ASA GRAY and the DEVELOPMENT of BRYOLOGY in the UNITED STATES Author(S): Norton G

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ASA GRAY AND THE DEVELOPMENT OF BRYOLOGY IN THE UNITED STATES Author(s): Norton G. Miller Source: Harvard Papers in Botany , December 2010, Vol. 15, No. 2 (December 2010), pp. 287-304 Published by: Harvard University Herbaria Stable URL: https://www.jstor.org/stable/41761699

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This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms ASA GRAY AND THE DEVELOPMENT OF BRYOLOGY IN THE UNITED STATES

Norton G. Miller1

Abstract. Asa Gray never expanded his knowledge of floristic bryology to the extent he developed expertise in flowering plant taxonomy. Nevertheless, he became experienced in bryological floristics early in his botanical career, and Gray absorbed new bryological information, both floristic and conceptual, throughout his life from wherever it was generated. He had plans to advance bryology in the United States, including an exsiccata and publishing a volume devoted to cryptogams as part two of the second edition of his Manual , but both never happened. His respect for the bryological talent and energy of William S. Sullivant, whose achievements Gray consistently encouraged and fostered, allowed Sullivant, a non-academic in Columbus, Ohio, to become a highly regarded bryologist of international stature and the designated Father of American Bryology. The growth of bryofloristic knowledge in the United States is traced from the earliest colonial period to later workers, including Dil- lenius, André Michaux, Palisot de Beauvois, Henry Muhlenberg, Lewis Schweinitz, Lewis Beck, and John Torrey, to Asa Gray, and eventually to William Sullivant. The bryological work and accomplishments of each of them show that all participated in a sophisticated international network of information exchange by letter or other con- veyance, thereby building important collections of bryophyte specimens and printed references. For some, this happened during the 1800s when improvements in compound light microscopy led to the resolution of morphology not before revealed with certainty in bryophytes and to conceptual advances in understanding the biology of these plants, which in turn allowed the discovery of the mesoscale structural uniqueness of them and continuing advancements in their systematics in the post-Sullivant era.

Keywords: Asa Gray, William Sullivant, John Torrey, Lewis Schweinitz, Lewis Beck, Henry Muhlenberg, Palisot de Beauvois, André Michaux, Dillenius, history of bryology, history of botany

" My desire to know every N. American plant with certainty is insatiable , as for foreign plants , let others explore them , there is room for all to study and admire the great variety. " - Henry Muhlenberg (Lancaster, Pennsylvania) to Dawson Turner (Yarmouth, England), 21 February 1803.

Asa Gray's botanical accomplishments are reading letters and journal entries he wrote profound, durable, and generally understood during his first trip to Europe, 1838-1839 (J. to pertain to the flowering plants. Yet, through- Gray, 1893). Gray's career bridged the earliest out his career, and especially during the time attempts to document the North American flora he trained to become a physician and early inby colonial plantsmen, botanical explorers, and his botanical work, Gray maintained a lively a small group of native-born botanists, some probing interest in all branches of floristic, of whom took as their goal securing names for structural, and physiological botany as these American bryophytes, and the later botanists subjects were then defined, including bryo- that carried bryofloristic knowledge beyond the phytes and other cryptogams. This is clear from pioneer stage.

I am grateful for information or assistance from the following: Robert Magill and Patricia Eckel (Missouri Botanical Garden), Ronald Stuckey (Ohio State University), Charles Sheviak (New York State Museum), Caryn Radick (Rutgers University Archives), Lynn Eaton (Rare Book, Manuscript, and Special Collections Library, Duke University), Stephen Sinon (LuEsther T. Mertz Library, New York Botanical Garden), and Daniel Weinstock, M.D., Geneva, New York. The last generously gave me permission to quote from unpublished letters of 19th century botanists in his collection. Sean P. Kilcoyne, Reference Archivist, Ohio Historical Society, helpfully searched the Sullivant family papers there for pertinent letters. Unless already published, all other quotations from letters are with permission of the institutional archive holding the original. Because this essay is in part about books and printed matter, it gives me much pleasure to acknowledge the great libraries I use regularly: New York State Library and its Manuscripts and Special Collection Division (Margaret Harris, Cindy Stark, Cara Janowsky, James Summa, and associates), University at Albany Library (Heather Swan Miller), and the Harvard University Botany Libraries (Judy Warnement and staff, especially Lisa DeCesare). Leigh Ann Smith and Patricia Kernan, New York State Museum, assembled the plate of images. 1 Biological Survey, New York State Museum, Albany, New York 12230-0001 , U.S.A. Email: [email protected]

Harvard Papers in Botany , Vol. 15, No. 2, 2010, pp. 287-304. © President and Fellows of Harvard College, 2010.

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In this essay I explore connections Lewis Beck, and associates, between as well as bryolo- Asa Gray's botanical activities gists and bryophyteand the collectors devel- that Gray later opment of bryology in mentoredthe Unitedand influenced. States. I have sought to find bryological Asa Gray's links published between contributions are num- and among Gray and his predecessorserous, nearly 800 of them and in fact,con- and much temporaries and to ascertain has been how written bryological about his life, associates, knowledge accumulated inand accomplishments.the United A listStates of Gray's writ- from the late 1700s to the middle 1800s. This ings is available (Watson and Goodale, 1888), was a time of transition, during which bryolog- as is an index (Seymour, 1888). Selections of ical expertise developed in the United States Gray'sin letters, chronologically arranged, and an native-born botanists. In particular I sought autobiographyto of his first 30 years, were pub- establish what bryological books and specimens lished by Jane Loring Gray (1893), his wife. were used by the early workers, which in ret- Many of Gray's scientific and other writings rospect serve as an assessment of how well were a reprinted by Sargent (1889). In addi- worker was connected to mainstream bryology tion, the historian A. Hunter Dupree (1959) has of the time, and also the names of authorities written a richly informative biography of Asa to whom specimens were sent for identifica- Gray, with attention to the national and interna- tion or other response. Such information and tional intellectual mileau in which he worked. examination of authentic specimens from an These sources, plus letters and other records expert's herbarium help understand the often in the Gray Herbarium archives at Harvard varying early application of the technical names University and the collection there of of bryophytes. Grayana, provide much information pertain- Steere (1977) has assembled much fasci- ing to the topic outlined above. Additional nating information pertaining to bryologists primary sources were consulted at or from the and bryological progress for a later period, Rutgers University Archives; Duke University mostly the last half of the 19th and first half Archives;of LuEsther T. Mertz Library (New the 20th centuries. By contrast, my focus is Yorkon Botanical Garden); library of the the initial development of bryology in the Academy of Natural Sciences (Philadelphia); United States and Gray's role in helping Ohio Historical Society (Columbus); and the move bryological science in America from itsDivision of Manuscripts and Special Collec- earliest period into a more modern era. I present tions of the New York State Library (Albany). information that explores bryological work Scans of pertinent unpublished letters in the of the botanical collaborators, Asa Gray and collection of Daniel Weinstock, M. D. (Geneva, John Torrey, and their predecessors, principally New York) were also read. Henry Muhlenberg, Lewis von Schweinitz,

Bryology in North America up to the Start of Asa Gray's botanical career Bryophytes had an almost inconsequential reached. My primary sources for names in this role in the work of plant collectors who ener- list were Dillenius (1741); Cardot (1899), who getically sought New World plant novelties for evaluated North American specimens and their European gardens and collections during the origins in the nomenclaturally important Hed- early colonial period in North America. At that wig-Schwägrichen Herbarium, which includes time, nevertheless, examples of North Ameri- specimens vouchering moss names in Hedwig can mosses and liverworts shipped to European (1801), the legislated starting point of moss taxonomie experts were numerous. The fol- nomenclature (excluding Sphagnum ), and its lowing list highlights botanical collectors that supplements (Sch wägrichen , 1811-1 842) ; gathered bryophyte specimens from various Hooker (1818-20); and the first major consol- parts of North America in this early period and idations of the published names and descrip- through about 1850 as members of exploring tions of mosses by Bridei (1826-1827) and expeditions, commercial ventures, royal and of liverworts by Gottsche et al. (1844-1847). other patronage, and a personal interest in the Before these last two publications such infor- identity of these plants, often stimulated by mation was available only in scattered and often the very European or American authority they obscure sources. My tallies of the frequency a

