Taxonomic Botany and Floristics in North America

North of Mexico: A Review

James L. Reveal

James S. Pringle

THE modern history of systematic botany and floristics in North America began when the first Europeans landed on these shores and began to collect objects of curiosity. It is imperative to use the term "modern," for long before colonization of the New World by Europeans, the Native Americans, who had arrived millennia earlier, had developed their own systems of classification, means of identification, and associated nomenclature. Unlike that of their European counterparts, their knowledge was transferred by the spoken rather than the printed word and was mostly lost as their civilizations fell to the invaders. To a great degree, it was not until the twentieth century that Native Americans were recognized as knowledgeable about their plants. By then, European thought dominated botany, and the Native American's botanical understanding was passed on only in an occasional native name retained in a Latinized form.

It was not until Columbus's second voyage, in 1493, that New World plants and animals were taken across the Atlantic. For the European scientific community, the unfamiliar specimens were a source both of great intellectual curiosity and of philosophical concern. The curiosities were clearly different from their Old World counterparts, and in some instances they were entirely novel. The likes and near-likes could be associated, but the distinctly different were philosophically troublesome.

The Spanish of the early sixteenth century were the first to describe the flora of the New World. Gonzalo Fernández de Oviedo y Valdes (1478--1557) visited several of the Caribbean islands and portions of Central America, trying to fit the tropical vegetation he observed into a classification scheme that recognized only six species of trees with persistent green leaves. Oviedo had become acquainted with native New World plants of equal or even greater value than those introduced to the New World by the Spanish, and he urged their use. He was ignored.

Nicolas Bautista Monardes (1493--1578) never saw the New World. His interests were the new medicines and new remedies he felt certain existed. He classified plants according to their medicinal properties, and for the American ones he often retained the native names. He accepted treatments recommended by the Amerinds, but as a firm believer in the Doctrine of Signatures, he occasionally modified them.

The missionary Jose d'Acosta (1539--1600) spent 20 years in Peru, returning to Spain in 1588 to publish various works on the New World. He urged scholars to regard the majority of living things in the New World as unique and not to assign them established European names. He described numerous native economic and medicinal plants and commented on the diversity of potatoes, tomatoes, and chili peppers he had found in the market; he also mentioned cacao and coca.

During this period, intellectual thought often was dominated by religious dogma. Scholarly investigations in the natural sciences began primarily in northwestern Europe. The first naturalists often had to flee the upheavals of the Protestant Reformation and, as a result, many traveled widely and learned from others. In this way, a more unified system of classification and nomenclature began to develop.

Herbals, those great tomes illustrated with woodcuts, were the primary botanical publications of the age. At first they were little more than restatements of Dioscorides or other classical authors, but as the herbals were developed over the next two centuries, new species and remedies were incorporated, including the wonders of the New World. Of equal importance was the development of botanic gardens, first established in Pisa in 1543. These soon became centers of scientific importance because not only could plants of faraway places be seen, but their medicinal properties could be determined also.

The herbalists of continental Europe were emulated by Englishmen such as William Turner (1510--1568), "Father of English Botany," and John Gerard (1545--1612) of London. Later John Tradescant (d. 1638) and his son, John Tradescant the Younger (1608--1662), set up a garden at Lambeth and, with royal patronage, established (1629) a natural history museum known as "Tradescant's Ark." To increase the holdings of the museum, the younger Tradescant repeatedly traveled to Virginia, beginning in 1637, to collect plants and animals for display. In 1621, Magdalen College granted Oxford University 5 acres to establish the Oxford Physic Garden, and although it did not reach prominence until after the Stuart Restoration of 1660, it became the one garden that received hundreds of Virginia plant species prior to 1700.

When the first English colonists arrived on the East Coast, they carried with them their English herbals. Although a few native American plants, such as potato, tobacco, maize, and Jerusalem artichoke, were accounted for in Gerard's Herball or Generall Historie of Plantes (London, 1597), the truly valuable American medicinal plants were either still unknown or considered inferior to European plants. Four decades later, in his Theatrum Botanicum (London, 1640), John Parkinson (1567--1650) accounted for some 3800 different kinds of plants, including many from eastern Canada, New England, and Virginia.

The French were equally active in the exploitation of New World plants. Samuel de Champlain (1567--1635), who was also an apothecary, made several voyages to North America and took special interest in the settlement of the St. Lawrence Valley. In 1610, he established a garden at Port-Royal (now Annapolis Royal in Nova Scotia) where he grew several American and European plants. He also sent seeds and living specimens to Jean Robin (1550--1629) in Paris.

The illustrated Canadensium Plantarum Historia (Paris, 1635), written by the Paris physician Jacques Philippe Cornut (1606--1651), accounted for 38 species from temperate eastern North America. It was this work that Linnaeus consulted a century later to understand better the plants of this region. Several French colonists contributed material to botanists working in Paris at the Jardin du Roy. Notable among these were two "Médecins du Roy," Michel Sarrazin (1659--1735), correspondent of Tournefort, and Jean-François Gaultier (d. 1756), correspondent of Duhamel de Monceau, who published Traité des Arbres et Arbustes... (2 vols., Paris, 1755). Gaultier received the active support of Interim Governor Barrin de la Galissonière, who encouraged explorers and officers to send specimens and seeds through him and Gaultier to Paris. Gaspard Bauhin (1560--1624) listed several North American plants in his Pinax Theatri Botanici... (Basel, 1623), the first attempt to summarize the already confusing array of names and synonymy.

The New England colonists learned from the Amerind the values of some plants, but little of scientific or medical value was recorded. Thomas Hariot, a member of Sir Walter Raleigh's company that founded the colony at Roanoke in 1585, wrote a florid description of the region (A Brief and True Report of the New Found Land of Virginia..., London, 1588) to encourage others to settle in the New World. Captain John Smith (A True Relation of...Virginia..., London, 1608) championed the region around Chesapeake Bay in equally glowing terms, describing the great forests, the abundance of wildlife, and the beauty of the land. While these publications were of limited scientific value, the maps and illustrations that accompanied them did much to demonstrate the bounty that awaited anyone willing to take the risk of settling the land.

To Europeans, the temperate New World was an opportunity awaiting exploitation. The sixteenth and early seventeenth centuries were a period of conquest, not scientific exploration. The first 150 years of exploitation of the New World set the scene for changes in Europe. The wealth so generated helped create a class of people that had free time, education, and an interest in the sciences. It also provided the means whereby individuals of talent, but not of wealth, could find the means to study the natural and physical world.

The philosophy of science that greeted the discovery of the New World was one hardened by centuries of total faith in the teachings of the ancient past. The immediate challenge the New World presented was that it called into question the accepted norm. As Francis Bacon wrote in Novum Organum (London, 1620), knowledge "must begin anew" from its very foundation. New means of evaluating human observations had to be devised to organize knowledge and to explain what exists.

Bacon realized that no one person could know or explain all, and as with the arts, which he deemed to be equal to the sciences, no one person could be fully accomplished in every aspect. He argued for corporate undertakings involving both those with the means to support the arts and sciences and those with the ability to be artists and scientists. Only by working jointly, and over generations, could humankind hope to comprehend all that exists in the world.

For the sciences, cooperation was slow to establish itself. The first science club was formed by Wadham College, Oxford, in 1648. A decade later the Oxford Club was founded with Robert Boyle, Robert Hooke, and John Evelyn among its members. By 1660 the concept of a Royal Society had taken shape, and soon thereafter formal scientific bodies were established in both England and France.

The societies provided a meeting place to witness experiments, hear papers, and generally discuss ideas of interest. They also became focal points where others, outside Europe, could send contributions in the form of specimens or objects of curiosity that were housed and maintained by the membership, or as manuscripts and letters reporting on observations. Through publications, patronage, and especially correspondence, the royal societies encouraged people to contribute. For a few individuals scattered along the Atlantic Coast of North America, this was a needed avenue to inform others of the curiosities of their new land.

By 1783, numerous men from North America had been elected to fellowship, and many hundreds more were correspondents. The societies not only provided a venue for science to be addressed, they actively promoted it. Governors, heads of major companies, and large landowners were elected to membership and then were encouraged to support individuals who could make observations on the natural and physical world. Tracts on natural products were written; plants, animals, and minerals were collected and described; the weather was recorded; and the movements of comets and planets were noted.

Still, important voices were heard outside the societies. John Josselyn (1608--1675) spent several years in New England, and in his New-England's Rarieties Discovered... (London, 1672), he described numerous new species of plants. Although Josselyn's drawings were crude, Linnaeus cited this work in 1753. This and Josselyn's later Account of Two Voyages to New England... (London, 1674) were the most complete summary of the North American flora for more than a century.

The 20-year period following the publication of the second part of Plantarum Historiae Universalis Oxoniensis (Oxford, 1680) by Robert Morison (1620--1683), was one of dramatic change in botanical thought in Europe. Nehemiah Grew (1641--1712) published his Anatomy of Plants... (London, 1682), and John Ray (1627--1705) summarized the known flora of the world in his three-volume Historia Plantarum... (London, 1686--1704), in which he applied the philosophy he had outlined in his Methodus Plantarum Nova... (London, 1682). John Ray and two competitors, Leonard Plukenet (1642--1706) and James Petiver (1663--1718), all members of the Royal Society as well as London's informal Temple Coffee House Botany Club, sought to obtain and study the new plants coming into England from throughout the world. In France, Joseph Pitton de Tournefort (1656--1708) toiled to organize the equally overwhelming wealth of new material arriving there, producing his three-volume work Eléments de Botanique... (Paris, 1694).

Early knowledge of the temperate eastern North American flora came primarily from Virginia. Foremost in providing specimens and new observations was John Banister (1650--1692). During his years at Oxford, Banister had attended Morison's lectures at the Botanic Garden. To complete his Plantarum Historiae Universalis Oxoniensis, Morison persuaded Bishop Henry Compton to send Banister as a minister to the James River area. Banister sailed to Virginia in 1678, where he began his descriptions, observations, and drawings of plants and animals. After Morison's accidental death in 1683, Jacob Bobart the Younger, with the help of William Sherard and Samuel Dale, completed the third part of Morison's Historia (Oxford, 1699), including over 400 Virginia species. Bobart shared Banister's catalog and plant specimens with John Ray, who published them in the second and third volumes of his Historia Plantarum. Plukenet also reproduced Banister's plant drawings in his Phytographia (7 parts, London, 1691--1705).

A wealth of information had been supplied by Banister and three collectors in Maryland: Reverend Hugh Jones (1671--1702), William Vernon (d. 1711?), and Dr. David Krieg (d. 1710), a correspondent of Petiver. This allowed Plukenet, in the last three parts of his Phytographia (1696--1705), and Ray, in the final volume of Historia Plantarum, to account for nearly a thousand species of temperate North American plants.

Petiver encouraged many of his other North American correspondents to send him seeds and specimens. Robert Steevens (flourished 1700), Edmund Bohun (fl. 1699--1703), and Robert Ellis (fl. 1700--1704), all of Charles Town and its environs, routinely sent Carolina plants. Surgeons and captains of ships also collected specimens for Petiver along the shores from Hudson Bay to the Florida Keys. Petiver soon had many plants in cultivation, describing and illustrating them in his publications.

