Page  101 ï~~2008 THE MICHIGAN BOTANIST 101 A NEW LECTOTYPIFICATION OF THE NAME MEDEOLA VIRGINIANA L. (LILIACEAE) James S. Pringle Royal Botanical Gardens, P.O. Box 399, Hamilton, Ontario, Canada L8N 3H8 jpringle@rbg.ca ABSTRACT The specimen previously designated the lectotype of the name Medeola virginiana L. actually represents Asclepias quadrifolia Jacq. and thus is significantly at variance with the protologue. Another element of Linnaeus's original material is compatible with the protologue as well as with historic and current usage. The latter specimen is designated the new lectotype, superseding the earlier choice and stabilizing the nomenclature of the species. The name Medeola virginiana L., published by Linnaeus (1753) in Species Plantarum, ed. 1, has consistently been applied to a liliaceous species commonly known as Indian cucumber-root, which is native to much of eastern North America (Utech 2002). This species has been accepted as the type species of the genus name Medeola L. (Jarvis et al. 1993; Jarvis 2007), following its designation by Hitchcock and Green (1929). Although Linnaeus (1753) included two other species in Medeola, one Neotropical and one African, the genus has for many years been restricted to the one species M. virginiana. The International Code of Botanical Nomenclature (McNeill et al. 2006), Art. 13.4, states that genus names first used by Linnaeus (1753) in Species Plantarum, ed. 1, are to be associated with the first subsequent descriptions given under those names in the fifth edition of his Genera Plantarum. In Genera Plantarum, ed. 5, Linnaeus (1754) placed Medeola in the Linnaean class Hexandria Trigynia (six stamens, three carpels), and described the genus as having a perianth of six equal parts, six stamens, three recurved stigmas, and a trilocular, baccate fruit. He distinguished M. virginiana from the two tropical species by its whorled leaves and non-spiny stems. In the synonymy of the name M. virginiana, Linnaeus cited two phrase-names from earlier publications. The word Medeola had been included only in the first of these, Gronovius's (1739) Flora Virginica, ed. 1, in which the cucumber-root was named Medeola foliis stellatis, fructu baccato. Gronovius quoted descriptive information from a manuscript by John Clayton, who had become acquainted with this species in Virginia and had sent a specimen of it to Mark Catesby in England, from whom it was acquired by Gronovius. That specimen, now in the John Clayton Herbarium at BM (cited below as the new lectotype; image at <http://internt.nhm.ac.uk/resources/research-curation/projects/claytonherbarium/lgimages/BM000032590.JPG>, accessed 6 Feb 2008), clearly represents the liliaceous species to which the name M. virginiana is now applied. An

Page  102 ï~~102 THE MICHIGAN BOTANIST Vol. 47 notations by Gronovius, one of which includes the same wording that he used in Flora Virginica, identify this specimen as the Medeola described in that work. Reveal (1983) has recognized this specimen as thereby having been "indirectly reported in Linnaeus' first edition of Species Plantarum." It has been annotated "Syntype specimen of Medeola virginiana L." The latter annotation is anonymous and undated, but antedates the current Linnaean Plant Name Typification Project. (Standard symbols for herbaria follow Holmgren and Holmgren 1998 et seq.) The other name cited by Linnaeus (1753) was from Plukenet's Almagestum Botanicum (1696, p. 401), in which the species was designated Lilium s[ive] Martagon pusillum Virginianum, floribus minutissime herbaceis. This phrasename was also cited by Gronovius (1739). Plukenet quoted it from, and attributed it to, a catalogue of Virginia plants by John Banister, which had been published in vol. 2 of Ray's (1688) Historia Plantarum. Banister's drawing of the plant to which he gave this phrase-name (his no. 82, original at BM; reproduced in Ewan and Ewan 1970, p. 15 of Banister's drawings, fig. 8), clearly represents the liliaceous cucumber-root. Plukenet also cited a figure alleged to represent this species in his Phytographia (1691, plate CCCXXVIII, fig. 4). This