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 2010 A TRIBUTE TO ASA GRAY 289 collector's name descriptions. appeared in For them the mostrevealed part their European a relatively small contacts impact took that role. of most of the earliest bryophyte collectors James Peti ver (Stearns, 1952), in in London, Colonial British America (John Banister, encouraged and sponsored John the collection Bartram, of Batch- elot de la Pylaie, natural Bosc history specimensd'Antic, in early colonial John Clayton, André Michaux, AmericaJohn (and from Mitchell); elsewhere in the world), the somewhat consistently greater and the huge influenceplant collection he assembled of and the next generation of workers publicized (Thomas eventually became part ofDrummond, the Hans Henry Muhlenberg, Palisot Sloane Herbarium de (Dandy, Beauvois), 1958). Apart from and greater impact still for collectorsa few attempts (Bellis, 2009) thatdirected mostly followed (Dennis Cooley, Chester atDewey, vascular plants, this herbarium,Lewis now invon the Schweinitz, John Torrey). It Britishbears Museum (Naturalemphasizing History), has not been that many of these botanists and examined bryophytesystematically for early 1700s Northcollectors did not themselves publish American bryophytenew records. names and bryophyte

Bryophyte collectors and workers in North America , north of Mexico , largely prior to the onset of Asa Gray's BOTANICAL CAREER AND THROUGH ABOUT 1850

Principal sources of publishing authors: (B) Bridei (1826-1827); (D) Dillenius (1741); (H) Hooker (1818-1820); (HS) Hedwig (1801) and Sch wägrichen (1811-1842); (GLN) Gottsche, Lindenberg, and Nees (1844-1847); Baldwin, William, 1779-1819: Georgia. (Darlington, 1843); Banks, Joseph, 1743-1820: Newfoundland, Labrador (B); Banister, John, 1605-1692: Virginia (B); Bartram, John, 1699-1777: Pennsylvania, eastern North America (D); Beck, Lewis Caleb, 1798-1853: New York State, Massachusetts, Vermont; Bigelow, Jacob, 1787-1879: New England (Mears, 1978); Boott, Francis M. B., 1792-1863: Massachusetts (H); Boykin, Samuel, 1786-1848: Georgia (Mears, 1978); Batchelot de la Pylaie, Auguste Jean Marie, 1786-1856: Newfoundland (B); Beyrich, Heinrich Karl, 1796-1834: eastern United States (Hampe, 1837, 1839); Bosc d'Antic, Louis Augustin Guillaume, 1759-1828: South Carolina (HS). (Harper, 1940; Rohde et al. ,2009); Brackenridge, William D., 1810-1893: United States Exploring Expedition under Capt. Charles Wilkes, northwest North America (Harpel, 2010); Chamisso, Ludolf Adelbert von, 1781-1838: Unalaska, Aleutian Islands (B); Clayton, John, 1693-1773: Virginia (D); Gronovius (1739-1743, 1762) (Patterson, 1965); Collins, Zaccheus, 1784-1831: Pennsylvania (Mears, 1978); Cooley, Dennis, 1787-1860: Massachusetts, Georgia, Michigan (B); Curtis, Moses Ashley, 1808-1872: North Carolina. (Curtis, 1867); Cutler, Manasseh, 1742-1823: New England; (Mosses sent to Muhlenberg; Cutler and Cutler, 1888; Mears, 1978); Dallman, Gustavus Heinrich, [dates unknown]: North Carolina. (Mears, 1978); Dewey, Chester, 1784-1867: Massachusetts, New York (B); Douglas, David, 1798-1834: west coast of North America; Drummond, Thomas, 1780-1835: western and southern North America (HS); Elliott, Stephen, 1771-1830: South Carolina. (Mears, 1978); Engelmann, George, 1809-84: Missouri and vicinity;

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Fraser, John, 1750-1811: southern and eastern North America (B); Gray, Asa, 1810-88: Massachusetts, New York, Appalachian Mountains (GLN); Greene, Benjamin D., 1793-1862: Massachusetts (B); Lawson, John, 1674-1711: North Carolina (Bellis, 2009); Lea, Thomas G., 1785-1862: Ohio, Indiana, Kentucky (GLN); Lesquereux, Leo Charles, 1806-1889: eastern North America; Lyon, Capt. George Francis, 1795-1832: arctic Canada (Lyon, 1825); Menzies, Archibald, 1754-1842: northwest North America on the Vancouver Expedition; Nova Scotia (H, HS); Michaux, André, 1746-1802: Canada, eastern United States (HS); Mitchell, John, 1676-1768: Virginia (B, D); Muhlenberg, Henry (Rev. Gotthilf Heinrich Ernst], 1753-1815: Pennsylvania (HS); Oakes, William, 1799-1848: New England (Reid, 1987; Sullivant, 1846a); Palisot de Beauvois, Ambroise Marie Françoise Joseph de, 1752-1820: New York to Ohio, south to Georgia (HS); Pickering, Charles, 1805-1878: United States Exploring Expedition under Capt. Charles Wilkes, northwest North America. (Harpel, 2010); Rafinesque-Schmaltz, Constantine, Samuel, 1783-1840: eastern North America (Merrill, 1943; Rafinesque-Schmaltz, 1808); Richardson, John, 1787-1865: northwestern boreal and arctic Canada on lst-3rd of Capt. John Franklin's Arctic Expeditions (HS) (Hooker and Taylor, 1827); Ross, John, 1777-1815: arctic British North America. (Ross, 1818; Simmons, 1913; Steere, 1947); Schweinitz, Lewis David von, 1780-1835: North Carolina, Pennsylvania (HS); Sabine, Capt. Edward, 1788-1883, and associates, including Capt. W. E. Parry: Melville Island, Arctic North America (HS) (Brown, 1823); Scouler, John, 1804-1871: British Columbia, 1824-1825 (H); Short, Charles Wilkins, 1794-1863: Kentucky and nearby areas (Sullivant, 1846a); Sullivant, William S., 1803-1873: Ohio, Appalachian Mountains (GLN); Torrey, John, 1796-1873: Massachusetts, New York, New Jersey (B, HS); Watson, Sereno, 1826-1892: western United States.

The frequency some of these bryofloras collectors of northwestern are Europe and northern cited in Bridei (1826-1827) North is revealing.America. For example, surprisingly more collections Comments on somefrom of more significant British Arctic America and Newfoundland bryological accomplishments of the early were available to European workers in the period early follow. 1 800s than from more southern parts of eastern The Historia Muscorum of Dillenius North America. This disparity is the result (1741), of containing polynomial moss and liver- successive sailing expeditions among the wortarctic names, synonymies, descriptions, and the islands in search of a Northwest Passage author's to numerous precise original illustrations, the Orient, overland expeditions in northwest- was a remarkable work for its time. It featured ern North America for the same purpose, mosses and sent to Dillen (1684-1747) from eastern Batchelot de la Pylaie's extensive collecting North on America by John Bartram, John Clayton, Newfoundland, which he visited in 1816 and and John Mitchell and cited the published bryo- 1819-1820 (Leroy, 1957; South, 1980). Hookerlogical contributions of others who contributed and Taylor (1827) published a brief account to the bryology of this region, including occa- of the rich moss collections John Richardson sionally generalized specimen origins (e.g., made as a member of Capt. John Franklin's Pennsylvania, Maryland, Virginia). Dillenius overland expeditions, including a list of species (1741 : 434) sometimes surprises the reader with that very early established similarities between details, for example, perhaps the earliest most