As the seventeenth century passed into the eighteenth, the natural sciences divided into a series of more specialized fields, botany diverging from zoology, and taxonomy from medicine. Many talented individuals became interested in understanding plant products and set to work at the task. Yet, the volume of novelties coming from India, southern Africa, Java, China, Japan, Turkey, and the American colonies was overwhelming the available resources. The value of Bauhin's Pinaxwas rapidly declining, so much so that at Oxford, William Sherard (1659--1728) began to draft a new edition. He was forced, however, to ask John Jacob Dillenius (1687--1747) to assist him as the volume of new names became too great. Bauhin had accounted for some 6000 names; Ray, in 1704, treated some 18,000. To summarize all of this nomenclature and render from it a sensible taxonomy required specimens from many sources and the cooperation of others. For Sherard and Dillenius, that cooperation was minimal. Tournefort and Sebastien Vaillant (1669--1722) in Paris and Paul Hermann (1646--1695), Herman Boerhaave (1658--1736), and others at Leiden all sent them specimens. Their greatest problem, however, was a fellow Englishman, Sir Hans Sloane (1660--1753).

Sloane had collected mainly in Jamaica from 1687 until 1689, amassing a large cabinet of curiosities that he began to study on his return to London. Sloane's marriage to a wealthy widow provided him with the income and social status to rise to a position of power as Physician to the Queen and secretary of the Royal Society. He outlived most of his contemporaries, and in time he acquired the libraries, correspondence, and collections of Plukenet, Petiver, and a host of others. The result was a collection so large that the British Museum was established to hold it.

The herbaria Sloane obtained were critical to the efforts of Sherard and Dillenius, but these herbaria were so unorganized, and Sloane had delayed access to them for so long, that even after Sherard's death in 1728, Dillenius was still unable to study them. Furthermore, William Sherard's brother James insisted that Dillenius divert his efforts to Hortus Elthamensis... (London, 1732), to the neglect of William's intended new Pinax. That manuscript was never finished, and the constant proliferation of names, together with the lack of a single source for all names, caused chaos for all who sought to identify and classify plants.

An Outline of Rationales

The modern taxonomic era began in 1753 with the publication by Carl Linnaeus (1707--1778) of a two-volume work entitled Species Plantarum... (Stockholm). The 20-year period leading to that publication was a time of intensive study for the young Swedish naturalist, isolated in a remote land far from the libraries, botanical gardens, and collections so vital to Dillenius. To expand his horizons, Linnaeus traveled to Holland in 1735 to obtain his doctorate and to study plants. His primary effort was to gain an appreciation for and a knowledge of the multitude of new plants being discovered in far-off lands.

The major innovation for which Linnaeus is credited today is the consistent use of binomial nomenclature. Yet the set of principles he formulated while yet a young man is of greater importance to taxonomy. His Fundamenta Botanica... (Amsterdam, 1736), with its 365 aphorisms, was an outline of the rationales he used to compose his Systema Naturae... (ed. 1, Leiden, 1735). These rationales matured over the following 15 years and formed the basis of Philosophia Botanica... (ed. 1, Stockholm, 1750). The concepts outlined by Linnaeus have come down to the present as some of the traditions and fundamental precepts in biological taxonomy.

At the time, the classification, identification, and naming of plants were, at best, nationalistic and personal, at worst, chaotic. Classification schemes were designed only to retrieve information, not to express relationships; identifications were based on regional features, not diagnostic characters; and nomenclature was a personal expression of opinion as to position, location, and features as compared to similar plants. Before Tournefort had developed his concept of the genus, plants often were grouped together by location, presumed affinities, or assumed virtues rather than by the possession of similar morphologic features.

Whereas taxonomic concepts and principles had been stated by previous authors, none seemed to carry sufficient weight to overcome national boundaries. Consequently, before Linnaeus, what was practiced by the English was not necessarily followed by the French or Spanish. Some authors, such as Ray and Plukenet, wrote descriptions based on individual specimens; Tournefort wrote his to fit an abstracted concept; but none had what might be regarded as a modern species concept. It was this innovation that Linnaeus contributed.

The modifications Linnaeus made to his aphorisms from 1736 to 1750 were mandated by experience gained from their use. Linnaeus's polynomials were diagnostic, each headed by a consistently used and uniformly applied generic name. The genera were arranged into orders and classes that reflected the system of classification Linnaeus had devised for the major groups of plants he accepted: algae, fungi, bryophytes, ferns, grasses, and the remaining herbs or trees. The Linnaean scheme made it possible to identify readily a given plant without an accompanying illustration. The idea that a given kind of plant should have but a single name was novel; but, of course, as other members of the same genus were found, the diagnostic features, and therefore the phrase name, had to be changed. To Linnaeus and his early followers, it was not the name that was consistent but the biological entity represented by the name.

In retrospect, Linnaeus's classification scheme was artificial in that it was based largely on the number and nature of reproductive structures; it was, however, practical. There was no evident concept of grouping organisms together that were related; that notion was to come later in the century. The value of Linnaean principles, however, was immediately obvious to many in Holland, and the name of Linnaeus preceded him as he toured France and England, visiting Bernard de Jussieu (1699--1776) in Paris, Dillenius at Oxford, and Philip Miller (1691--1771) at Chelsea. Boerhaave, Jan Fredrik Gronovius (1690--1762), and Adriaan van Royen (1704--1779) championed Linnaeus's work in Holland, as did Peter Collinson (1694--1768) in England. Through these individuals, Linnaeus's ideas were soon widely adopted, and the published works of the young Swede were dispersed widely, even to the American colonies.

The Clerk from Gloucester County

Botanical explorations along the Atlantic Coast of North America became limited as interest in this flora diminished with the introduction starting in 1700 of the diverse Chinese flora. Correspondents of Petiver continued to supply the London apothecary with plants from the Carolinas and Florida. Surprisingly rich collections of Hudson Bay plants were collected by Richard Tilden (fl. 1700--1707) in 1700, and by Dr. John Smart, a ship's surgeon, in 1708. Hannah Williams, perhaps the first woman collector in temperate North America, who sent mainly butterflies to Petiver, encouraged Joseph Lord (1672--1748) to collect. Lord became the foremost collector in Carolina. From 1701 to 1710, Petiver accounted for many of Lord's new species. John Lawson (d. 1711), a surveyor, also collected plants for Petiver, traveling far to the west and into many unexplored areas.

Mark Catesby came to Williamsburg, Virginia, in 1712 and collected throughout much of the area. The specimens were misplaced, however, and unstudied until 1981, although Catesby's collections in the Sherardian Herbarium at Oxford had been studied by Frederick Pursh, who based some new species on his collections. In 1719, when Catesby returned to England, Samuel Dale (1659--1739) remarked to Sherard that of the 70 specimens he had examined, at least half represented new species. When Catesby came back to the New World in 1722, he resided at Charles Town and traveled widely over the next four years, going as far south as northern Florida and the Bahama Islands. He was able to amass a huge collection of plants, animals, fossils, and other items of interest. He also skillfully sketched and painted many of the plants and animals he had seen in their natural habitats. When Linnaeus came to Oxford in August 1736, he probably examined Catesby's specimens, for they were the foundation for Catesby's Natural History of Carolina, Florida and the Bahama Islands... (2 vols, London, 1730--1747). Even so, Catesby had a lesser impact on Linnaeus than did his successor, John Clayton (1693--1773).

Clayton arrived in Virginia around 1720 and took up residence near Williamsburg where he served as the clerk of Gloucester County while maintaining a productive plantation and a lively correspondence with fellow botanical enthusiasts. Catesby and Clayton knew one another, and after Catesby returned to England, Clayton began to botanize actively, sending to Catesby at Oxford a large shipment of dried specimens collected in 1734. Catesby was ill prepared to identify the specimens, and reluctantly he sent this collection, and another received in 1735, to Gronovius in Holland, who was equally unable to cope with the problem.

Unaware of their fate, Clayton kept sending specimens to England. Catesby kept a few of the dried specimens for the herbarium at Oxford, and Collinson went through the seeds and retained for himself what he wished, but the bulk of the material continued on to Holland. Gronovius soon asked the young Linnaeus, then at the estate of George Clifford (1685--1760) at Hartekamp, to examine those specimens Gronovius was unable to place or to confirm those he recognized to be new.

Meanwhile, Clayton drafted a catalog of the plants of Virginia, following Ray's method, and sent it to Catesby, asking that he find a publisher. The manuscript went to Gronovius who, with the assistance of Linnaeus, completely revised the text according to the Linnaean method and published the work in 1739 as the first part of Flora Virginica... (2 vols., Leiden, 1739--1743). When Clayton finally saw this work in print, the authorship had been attributed to Gronovius, who had often used his own phrase names, those of Linnaeus, or those from earlier works (mostly Banister's specimens as rephrased by Morison, Ray, Plukenet, or Petiver) for Clayton's specimens. Moreover, Gronovius had cited Clayton's phrase names only as synonyms. At the urging of Collinson and Catesby, Clayton obtained most of the available Linnaean works and began to study them, all the while sending more specimens to Catesby, now arranged in the Linnaean fashion. Once again Clayton was disappointed, for the second part of the flora, published in 1742, still carried Gronovius's authorship.

Linnaeus was interested in Clayton's fine specimens, for with them he was able to evaluate many of the names for American plants proposed previously by Ray, Plukenet, Petiver, Hermann, and others. To what extent he called on Dillenius to resolve the nomenclatural morass, or actually examined specimens at Oxford in 1736 to do so, is unknown. But resolve nomenclatural problems he did, and in Flora Virginica one finds signs of the same care that Linnaeus would give to the flora of the whole world two decades later.

Another of Catesby's English friends was the physician John Mitchell (1711--1768), who came to Virginia in 1735 and remained until 1746. Unfortunately for Mitchell, Spanish pirates plundered his England-bound ship, and most of his collection was lost. With the help of Collinson, Mitchell published several new genera, but Linnaeus never saw any of the surviving specimens. In New York, Cadwallader Colden (1688--1776) became a correspondent of Linnaeus via Collinson and sent him a manuscript on the plants of that colony. This was published in 1749 and 1751. Unfortunately, the same pirates took his specimens and none were recovered. John Bartram (1699--1777) of Philadelphia sent seeds and dried specimens to Sloane, Sherard, Collinson, Dillenius, and Gronovius, and through them to Linnaeus, promoting his botanical garden (the first in the United States, founded in 1731) and his desire for a royal appointment as the King's Botanist.

Pehr Kalm (1715--1779), who had studied with Linnaeus, arrived in New York in September 1748 and was soon collecting seeds and plants in New to southern Canada; in 1750 he went across western Pennsylvania to the Great Lakes. When Kalm left the United States in October of that year, he was heavily laden with all kinds of plants, not just those of interest to the gardener or physician.

