of the botany of the islands was published in a paper in the Shattuck's Vegetation of the Bahama Islands (Coker, 1905). Coker discussed the vegetation types, and provided an overview of botanical exploration of the Bahamas, its useful plants, and descriptions of the flora on the various islands visited during the expedition. He concluded with a list of species collected and photographs of habitats and plant species encountered.

Emphasis on plant collection in the Bahama archipelago turned to a new institution, the New York Botanical Garden, in the first decade of the 20th century. Under the guidance of Nathaniel Lord Britton (1859-1934; Merrill, 1938), director of the New York Botanical Garden, and Charles F. Millspaugh (1854-1923; Sherff, 1924) curator of the department of botany of the Field Museum of Natural History, expeditions were undertaken to collect plant specimens in the Bahama archipelago with the ultimate goal of producing a flora of the archipelago. Many collectors were involved in the project were, in addition to Britton (and his wife, Elizabeth) and Millspaugh (and his wife, Clara). Allen Hiram Curtiss (1845-1907) collected on New Providence in 1903; George Valentine Nash (1864-1921; Britton, 1921) and Norman Taylor (1883-1967) made trips to the Bahamas in 1904 and the Turks Islands in 1905; Alexander E. Wight collected in 1904-05 for the Gray Herbarium, Harvard University (Robinson, 1906); Glover M. Allen (1879-1942; Barbour et al., 1943), Thomas Barbour, and Owen Bryant collected widely in the Bahamas in 1904 (Allen & Barbour, 1904; Britton & Millspaugh, 1920), and their specimens are at GH; John Kunkel Small (1869-1938; curator at NY; Core, 1938) and Joel J. Carter (1843-1912; Pennell, 1943) collected on Andros Island in 1910 (Small, 1910); Marshall A. Howe (1887-1936; Setchell, 1938) and Percy Wilson (1879-1944; Gleason, 1944), collected in 1907 and 1909 (Howe & Wilson, 1908). Much more could be written here on these expeditions leading up to the publication of The Bahama Flora in 1920, but itineraries are already included in Britton & Millspaugh (1920), and this history is discussed in detail in this proceedings volume by Brian Boom (2014). According to Boom, Britton and Millspaugh provided coverage of "1,982 species composed of 995 spermatophytes, 33 pteridophytes, 69 bryophytes, 150 fungi, 197 lichens, 519 algae (including diatoms), and 11 myxomycetes". Based on data in this flora, Taylor (1921) claimed over 14 % endemism for the Bahama flora.

B 1920 to 1984 and the Publication of the Correll & Cornell Flora

After the publication of Britton & Millspaugh's flora, botanical field work in the Bahamas slowed dramatically, essentially ceasing until about 1950 (Correll & Correll, 1982; Fang & Harrison, 1972), perhaps in part due to the world-wide effects of the Great Depression in 1929 and World War n. In the early 1920's, Paul Bartsch, a conchologist from Washington, D.C. (Rehder, 1961; Seidenschnur & Shetler, 1965) collected a few specimens of vascular plants and algae from the Bahamas; his specimens are housed in herbarium US. In 1950, Richard A. Howard (1917-2003), a graduate of the Botany Department at Miami University (1938) and recipient of a Ph.D. from Harvard University (1942), was employed by Harvard University as professor and director of the Arnold Arboretum (Warnement & Wood, 2004; Warnement, Wood, and Tomlinson, 2004). In 1948, Howard and his wife spent several months collecting in the Biminis, gathering 325 specimens. As a result, he published Vegetation of the Biminis (Howard, 1950), a thorough ecological survey and species list. His specimens are in the Harvard University herbaria (A, AMES, GH), with duplicates elsewhere. Howard spent much of his research career working in the Caribbean region, completing, among other major works, Flora of the Lesser Antilles (Howard, 1974--1988).