figure, from a drawing by John Collins, is not adequate for positive identification. As noted by Ewan and Ewan (1970, p. 209), it does not correspond closely to Banister's drawing, differing conspicuously in the numbers of leaves per whorl and in the depiction of the venation of the leaves, as well as in details of the flowers, which are shown as having five stamens each. A plant of a different species may have been used as a model. Because the phrase-name was of Banister's authorship, his own clearly identifiable drawing, rather than this second figure, indicates the identity of the plant he thus named. In designating a lectotype for the name Medeola virginiana, Reveal (in Jarvis et al. 1993) chose neither Clayton's specimen nor Banister's drawing but instead selected a specimen from Linnaeus's herbarium that had been collected by Pehr Kalm (Herb. Linn. 468.1, LINN; image at <http://www.linnean-online.org/ 4158/>, accessed 6 Feb 2008). Linnaeus did not mention Kalm in connection with Medeola virginiana, but his annotations "virginica" [sic] and "2" indicate that he probably did identify this specimen as that species. The specimen is in the Medeola genus file in the Linnaean herbarium, and M. virginiana is species no. 2 under Medeola in Species Plantarum, eds. 1. Although Linnaeus had acquired Kalm's specimens of North American plants prior to writing Species Plantarum, ed. 1, there is no indication as to whether he thus identified this specimen before or after that work was published. The Kalm specimen currently designated the lectotype is actually a specimen of the four-leaved milkweed, Asclepias quadrifolia Jacq. (Asclepiadaceae), and had been identified and annotated as Asclepias quadrifolia by J.E. Smith. (Authorship of annotations is attributed in accord with the Web site of the Linnean Society of London, cited above). This specimen has pentamerous sepals, petals, and stamens, as is usual in A. quadrifolia. It is, therefore, in conflict with the generic characters given by Linnaeus (1754) for Medeola, including its placement in the Hexandria Trigynia. In equating his M. virginiana with Gronovius's

Page  103 ï~~2008 THE MICHIGAN BOTANIST 103 Medeola, Linnaeus implicitly accepted Clayton's description of the species as having a hexapetalous flower, with a purple style composed of three long filaments [actually three stigmas], and Gronovius's further description of its leaves as being in whorls of six or seven (as they are, respectively, in the two plants in Clayton's collection) and its fruit as being baccate. None of the foregoing is compatible with A. quadrifolia, nor, considering the conspicuously pinnate leaf venation of the milkweed, is Banister's interpretation of his plant as "Lilium sive Martagon." All of this descriptive material, and the placement of the genus in the Hexandria Trigynia, is in accord with the morphology of the cucumber-root. It does not appear that A. quadrifolia was significant in the formation of Linnaeus's concept of the genus Medeola or the species M. virginiana or in his preparation of the descriptions. The specimen in the Clayton herbarium at BM can be considered a part of Linnaeus's "original material" of M. virginiana. Like the other specimens in that assemblage, it is from the herbarium of J.F. Gronovius, much of which was studied by Linnaeus prior to the publication of Species Plantarum, ed. 1, and from which the lectotypes of many Linnaean plant names have been selected (Steamrn 1957). With Medeola having been described toward the beginning of Flora Virginica, in the pages presumably earliest written, it is especially likely that this specimen was seen by Linnaeus when he was working with Gronovius on that publication. Wording from Flora Virginica, quoted by Linnaeus in Species Plantarum, was based on this specimen. The morphology of the cucumber-root, including this specimen, is compatible with the protologue and the synonymy. The specimen in the Clayton herbarium cited below, therefore, is here designated the lectotype of the name Medeola virginiana, superseding the earlier choice by Reveal, in accord with the provisions of Art. 9.13 of the International Code of Botanical Nomenclature. This designation is consistent with traditional and current usage of the name. This action also stabilizes the typification of the subfamily name Medeoloideae (Benth.) M.N. Tamura and the family name Medeolaceae (S. Watson) Takht., and permits the continued use of the long-established name Asclepias quadrifolia for the four-leaved milkweed. Medeola virginiana L., Sp. P1. 