The Prince of Botany

The stature of Linnaeus had grown significantly in the eyes of his fellow naturalists after he had left Holland for Sweden in 1738. His writings were eagerly awaited and carefully studied by naturalists. Specimens were sent to him, often via intermediaries such as Collinson or Gronovius, for identification and naming. The simplicity of his system attracted many, and anyone could readily determine if a given plant was known to the great man. His production was prodigious. Edition after edition of Systema Naturae and Genera Plantarum... (ed. 1, Leiden, 1737) was published. A summary of the world's flora at the species level, however, was more difficult to achieve.

Three problems appear to have troubled Linnaeus: the morass of names created by previous writers, the increasing number of new species being found, and the vast array of existing specimens and published literature. He had already developed a concise style of presentation, which he had used in Hortus Cliffortianus...(Amsterdam, 1737), and specimens and literature were being sent to him from all over the world, notably duplicates of Clayton's Virginia plants from Gronovius, and seeds and dried specimens from Collinson and Miller, and from Madrid, Paris, and Vienna. The most recent items were less troublesome. The older literature and collections, however, remained difficult. Linnaeus studied the herbarium of Joachim Burser (1583--1639) to understand the dispositions of names according to Bauhin's Pinax, consulted the herbarium and drawings of Hermann's Ceylon plants, and examined copies of the original drawings made by Father Charles Plumier (1646--1704) of plants observed in the West Indies.

Preventing Linnaeus from fully comprehending the past were the names associated with the specimens scattered among the vast holdings of Hans Sloane. Through his many friends, Linnaeus sought to obtain the older literature so that he might study it. Even so, as he struggled with writing a species plantarum, he encountered too many unanswered questions, and in 1748, with the manuscript only half finished, he gave up.

In June 1751 Kalm arrived at Uppsala with his American discoveries. He had kept a diary and wanted to publish therein the names of the plants he had found. He called on his mentor to identify the remainder of his plants because he was certain several were new. Earlier, in 1749 and 1751, Linnaeus had described several of Kalm's new genera, but the task of naming all of the species was a challenge.

To what degree Kalm's arrival prompted Linnaeus to attempt again a summary of the world's flora is not known, but the two events coincide so closely that it is likely that Kalm provided at least some impetus. This time, however, instead of trying to account for every species and all names, Linnaeus treated only those species he felt he had some degree of confidence to address, and he cited only a selected synonymy. Although he had not adopted the use of binomials in his first draft, the logic of using them became obvious in 1751: each species would have a simple name that the uninformed could use, and by making each unique within a genus, indexing was simplified. Only 13 months after he had started the new draft, and in spite of an extended illness, he completed the task.

Some last-minute additions were inevitable. Pehr Osbeck (1723--1805), chaplain of a ship, returned from China with numerous boxes of plants just as the book was being readied for the printer, and garden material (mainly from Bartram via Collinson or Miller) continued to arrive. The novelties were inserted, and Species Plantarum, the long-sought summary of the world's flora, was published in two volumes at Stockholm in 1753. Linnaeus had accounted for some 8000 species from all parts of the known world. He revolutionized the science of taxonomy and ushered in a new era. A milestone had been set in the progress of knowledge, and no one knew it more than Linnaeus himself. He would thereafter be, as he christened himself, the Prince of Botany.

"A Child of Fortune"

The majority of botanists, although not all, readily adopted Linnaeus's innovative nomenclature and classification scheme, and modeled their work accordingly. The adoption of his system by the North American community was championed by Collinson and John Ellis (1710--1776) in London. From southeastern Canada to Georgia, plants suspected to be new were sent to Europe in the hope that they would prove to be novelties and that the great Linnaeus himself would name them. Some, such as Jane Colden of New York, Cadwallader Colden's daughter, labored in obscurity. Others, such as Alexander Garden (1730--1791), became famous, although he waited a long time before hearing that some of his new plants would be given names.

With Collinson's help, John Clayton prepared his own edition of a Virginia flora, but his hopes were ended when Gronovius published a second edition of Flora Virginica in 1762 (Leiden). Clayton's manuscript and associated drawings by Georg Ehret (1708--1770) lost when Clayton's clerk's office was set ablaze by an escaped prisoner; the remainder of his correspondence and library were destroyed in a house fire around 1906.

The collections of John Bartram and his son William (1739--1823) fared better. They had been collected during John Bartram's travels from Lake Ontario to Florida. As they arrived in England, they were dispersed among Sloane and his friends. Collinson sent seeds and specimens to Linnaeus, but many of their new species remained undescribed. In 1768 Philip Miller named many of the Bartrams' curiosities, and Nicolaus Jacquin (1727--1817) named others from his gardens in Vienna.

Younger men were entering the field, especially in England, to carry on the botanical tradition started by Sloane and furthered by Collinson and Ellis. Foremost among them was Sir Joseph Banks (1743--1820). After his father's death, the young Banks settled in London where, as a "child of fortune," he became acquainted with Philip Miller and others associated with the Chelsea Physic Garden. In 1766 Banks sailed for Labrador and Newfoundland, where he made numerous collections that allowed him, as a newly elected member of the Royal Society, to become acquainted with others interested in the plant kingdom. This resulted in his voyage to Australia for botanical discoveries that were to earn him lasting fame.

Sailing with Banks in 1768 was Daniel Solander (1736--1782), a Linnaean student who had become the assistant librarian at the newly established British Museum in 1762. As Banks's personal librarian, Solander had access not only to the specimens the two had collected while traveling around the world from 1768 to 1771, but also to those available to him as Keeper of the Natural History Department at the museum. As a result, Solander finally gained access to the collections Sloane had acquired over his 93 years, and for the first time it was possible to study the original specimens used by Plukenet, Petiver, and others to describe species that had long puzzled Linnaeus. Because of his own great wealth and interest, Banks too obtained herbaria, including those of Gronovius and Jacquin.

As president of the Royal Society, a position he assumed in 1778 and retained until his death, Banks was able to direct the interests of the society into many corners of the world in search of botanical novelties. The world's largest collection of plants lay before Solander, and he set to work describing the thousands of new species he was finding. His early death prevented his publishing much, but through the work of his successors as Banksian librarians, Jonas Dryander (1748--1810) and Robert Brown (1773--1858), many of Solander's names were ultimately published. Unfortunately, the authorship of most of his names for American species must be credited to William Aiton (1731--1793), who, as head gardener for Princess Augusta of Saxe-Gotha at Kew, was the author of the three-volume Hortus Kewensis... (London, 1789), in which most of the names were formally established. Even the efforts of Francis Masson (1741--1805), one of Kew's foremost collectors, who worked briefly in Canada, were relegated to obscurity in a similar manner.

Views of Enlightenment

The Linnaean Society of London, established in 1788, had as its first president James Edward Smith (1759--1828), who, with Banks's encouragement, had purchased Linnaeus's library and herbarium in 1784. Smith championed the Linnaean system for the next half century, after it had outlived its usefulness. In England, Robert Brown (1773--1858) and John Lindley (1799--1865) led the opposition to it. In France, the changes in social values brought about by the Revolution of 1789 coincided with the acceptance of a natural system of classification. Antoine Laurent de Jussieu (1748--1836), nephew of Bernard de Jussieu and friend of Linnaeus, in his Genera Plantarum... (Paris, 1789), arranged the genera of the world into 100 families (ordines naturales) based on concepts developed by his uncle Bernard, in a continuation of the ideas proposed a generation before by Michel Adanson (1727--1806) in his Familles des Plantes (2 vols., Paris, 1763[--1764]).

Adanson's views were unconventional. As had Pierre Magnol (1638--1715), Sloane's professor, long before him, Adanson believed that plants could be arranged into natural families and genera. It was a classification free of a priori weighting and metaphysical themes, all entities circumscribed so as to reflect their affinities as determined by the sum total of their features, and a system derived only from an empirical search for similarities and discontinuities.

The differing views of classification had little impact in America, and new species from there continued to be described by Europeans. Essentially, no one in America was independent enough of European authority to risk expressing his own opinion. Linnaeus continued to revise his Species Plantarum and to add new American species to its pages. He occasionally recognized names proposed by others, but the many names of John Hill (1716--1775) and Miller, each of whom had described several North American species, were not taken into consideration. In Linnaeus's own view, binomials were trivial names, only for the uninitiated, and were never intended to be used by the knowledgeable naturalist. Therefore, when Gronovius published the second edition of Flora Virginica in 1762, he used polynomials, not binomials. Consequently, this work is now nomenclaturally irrelevant by virtue of the current code of botanical nomenclature.

To describe their new species, the Bartrams clearly depended on others, e.g., Aiton in Hortus Kewensis, Jean-Baptiste de Lamarck (1744--1829) in his 13-volume Encyclopédie Méthodique (Paris and Liège, 1783--1817) and Charles-Louis L'Héritier de Brutelle (1746--1800) in his Stirpes Novae. The first flora authored by an American, Flora Caroliniana... (London, 1788) by Thomas Walter (1740--1789), was published and heavily edited in England by John Fraser (1750--1811). The first wholly American contribution was Arbustum Americanum..., by Humphry Marshall (1722--1801), published in Philadelphia in 1785. Both floras were in the Linnaean style. Some early American taxonomic works appeared in the Transactions of the American Philosophical Society. Unfortunately most of the names proposed by H.Muhlenberg (1753--1815) are invalid, according to the modern code of nomenclature (W.Greuter et al. 1988). Likewise, the validity of the names mentioned by the younger Bartram in his 1791 Travels... (Philadelphia) is still in doubt.

The year 1789 marked the end of the domination of the Linnaean classification. Linnaeaus's herbarium and library were in London, and binomial nomenclature was widely adopted. New editions of Species Plantarum were being written by others. Nevertheless, the Linnaean manner of presentation continued into the 1830s. Importantly for American naturalists, however, the War of 1812 with England broke the bond that demanded their unquestioned acceptance of the Linnaean method. This allowed North American naturalists to be influenced by the radical views of the French and the French naturalists. Also in 1789, Benjamin Smith Barton (1766--1815) began teaching natural history at the University of Pennsylvania, and an American flavor was added to taxonomy. Others soon followed, notably Samuel L. Mitchell (1764--1831) at Columbia University in 1792. The center of American botany was, nevertheless, firmly established in Philadelphia.

The Continent Is Opened

The establishment of the center of botany in Philadelphia was effected through the efforts of the Bartrams, their garden, and a host of others with scientific backgrounds who resided in that city after the Revolutionary War. Students interested in the native flora flocked to use Barton's large library and herbarium. Ultimately, Barton sponsored the fieldwork of Fredrick Pursh (1774--1820) and Thomas Nuttall (1786--1859) as part of his dream of writing a flora of North America. He also introduced William Baldwin (1779--1819), William Darlington (1782--1863), and Yale's Eli Ives (1779--1861) to taxonomy and aided them in their early careers.

While the American scene was developing, the French André Michaux (1746--1802) and his son, François André Michaux (1770--1855), began their explorations of temperate North America. In 1785, they traveled from Lake Mistassini to Florida in search of plants, with special emphasis on potential garden introductions. In 1803, a year after the elder Michaux had died, Flora Boreali-Americana... (2 vols., Paris and Strasbourg) was published with the aid of his Paris colleague Louis Claude Richard (1754--1821). At last, temperate North America had an account of its native plants.