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complete locality botany. citations Muhlenberg began tofor assemble a theher- moss now known as Pogonatum barium of North pensylvanicum American plants, including (Hedw.) P. Beauv.: "gathered bryophytes, [by in the John 1770s (Harshberger, Bartram] 1899; on the Jersey Side at the Smith,Memesinks 1962), and he actively exchanged[Minisink] bryo- , at the Upper Inhabitants phyte specimenson thewith European Delaware specialists over [River]." Minisink is the name many years. given When Muhlenberg to beganthe his bry- region where Pennsylvania, New ological York, studies, the mossand and liverwort New flora Jersey join above the Delaware of eastern Water North America Gap. was nearly It unstud- was in New Jersey before the iedboundary in an integrated way. To betweenhelp in the iden- New York State and New Jersey tification wasof unknowns, set only in a handful 1800 of but now is in New York. Dillenius bryophyte descriptions (1741) were availableand in alater few editions were widely used in books, bryophytesuch as those of Gronovius (1739-1743,identification in the late 1700s before 1762) and theWalter (1788) Hedwig-Schwägrichen that focused mainly on era, substituting vascular binomial plants of small parts equivalents of a large conti- intro- duced by Linnaeus nent. Therefore,(1753). Muhlenberg turned for help André Michaux ( 1to the748- leading European 1 802) bryophyte , experts a great of botanical explorer, traveled the period (Dillenius, widely J. Hedwig, Palisotin de the eastern United States and Beauvois, O.into Swartz, J. E. Canada,Smith, K. Sprengel, amassing large collections, particularlyD. Turner, among others; Harshberger, from 1899; the southern Appalachian MerrillMountains. and Hu, 1949), and in timeHis he also Flora Bore- ali- Americana (Michaux, found and encouraged 1803), many American treatingbotan- both vascular plants and ical collaborators bryophytes, and correspondents, includ- became the standard identification ing William source Baldwin (Darlington for 1843), eastern from North America in the early whom he acquired1800s. specimens Itof taxonomie seems, and however, it was the Parisian botanist Louis Claude Marie geographic importance. Richard who brought taxonomie order to Mich- The way Muhlenberg went about accumulat- aux's bryophyte collections, as is acknowledged ing bryological intelligence is summarized by in Bridei (1826-1827) and Hooker (1825) (see the following quotations from a letter he wrote also Sayre, 1976). to Dawson Turner in England on 21 February During this period, another Frenchman, 1803. "Mosses I have gathered pretty generally. Palisot de Beauvoìs (1752-1820), centered I send the most of them to John Hedwig my in Philadelphia (Merrill, 1936) but traveling good and faithful friend. He promised me the widely in the area from New York to Ohio, names of each, but unfortunately died before he south to Georgia, between 1793 and 1798, col- could fulfill his promise. His Species Musco- lected and eventually described numerous new rum were sent for me but were lost, so I am yet bryophytes from his studies of eastern North unable to say how exact his Figures and Descrip- American examples of these plants (Palisot de tions are... I have since sent a great number of Beauvois, 1805; Crum, 1952). His work was an specimens to his son Romanus Hedwig and improvement in floristic documentation over his Editor D. Schwagrichen, because I heartily Michaux (1803). Most of the Palisot de Beau- wish our mosses to be described... As I wish vois' North American herbarium is known to not any of my letters to go quite empty, I will have been lost at sea, and therefore authentic try to inclose 34 specimens of Cryptogams of specimens of his are generally considered no which I suppose none is common to Europe and longer available for study. In fact some survived America, if I except the Neckera [Climacium] in at least one collection (Arnott, 1826), with dendroides with fructifications. The specimens fragments now in the herbarium at Glasgow must therefore be small. I will number each and University and perhaps elsewhere. Palisot de continue number whenever I send more. So Beauvois (1822) later elaborated his proposed in the future any animadversion, observation, system of mosses in an illustrated monograph. or desideratum may be facilitated. I will add Henry Muhlenberg (1753-1815), of Lan- candidly my name without insisting upon it that caster, Pennsylvania, a native-born American, I am positive" (from an unpublished letter in attended school and briefly university in Halle, the D. Weinstock, M. D., Collection, Geneva, Germany, but was essentially self-trained in New York).

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Muhlenberg included results Schweinitz of his returned exten- to the United States in sive bryological studies in both 1812, editions but church (1813, business took him back to 1 8 1 8) of his catalogs of North Germany American two more plants, times. His German botan- two works that had at the time considerable ical connections and those cultivated elsewhere influence on taxonomie botany in the United at European centers ensured an abundant and States. Prefatory information in these works constant flow of botanical information- books, included lists of books he consulted in assem- specimens, conceptual advancements- from bling the catalogs and the names of colleagues Europe to North America to him and his col- assisting him. He was linked to the best sources laborators. Much is known about the contents of European bryological information. Moss of Schweinitz' botanical library and his many genera listed in the catalogs followed closely correspondents, particularly those who con- Hedwigian names and concepts, but with a few tributed specimens to his herbarium (Barnhart, additions, namely Hedwigia P. Beauv., Pterigo- 1934; Benedict, 1934), which contained speci- nium Sm., Diphyscium Mohr, and Climacium mens of about 23,000 species of higher plants F. Web. & Mohr. Moss and liverwort names (Pennell, 1934) and many of cryptogams. Sch- in the Muhlenberg catalogs are mostly without weinitz owned a comprehensive bryological authors and descriptions, and the new names library, and, as was true of Muhlenberg, his introduced there are nomina nuda and therefore correspondents included most of his European without nomenclatural standing. Descriptions bryological contemporaries. The Schweinitz of some of them, however, have been supplied Herbarium is at the Academy of Natural Sci- by his European correspondents, for example, ences of Philadelphia. Hedwig (1797, 1801) and Turner (1806). John Torrey, whom I discuss in more detail Muhlenberg's widely distributed bryophyte col- next, first wrote to Schweinitz on 29 Dec 1819, lections continue to be important in typification and their correspondence continued until just (e.g., Orthotrichum strangulatum P. Beauv.; before Schweinitz' death in 1834 (Shear and Atwood & Allen, 2009). The Muhlenberg bryo- Stevens, 1921). Their letters are full of interest- phyte herbarium is in the Academy of Natural ing bryological information. In reading them, Sciences of Philadelphia, where it is preserved one develops a sense that the path to floristic with supporting archival information that aids illumination, which both eagerly sought, was in its interpretation. Merrill and Hu (1949) con- never straight and always onward. For example, sidered the Muhlenberg Herbarium "apparently Torrey wrote Schweinitz on 22 March 1821 the first really large one assembled by a native- thanking him for remarks on the cryptogamia born American botanist." Torrey had sent to him earlier and noting, "You Lewis David von Schweinitz (1780-1834; do not always agree with Sprengel [Kurt Poly- Figure 1A), born at Bethlehem, Pennsylva- carp Joachim Sprengel, 1766-1833, Profes- nia, and in later years a pastor and administra- sor of Botany in Halle, Germany] to whom I tor in the Moravian Brethren, initially placed by have at different times sent many of the spec- his church in Salem, North Carolina, and later imens I send you... . Indeed this great diver- in Bethlehem, Pennsylvania, pursued a broad sity in opinion among equally great botanists range of botanical interests (Harshburger, 1899; almost discourages me from pursuing the study Pennell, 1934). He also assiduously focused of Cryptogamia. I often find as many differ- on bryological study and published (Schwein- ent names given to the same plant as persons itz, 1821) the first work solely about liverworts to whom I send it" (Shear and Stevens, 1921: of eastern North America, in which 78 species 137). To which Schweinitz replied on 19 April were described, some newly revealed to science 1821, "There is to be sure too much truth in through his studies. your observation concerning the great differ- Schweinitz, starting at age 18, was a student ence of opinion concerning certain Cryptogam. at a Moravian theological institute at Niesky, Plants- I believe it arises chiefly from this cir- Silesia, Germany, and later, after gradua- cumstance, that many Botanists, & especially tion, he preached at Moravian settlements in such as Sprengel (who by the by is rather noted Germany. During studies at Niesky he worked for his superficiality in this respect) do not take with a highly regarded botanist, Prof. J. B. the trouble really to investigate closely the spec- von Albertini, and developed a foundation in imens sent, but hazard an opinion at first blush mycology and other botanical specialties. without accurate comparison & examination;

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Figure 1. Late 17th and 18th century American botanists having a strong interest in bryology. 1A, Lewis David von Schweinitz (1780-1834); IB, John Torrey (1796-1873); 1C, Asa Gray (1810-1888); lD,WilliamS. Sullivant (1803-1873). The images were selected to represent them in middle age. Courtesy of the Archives of the Gray Herbarium, Harvard University.

which is but too natural mistaken- considering the smallness the minute of the specimens may exam, that is often be necessary an excuse. to beBut certain I candidly of the believe that many identity of any moss of Sprengel's&c. I am far determinations from think- ought not to be ing my determinations depended altogether upon on free account from of his inclination to this reproach- However make short as towork. the Dr.Musei Sch wägrichenyou is certainly sent me- I took greatthe more correct &pains acute of the two & & I intendwhenever I am