George Richardson Proctor (born 1920), American botanist whose research has centered in the Caribbean islands, took up residence in Jamaica in 1949 to complete an unpublished manuscript on the ferns of that island begun by W.R. Maxon. Proctor was employed by the Institute of Jamaica for 29 years (1951-1980), where he developed the herbarium and served as head of the Natural History Division. He subsequently held posts of herbarium supervisor at the National Botanic Garden in Santo Domingo, Dominican Republic (1982-1983), and biologist and director of the herbarium at the government's Department of Natural and Environmental Resources, San Juan from 1983 to 1998 (http://plants.jstor.org/person/bm000006736, accessed 13 Nov 2013). Proctor made several collecting excursions to the Bahamas and Turks & Caicos over many years, since the early 1950's (Proctor, 1954a, b; Proctor 1955). Many of Proctor's specimens are in herbarium IJ.

John Popenoe (born 1929) was director of the Fairchild Tropical Botanic Garden from 1963 to 1989. The garden had been conducting horticultural research in the Caribbean, and had done some work in the Bahamas (see, for example, Anonymous, 1959). In 1963, Popenoe received a grant from the American Philosophical Society to establish a "Bahamian Plot" in the garden (Jestrow et al., 2014). Popenoe's interests in the Bahamas were primarily horticultural (e.g., Popenoe, 1980), but this led to an ongoing study of the Bahama flora that ultimately led to the hiring of W.T. Gillis by the garden to work toward a flora, and then the hiring of Donovan Correll to write it, after Gillis left the garden.

William T. Gillis (1933-1979; Cowan, 1979) worked at the Fairchild Tropical Garden from 1968 to 1972 (Anonymous, 1968; Kass & Eshhbaugh, 1993), after which he moved to Michigan State University. Gillis conducted extensive field work in the Bahamas as preparation for writing a new Bahama flora. Between 1963 and 1978, Gillis made 22 trips, gathered 3,060 specimens, and published at least 24 papers on the Bahamian flora. These papers were often co-authored by R. A. Howard, G.R. Proctor, and other experts on the flora of the region (Kass & Eshbaugh, 1993). His research was funded, at least in part, by the National Geographic Society (Beaman, 1982), and by the American Philosophical Society (for work on the herbarium of William Hamilton; Gillis, 1979). Gillis' specimens are housed primarily at FTG and MSC with duplicates at many other herbaria. Sadly, Gillis died in 1979, at the age of 45, before he could see his work completed.

In addition to W.T. Gillis, many other botanists conducted field work in the Bahama archipelago prior to the publication of Correll & Correll (1982). Steven R. Hill, American botanist, collected 870 specimens (FTG) in the Bahamas in the early 1970's, and published 4 papers on the flora (Hill 1974, 1976a, b, c), including a new species, Euphorbia longinsulicola S. R. Hill. Other individuals conducting field research in the archipelago, include (listed with the major herbaria where their specimens are held): Daniel F. Austin (FTG), Roger Byrne (WIS), Walter H. Lewis (MO), Norton Hart Nickerson (1926-1999; TUFT, original herbarium destroyed by fire in 1975, balance transferred to MASS in 1998 [Thiers, continuously updated]), Robert Roy Smith (1934-1995; Rabeler, 1996; HHH), Dennis W. Stevenson (NY), Grady L. Webster (1927-2005; Beaman, 2006; DAV).

Donovan S. Correll (1908-1983; Schubert, 1984), and Helen B. Correll, American botanists, explored the Bahamas from 1973 to 1980 with the goal of writing a new flora of the archipelago (see the paper by Stevenson and Stevenson (2014), in this symposium volume). Upon the departure of W.T. Gillis from Fairchild Tropical Garden in 1973, D.S. Correll was hired to write the new flora (Anonymous, 1973). The flora of the Bahama archipelago project was funded by 3 grants from the National Science Foundation, and over the course of 7 years, the Cornells made 83 trips and collected approximately 11,000 specimens (from itinerary, pp. 1605-1607, Correll & Correll, 1982), most of which are deposited in FTG. During the course of the research, the Cornells and other botanists who accompanied them visited most major islands in the archipelago. This field work resulted in a series of scientific papers, and, of course, the flora itself. The flora opens with an introduction to the archipelago, including discussions of its geography, geology, hydrology, the origin of the flora, endemism, plant communities, natural resources, agriculture, plant products, bush medicine, and edible and poisonous plants. The Flora covers 1371 vascular plant taxa, in 663 genera, and 144 families. These are arranged in a phylogenetic system generally following the system of Engler & Prantl. The flora is illustrated beautifully by Priscilla Fawcett (1932-2010; Korber et al., 2012), whose original illustrations are now archived at FTG (Mosely, 2013). The flora is based on the Corrells' extensive collections, a thorough review of literature, and a review of herbarium specimens from many herbaria. Reviews of the flora were quite positive. Nicolson and Wasshausen (1983) stated that "the work is solid, quietly authoritative and conservative." Schmid (1997), reviewing a 1996 facsimile reprint, state that it was "a gorgeous facsimile reprint of a magnificently done flora."