Michaux's book used a modification of the Linnaean and Jussieu styles as promoted by Augustin Pyramus de Candolle (1778--1841) in his revision of Lamarck's Flore Françoise... (ed. 3, 5 vols., Paris, 1805). Michaux illustrated some of his new genera and gave precise habitat information. He did not, however, include keys, the shorthand method for identification that Lamarck had developed in an earlier edition of the Flore. Michaux had talked with Barton and visited his garden. Consequently, unlike the works of Linnaeus that were still in common use, Michaux's was based on firsthand observation and a careful comparison of his specimens against those used by others who had named plants previously. This work prodded Barton to proceed with his flora, a small part of which had been published in London in 1787 as Observations on Some Parts of Natural History..., Part 1.

Criticisms of Michaux's work abounded. Barton was not satisfied, and he urged his students and colleagues into the field to collect the new species that he knew existed to the west. Pursh, then in London, published Flora Americae Septentrionalis... in 1814 (2 vols., London). Similar in format to Michaux's work, this one included the new species collected by Barton, by Pursh for Barton, and by Meriwether Lewis (1774--1809), President Jefferson's personal secretary, who was in the Far West as part of the Lewis and Clark expedition of 1804--1806. A few of the expedition's early specimens had reached the president, who sent them on to Barton for naming. All the specimens found in 1805 were lost, and only those collected on the hasty return trip of 1806 reached Philadelphia. Even so, these specimens were ignored, and had Pursh not purloined snippets of the plants, they might not have been consulted for years.

The Lewis and Clark specimens represented the first American contribution from the western part of North America. But Lewis was not the first to collect specimens from that area. Georg Steller (1709--1846) had managed, in six hours, to collect 141 specimens on a small island off the coast of Alaska in 1741. After his death, these specimens were sent to Linnaeus, who accounted for about 30 of them (all from Russia) in his 1753 work. Plants from coastal California were first collected in 1786 by a French gardener, Jean-Nicolas Collignon (1762--1788), and Lamarck described the first species from west of the Mississippi, Abronia umbellata, five years later.

The English, Spanish, and Russians all sent out early expeditions that touched land from San Diego to Alaska. In 1791, the Malaspina expedition visited Monterey in California, Nootka Sound in British Columbia, and Yakutat Bay in Alaska. Its naturalist, Thaddeo Haenke (1761--1817), made the first large collection of dried specimens to reach Europe from this part of the world. Many of his new species were described by Karl Presl (1794--1852) in the two-volume Reliquiae Haenkeanae... (Prague, 1825--1835). Archibald Menzies (1754--1842) was the first of many British naturalists to explore the Far West. He was with the Vancouver expedition when it surveyed the Pacific Coast from Chile to Alaska from 1792 to 1794. Many of Menzies' new species were named by James Edward Smith in articles written between 1802 and 1820 for Abraham Rees's Cyclopaedia (39 vols., London).

Alaska was then a possession of the Russians, and when the von Krusenstern expedition arrived in Sitka in July 1805 during their round-the-world voyage, they found at anchor an American ship, the Juno. The Russians purchased the ship, and in March 1806, a small company set sail for the coast of California to obtain supplies. On board was Georg von Langsdorff (1774--1852), surgeon-naturalist, who botanized briefly near San Francisco in April. In his published remarks (Bemerkungen auf einer Reise um die Welt, London, 1813), however, Langsdorff did not account for a single California plant.

The first specimens from Greenland were collected by Hans Egede (1686--1758) and his son Poul (1708--1789), starting in 1728. Unknown to Linnaeus, these specimens went unreported. The first major collection was made by Morten Wormskjold (1783--1845), who led a naval expedition to Greenland in 1813. In the 1830s, Jens Vahl (1796--1854) made a large collection there.

The first botanical explorer of Newfoundland and St. Pierre and Miquelon was Bachelot de la Pylaie (1786--1856) who collected plants on trips in 1816 and 1818. His specimens are housed in Paris. The Moravian missionaries, compatriots of the Egedes of Greenland, were active along the coast of Labrador. They sent carefully prepared specimens across the ocean to Banks in the 1780s, and then to others in England and the United States in the nineteenth century. John Goldie (1793--1886) collected near the lower Great Lakes in 1819, and James McNab (1810--1878) collected several hundred specimens in southeastern Canada in 1834. Both men were mainly interested in plants of horticultural significance. Through the nineteenth century, many collectors sent specimens to German botanists, and until the 1890s, American recipients often obtained materials from German intermediaries.

The first Arctic expedition of botanical significance was led to Baffin Bay by John Ross in 1818, and it included the naturalist Edward Sabine (1788--1883). The three expeditions of Edward Parry (1790--1885), between 1819 and 1825, resulted in Robert Brown's Chloris Melvilliana (London, 1823), so named because many of the plants were collected on Melville Island.

Exploration of the interior of North America accelerated in the early nineteenth century. Thomas Nuttall was 22 years old in the spring of 1808 when he first met Barton in Philadelphia and began to make use of his extensive library. Nuttall proved to be a willing student and an even more willing explorer. He actively sought plants near Philadelphia, and in April 1810, when Barton offered him eight dollars a month plus expenses to collect in the Old Northwest, Nuttall eagerly accepted. Within days he was heading for the upper reaches of the Mississippi River. Later, Nuttall proceeded to St. Louis where he accepted an invitation to travel up the Missouri River the following March with a fur-trading expedition led by Wilson Price Hunt of the American Fur Company.

Also in St. Louis was a Scottish naturalist, John Bradbury (1768--1823). He too had accepted an offer to join Hunt, and for a while he and Nuttall traveled jointly but separately, up the Missouri, each collecting wherever possible. In June, Manuel Lisa and his Missouri Fur Company caught up with the Hunt party. Lisa was trying to reach the Arikara Indians first, and Bradbury sensed that if he joined Lisa, he could beat Nuttall to the botanical treasures. Both groups traveled their separate ways to the Mandan villages in what is now North Dakota before returning to St. Louis in the fall of 1811, loaded with a large number of new and exciting plants.

Shortly thereafter, Nuttall went to New Orleans, where he sailed for England, rather than Philadelphia as he had agreed with Barton. The War of 1812 was imminent, and Nuttall felt it was prudent to return to his native England. Meanwhile, Bradbury had remained in Louisiana to collect, although he had sent his Missouri specimens to London. Bradbury was trapped in America by the War of 1812, while Nuttall, his plants, and those of Bradbury sailed peacefully away.

The consequences were devastating for Bradbury. In England, Pursh gained access to Bradbury's collection and described most of the new species in 1813. Nuttall technically avoided being scooped by Pursh when some of his species were described in a catalog of new plants from "the river Missourie," issued by John Fraser earlier in 1813. Shortly after, both Pursh and Nuttall published new species in Curtis's Botanical Magazine, each description augmented by a colored plate.

Nuttall returned to the United States in 1815 and collected throughout much of the Southeast. He worked feverishly in Philadelphia to update the Michaux and Pursh floras in his Genera of North American Plants... (2 vols., Philadelphia, 1818).

Other works also appeared. From Charleston, John Linnaeus Shecut (1770--1836) published his Flora Carolinaeensis... in 1806. A decade later Stephen Elliott (1771--1830) followed with A Sketch of the Botany of South-Carolina and Georgia (2 vols., Charlestown, 1821--1824). In 1814 Jacob Bigelow (1787--1879) published Florula Bostoniensis (Boston). All of these works followed the Linnaean method. Amos Eaton (1776--1842) included a section on the natural system as envisioned by Jussieu in his Manual of Botany... (ed. 1, Albany, 1817), but he retained the plants in the familiar Linnaean sequence. Even Nuttall's work retained traces of the Linnaean era, although as early as 1808 he was using Jussieu's ordinal names. The first author of an American text to follow Jussieu's natural system was J.F. Correa da Serra (1750--1823), who published a Reduction of All the Genera...in the Catalogues...of Muhlenberg... in 1815 (Philadelphia). C.S. Rafinesque (1783--1840) also used a natural system in his 1817 Florula Ludoviciana... (New York). Two years earlier, he had recognized more than 300 natural orders, many of which were adopted by others over the next thirty years without, as Rafinesque complained, giving him credit for their authorship.

As the second decade of the nineteenth century closed, the center of American systematics moved again, this time from Philadelphia northeastward. In New York, David Hosack (1769--1835) established the Elgin Botanic Garden, the first public garden in the United States, and published a catalog of its plants in 1806 (New York). A likable person, Hosack attracted several young people into botany, the most notable being Amos Eaton, then a student of law. While he was jailed for his dealings in a troublesome land sale, Eaton taught the young son of the prison's fiscal agent his first lessons in the Linnaean method; the youngster was John Torrey (1796--1873).

Only 10 Feet Less

Torrey had been educated and trained in the usual haphazard way of his generation. His training in medicine easily allowed him to assume teaching positions in chemistry, botany, and mineralogy. Torrey was an excellent teacher as well as a skilled botanist. Both he and Eaton were to share in the labors of the one person who came to dominate American taxonomy more than any other, Asa Gray (1810--1888). A half century later, two peaks in the Colorado Rockies were named for Torrey and Gray, with Torrey's Peak only 10 feet less in elevation than Gray's---a fitting tribute.

Eaton held fast to the Linnaean way; Torrey faltered, then adapted. Torrey held firmly to tradition through his 1826 Compendium of the Flora of the Northern and Middle States (New York) only to change that very year in an article on Rocky Mountain plants. When the first part of Torrey and Gray's Flora of North America... (2 vols., New York, London, and Paris, 1838--1843) appeared, the Linnaean era in America was ended, and Lindley's natural system was promoted by America's new voice of authority. Even Eaton gave in, in a fashion, urging Gray to take over his regional flora for the northeastern United States, which Gray eventually did. In many ways, the future of American botany was now firmly in the hands of Asa Gray.

When Gray went to Harvard University in 1842, following Nuttall, and Torrey to Columbia and Princeton, after years at West Point, the simple fact of distance lessened their joint efforts. Westward exploration and collecting intensified during this time, and the resulting specimens sent to them for naming overwhelmed them with responsibilities and an abundance of new species.

The Age of Exploration

The increase in plant collecting was stimulated in part by the preparation of a flora that addressed a significant portion of temperate North America, William Jackson Hooker's (1785--1865) Flora Boreali-Americana... (2 vols., London, Paris, and Strasbourg, [1829--]1833--1840). This work was devoted primarily to the British-dominated portions of the continent and was based on extensive field studies by several explorer-naturalists, the foremost being David Douglas (1799--1834) of the Royal Horticultural Society of London.

Douglas, who was particularly interested in oaks, first came to North America in 1823, confining his botanical activities mainly to the middle Atlantic region and as far west as Michigan and southern Ontario. He no sooner had returned to London in January 1824 than he was urged to proceed to western North America. On 26 July 1824 Douglas left Gravesend for the mouth of the Columbia River. He collected extensively in the Oregon Country and crossed Canada to reach York Factory on Hudson Bay in 1827 for a brief trip back to England. In 1829 he returned to Fort Vancouver and made side visits to California and the Hawaiian Islands, where he died in July 1834.