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 294 HARVARD PAPERS IN BOTANY Vol . 1 5 , No. 2 to send him all those of which Schweinitz I andentertain Torrey, or possibly any a joint effort doubts" (Shear and Stevens, for each1921: group. 142). In the preface These to the Flora of comments reflect the letter thewriter's Northern andfrustration, Middle Sections of the United but at the same time they Statesestablish , Torrey (1823-1824:that Sch- vi, vii) wrote "Of weinitz and Torrey were in contact the Cryptogamia, with it isthe concluded best at this time of the European specialists of to describethe time, only the ferns,and leavingthat the other useful information was transmitted. orders for a future period; when the author, in Schweinitz and Torrey discussed plans to connexion with his friends, the Rev. L. D. Sch- prepare a cryptogamic flora of the United States weinitz, and A. Halsey, Esq., proposes to give as a companion to Torrey 's vascular plant flora a cryptogam flora of the United States. Large of northern and middle United States (1823- collections have already been made towards 24). Torrey wrote on 22 March 1821, "It gives this work." me great pleasure to hear that we are at last Near the end of Schweinitz's life a letter in hope of having a Cryptogamic flora of the arrived from Asa Gray who was starting a career United States. You are probably in posses- in botany (Shear, 1905). This letter, written on sion of a greater quantity of materials for this 8 March 1833 from Utica, New York, where purpose than any person in this country. I hope Gray was teaching in Bartlett's School, did you are well acquainted with all the species not mention bryophytes but instead focused on which Muhlenberg mentions in his catalog. Did sedges, willows, and other vascular plants, and you correspond with him? He has some species Gray's plans for field studies later in the year. whose names I can not find in my books. You Thus, it appears that Schweinitz and Gray did may depend on receiving everything from me not exchange information about bryophytes, which will be of any assistance in your work, except for what may have been transmitted to & I believe you will then have all that has been Gray from Torrey 's extended correspondence observed in the States north of Pennsylvania. with Schweinitz. All of the botanists here send me everything John Torrey (1796-1873; Figure IB) repre- they collect; so that I can thus do for you a great sents the next generation of botanists pursuing deal more than I could do alone" (Shear and bryological study in the United States. As we Stevens, 1921: 137). Schweinitz's response was, have seen, there was a substantial overlap in "... I am glad you seem to favor and encourage information exchange among him and Schwein- my undertaking of a Cryptogamic flora. I shall itz and Muhlenberg. Torrey dominated floristic however not proceed to the execution before I botany in the United States for a long period have made larger collections & more accurate of time (Rogers, 1942). It was he who helped observations. Unhappily the death of Muhlen- launch Asa Gray by hiring him to fill what we berg deprived me of the advantage of his com- would today categorize as a Research Assistant- munications. He had begun & about half finished ship in Torrey 's herbarium, which was located a letter to me commenting upon my Musei & in New York City in Torrey 's home, an event Lichens sent to him- & all my endeavors to that marked the beginning of a fruitful period regain possession of the specimens (it was the of collaboration between them. From corre- whole of my then collection, so that I do not spondence with Schweinitz it becomes clear know to which species his observation apply) that Torrey at age 23 already had accumulated after his death prove unsuccessful. He has a much bryological knowledge, and, as Shear and number of Species- to which Swar[t]z is sub- Stevens (1921) noted, he had early resolved to joined in his Catalogue of which it seems impos- devote most of his time to the study of mosses. sible to know what was meant but by examining The Schweinitz-Torrey correspondence con- his Herbarium..." (Shear and Stevens, 1921: tains ample evidence of this, and also that 142). Schweinitz and Torrey 's long-discussed Torrey was a master at working simultaneously, United States cryptogam flora was never com- and productively, at many disparate scientific pleted, although Schweinitz (1832) published tasks. His income came largely from teach- the equivalent of the mycological part, and Tor- ing chemistry and mineralogy and demonstrat- rey 's friend, Abraham Halsey (1824), of New ing aspects of the physical world to college and York City, was at work on lichens. It is not clear medical students, but there was always time for who was to have responsibility for mosses and botany. There too he maintained diverse inter- liverworts, most likely some combination of ests, in reality all aspects of phanerogamic and

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 20 10 A TRIBUTE TO ASA GRAY 295 cryptogamic botany, bryophytes from even at least Georgia. though Torrey for- as his work progressed seed-plant warded parts of thesetaxonomy collections to Bridei became his primary focus. (1826-1827) who used some as the basis of Torrey was a prolific letter writer, and the new species or in citations of occurrence. Dis- network of correspondents he assembled in the tributional ranges of bryophytes were for most United States and beyond was remarkable for its regions of the world incompletely known breadth and intensity. Torrey knew that progress early in the 1800s. Thus, such early informa- in documenting the flora of the United States tion is useful in tracing how quickly species would come from new exploration throughout ranges became defined as botanical exploration the North American continent, access to authen- continued. tic specimens used by European experts in pro- Torrey never published on bryophyte tax- posing new species, and the constant and timely onomy. Although his name is associated with exchange of printed matter and specimens. He bryophyte species described in works of his con- encouraged Europeans to publish in American temporaries, he did not alone publish any new journals, including the Annals of the Lyceum of species of mosses or liverworts. His catalog of Natural History of New York , in which an early plants (1819) occurring within 30 miles of New bryological paper by Greville (1824) appeared. York City, completed at age 21 , was for its time Torrey amassed a large botanical library, and he an advancement over similar lists that relied constantly endeavored to increase it. He consid- principally on the Hedwigian genera. Torrey ered his book collection to be a lending-library, adopted names and taxonomie concepts for with needed items dispatched to correspon- mosses in Palisot de Beau vois (1805), and the dents on loan, and he was not reluctant to ask catalog of 62 mosses and 20 liverworts Torrey others to send him a book he had not seen. presented is noteworthy for its citational com- Torrey 's herbarium served as a clearing house pleteness (synonyms, habitat and locality data, to which boxes of specimens and printed month or season of fruiting included), in com- matter were shipped from places in the United parison to Muhlenberg's (1813, 1818), Green's States and Europe and then distributed by Torrey (1814), or Barton's (1815) sparse contempo- to the addressee or to whomever he thought rary catalogs, for example. Torrey 's work in would most benefit by sharing the contents. bryology in retrospect can be seen as genuine This made Torrey an exchange hub at a time and well grounded, but he never developed as when mail service and the shipment of boxes an authority on the taxonomy and identification and parcels were poorly developed, costly, andof bryophytes. He moved bryology forward in often by different sometimes unreliable carriers the United States by encouraging collectors, (Fuller, 1972). building a personal herbarium of authentically Specimens flowed into Torrey 's Herbarium named specimens, and shipping specimens to from diverse sources. As he was particularly European authorities who often brought into interested in the bryophyte flora of the United print bryological information about the United States, he encouraged numerous collectors. StatesFor flora. His encouragement of Asa Gray, example, Dennis Cooley, M.D., and Chester as we shall see, ultimately had a powerful Dewey, D.D., both initially collecting effectin on the further development of bryology Massachusetts, transmitted bryophytes into the United States, and through Gray to him from that state, and Cooley also sent himWilliam S. Sullivant.

Asa Gray

Gray (1810-1888; Figure 1C) was born botany in and mentored him as his professional the Mohawk Valley, New York, in Sauquoit, goalsand matured and moved from medicine to educated at schools close to his home. He was botany. Gray wrote that his botanical inter- introduced to academic botany as part of the cur- ests began during the winter of 1827 (J. Gray, riculum at College of Physicians and Surgeons 1893). He purchased a copy of Amos Eaton's of the Western District of the State of New York A Manual of the Botany of the Northern and (the Fairfield Medical College), graduating Middle States , ed. 4 (1824), and absorbed its from it with an M.D. in January 1831 at age 21 contents. (Dupree, 1959). This led to field explo- The Professor of Materia Medica there, James ration the following spring and collections that Hadley, nurtured Gray's developing interest in began his herbarium. The Amos Eaton floras