C Botanical Exploration in the Bahama Archipelago since 1982

The pace of botanical exploration in the Bahamas, post-Correll, has picked up considerably, in rather stark contrast to the pace that followed the publication of the Britton & Millspaugh's flora of 1920. While it will not be possible to enumerate here all of the individuals who have conducted research on the flora of the Bahama archipelago, we will attempt to enumerate some important contributions. This enumeration is by no means complete, but we hope it includes most of the major contributions.

Lee Kass, American botanist, formerly of Elmira College, has been conducting research on San Salvador since 1982, and has published 8 books on the San Salvador flora, as well as 37 papers or notes, and many abstracts of talks at meetings. Several of her papers, as well as those of Robert Smith and several other botanists mentioned in this review, are in publications of the Gerace Research Centre (formerly the Bahamian Field Station) on San Salvador, at which research conferences and symposia have been held since 1982. Conference and symposia proceedings that were published are available on the Gerace Centre web site, as are books on the natural history of the archipelago.

Ted R. Bradley, George Mason University (GMUF), took classes to Andros Island for many years, collecting many specimens during those trips.

Many other institutions have been involved with research in the Bahamas archipelago, including the Nature Conservancy, the Bahamas National Trust, university groups, and so on. Among the universities from which researchers have visited the Bahamas are the University of South Florida, the University of Florida, the University of Miami, and Harvard University.

Botanists from Miami University in Oxford, Ohio, have been actively working in the Bahamas since 1973. W. Hardy Eshbaugh and Thomas K. Wilson taught field classes on Andros Island for many years, and as a result, researchers from the university published a large number of papers and gave many public scientific presentations. Among them are R. James Hickey and Michael A. Vincent, and many students, including Ethan H. Freid, Logan R. Randolph, Eric Kjellmark, Laurel Richey, etc.... As a result, 22 theses and dissertations were written, as well as 90 papers, and many abstracts and presentations (Eshbaugh, 2014). Also resulting from field work associated with these projects are about 20,000 herbarium specimens that are housed at the W.S. Turrell Herbarium (MU), with duplicates in the Bahamas National Herbarium in Nassau, and herbarium FTG, and others.

V Towards a new Flora of the Bahama Archipelago

The Flora of Cornell and Cornell is a thorough and extremely useful work which has served well for 30 years. No living flora (the plant life of a given geographical or geopolitical region) is static, however; it changes through time. For this reason, some (e.g., Reznicek et al., 2011) say that any floristic treatment is outdated almost as soon as it is published.

Why are floras so quickly outdated, and why do the taxa included, nomenclature, and numbers of species change from treatment to treatment? As exemplified by the long history of collecting in the archipelago, additional field work results in discovery of previously unseen species in the region. These may be plants native to the flora region, or may be introductions. These new introductions may be casual introductions, or may become invasive species (Carey et al., 2014).

Changes in nomenclature may result from revisionary studies in which taxa previously recognized as distinct are lumped under another name, or where a taxon is split into smaller taxa, resulting in more names. These kinds of changes may result from morphological or phylogenetic studies. Changes in understanding of relationships resulting from new morphological or phylogenetic studies may cause realignment of species within genera, or genera within families. Extinctions may also result in changes in a published flora.