Douglas's enthusiasm inspired others. Traveling with him to Northwest America in 1824 was a fellow Scot, Dr. John Scouler (1804--1871), a surgeon-naturalist. Scouler collected from Fort Vancouver on the Columbia River to Nootka Sound and made his specimens available to W.J. Hooker, his former professor at Glasgow. A Hudson's Bay Company surgeon, William Fraser Tolmie (1812--1886), collected mainly in Washington and perhaps British Columbia, climbing Mt. Rainier in August 1833. Later, a fur trapper, John McLeod, collected along the Oregon Trail from Fort Vancouver to the 1837 rendezvous site on the Green River in Wyoming. He is "Tolmie's friend" to whom Hooker assigned so many unusual species in his flora.

In Canada, Dr. John Richardson (1787--1865) and Thomas Drummond (1780--1835) were associated with the second Franklin "Land Expedition to the Polar Sea" of 1825--1827, a part of the great search for the Northwest Passage involving the Parry and the Beechey expeditions. Both Richardson and Drummond made large collections from the Mackenzie River region south and eastward to the Rocky Mountains and Hudson Bay. The collections of Douglas, Richardson, and Drummond were significant for Hooker's flora.

Those made by Alexander Collie (1793--1835) and George Lay (d. 1845) on the Blossom expedition, commanded by Frederick Beechey, proved to be of lesser significance, although several new species were found in California, Mexico, and other regions of the world. W.J. Hooker and George Walker-Arnott (1799--1868) described the plants made on this 1825--1828 voyage over many years. Even so, most of the new species from North America for which they accounted in their Botany of Captain Beechey's Voyage... (10 pts., London, [1830--]1841) were based on specimens in the collections of Menzies and Douglas.

In Alaska, Carl Mertens (1796--1830), naturalist with the Russian Lutke expedition, visited Sitka in 1827 aboard the Senjavin. His plants were described by the German-born Russian botanist August Bongard (1786--1839) in 1833. Ilja G. Vosnesensky (1816--1871), who had collected at Fort Ross in California in 1842, collected plants during his visits to Alaska between 1840 and 1849, often in collaboration with others of the Russian-American Company.

In the frozen Arctic, John Rae (1813--1893) and David Lyall (1817--1895), among others, made many plant collections from previously unexplored islands. Meanwhile Richardson and others searched from 1847 to 1850 for some trace of the third Franklin expedition, which had ended in tragedy. Joseph Dalton Hooker (1817--1911) subsequently accounted for the Rae and Lyall specimens.

The success of the Lewis and Clark expedition was not repeated by the United States government for some 30 years. The Freeman expedition up the Red River of the South in 1806 was repulsed by the Spanish, as was the one led by Zebulon Pike into the Colorado Rockies the same year. In 1819, however, Nuttall ventured up the same Red River into what is now Oklahoma, and his findings were numerous. The same year, Dr. William Baldwin (1779--1819), who had studied with Barton, traveled up the Missouri River on the Yellowstone expedition, which included Stephen H. Long of the topographical engineers. Delays and the death of over a hundred men due to scurvy prompted Congress to halt the expedition's funding. Baldwin died at Franklin, Missouri, during his own attempt to reach St. Louis, with little to show for his efforts.

Edwin James (1797--1861) replaced Baldwin as surgeon-naturalist in 1820, joining a new Long expedition in St. Louis. With him was Thomas Say (1787--1834), a self-taught naturalist who became one of America's best-known zoologists and a distinguished entomologist. The route was westward to the Rocky Mountains of central Colorado; it was direct and rapid, and both men collected whenever they could. Once in the Rockies, James climbed Pikes Peak in June, where he found many alpine species. By September, the expedition was at Fort Smith in the Arkansas Territory on its return leg to St. Louis.

Baldwin and James's specimens were sent, eventually, to Torrey, who accounted for them in 1824 and 1827. Less delayed in their description were the plants collected by Captain David Douglass (1790--1849), then at West Point, who accompanied the Cass and Schoolcraft expedition on their search for the headwaters of the Mississippi River in 1820. Torrey published his summary of Douglass's plants in 1822.

Lewis David von Schweinitz (1780--1835), the noted mycologist and monographer of North American Carex, wrote a catalog of the small number of plants Say found along the Red River of the North in 1823 while on the Long and Keating expedition. Just before he died, von Schweinitz purchased Baldwin's herbarium from the latter's widow; this was eventually deposited at the Academy of Natural Sciences in Philadelphia, and many of its new species were described by Torrey or by Nuttall.

Westward Expansion

Torrey and Gray realized the importance of collectors exploring remote regions of North America, and if they were to account for its plants, skilled botanists had to be in the field whenever possible. The increasing number of government expeditions provided them with a potential source of novelties, and Torrey lobbied hard to ensure that naturalists loyal to him and to Gray were attached to as many expeditions as possible. Even when Congress seized on the notion of a grand, worldwide expedition, such as the British had conducted a half century earlier, Torrey saw to it that William Brackenridge (1810--1893) was on Wilkes's round-the-world voyage.

To be sure, private collectors, friends, and even remote acquaintances provided Torrey and Gray with exciting specimens, especially from the southeastern part of the United States. Nuttall's friends, Hardy Croom (1797--1837) and Harris Loomis (d. 1837), indirectly contributed plants, as did Rafinesque's replacement at Transylvania University, Charles Short (1794--1863), also one of Barton's students. Torrey and Gray's cadre of collectors in the 1830s was impressive, and specimens came to them from many who were to make substantial contributions in their own right: Lewis Beck (1798--1853), John Blodgett (1809--1853), Samuel Boykin (1786--1848), Samuel Buckley (1809--1884), Elias Leavenworth (1803--1887), John LeConte (1784--1860), Charles Pickering (1805--1878), Zina Pitcher (1797--1872), John Riddell (1807--1865), William Sullivant (1803--1873), and Edward Tuckerman (1817--1886).

In the still unexplored American West, other adventurers and naturalists were finding a wealth of novelties. Nathaniel Wyeth (1802--1856), a Boston fur trader, followed the Oregon Trail from St. Louis to Fort Vancouver in 1832. He returned the following year via the Clark River of northern Idaho to the Flathead Post in Montana, then south to Fort Bonneville in Wyoming. He submitted his large collection to Nuttall, then at Harvard, to name. This Nuttall did, but before the paper was published, Nuttall was headed west himself with Wyeth. Nuttall collected thousands of specimens during the next three years in the Northwest, California, and Hawaii. Several hundred species were described ultimately by Nuttall or by Torrey and Gray in Flora of North America.

During this time, Thomas Drummond left the Canadian north and collected from Florida to Texas from 1831 to 1835, his specimens going to W.J. Hooker in London. The specimens collected earlier in Texas by Jean Berlandier (1805--1851) had been sent to Alphonse de Candolle (1806--1893) in Geneva. Those of a later immigrant, the German Ferdinand Lindheimer (1801--1879), were studied by Gray, and the novelties were published with George Engelmann (1809--1884) of St. Louis between 1845 and 1850. Adolf Scheele (1808--1864) also published seven articles on Lindheimer's Texas plants, from 1848 to 1852, in the European journal Linnaea.

The westward movement of Americans began in earnest in the 1840s, but the challenge of the great grassy plains (termed the "Great American Desert" by Pike) was one few wished to risk. Missouri Senator Thomas Hart Benton knew that to open the West, the United States must have a safe route to the Pacific Coast. Who better to find it than his son-in-law, John Charles Frémont (1813--1890)?

Frémont's expeditions are now schoolboy legends, complete with the exploits of Kit Carson and "Broken-hand" Fitzpatrick, the lost cannon, and the battle of San Pasqual. The name of Frémont is found in every western state, but few know that many of the plants on the peaks, near the rivers, or in the counties that bear his name were also found or named by Frémont. Yet he was but one of many explorers, naturalists, soldiers, and scientists for the U.S. Army's Corps of Topographical Engineers that Torrey and Gray urged to collect plants.

The war with Mexico saw Frémont in the West once again, conquering California and collecting plants. Even the military commander, Major William Emory (1811--1887), and his "Army of the West" collected plants as they crossed New Mexico and Arizona to reach the Pacific Coast and claim California in the name of the U.S. Army from Frémont, self-proclaimed military governor. After the war, the Mexican boundary survey, led by Emory, collected more western specimens with the assistance of Charles Parry (1823--1890), George Thurber (1821--1890), and Charles Wright (1811--1885).

The fulfillment of Benton's westward vision brought to the United States more territory than just that resulting from the war with Mexico. With the acquisition of the Oregon Country in 1846, the United States stretched across a continent. The activity associated with this expansion brought more new plants to the attention of the small American botanical community around Torrey and Gray.

While Torrey and Gray labored on their continental tome, manuals and regional floras were published. The eight editions of Eaton's Manual informed naturalists about the plants in the northern states. In 1841, John Darby (1804--1877) of Georgia published a companion Manual (Macon) for the southern states, followed by his Botany of the Southern States in 1855 (New York, Cincinnati, and Savannah). After Eaton's death in 1842, Gray began to work on his own regional flora, A Manual of the Botany of the Northern United States. It was first published in 1848 in Boston, Cambridge, and London, and went through numerous editions.

Alvan Chapman (1809--1899) published the first edition of his Flora of the Southern United States in 1860 (New York), and it went through two more editions, the last appearing in 1899. Journals established in different parts of the United States allowed American naturalists, even amateurs, to publish their new discoveries. The results were overwhelming, and it was all Torrey and Gray could do to keep up with teaching and with writing their government reports. The long desired flora of North America was an impossibility as long as novelties arrived almost monthly from the West.

At Harvard, Gray's influence grew as that of Torrey's waned, and the center of North American botany moved from New York to Cambridge. Gray, the modernist, had readily adopted the arrangement of natural orders proposed by Lindley in 1830 (An Introduction to the Natural System of Botany..., London) and had developed keys to the families similar to those in Lindley's Key to Structural, Physiological and Systematic Botany..., published in 1835 (London). These were now standard in most American floras. Gray also included keys to the genera, taking the idea from the British naturalist Samuel Frederick Gray (1766--1828), who first used them in 1821 in his Natural Arrangement of British Plants (London). Asa Gray did not include keys to species in his works. Rather, species were arranged into groups under diagnostic headings, following the format designed by de Candolle for his Prodromus... (17 vols., Paris etc., 1823--1873).

As a professor, Gray realized that students needed a useful, concise textbook in botany. Many were available, but those in English were nearly always directed toward the British student. American textbooks were few, and Barton's Elements of Botany... (ed. 1, Philadelphia, 1803) was outdated. Once again, Lindley provided a model, and in 1836 Gray published his own Elements of Botany (New York). Its major competitor was Class-book of Botany..., by Alphonso Wood (1810--1881), which first appeared in 1845 (Boston and Claremont, N.H.). Almira Hart Lincoln (later Phelps; 1793--1884), better known as "Mrs. Lincoln," began publishing her Familiar Lectures on Botany in 1829 (Hartford, New York, Boston). The three works sold over two million copies!