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 296 HARVARD PAPERS IN BOTANY Vol. 15, No. 2 were used widely in schools I am and unable collegesto send specimens in ofthe all the plants United States (McAllister, you 1941). wished- Eight whenever editions I obtain them I will appeared, starting with Eaton supply you(1817) with pleasure. and Iending have also enclosed with Eaton and Wright a(184b), few Grasses and &c and althoughsome Mosses which the earliest ones contained were collected only last the fall. barestThey are numbered and rudiments of bryology, Eaton when and you Wrighthave leisure ( to1 examine840) them you advertised that their presentation will oblige meof bythe transmitting crypto- their names with gams was "the only general those system of the undetermined in English, ones in package of hitherto attempted, on the last fivesummer. lower I intend ordersto pay some attention to of American Cryptogamia..." Cryptogamous (1840: plants in97). the ensuingAll season, and editions adhered to the artificial Linnaean shall be pleased to supply you with such speci- system of flowering plant classification, which, mens as you may be desirous of obtaining. The though having an utilitarian advantage in facil- want of proper works on the American cryp- itating identification, was quickly superseded togamia renders their study difficult, and I am by the more biological natural system cham- looking forward anxiously for the publication of pioned in the United States by John Torrey, your work on the Ferns & Mosses" (L. C. Beck Lewis C. Beck (Sebring and Sebring, 1934), Papers, Rutgers University Archives). Beck and their successors. replied on 3 August 1831, "As to your mosses, Gray initiated correspondence with Beck of I have not yet had time to do any thing in that Albany, New York, who had described a new department this summer, except in the way of moss, Grevillianum serratura (= Timmia mega- collecting. I shall, however, very soon take them politana Hedw.; Beck and Emmons, 1826), and up & as my means of comparison with authen- in a later paper (Beck, 1829) announced a plan tic European & American specimens have been to publish a fern and moss flora of the United greatly increased during the past season, I shall States. For reasons unknown, Beck never fin- probably be able to determine most of them sat- ished his announced moss flora, although his isfactorily" (Archives of the Gray Herbarium). bryophyte herbarium, a draft manuscript, and Beck presented summer courses of botanical letters attest to how he proceeded to summarize lectures at Fairfield Medical College, but I can bryological information available to him. I pres- find no evidence that Gray enrolled in any of ently have these sources under study. them. In the summer of 1831, however, when Gray first wrote Beck on 6 February 1930, Beck was unable to teach at Fairfield, James "By the advice of Prof. Hadley, I have taken Hadley obtained Gray as the replacement, no the liberty to send you a small package of dried doubt an acknowledgment of Gray's advanced plants. Will you have the goodness to return to botanical knowledge. me the name of the plants with their numbers, A large part of the Lewis C. Beck bryophyte by mail. The specimens if they will be of any herbarium, kept in 19 small books in the way use to you will please to retain" (L. C. Beck he arranged the specimens, is in the Bryophyte Papers, Rutgers University Archives). The entry Herbarium of the New York State Museum. The for L. C. Beck in the Jane Loring Gray auto- Beck Herbarium includes three bryophytes col- graph collection (Archives of the Gray Herbar- lected by Asa Gray, all from his home terri- ium, Harvard University) carries the annotation, tory (Oneida County) in east-central New York. "Asa Gray's first botanical correspondent." They are Conocephalum conicum (L.) Lindb., Beck responded on 3 May 1830, commenting s. lat. (as Marchantía conica ), Pohlia wahlen- at length on the identity of ferns and seed plants bergii (F. Web. & Mohr) Andrews (as Bryum Gray had sent him, and concluded, "I wish you androgenům Hook.), and Rhytidiadelphus tri- would take up the mosses of your region. If you quetrus (Hedw.) Warnst, (as Hypnum trique- desire it I will put up for you a few from this trum L). Early Asa Gray bryophyte collections region..." (Archives of the Gray Herbarium). in the Steere Herbarium (New York Botanical Gray wrote again to Beck on 5 March 1831, Garden) include specimens from Bridgewa- "I embrace this opportunity to transmit a few ter, Fairfield, and Penn Yan, New York; White specimens of those plants of which you desired Mountains, New Hampshire; and Allegh- an additional supply last summer. I regret that any and Appalachian Mountains (New York

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Botanical Garden, circumstanceC. V. canStarr be seen as pivotalVirtual in Gray's Herbar- ium, 2010). Duplicates botanical career. of Torrey somewrote to Schweinitz of on these and others of Gray's bryophyte 2 November 1833, "[Gray]... collections is now working are in the Sullivant Herbarium daily inat my herbarium.the Farlo My whole collectionw Herbarium, Harvard University. will soon be arranged according to the Natural Asa Gray and John method, Torreyand in the spring (D.V.),were I shall attackin contact. Torrey, in a letter with to zeal, myGray's Flora Synopsis homeof North Ameri- in Bridge- water, New York, can dated plants. Dr. G. will19 devote April part of his time1831, to invited Gray to visit him his ownin concerns New (according York to our agreement), City "in the present season," and & has madewrote arrangements the for publishing following collec- about work Torrey had accomplishedtions of dried plants of the more difficultrecently genera in bryology. "I have the & presentfamilies:- such as Gramineae,season Cyperaceae, resumed the study of mosses & Aster, have Musei, &c. commenced He hopes to publish the 1st a general account of the species No. of hishitherto N. Amer. Grasses in obtainedthe Spring & the in North America. The skeleton 1st No. of his N.of Amer. the Mosses about work the same is already prepared & a place time" provided (Shear and Stevens, for1921: 280). anyGray did new materials which may come in fact issue toan exsiccata hand. of grasses andYou sedges will oblige me very much by collectingto acclaim (Gray, 1833-35), for but heme never fol-all the Musei which you observe. lowed Ithrough wish on his plana tolarge do the same forquantity of each species so that the mosses. when A reason forI thishave is not available, determined a number I may have but enough Torrey 's comment to suggests last Gray hadfor been some time. The specimens should working onbe mosses in and fruitaccumulated a substan-if it is possible to obtain them in that state. I am sure we have tial, dividable collection, or, possibly Torrey many new species in the middle and northern encouraged Gray to carry forward the Torrey- States for I showed my collection to Mr. Drum- Schweinitz plan for the moss portion of the mond, the great Scotch Muscologist, a few days much-discussed cryptogam flora of eastern ago & there were many in it with which he was North America. Torrey 's knowledge of cryp- unacquainted. This gentleman, as you are prob- togams and his intentions were well known. ably aware, accompanied Capt. Franklin & Dr. Hooker (1825: 280), for example, wrote, "... Richardson in the last exped" to the north. He Dr. Torrey has made a most ample collection spent two years (most of the time among the of the cryptogamic plants of the United States; Rocky Mountains) & and made extensive col- that he is well acquainted with the species and lections, especially of the mosses. On his return their characters; and we may confidently hope he published 50 sets of dried specimens each that this department of botany will now find a containing about 280 species of mosses- one place in the Floras of North America." Alas, set I have just received. This gentleman is just that was not to be. With a few early excep- now on his way to the regions west of the Mis- tions (e.g., Darlington, 1853) and the start of sissippi for the purpose of collecting plants. an effort to document floras of specific locali- He sent out two tons of paper round by the way ties (e.g., Hitchcock, 1829), there was usually a to New Orleans! So that you may judge how sharp divide between floras and catalogs treat- intensively he intends to collect" (Archives ing cryptogamic groups and vascular plants. of the Gray Herbarium). Later in 1831, Gray Gray secured in 1842 a professorship at had sent mosses and gained sufficient experi- Harvard University where he spent the rest of ence that Torrey wrote, "The mosses which his career. There he developed an important you sent me have been laid aside for a future herbarium and a botanical library, in addition study, but I would mention that such as you to teaching and performing other university have labeled are generally correctly named" duties, including supervisory care of the uni- (Archives of the Gray Herbarium, Torrey toversity's botanical garden. Before moving to Gray, 6 October 1831). Cambridge, Massachusetts, late in the 1830s, The relationship between Torrey and Gray and when he was still associated with Torrey in blossomed, and in 1833, Asa Gray became John New York City, Gray (but probably Torrey first) Torrey 's assistant in botany and chemistry. This began to correspond with William S. Sullivant