Changes in understandings of nomenclature, relationships, and so on are reflected in a recent publication, Catalogue of Seed Plants of the West Indies (Acevedo-Rodriguez & Strong, 2012), in which the authors have derived a checklist based on a review of available literature, specimens at US, and on-line resources. In this work, the checklist for the Bahama archipelago (excluding pteridophytes) includes 1337 taxa, in 507 genera in 127 families (Acevedo-Rodriguez & Strong, 2010; from Table 1, page vi). This checklist provides yet another resource for the creation of a new flora of the Bahama Archipelago.

In our efforts to create a new Flora of the Bahama Archipelago, we have been reviewing literature on the nomenclature and relationships of taxa known previously to occur in the flora range, and have been conducting field research throughout the archipelago. We have also reexamined many specimens from herbaria in the Bahamas, the United States, and Europe. As a result, many distributional and nomenclatural changes will be reflected in our new flora. As an example, reexamination of the fern flora has resulted in major changes in our understanding of the fem species in the region. Correll & Correll (1982) recognized 7 families of ferns, encompassing 24 genera and 45 species. In our new treatment, we accept 16 families, encompassing 29 genera and 51 species (Table 1).

A new flora will need to meet new demands and expectations as we move forward into the 21st Century, so our flora will be designed to meet those needs, and will be designed to be an on-line product, in addition to the traditional print volume. In this way, we hope to build on what has gone before, and also to look forward to a new generation of a study of the flora of the Bahama Archipelago.

DOI 10.1007/s12229-014-9143-1

Acknowledgments We thank W. Hardy Eshbaugh, Javier Francisco-Ortega Brett Jestrow, and Lee Kass for providing assistance and information pertinent to this paper. This research was supported in part by the Willard Sherman Turrell Herbarium Fund, Miami University, Oxford, Ohio.

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Michael A. Vincent (1,2) * R. James Hickey (1)

(1) Department of Biology, Miami University, Oxford, OH, USA

(2) Author for Correspondence; e-mail: vincenma@miamioh.edu

Published online: 7 August 2014
Table 1 Comparison of the fern treatment in Correll & Cornell
(1982) and our current understanding

Correll & Correll                     Current classification

Selaginellaceae
  Selaginella ealonii Hieron.         Selaginella ealonii Hieron.
  ex Small                              ex Small
                                      Selaginella umbrosa Lem.
                                        ex Hieron. (a)