The Patriarch

The trip to Europe that Gray took in the winter of 1839 was fundamental for many reasons besides allowing him to study critical type material. Although Gray paid his compliments to Robert Brown and met the elder Hooker and his son Joseph at the British Museum, it was George Bentham (1800--1884), who was closer to his age and temperament, with whom he felt a companionship. Bentham was rapidly becoming England's foremost botanist. He had recently completed a study of the North American eriogonums, a favorite group of Torrey's, and was interested in mints throughout the world.

Gray had been following established tradition, working on regional and national floras, reviewing only those species of a group as he came to them. Bentham, however, tended to study a group throughout its range, thereby gaining a greater understanding of it. The idea of examining a group throughout its range was reasonable, providing one had access to libraries and collections. This was not a realistic expectation for Gray who, at the time, was considering a position at the University of Michigan. Still, several wholly American genera could do with his attention, and most were in Asteraceae. Therefore, like Bentham, Gray began to write both monographic and floristic treatments, and he encouraged others to do the same.

When Gray returned to England in June 1850, he was fully established as America's leading plant taxonomist and perhaps its leading botanist, and Bentham's position was no less secure. With Bentham, Gray worked over the plants from the Wilkes expedition and continued his studies of type material. In Dublin, Gray visited William H. Harvey (1811--1866), the algologist who had accompanied Wilkes. When Joseph Hooker returned to London in the spring of 1851 with his collection from the Ross expedition to India, Gray was at hand to glimpse the wonders from another part of the world. Here, too, Gray met Richard Owen and Thomas H. Huxley, both of whom played important roles at the end of the decade in the debates on evolution.

Awaiting Gray at home in the fall of 1851 were specimens from more expedition botanists, friends, and colleagues. He published on the specimens collected by Wright and Thurber on the boundary survey; the publications from those of Wilkes and Emory were delayed; and no sooner had he started again on the flora than the railroad survey specimens began to accumulate.

When the theory of evolution began to have an impact in 1859, Gray was in the thick of it, and the time to devote to the flora was limited even more. Bentham was no less overwhelmed with his tasks. Bentham, however, persevered in his labors, and his tasks were completed while Gray's rarely were.

Other events conspired to slow the work both of Gray and the plant collectors. Aside from the fiscal problems at Harvard and the demands of students, the American Civil War was raging. Torrey was nearing 70, and Daniel C. Eaton (1834--1895) at Yale, the only other full-time botanical professor in the United States, was heavily committed to the study of ferns and their allies. Gray attempted to turn again to his much-neglected Flora of North America with the hope of producing another volume. It was not to be.

Decentralization of American botany had now begun. In Washington, D.C., the new Smithsonian Institution was considering the establishment of a botany section, and the newly instituted Department of Agriculture was supporting the idea of a National Herbarium. Businessman Henry Shaw (1800--1889) established a botanical garden in St. Louis, with the advice of George Engelmann, Asa Gray, and J.D. Hooker. On the West Coast, Albert Kellogg (1813--1877) and his friends had established the California Academy of Sciences in San Francisco in 1853, with the intent that their institution would be equal to any on the East Coast.

The Morrill Act of 1862, signed by President Lincoln during the war, said that the curricula of all land-grant colleges (i.e., state-supported colleges emphasizing agriculture and applied sciences) were to include botany, and in particular taxonomy. This resulted in numerous opportunities for young botanists to find work in some of the still relatively unexplored regions of the West. Therefore, as Gray considered retirement, the botanical community became fragmented, and the patriarch began to lose control.

Branching of Talent

Botanical exploration in western North America after the Civil War increased, albeit with limited governmental support. In Canada, John Jeffrey (1826--1854) arrived at York Factory on Hudson Bay in August 1850 and worked his way westward to British Columbia and eventually California, collecting for botanical gardens in Scotland. Eugène Bourgeau (1813--1877), with backing from the Royal Geographic Society, collected some 60,000 specimens from the Great Lakes to the Rocky Mountains over a two-year period (1857--1859) before going to Mexico, where he made even more extensive collections.

In 1858 George Lawson (1827--1895), the "father of Canadian botany, " moved to Canada from Edinburgh and soon thereafter established the Botanical Society of Canada. In 1860 the first issue of the Society's Annals was published. Lawson also proved to be an inspiration for amateur botanists throughout the country, encouraging many to publish their observations. One such amateur, the Abbé Léon Provancher (1820--1892), published Flore Canadienne... (2 vols., Quebec, 1862[1863]), which treated mainly the plants of eastern Canada, although he accounted for the species from Ontario reported by W.J. Hooker. Provancher founded the journal Le Naturaliste Canadienne in 1869.

To the north and in Alaska, Robert Kennicott (1835--1866) of the Chicago Academy of Sciences led a series of collecting forays from the Great Slave Lake region to Fort Yukon. On other trips he ventured to British Columbia, the Mackenzie River, and, in 1864, Sitka. Accompanying him on various trips were Joseph Rothrock (1839--1922), who later collected in the southern Rocky Mountains, and William H. Dall (1845--1927), who included a list of "useful plants" in a book he wrote in 1870 on Alaskan resources.

Geological surveys became the modern means for botanical explorations. In the United States, the survey of the fortieth parallel, led by Clarence King (1842--1900) from 1867 to 1874 across Nevada and Utah, resulted in the discovery of numerous new plants, most of which were found by the camp cook turned naturalist, Sereno Watson (1826--1892), in 1868 and 1869. A shy man with a checkered past, Watson took his collections to Cambridge where, with Gray's assistance, he worked up the results. By 1872 Watson was a permanent fixture, taking over much of Gray's correspondence with western botanists and serving, essentially, as curator of the herbarium.

The following year Gray formally retired from Harvard with the intent of devoting the remainder of his years to taxonomy. Unfortunately, Torrey had died shortly before Gray retired, and so had Louis Agassiz (1807--1873), Harvard's famed zoologist and fellow professor; Gray was now free to concentrate on the North American flora, but very much alone.

After the American Civil War, many ventured west and began to collect plants. Charles L. Anderson (1827--1910) collected in western Nevada from 1862 to 1867. He was amused that the clover by the privy proved to be a new species that Gray named for him. In 1871 Anderson wrote a catalog of Nevada flora for the state mineralogist's report. John Gill Lemmon (1832--1908) moved to California in 1866 and began his long botanical career there; most of his plants were named by Watson. The state-sponsored geological survey program in California had two fine collectors, William Brewer (1828--1910) and Henry N. Bolander (1831--1897). With Watson and Gray, Brewer published a two-volume flora of California (Cambridge, Mass., 1876--1880), the first modern flora for a western state.

The last military expeditions, as well as the expeditions made in the early years of the U.S. Geological Survey (established in 1879), provided Gray and Watson with a constant flow of new species from the Rocky Mountains. Joseph Rothrock, who later had a distinguished career in forestry, served as botanist/surgeon for the Whipple expedition from 1873 to 1875. The several expeditions of Ferdinand Hayden (1829--1887) and John Wesley Powell (1834--1902) employed many botanists. Engelmann generally published the new plants found by Hayden, and Watson published those found by Powell.

To the north, the Geological Survey of Canada began its boundary survey in 1873. George Dawson (1849--1901) served as geologist/botanist, although most of the plants were collected by Thomas J.W. Burgess (1849--1925). At this point, Canada's foremost collector, John Macoun (1831--1920), began his botanical career in earnest. From 1869 until his death, Macoun collected thousands of specimens throughout much of Canada. He summarized these records in his Catalogue of Canadian Plants, published between 1883 and 1902 (7 pts., Montreal and Ottawa).

For his part, Gray concentrated on writing his Synoptical Flora of North America (2 vols., New York etc., 1878--1897), the project of his retirement. The years between 1842, when the last part of Flora of North America appeared, and 1873, when Gray retired, had been expansive in terms of growth in the number of new species and knowledge of the continent's flora. Gray turned to others for help, asking several younger men to assist him while retaining the cooperation of his old friend Engelmann and his newly acquired assistant Watson. Added to Gray's group were John Merle Coulter (1851--1928), William Farlow (1844--1919), George Goodale (1839--1923), and Charles Sprague Sargent (1841--1927), all famous later. Likewise, many of Gray's students began to fill the botanist posts at the newly established land-grant colleges, thereby providing him with capable collectors in critical regions. The goals were set, the players in position, and the work commenced, but Gray had not anticipated the rise of western botanists whose work would only make his more difficult.

Systematic botany had been centered in Cambridge for 30years, and its position had never been challenged. Few botanists attempted to work on the plants of North America without Gray's prior approval, and consequently he could influence what was done. To some extent, Gray dominated what was published by controlling the publications. By having one of the largest libraries and collections of American plants immediately at hand, Gray quickly settled identifications.

To many working in the American West, however, there was no joy in collecting a novelty if its publication was delayed because Gray or Watson already had specimens but had not yet published the name. Equally troublesome was the assumption that those in the East could somehow prevent those in the West from publishing their findings. At the forefront of the western dissidents was Edward L. Greene (1843--1915), minister, college professor, and protagonist. Greene had a keen eye for botanical novelties and a good knowledge of the classical literature. He was a "splitter," the antithesis of Gray and his followers, and he was not opposed to taking up the earliest available, validly published name, particularly names that Gray had chosen purposely to ignore.

Another botanical irritant was Marcus Eugene Jones (1852--1934), mining engineer, botanist, and Latinist. Jones collected widely in the Intermountain West, finding many locally endemic plants among others.

Individually and in the problems they jointly caused, Greene and Jones initiated the end of Gray's domination. Their defiance took two forms: first, they worked with the Smithsonian and the California Academy of Sciences, both institutions trying to proclaim their own independence; and second, they established their own journals. They therefore had means to publish names, and they had institutions from which to work.

Gray's efforts were focused. He wrote his Synoptical Flora and let Watson and others deal with the troublemakers. As for the expanding influence of other institutions, this Gray encouraged. North America needed more botanists and botanical institutions, but he felt these should be dispersed among universities, with only the difficult problems coming to Cambridge for final review. The feelings of others to the contrary, Gray did not discourage others from publishing their results; he simply felt that editors should not needlessly let new synonymy be created. The influence Gray possessed was substantial, and his authority was final. Watson, Farlow, Sargent, and Coulter were not equal to Gray in this, and when Gray died in 1888, still working on his flora, no one among them was capable of dominating taxonomy.

"Bughole Botany"

When John Torrey died in 1873, his large library and rich collection were at Columbia University. In 1896, when Nathaniel Lord Britton (1859--1934) became the first director of the newly established New York Botanical Garden, he arranged to have transferred the university's herbarium of some 400,000 specimens, along with Torrey's library. Both arrived in 1899. New York was suddenly able to take a botanical lead.