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( 1 803-73; Figure ID), a wealthy of the surveyor, world. Heland- became, as Asa Gray called owner, banker, and businessman him, the of Father Columbus, of American Bryology (Steere, Ohio (Rogers, 1940). Although 1977). Rogersnearly (1940),seven Gray (1873), and others, years older than Gray, Sullivant have summarized became a close why such a title is justified, friend and remained so throughoutso only a brief his life,synopsis of Sullivanťs many which, starting about 1840, bryological became ever accomplishments more follows. productively devoted to bryology, As we have seen, thereespecially was no substan- mosses. Gray played a major tial publishedrole in flora this, (identification recog- book) that nizing his friend's talent, couldand, be asused ato consequence,name American bryophytes. sending specimens and books, Therefore, floraspurchasing written for parts at of Europe Sullivanťs behest the best were microscope usually employed. available Hooker and Taylor in Europe, and eventually (1818, putting 1827) are good Sullivanťs examples of books that name in front of the most found accomplished utility in America for Euro- this purpose. That pean bryologists. The last changed resulted with Sullivant. in a cascadeThe first edition of of authentically identified Gray's reference Manual of the specimens Botany of the Northern arriving at the Sullivant laboratory United States appeared and in library1848 and contained in Columbus. William J. Hooker, accounts of Musei a bryological and Hepaticae by Sullivant. leader at the time, named This specimens is the first flora devoted for to Sulli- North Ameri- vant in 1841 (Rogers, 1940), can bryophytes, the yeareven though Hooker it treated species left a professorship at Glasgow occurring only University in a portion of the to continent- begin as Director of the Royal New England, Botanic west to Wisconsin, Garden, and south to Kew. Hooker maintained a Ohio strong and Pennsylvania. interest Gray had in intended his the North American bryoflora new book to and be a "compendious generously flora," by which encouraged Sullivant (and hemany meant all theother phanerogams novice and vascular and bryologists) over a long period other cryptogams of time. of the region. In the preface As a result of native ability Gray announced and member- that Edward Tuckerman, at ship in Gray's network Gray'sof invitation,correspondents, had completed an account of Sullivanťs progress in bryology lichens for thewas Manual rapid. , but at overIn 700 pages 1846 he communicated long(and already, probably more could not bepub- added to it. The lished; Sayre, 1981) his first second bryologicaledition of the Manual paper , Gray (1848: vi) to the American Academy stated, of would Arts have a supplementaryand Sci- volume ences (Sullivant, 1848a). The treating first algae, bryophytes,of the lichens,four and fungi. exsiccatae he distributed, However, Musei edition twoAlleghani- did not fulfill that plan, enses , was issued in the samebut instead year contained (Sullivant, an expanded bryophyte 1846a). It appeared under thesection joint by Sullivant, sponsorship including elegant illus- of Asa Gray and W. S. Sullivant trations from and copper consisted engraving plates based on of 292 dried moss and liverwort original Sullivant drawings.specimens, Thus was born the with printed labels containing first illustrated synonymies, manual of mosses and liverworts habitat information, and descriptions for any part of North of America, the whichnew in edition species. A separate publication 2 was expanded of the to includelabels Virginia (the and Ken- schedae) also appeared that tucky and year westward (Sullivant, from Ohio and Wisconsin 1846b). In its preparation theto the MississippiMuhlenberg River. The bryophyteher- por- barium was consulted, Thomas tions of editions Drummond's 1 and 2 of the Manual were specimens examined critically, also issued as and separate informa- small books (Sullivant, tion from Torrey's herbarium 1848b; Sullivant, incorporated. 1856), thereby providing an Musei Alleghanienses won unencumbered Sullivant focus immediate on unique morphologi- attention wherever bryology cal characteristics was under of mosses serious and liverworts and study, and its completion theircan taxonomy.be seen Sullivanťs as fulfill- floristic summary in ing Gray's plan, announced edition in 2, 1833,and no doubt to the issue illustrations, a encour- study set of American mosses. aged many From in the United these States early to work on bryo- works, Sullivant went on phytes.to accomplish The demand for this much book continued for more in the documentation decades, of theand there North was a second Amer- later printing ican bryoflora and that of by amany different otherpublisher (Sullivant,parts 1871).

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As successful as these two American floras leaf cells and generalized peristomial struc- were, Sullivanťs Icones Muscorum (1864) and ture, the latter providing important criteria for its Supplement (1874), the latter a posthumous the classifications they proposed, but even so work, were his supreme achievements. They the drawings lacked the detail achievable later. also were primarily about eastern North Amer- With improved compound microscopes, fine ican mosses, and they ambitiously set out cellularto and peristomial details became clearer, describe and present all species for which anallowing newly revealed information to enter acceptable illustration was not already available classifications, as documented by Sullivant, in another source, including some mosses from his contemporaries, and those who followed. western North America. Illustrations in these Frahm (2000), in discussing Hedwig 's bryolog- two books, indeed those in all of Sullivanťs ical achievements, emphasized that by employ- publications, were executed at the highest ing compound microscopy at magnifications level of achievement, and therefore equal, or of 170-290X Hedwig was the first to introduce often better, than those published at the time microscopic detail into bryological systematics. in Europe. Sullivanťs drawing skills, and Sullivanťs labors in bryology began at a those of Eliza Wheeler Sullivant, his second time when major conceptual advances were wife and longtime bryological assistant (Rogers, achieved in understanding the biology of 1940), were the foundation for this, as was mosses and liverworts. These concepts included his willingness to pay for the best engraving the final, definitive elucidation of the bryo- and printing, but advances in microscopy, par- phyte life cycle and tracing the development ticularly increased resolution provided by of the bryophyte plant body back to the activ- advances in compound microscope design ity of single apical cells (Sachs, 1906). Taken contributed as well. broadly, these advances linked bryophytes to The historian, Nathan Reingold (1964: 41), other groups of green plants- the green algae, in a discussion about scientific progress in vascular cryptogams, seed plants- in a mean- botany and internationalism in science in 19th ingful way that was compatible with the theory century America, wrote, "Identification and of evolution, and at the same time established classification of the plants was the principal their uniqueness. object of the international network. To properly Sullivant, following Torrey and Gray, devel- accomplish this, a botanist needed only refer- oped his own network of correspondents. Fifty- ence works giving past descriptions of plants three of them are named by Rogers (1940). He but also a herbaria [sic] containing preserved acquired Dennis Cooley, then living in Macomb specimens." However, for bryophytes and other County, Michigan, who earlier had extensive cryptogamic groups, there was a third compo- bryological exchanges with John Torrey. Sul- nent, microscopy, which provides the means livant wrote Cooley on 25 August 1845, con- to see and interpret small, botanically impor- firming when he began to work actively with tant, structures. Sullivant worked in bryology bryophytes, and requesting an exchange, "For at a time when compound light microscopy the last 3 or 4 years I have been attending to became greatly refined in resolving power. By cryptogamic botany in part, i.e., Musei & comparison, Dillenius' engravings on copper Hepaticae- hence my duplicates of Phaenog. plates (1741) were of images seen through a are much reduced, still I can get together a few simple (one lens) microscope and were ren- yet- some carices, formerly a favorite study dered at actual size. Therefore, only rarely were with me. . . Should be very glad to have Mosses cells resolved at the magnifications then avail- & Hepaticae (copious specimens) of your neigh- able, for example, in his renderings of leaves borhood-I can make you satisfactory returns for Plagiomnium and Hookeria. As compound I think" (from an unpublished letter in the D. (two-lens) microscopes improved, cellular and Weinstock, M. D., Collection, Geneva, New epicellular structure became observable, and York). Specimen exchanges and correspon- illustrations of such morphological features, dence with T. G. Lea of Cincinnati, Ohio (Lea, which became known to be rich in taxonomi- 1849), Charles Wilkins Short of Lexington, cally important characteristics, were illustrated. Kentucky, and other botanists with bryological Hedwig (1801) and Hooker and Taylor (1818), interests in eastern and middle-western North and others after them published renderings of America were also initiated. Gray encouraged

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 300 HARVARD PAPERS IN BOTANY Vol. 15 , No. 2 his collectors, notably Charles in the United StatesWright during the and 19th century Augustus Fendler, to gather (Stuckey, bryophytes 1997). Gray hadfor special Sul- plans for livant, and Gray's work with the 6th aedition cadre of the of textbook, plant which was to explorers in the American include West a separate (McKelvey, volume for the cryptogams, 1955) led to significant collections both their structure coming and systematics, to written by Sullivant for study. Bryophytes his Harvard colleague(Say re,W. G. Farlow,1975) but only from the United States twoExploring of a projected four Expedi- volumes came into print tion, 1838-1842, under the (Gray, command 1879; Goodale, of 1885). Capt. Most editions Charles Wilkes, came to him for taxonomie of the textbook contain brief statements about study, and an imperial folio with 26 sumptuous bryophytes, but the first, second, and last of the plates (Sullivant, 1859) was one of several pub- series (Gray, 1887b) have longer descriptions lished results. My census of the Sullivant Her- of them as plant classes, but floristic detail is barium, maintained separately at the Farlo w not present. Prior to Gray's textbook series, as Herbarium (Harvard University) shows that abbreviated as most were in bryological matters, Gray systematically sent Sullivant most (all?) there was little published in America present- of the major bryological holdings in Gray's her- ing introductory information about bryophytes barium. These are now returned to Cambridge. and bryology. The Amos Eaton floras (1817- Sullivant's herbarium was sufficiently rich in 1840) have already been mentioned. Encyclo- duplicate specimens from publishing bryolo- pedias were authoritative widely read sources gists worldwide that he completed taxonomie of information in the early 1800s. In the Edin- accounts of collections from Tierra del Fuego, burgh Encyclopaedia , an American Edition of the Pacific Islands (including Hawaii), Cuba, which was published (Brewster, 1832), there and Venezuela (Schräder, 1879). was a comprehensive article by John Stewart of Gray continued to support Sullivant's bry- Edinburgh on Musei (mosses), including Musei ological work throughout his life. Gray's edi- Hepaticae (liverworts), and three plates of illus- torial expertise and knowledge of the printing trations by R. K. Gre ville, and Abraham Rees' industry was applied to his friend's publica- Cyclopaedia (1819) had similar entries. Musei tions, a practice that continued at the Gray Her- and Hepaticae are treated in a general way in barium through the publication of Lesquereux edition seven of the Encyclopaedia Britan- and James' Manual of the Mosses of North nica (1842) as part of a lengthy article about America (1884), which Sereno Watson helped botany. Elements of Botany by Benjamin Smith to guide into print. Gray continued until the Barton (1803), a physician and botanist of Phil- year before his death (Gray, 1887a) to write adelphia, continued in two subsequent editions/ influential reviews of significant bryological issues (Barton, 1814, 1827), contained brief contributions published abroad and in North summaries of the bryophytes and illustrations America. This kept bryology in the view of all of mosses and a liverwort, as did the fourth revi- botanists. Gray and Sullivant conducted joint sion by William P. C. Barton (Barton, 1836), field work in 1843 in the Alleghany Mountains his nephew. John Torrey (1831) in his Amer- in the southeastern United States, discovering ican edition of Lindley's Botany , a book that there species of significant taxonomie and bio- helped bring the natural system of plant clas- geographic importance. Gray in 1848 sent to sification into regular use in America, added Sullivant, Leo Lesquereux, a meritorious Swiss to Lindley's introductory material about bryo- immigrant bryologist, initially employed by phytes, an appendix entitled, "Catalogue of Gray as an assistant in his herbarium. Sullivant the North American Genera of Plants." In it hired Lesquereux to work with him in Colum- were listed in a table 45 genera of mosses and bus. Lesquereux collected mosses widely in 8 of liverworts, with the number of species in eastern North America, making new discoveries each genus. The American edition of Smith's A and later distributing the specimens with Sulli- Grammar of Botany (1822) also presented an vant or by himself (Miller, 2008). abbreviated treatment of bryophytes. Familiar Asa Gray's many textbooks, written for stu- Lectures on Botany (Lincoln, 1829) and dents of botany of different ages and published succeeding editions through 1869 (Rudolph, between 1836 and 1887 (the last continuing 1984), Sumner (1820), and somewhat later, in print after his death), greatly influenced the Alphonso Wood's Classbook of Botany (1845; teaching of botany in high schools and colleges also with numerous new editions and printings)