Psilotaceae
  Psilotum nudum (L.) P. Beauv.       Psilotum nudum (L.) P. Beauv.
Osmundaceae
  Osmunda regalis L. var.             Osmunda regalis L. var.
  spectabilis (Willd.) A. Gray          spectabilis (Willd.) A. Gray
Hymenophyllaceae
                                      Trichomanes punctatum Poireta
Schizaeaceae
  Anemia adiantifolia (L.) Sw.        Anemia adiantifolia (L.) Sw.
  Anemia cicutaria Kunze ex Spreng.   Anemia cicutaria Kunze ex Spreng.
  Anemia wrightii Baker               Anemia wrightii Baker
  Schizaea poeppigiana Sturm          Schizaea poeppigiana Sturm
Marsileaceae
  Marsilea nashii Underw.             Marsilea nashii Underw.
  Marsilea vestita Hook. & Grev.      Marsilea vestita Hook. & Grev.
Polypodiaceae                         Lindsaeaceae
Sphenomeris clavata (L.) Maxon        Sphenomeris clavata (L.) Maxon
Dennstaedtiaceae
Lindsaeaceae                          Pteridium caudatum (L.) Maxon.
Pteridaceae
  Acrostichum aureum L.               Acrostichum aureum L.
  Acrostichum danaeifolium            Acrostichum danaeifolium Langsd.
    Langsd. & Fisch.                    & Fisch.
  Adiantum capillus-veneris L.        Adiantum capillus-veneris L.
  Adiantum melanoleucum Willd.        Adiantum melanoleucum Willd.
                                      Adiantum raddianum C. Presl (a)
  Adiantum tenerum Sw.                Adiantum tenerum Sw.
  Cheilanthes microphylla (Sw.)       Cheilanthes microphylla (Sw.) Sw.
    Sw.
  Pityrogramma calomelanos            Pityrogramma calomelanos
    (L.) Link                           (L.) Link
  Trismeria trifoliata (L.) Diels     Pityrogramma trifoliata (L.)
                                        R. M. Tryon
  Pteris longifolia L. var.           Pteris bahamensis (J. Agardh) Fee
    bahamensis (J. Agardh)
    Hieron.
  Pteris vittata L.                   Pteris vittata L.
  Vittaria lineata (L.) J. E. Sm.     Vittaria lineata (L.) J. E. Sm.
Aspleniaceae
  Asplenium dentatum L.               Asplenium trichomanes-dentatum
                                        L. (b)
  Thelypteridaceae
  Macrothelypteris lorresiana         Macrothelypteris torresiana
    (Gaud.) Ching                       (Gaud.) Ching
  Thelypteris augescens (Link)        Thelypteris augescens (Link)
    Munz & I. M. Johnst.                Munz & 1. M. Johnst.
  Thelypteris cordata (Fee)           Thelypteris abdita Proctor (b)
    Proctor
  Thelypteris denlala (Forssk.)       Thelypteris dentata (Forssk.)
    E. P. St. John                      E. P. St. John
  Thelypteris kunthii (Desv.)         Thelypteris kunthii (Desv.)
    Morton                              Morton
  Thelypteris ovata R. P. St.         Thelypteris ovata R. P. St.
    John in Small var. ovata            John in Small var. ovata
  Thelypteris reptans                 Thelypteris reptans (J. F. Gmel.)
    (J. F. Gmel.) Morton                Morton
Blechnaceae
  Blechnum serrulatum L.              Blechnum serrulatum L.
Dryopteridaceae
  Ctenitis hirta (Sw.) Ching.         Ctenitis hirta (Sw.) Ching.
Lomariopsidaceae
  Nephrolepis biserrata (Sw.)         Nephrolepis biserrata (Sw.)
    Schott                              Schott
                                      Nephrolepis cordifolia L. C.
                                        Presl (a)
  Nephrolepis exaltata (L.)           Nephrolepis exaltata (L.) Schott
    Schott
  Nephrolepis mutiflora (Roxb.)       Nephrolepis brownii (Desv.)
    Jarrett ex Morton                   Hovenkamp & Miyam. (b)
Tectariacaeae
  Tectaria coriandrifolia (Sw.)       Tectaria coriandrifolia (Sw.)
    Underw.                             Underw.
  Tectaria heracleifolia (Willd.)     Tectaria heracleifolia (Willd.)
    Underw.                             Underw.
  Tectaria lobata (Presl) Morton      T. fimbriata (Willd.) Proctor &
Polypodiaceae                           Lourteig (b)
  Neurodium lanceolatum (L.) Fee      Neurodium lanceolatum (L.) Fee
  Polypodium aureum L.                Phlebodium aureum (L.) J. Sm. (b)
                                      Phymatosorus grossus (Langsd. &
                                        Fisch.) Brownlie3
  Polypodium heterophyllum L.         Microgramma heterophylla (L.)
                                        Wherry (b)
  Polypodium phyllitidis L.           Campyloneuumn phyllitidis (L.)
                                        C. Presl
  Polypodium plumula Humb. &          Pecluma plumula (Humb. & Bonpl.
    Bonpl. ex Willd.                     ex Willd.) M.G. Price
                                      Pleopeltis michauxiana (Weath.)
                                        Hickey & Spruntc
  Polypodium polypodioides            Pleopeltis polypodioides (L.)
    (L.) Watt.                          E. G. Andrews & Windham (b)
  Polypodium squamatum L.             P. squamata (L.) J. Sm. (b)

(a) New to the flora

(b) Replacement name for misapplied name

(c) Pleopeltis michauxiana (Weath.) Hickey & Sprunt, comb. nov.
Basionym = Polypodium polypodioides var. michauxianum Weath.,
Contr. Gray Herb. 124: 31. 1939
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