In addition, with the backing of Addison Brown (1830--1913), Britton published An Illustrated Flora of the Northern United States... (ed. 1, 3 vols., New York, 1896--1898) for the very region for which Gray's Manual had long been the primary authority. The sixth edition of the Manual (New York and Chicago, 1890), by Watson and Coulter, continued Gray's format of a single, tightly concise volume with keys to all taxa, brief descriptions, limited synonymy, habitat data, and distribution notes. Instead, Britton provided expanded keys, full descriptions, rather complete synonymy, and illustrations of each species. Furthermore, Britton produced his own one-volume Manual... in 1901 (New York). Unlike his competitors at Harvard, Britton accepted many of Greene's reestablished genera and used species names long since ignored by Gray. Britton's concept of species was similar to that of Gray, but wherever a split might be possible, Britton would make it, whereas Gray would not.

The publication battle was fierce. Britton issued new editions of his Manual in 1905 and 1907 (both New York); Benjamin L. Robinson (1864--1935) and Merritt Lyndon Fernald (1873--1950) answered with the seventh edition of Gray's Manual in 1908 (New York, Cincinnati, and Chicago). This was countered in 1913 by a second edition of Britton and Brown's Illustrated Flora (New York). Likewise, both sides began to marshal supporters at other institutions and began to stake claims to portions of the United States. In Washington, D.C., the Smithsonian Institution was doing likewise through its flora program, considering itself the sole authority on the flora of much of the West.

Under Gray's influence, Coulter had published his Manual of the Botany... of the Rocky Mountain Region... in 1885 (New York and Chicago), but by the time Aven Nelson (1859--1952) completed a revision in 1909 (New Manual of Botany of the Central Rocky Mountains..., New York, Cincinnati, and Chicago), the result was more a work of New York than of Cambridge. Smithsonian efforts on the West appeared as whole issues of the Contributions from the United States National Herbarium. Coulter published his flora of western Texas in 1891. Frederick Coville (1867--1937) published his on Death Valley in 1893. State floras followed for Washington (1906), written by Charles Piper Smith (1877--1955), and for New Mexico (1915), by Elmer O. Wooton (1865--1945) and Paul C. Standley (1884--1963). In 1925, Ivar Tidestrom (1864--1956) published his flora of Utah and Nevada. Most of these works consisted of little more than keys to species and genera, with annotations reporting distributions.

The group at New York was the most productive. John Kunkel Small (1869--1938) completed the first edition of his 1400-page Flora of the Southeastern United States... (New York, 1903), with full keys, good descriptions, habitat and distribution data, and an occasional illustration. Per Axel Rydberg (1860--1931) produced catalogs of the plants of Montana (New York, 1900) and Colorado (Fort Collins, 1906) in the abbreviated style of the Smithsonian publications. They were much in the manner of his first two floristic efforts, on the Sand Hills of Nebraska (1895) and on the Black Hills (1896), published in the Contributions. His Flora of the Rocky Mountains and Adjacent Plains... (New York, 1917) and his posthumous Flora of the Prairies and Plains of Central North America... (New York, 1932), however, were more akin to Small's works.

Small and Rydberg were both splitters, which appalled many, especially those of the Harvard tradition. Not only did Small and Rydberg recognize numerous species, they fragmented genera and families to a degree never before seen in American botany. Marcus E. Jones called Rydberg's effort "bughole botany" because, to him, Rydberg seemed to recognize species based on the number of bugholes in each leaf.

LeRoy Abrams (1874--1956) was in the New York camp after spending a year at the garden while finishing a flora for the Los Angeles area (ed. 1., Stanford, 1904). Under Britton's influence, Abrams began An Illustrated Flora of the Pacific States (Stanford). The first volume appeared in 1923, but the depression and war years delayed the second until 1944; the remaining two volumes were published in 1951 and 1960.

California still proved to be resistant to eastern domination. Certainly its isolation contributed, but so too did its long history of local botanical efforts dating to the 1850s. Following publication of the state flora by Brewer and Watson, a series of local floras began to appear. Volney Rattan (1840--1915) published A Popular California Flora in 1879 (ed. 1, San Francisco), and Greene, then at Berkeley, followed with Flora Franciscana (4 pts., San Francisco, 1891--1897). Willis Linn Jepson (1867--1946), the state's most acclaimed taxonomist, began his efforts in 1901 with A Flora of Western Middle California... (ed. 1, Berkeley), followed by his statewide Manual... (Berkeley, [1923--1925]). Jepson began the monographic style of flora writing in 1909 when he published the first volume of A Flora of California... (3 vols., San Francisco etc., 1909--1943).

Monographs and revisions began to appear more frequently, many the works of newly trained students. Several authors presented treatments on specific groups of plants, such as Sargent's The Silva of North America..., published between 1890 and 1902 (14 vols., Boston and New York), and various treatments of grasses beginning with George Vasey (1822--1893) and ending with Albert S. Hitchcock (1865--1935), with his Manual of the Grasses of the United States (ed. 1, Washington, 1935).

More state floras gradually became available as well. Tidestrom and Sister Mary Teresita Kittell (1892--?) treated Arizona and New Mexico in 1941 (Washington), followed by Thomas H. Kearney (1874--1956) and Robert H. Peebles (1900--1956), who wrote on the plants of Arizona (Washington, 1942; Berkeley and Los Angeles, 1951). Morton E. Peck (1871--1959) authored the first edition of his Oregon flora in 1941 (Portland). One for Idaho by Ray E. Davis (1895--1984) followed in 1952 (Dubuque). Events in the East were similar. In 1901 Charles Mohr (1824--1901) completed a flora of Alabama (Contr. U.S. Natl. Herb. 6), and Augustin Gattinger (1825--1903) published one for Tennessee (Nashville, 1901). In 1910 Forrest Shreve (1878--1950) wrote an account of the plants in Maryland (Baltimore).

On the Road

The period from 1870 to 1945 had many collectors, amateur and professional. They roamed widely in search of novelties, some publishing their own findings, some handing over their specimens for others to name. Local societies and clubs were formed, proceedings published, and activities sponsored. Institutional and university faculty were encouraged to be in the field, and students particularly so.

Botanical exploration of Alaska accelerated. Albert Kellogg of the California Academy of Sciences collected about 500 specimens in Alaska in 1867 while assigned to a survey party of the new U.S. Coast and Geodetic Survey. Other government officials began to collect plants as well. Mark W. Harrington (1848--1926) and Lucien M. Turner (1847--1909) sent specimens to Cambridge and to the U.S. National Herbarium in the 1870s. During the same decade, Edward W. Nelson (1855--1934), then a weather observer at St. Michael but eventually chief of the Bureau of Biological Survey, made a small collection. The foremost collectors of the period, however, were Frans Kjellman (1846--1907) and Ernst Almquist (1852--1946), both of Sweden, who made extensive collections along the coast of the Bering Sea.

James M. Macoun (1862--1920) and John Macoun (1831--1920), with the Geological Survey of Canada, explored much of western Canada, depositing their specimens at Ottawa. William C. McCalla (1872--l962), photographer, collector, and author of a wildflower text for high schools, also collected with J.M. Macoun.

Amateur botanists were also busy throughout Canada. David Watt (1830--1917), a businessman, amassed a large herbarium mainly through exchange. His dream, unfulfilled, was to write a flora of Canada. Braddish Billings Jr. (1819--1871), who worked for the St. Lawrence and Ottawa Railroad in the 1860s, published a local list of Ontario plants. In British Columbia, Joseph Kaye Henry (1866--1930), a professor of English, published a descriptive flora of the province in 1915 (Toronto). Robinson and Fernald from Harvard took an interest in the flora of eastern Canada in the 1890s, and they botanized there well into the next century. Fernald himself remained active in Canada into the 1930s, discovering numerous new species, most of which he described in Rhodora, the journal of the New England Botanical Club.

The foremost collector in eastern Canada was Frère Marie-Victorin (Joseph-Louis Conrad Kirouac) (1885--1944). In 1920 he established the Institute Botanique at the Université de Montréal, and despite suffering a heart attack in 1923 while in the Shickshock Mountains on the Gaspé Peninsula, he was in the field for the next five years in some of the harshest environments. In 1931 he established the Jardin Botanique de Montréal. In 1935 he published Flore Laurentienne... (Montreal).

The northern part of the continent was also becoming accessible. Another Cambridge-based collector was Hugh Raup (b. 1901) of the Arnold Arboretum. With support from the National Museum of Canada, Raup collected well to the north of most, working mostly in the Northwest Territories and along the Alaska Highway. Malte Oskar Malte (1880--1933), originally an agrostologist with the Canadian Department of Agriculture, moved to the National Museum in 1920. He concentrated on the prairie flora as well as on the flora of boreal and arctic Canada. Morton P. Porsild (1872--1956) was then actively collecting in Greenland. Alf Erling Porsild (1901--1977), his son, joined the National Herbarium of Canada and was there from 1936 to 1966. He reported on collectors for Flora of the Canadian Arctic (1955, 1957). Nicholas Polunin (1909--1991) covered the eastern Canadian Arctic (Bull. Natl. Mus. Canada 92, 97, 104, 1940--1948).

The Abbé Ernest Lepage (1905--1981) began his botanical career with an interest in bryophytes and lichens. In 1943 he began to collect with Père Arthème-Antoine Dutilly (1896--1973), and together they botanized the arctic and boreal regions from Alaska to Labrador.

In Alaska, Frederick Funston (1865--1917), who had previously collected plants in Death Valley, made a large collection of plants from 1892 to 1894 that Coville described in 1895. At the same time, but continuing to 1902, Martin Gorman (1853--1926), who also collected in Mexico, made several trips to Alaska. Most of his specimens went to the U.S. National Herbarium. Thomas Howell (1842--1912), who is better known for A Flora of Northwest America... (Portland, 1897--1903), was with Gorman in 1895. The Alaskan collections of Walter H. Evans (1863--1941), Arthur L. Bolton (1876--?), Frank C. Schrader (1860--1940), and botanical artist Frederick Walpole (1861--1904) also eventually reached Washington, D.C.

The 1899 Harriman expedition to Alaska had Coville, William Trelease (1857--1945), Brewer, and Thomas Kearney collecting plants (later described in vol. 5 of the 1904 report on the expedition). The most active of the Alaskan collectors, however, was Jacob P. Anderson (1874--1953), who collected extensively throughout the state from 1914 to 1940 while employed by the federal government. Also active was Eric Hultén (1894--1981) of the University of Lund, who amassed several thousand specimens in preparation for his many publications on the Alaskan flora.

Across the conterminous United States, numerous collectors were in search of novelties. Fernald was active in the Northeast until the late 1940s, making a series of expeditions with friends, all duly reported in Rhodora. Small continued to collect in the Southeast, especially in Florida after 1903, discovering several new species that were incorporated into a 1933 revision of his flora (Manual, New York).

Many came to the Rocky Mountains, some even collecting an occasional plant on the Great Plains portion of their trip. By the 1870s, Charles Parry and Edward Palmer (1831--1911) were well established in the West, collecting throughout the Rockies and the American Southwest for the Department of Agriculture and Smithsonian Institution, respectively.