This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:58:36 UTC All use subject to https://about.jstor.org/terms 2010 A TRIBUTE TO ASA GRAY 301 provided distillations Gray fostered of the development general of an accom- bryophyte morphology for beginning plished, well-regarded, students, American-born bry- but no information on the floristic ologist of international diversity fame, and in America of mosses and liverworts. With a tradition all ofof high qualitythis, bryological entry work. into bryology must still have Sullivanťs been organizational difficult skills, bryologi- for Americans until the publication cal talent, ambition, of andeditions wealth enabled him one and two of Gray's Manual . to achieve rapid taxonomie progress in North Gray maintained American bryophytea lifelong floristics and taxonomy. interest in bryology, but Sullivanthe wasdeferred the first botanist in America toto Sullivant in bryological matters devote most ofinvolving his scientific work to mosses specimen identification, taxonomy, and liverworts. Together, and Sullivant research and Gray conclu- sions. Gray helped Sullivant format his advanced bryological achievement in the publications so that they were the equal to United States, and with the foundation they those of the best bryological taxonomists and built others were encouraged to expand equal also to the best contemporary papers in bryological knowledge, in floristics, and in taxonomie botany in general. In these ways many other directions. Literature Cited Arnott, G. A. W. 1826. Tour to the south of France Bridel, S. E. 1826-27. Bryologia Universa 1-2. and the Pyrenees, in 1825. Edinburgh New Phil. Barth, Lipsiae. Jour. 1: 66-78, 268-275; 2: 241-255. [Arnott Brown, R. 1823. Chloris Melvilliana. Printed by listed results of his studies of Palisot de Beauvois W. Clowes, London. moss specimens in Herb. Delessert that enabled Cardot, J. 1899. Etude sur la flore bryologique him to "make out what many of his hitherto doubt- de L'Amérique du Nord. Revision des types ful mosses are."] ď Hedwig and de Schwaegrichen. Bull. Herb. Atwood, J. H. and B. Allen. 2009. A synopsis of the Boisser 7: 300-380, plates 7-10. Orthotrichum strangulatum complex (Orthotrich- Crum, H. A. 1952. Bibliographic notes on Palisot aceae). Bryologist 112: 208-220. de Beauvois' prodrome. Bull.Torrey Bot. Club 79: Barnhart, J. H. 1934. The botanical correspondence 407-409. of Schweinitz. Bartonia 16: 19-36. Curtis, M. A. 1867. Geological and Natural History Barton, B. S. 1803. Elements of Botany, parts 1-2. Survey of North Carolina, Part III, Botany; Printed for the author, Philadelphia. Containing a Catalogue of the Indigenous and

Naturalized Plants of the State. Printed at N. parts 1-3. Printed for the author, C. Institution Philadelphia.for the Deaf and Dumb and the Blind, Raleigh, North Carolina. [Bryophytes on Desilver. Philadelphia. [Reissue pages 69-76.]of edition 2, with a new title page.] Cutler, W. P. and J. P. Cutler. 1888. Life, Journals and Correspondence of Rev. Manasseh W. P. C. Barton. Robert Desilver, Cutler, LL.D., Philadelphia. 1-2. Robert Clarke & Co., Barton, W. P. C. 1815. Florae Philadelphicae Cincinnati. Prodromus. Printed for the author by J. Maxwell, Dandy, J. E. [editor]. 1958. The Sloane Herbarium, Philadelphia. an Annotated List of the Horti Sicci Composing Beck, L. C. 1829. A synoptical table of the ferns and it, with Biographical Accounts of the Principal mosses of the United States. Am. Jour. Sci. Arts Contributors. British Museum (Natural History), 15: 287-297. London.

Darlington, W. [compiler]. 1843. Reliquiae Grevillianum serratum, a Baldwinianaenew genus, : Selections belonging from the Correspon- to the order Musei. Am. Jour. dence Sci. of the Arts Late William 1 1 Baldwin,: 183, M.D. plate Kimber 1A-E. and Sharpeless. Philadelphia.

Bellis, V. 2009. John Lawson's plant collections from North Carolina, 1710-1711. Castanea 74: Blakiston. Philadelphia. 376-389. Dillenius, J. J. 1741 [published 1742]. Historia Benedict, D. M. 1934. Schweinitz' botanical library. Muscorum. E theatro Sheldoniano, Oxonii. Bartonia 16: 14-19. Dupree, A. H. 1959. Asa Gray, 1810-1888. Belknap Brewster, D. 1832. The Edinburgh Encyclopaedia. Press of Harvard University Press. Cambridge, Volume 14. Joseph and Edward Parker, Massachusetts. Philadelphia. [Musei, by John Stewart, illustrations Eaton, A. [Members of the Botanical Class in by R. K. Greville, pages 1-36, plates 401^03.] Williams' College (Massachusetts).] 1817.

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A Manual of Botany for theGreville, Northern R. K.States. 1824 [pub. 1825]. Descriptions of Websters and Skinners. Albany, some New new York. plants belonging to the orders Musei and Hepaticae. Ann. Lyceum Nat. Hist. New York 1: and Middle States of America, 271-278, edition 4.plate Websters 23. and Skinners. Albany, New York.Grovovius, J. F. 1739-1743. Flora Virginica parts

1-2. Cornelius Haak. Leiden. Botany; Comprising the Native and Common Cultivated Plants, North Halsey, of A. Mexico, 1824. A synopticaledition 8. view or the lichens Elias Gates. Troy, New growingYork. in the vicinity of the City of New York. Encyclopaedia Britannica. Ann. 1842. Lyceum Botany. Nat. Hist. Volume New York 5:1 : 3-21 , plate 1 . 30-141 , plates 1 12-126. Hampe, Adam E. 1837.and CharlesGenus novum Black, muscorum fron- Edinburgh. dosorum phascaceorum. Linnaea 11: 279, 280. Frahm, J. -P. 2000. The life and work and Johannes [Spor lederà beyrichiana.] Hedwig. Nova Hedwigia 70: 1-13. Fuller, W. E. 1972. The American Mail, Enlarger of C. Beyrich auf seiner letzten Reise in Nordamerika the Common Life. University of Chicago Press, gesammelten Laubmoose. Linnaea 13: 39-48. Chicago and London. Harpel, J. A. 2010. A history of bryology in the San Goodale, G. L. 1885. Gray's Botanical Text-Book, Juan Islands, Washington State, USA. Botany edition 6, volume 2. Physiological botany. Part I. 88: 329-335. Outlines of the Histology of Phaenogamous Plants; Harper, F. 1940. Some works ot Bartram, Daudin, part II. Vegetable Physiology. I vison, Blakeman, Latreille, and Sonnini, and their bearing upon Taylor & Co., New York. North American herpetological nomenclature. Am. Gottsche, C. M., J. B. G. Lindenberg, and C. G. Midi. Nat. 23: 692-723. Nees ab Essenbeck. 1844-1847. Synopsis Hepati- Harshberger, J. W. 1899. The Botanists or carum. Meissner, Hamburg. Philadelphia and their Work. T. C. Davis & Gray, A. 1833-1835. North American Gramineae Sons, Philadelphia. and Cyperaceae, parts 1-2. Printed by J. Post. New Hedwig, J. loUl. Species Muscorum tronaosorum. York. [Exsiccata.] Opus Posthumum Editum a Friderico Schwae-

2 rie hen. Barth, Lipsiae. Co., New York. ico-Analytica Frondosorum. Vol. 4. Bibl. Gledits- Part I: An Introduction chiano.to Structural Lipsiae. and [Hypnum Physio- delicatulum and H. logical Botany; part II: Theminutulum Principles from of Pennsylvania, System- leg. Muhlenberg.] atic Botany. Wiley and Hitchcock,Putnam, New E. 1829.York. Catalogue of Plants Growing

Without Cultivation in the Vicinity of Amherst Northern United States. James Monroe and Co., College. Published by the Junior Class in that Boston and Cambridge. [Musei and Hepaticae by Institution. J. S. and C. Adams, and Co. Printers, W. S. Sullivant.] Amherst, Massachusetts.

Hooker, W. J. 1818-1820. Musei Exotici. Longman, ern United States, edition 2. Hurst, George Rees, P. Putnam Orme, & and Brown. London. Co., New York. [Musei and Hepaticae by W. S. Sullivant.] Sci. Arts 9: 263-284.

Acad. Arts Sci. 9: 271-276. nica ; Containing the Mosses of Great Britain and

Ireland, Systematically Described and Arranged. 6, volume 1 . Structural Botany, Longman, or Organography Hurst, Rees, Orme, and Brown, London. on the Basis of Morphology; to Which is Added the Principles of Taxonomy and Containing Phytography, and the Mosses of Great Britain and a Glossary of Botanical Terms. Ireland, American Book Systematically Described and Arranged, Company, New York, Cincinnati, edition Chicago. 2. Longman, Hurst, Rees, Orme, Brown & Green, London. 34: 490^494. [Includes reviewLea, ofT. Braithwaite's G. 1849. Catalogue of Plants, Native and British Moss Flora .1 Naturalized, Collected in the Vicinity of Cincin-

nati, Ohio, During the Years 1834-1844. Printed Book Company, New York. by T. K. & P. G. Collins, Philadelphia. Gray, J. L. [editor]. 1893. Leroy,Letters J. F. 1957. of Le Asa premier Gray explorateur 1-2. de Houghton, Mifflin, Boston. Terre-Neuve et des îles Saint-Pierre et Mique- Green, J. 1814. An address lon: on Bachelot the de botany la Pylaie (1786-1856). of the Pages United States... to which is 203-230, added, plates a 17-22, catalogue in Les Botanistes of plants indigenous to the State Français of en New Amérique York. du North Trans. avant 1850. Soc. Promot. Useful Arts 3: Colloques 65-136. Internationaux du Centre Nationale de la Recherche Scientifiques 63. Paris.

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Lesquereux, L. and T. P. James. 1884. Manual of faisant partie du primier volume des Mém. Soc. the Mosses of North America. S. E. Cassino & Linn. Paris" (1822)]. Co. Boston. [Includes plates 1-6 from Sullivant Patterson, P. M. 1965. John Clayton's collection of (1856).] Virginia mosses. Bryologist 68: 105-109. Lincoln, A. H. [A. H. Lincoln Phelps]. 1829. Famil- Pennell, F. W. 1934. The botanist Schweinitz and his iar Lectures on Botany. H. and F. J. Huntington, herbarium. Bartonia 16: 1-36. Hartford, Connecticut. Rafinesque-Schmaltz, C. 1808. Prospectus of Mr. Linnaeus, C. 1753. Species Plantarum , volume Rafinesque-Schmaltz's 2. two intended works on Impensis Laurentii Salvii , Holmiae. [Musei pages North- American botany. Medical Repository II. 1100-1130, Algae (including most Hepaticae, 5: 350-356. [" Heterodon , ( bryoides ) small moss pages 1131-1140.] with peristome, dentatures unequal. It grows in the Lyon, Cj. h. 1 ö25. A brier Narrative or an Unsuccess- waters of New-Jersey."] ful Attempt to Reach Repulse Bay. John Murray, Rees, A. 1819. The cyclopaedia; or universal dic- London. [Botanical appendix by W. J. Hooker, tionary of arts, sciences, and literature. Hepaticae, pages 185-198.] volume 17 [1 page], Musei, volume 24 [8 pages]. McAllister, E. M. 1941 . Amos Eaton, Scientist and Longman, Hurst, Rees, Orme & Brown, London. Educator, 1776-1842. University of Pennsylvania Reid, A. M. M. 1987. Pioneer New England bryolo- Press, Philadelphia, Pennsylvania. gists- a prosopography. Occas. Pap. Farlo w Herb. McKelvey, S. D. 1955. Botanical exploration of Cryptog. Bot. Harvard Univ. 19: 1-71. the Trans-Mississippi West 1790-1850. Arnold Reingold, N. [editor]. 1964. Science in nineteenth- Arboretum, Jamaica Plain, Massachusetts. century America, a documentary history Hill and Mears, J. A. 1978. Sources of the herbarium of Wang. New York Henry Muhlenberg (1753-1815). Proc. Am. Phil. Rogers, A. D. III. 1940. "Noble Fellow" William Soc. 122: 155-174. Starling Sullivant. G. P. Putnam's Sons, New York.

Merrill, E. D. 1936. Palisot de Beauvois as an overlooked American botanist. Proc. Am. Phil. can botany. Princeton University Press, Princeton, Soc. 56: 899-920, 1 plate. New Jersey. Rohde, F. C., R. G. Arndt, J. W. Foltz, and J. technical names for the M. cellularQuattro. 2009. Freshwater cryptogams. fishes of South Farlowia 1: 245-262. Carolina. University of South Carolina Press, Columbia, South Carolina. of Henry Muhlenberg, with Ross, J. special 1819. Voyage attention of discovery made to under unrecorded or incorrectly the ordersrecorded of the Admiralty binomials. in His Majesty's ships Bartonia 25: 1-66, plate 1 . Isabella and Alexander for the purpose of explor- Michaux, A. 1803. Flora boreali-americana ing Baffin's Bay and a north-west , volume passage [Botany, 2. Caroli Crapelet. Parisiis incl.et bryophytes,Argentorati. by Robert Brown, [Hepat- cxxxvi-cxliv.] icae, pages 276-280; Musei, Rudolph, pages E. 1984. 285-319.] Almira Hart Lincoln Phelps Miller, N. G. 2008. A new biography of Leo (1793-1 884) and the spread of botany in nineteenth Lesquereux. Review of: Lesquereux, L. 2006. century America. Am. Jour. Bot. 71: 1161-1167. Letters written from America 1849-1853. Trans- Sachs, J. von. 1906. History of botany (1530-1860). lated from the French by H. D. Page. Includes a English transi, by H. E. F. Garnsey, rev. by I. B. biography of Lesquereux by D. G. Tritt. Picton Balfour. Clarendon Press, Oxford, England. Press. Rockville, Maine. Bryologist 111:1 34-1 36.Sargent, C. S. 1889. Scientific papers of Asa Gray Muhlenberg, H. 1813. Catalogus plantarum Ameri- selected by Charles Sprague Sargent. Volume cae septentrionalis . Printed by William Hamilton, I: Reviews of works on botany and related sub- Lancaster, Pennsylvania. jects, 1834-1887; volume II: Essays; biographical sketches, 1841-1886. Houghton, Mifflin and septentrionalis , edition Company, 2. BostonSolomon and New York. W. Conrad. Philadelphia, Pennsylvania. Sayre, G. 1975. Cryptogamae exsiccatae- an New York Botanical Garden, C. V. Starr Virtual annotated bibliography of exsiccatae of Algae, Herbarium. 2010. List of Asa Gray's moss and Lichenes, Hepaticae, and Musei. Mem. New York liverworts specimens in Steere Herbarium. Bot. Gard. 19:277-423.

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