Theodore D.A. Cockerell (1866--1948) collected plants on all of his travels. In 1889 he met Alice Eastwood (1859--1953), a Denver school teacher, collecting in the Rockies. Three years later she was an assistant curator at the California Academy of Sciences and returned to Colorado, searching the western Mancos clay hills for undiscovered wonders. Over the next 60 years, Miss Eastwood, in her long dresses and broadbrim hats, looked for novelties to collect. In the 1930s and 1940s her traveling companion was her colleague, John Thomas Howell (b. 1903), who has continued the study of California plants.

Across the Intermountain West came collectors like Carl Purpus (1851--1941), who also collected plants in Canada and Mexico, and the team of Dwight Ripley (1908--1973) and Rupert C. Barneby (b. 1911), who discovered many narrowly endemic species. H. Theodor Holm (1854--1932) collected in Greenland, Canada, and the Rocky Mountains, ending his days concentrating on grasses and sedges at the Catholic University of America in Washington. Isaac Martindale (1842--1893) amassed a huge herbarium through exchange and purchase, as well as through his own efforts. The collection is now at the U.S. National Arboretum.

Many students went on to distinguished careers. Aven Nelson encouraged many students during his long tenure at the University of Wyoming (1887--1952): Elias Nelson (1876--1949), Leslie Goodding (1880-1967), James Macbride (1892--1976), Edwin Payson (1893--1927), Louis Butler Payson (1895--1969), George Goodman (b. 1904), Louis O. Williams (1908--1990), and Reed C. Rollins (b. 1911). Jepson, at the University of California, fostered the careers of Herbert Mason (b. 1896), Ivan Johnston (1898--1960), David Keck (b. 1903), Conrad V. Morton (1905--1972), and Lincoln Constance (b. 1909). Many students took their degrees in St. Louis, working through a graduate program at the Missouri Botanical Garden. The dissertations of Carl C. Epling (1894--1968), Robert E. Woodson (1904--1963), Mildred E. Mathias (b. 1906), Julian A. Steyermark (1909--1988), C. Leo Hitchcock (1902--1986), and many others were published in the Annals of the Missouri Botanical Garden.

By the turn of the century, Cornell University was attracting many students, who on finishing went far and wide to educate the next generation. One was Bassett Maguire (1904--1991). He went to Utah State University in 1931 and remained until 1943, collecting from Montana to Arizona, taking a year out in 1938 to attend Cornell and complete his doctoral degree. Although he continued to collect in the West after he joined the staff of the New York Botanical Garden, his enthusiasm for the tropics was greater, and so his skill and drive were directed there.

The combination of world wars and economic depression had slowed systematic botany. Publications were less frequent, and collecting was limited. Also fewer in number were the small, privately published journals that were largely for the exclusive use of their editor, such as those promoted by Greene, Jones, and Amos A. Heller (1867--1944), and those restricted to friends, such as those published by Charles Orcutt (1864--1929) and the team of Mary Katharine (Curran) Brandegee (1844--1920) and her husband, Townshend Brandegee (1843--1925). Journals associated with institutions and societies, however, became much more numerous.

The growth of university-based systematics increased significantly, even during the difficulties of the Great Depression. When World War II ended, universities welcomed a generation of mature students, and for the next three decades systematic botany in the United States and Canada was dominated by individuals trained during or shortly after the war.

Where Have All the Flowers Gone?

With the end of the war, taxonomy as understood by Torrey and Gray was gone. Searching for novelties or trying to find unreported species was no longer the primary focus of the field. Taxonomy broadened into systematics and was rapidly becoming a laboratory science. Walking across a meadow enjoying the flowers was often replaced by white coats, laboratory walls, and the thrill of a good chromosome squash or a well-resolved chromatograph.

This was to be expected. Since the 1930s, more and more taxonomists had become interested in experimental taxonomy. The long-debated questions of relationships could be addressed and reasonable conclusions could be drawn using the new technology, whereas earlier any solution was open to doubt. Transplantation studies, championed in the New World by Clausen, Keck, and Heisey, proved informative, and new insights were possible concerning the relationship of a plant to its habitat.

Over the past half century, the working lifetime of many still very active systematists, new methods have evolved, and problems once thought impossible to address are being routinely resolved. Even such mundane things as the rules of nomenclature have changed. Significant changes may still come, as today's cladists and molecular systematists challenge the fundamental way all systematists think, much as Darwin did in 1859.

Systematic centers are now more dispersed, although Cambridge, New York City, Ithaca, Washington, St. Louis, and Berkeley are still major players, and now many other institutions, from Austin to Ann Arbor, and Chapel Hill/Durham to Seattle have joined them, each offering its own particular brand of systematics.

Fieldwork is still needed, but now few remote areas remain in North America north of Mexico where one might find large numbers of undescribed vascular plants. New genera, such as Apacheria, Shoshonea, or Dedeckera, are increasingly less likely to be found, although new genera carved from older groups will continue to be proposed.

Many new floras and revisions of older ones have been completed since 1945. The style of manuals has improved over that of Britton's in 1901. Jepson's use of detailed descriptions is now the norm, as are full distribution and ecological statements, data on flowering times, and notes on uses. Distribution maps are more common, and illustrations are becoming standard. The style of regional floras has changed too, most being illustrated and monographic.

And funding has changed. Research support for floristics was minimal before World War II, but in the early 1950s federal funding in both the United States and Canada became available to study the plants of large portions of North America. Floristic projects outside of North America now share the available funds with North American projects, which still play a vital role.

Even the reasons for doing taxonomy have changed. While the identification, naming, and classification of plants are still fundamental needs for all botanical science, they have, in recent years, become important to governments also. Concern for endangered plant species, regulation of trade and commerce, and the ecological well-being of the environment have awakened a broad array of citizens who truly are concerned about where all the flowers have gone.

The dream of Barton, Torrey, and Gray to have a flora of North America has never left the botanical community. Over the past 50 years various attempts have been made to accomplish the task. At least now we are at a stage familiar to Torrey and Gray---one volume is completed. Certainly when Torrey talked to Hooker about the task in 1833, the time frame was realistic relative to what was known. He and Gray did not realize they might be overwhelmed with new species. We know that we may not find many new species; rather, we now recognize the need to understand more fully what species we have.

No modern flora can be written without acknowledging the legacy of those who went before us. We must try to make sense of the diversity that exists; that is our duty as systematists. Those devoted to the task must be willing to set aside the multitude of competing requirements and spend their lives at it; no doubt, Asa Gray would understand.

Review

A detailed history of botanical explorations and the development of floristics in North America north of Mexico has yet to be written. The literature is very scattered, and as with most history, much of it is hard to find. Nonetheless, some portions of the subject have been reviewed (E.B. Davis 1987), and Reveal (1990, 1992) has broadly covered many of the events.

The colonial era is summarized in R.P. Stearns (1970), and this is the basic source for the period. Major bibliographies exist on many of the Old World naturalists who described American plants, and these may be found easily in F.A. Stafleu and R.S. Cowan (1976--1988) and F.A. Stafleu and E.A. Mennega (1992). As for those who actually collected in North America, works are available on John Banister (J. Ewan and N. Ewan 1970), Mark Catesby (G.F. Frick and R.P. Stearns 1961), Alexander Garden (E.Berkeley and D.S. Berkeley 1969), and John Clayton (E.Berkeley and D.S. Berkeley 1963). John and William Bartram have been the subject of numerous books, but the one by E. Berkeley and D.S. Berkeley (1982) is most readable. An essential reference on French botanists in North America before 1850 is J.-F. Leroy (1957). A good bibliographic and biographic source on Marie-Victorin is D.Barabé (1985). An augmented autobiography of Macoun is now available (J.Macoun 1922). J.S. Pringle (forthcoming) will provide access to Canadian floristic history.

Linnaeus's contribution is complex, and works are available on nearly every aspect of his life and times. F.A. Stafleu (1971) wrote one that is perhaps the most useful to systematists trying to understand eighteenth-century taxonomy, but that by W.Blunt and W. T. Stearn (1971) is most informative on Linnaeus.

B.Henrey's (1975) three-volume work on British botanical and horticultural literature before 1800 is indispensable and wonderful reading. More than in any other single source, one may find here the magic of discovery.

An excellent recent work by H.B. Carter (1988) on Joseph Banks covers not only the man but his times as well. This is a nice companion to D.J. Mabberley's study (1985) of Robert Brown.

A biography has finally appeared on André and François André Michaux (H. Savage Jr. and E.J. Savage 1986), but none on Pursh has yet been published. The standard references for Nuttall (J.E. Graustein 1967), Torrey (A.D. Rodgers III 1942), and Gray (A.H. Dupree 1959) remain. Likewise, the brief bibliographic sketches given by H.B. Humphrey (1961) for several dozen of North America's botanists provide a useful introduction. The older book on Philadelphia botanists by J.W. Harshberger (1899) contains odd bits about personalities and characteristics that more recent writers tend to ignore or, out of respect, not mention.

The classical work on western explorations in the United States remains S.D. McKelvey's opus (1955). This work is basic for an understanding of the early collectors and botanists. For an overview of western American history, the texts by R.A. Billington (1982) and L.R. Hafen et al. (1970) are the best. A more detailed study of westward expansion, and the role played by scientists, may be found in W.H. Goetzmann (1966).

Publications on the history of botanical explorations of our northern regions are extremely scattered. E.Hultén's 1940 article on Alaska is still the best, but for Canada, the forthcoming articles by J.S. Pringle will improve the situation immeasurably.

L.W. Lenz (1986) has published an excellent work on Marcus E. Jones, and R.L. Williams (1984) examined the life of Aven Nelson. Both provide a broad overview of taxonomy at the end of the last century and the first few decades of the present one. With the nice biography on Rydberg by A.Tiehm and F.A. Stafleu (1990), the chronicling of Greene remains to be tackled.

Important information on botanical history is available. First, most floras have, in their introductions, a section on the individuals who have collected in the area, and these are often very informative. Second, botanical journals are a rich and never-ending source of articles and tidbits on the past, and even revisions and monographs can be well worth perusing because historical items are frequently scattered throughout them.

Two recent systematics symposia at the Missouri Botanical Garden have concentrated on historical subjects. In 1984 George Engelmann served as the focal point; in 1989, the period from 1889 through 1989, only briefly mentioned in this volume, was examined in some detail. Papers delivered at those symposia were published in the Annals of the Missouri Botanical Garden in 1986 and in 1991.

Finally, there are the contributions of a few individuals who, as a result of their life's work, have made understanding the past that much easier. First and foremost is Joseph Ewan (K.Crotz 1989). Anyone wishing to know something about American botanical history should do a computer search on his name. The results are staggering. Likewise, search on the name William T. Stearn. His forte has been the Linnaean era. And then there is Ronald L. Stuckey. His name is less commonly seen, but his subject---botanists in the East during the Torrey and Gray years---is more difficult.

Most who pick up this and the subsequent volumes will find the history of systematics and floristics on every page, for each plant name has a story to tell. Those who look into that story will find wonderful rewards and an even greater appreciation of systematics.