1: 339. 1753. Lectotype, designated here: U.S.A.: Virginia: Clayton 22 (BM No. 000032590). ACKNOWLEDGMENTS I am grateful to the library staff of the Gray Herbarium and Arnold Arboretum of Harvard University for the opportunity to examine historic publications. I am also grateful to Dr. Charles E. Jarvis of the Natural History Museum (London) for helpful comments and suggestions. This paper is Contribution No. 184 from the Royal Botanical Gardens, Hamilton, Ontario, Canada. LITERATURE CITED Ewan, J., and N. Ewan. (1970). John Banister and his Natural History of Virginia, 1678-1692. University of Illinois Press, Urbana and Chicago, IL. Gronovius, J.F. (1739-1743). Flora Virginica Exhibens Plantas Quas V.C. Johannes Clayton in Vir

Page  104 ï~~104 THE MICHIGAN BOTANIST Vol. 47 ginia Observavit atque Collegit. Cornelius Haak, Leiden, The Netherlands. Facsimile published 1946. Arnold Arboretum, Jamaica Plain, MA. (Medeola in Pars Prima. 1739.) Hitchcock, A.S., and M.L. Green (1929). Nomenclature: Proposals by British botanists. H.M. Stationery Office, London, U.K. Holmgren, P.K., and N.H. Holmgren. (1998 and continually updated). Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden's Virtual Herbarium. <http://sweetgum.nybg.org/ih/>, accessed 23 Jan 2008. Jarvis, C.E., F.R. Barrie, D.M. Allan, and J.L. Reveal, compilers. (1993). A list of Linnaean generic names and their types. Regnum Veg. 127. Jarvis, C.[E.] (2007). Order out of Chaos: Linnaean Plant Names and their Types. The Linnean Society of London, London, U.K. Linnaeus, C. (1753). Species Plantarum Exhibitentes Plantas Rite Cognitas ad Genera Relatas, cum Differentiis Specificis, Nominibus Trivialibus, Synonymis Selectis, et Locis Natalibus, Secundum Systema Sexuale Digestas, ed. 1. Laurentius Salvius, Stockholm, Sweden. Facsimile published 1957-1959 as Ray Soc. Publ. 140 and 142. The Ray Society, London, U.K. Linnaeus, C. (1754). Genera Plantarum eorum Characteres Naturales Secundum Numerum, Figuram, Situm, et Proportionem Omnium Fructificationis Partium, ed. 5. Laurentius Salvius, Stockholm, Sweden. Facsimile published 1960. H.R. Engelmann (J. Cramer), Weinheim, Germany, and Wheldon & Wesley, Codicote, U.K. McNeill, J., F.R. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, R. Marhold, D.H. Nicholson, J. Prado, P.C. Silva, J.E. Skog, J.H. Wiersema, and N.J. Turland, eds. (2006). International Code of Botanical Nomenclature (Vienna Code). Regnum Veg. 146. Plukenet, L. (1691). Phytographia, sive, Stirpium Illustriorum & Minus Cognitarum Icones, Tabulis Aeneis Summa Diligentia Elaboratae, Quarum Unaquaeque Titulis, ex Notis suis Propriis, & Characteristicis Desumptis, Insignita ab aliis Sortis Facile Discriminatur. Pars Altera. Published by the author, London, U.K. Plukenet, L. (1696). Almagestum Botanicum sive Phytographiae Pluc'netianae Onomasticon Methodo Synthetici Digestum Exhibens Stirpium Exoticarum, Rariorum, Novarumque Nomina, quae Descriptiones Locum Supplere Possunt. Published by the author, London, U.K. (Gronovius's and Linnaeus's citations give the impression that "401" and "t. 328 f. 4" were in the same publication. In the original printings they were in two separate, independently titled works. Gronovius and Linnaeus probably saw the 1720 reprint of the combined works.) Ray, J. (1688). Historia Plantarum: Species Hactenus Editas Aliasque Insuper Multas Noviter Inventas & Descriptas Complectens. Tomus Secundus. Typis Mariae Clark, London, U.K. Reveal, J.L. (1983). Significance of pre-1753 botanical explorations in temperate North America on Linnaeus' first edition of Species Plantarum. Phytologia 53: 1-96. Stearn, W.T. (1957). An introduction to the Species Plantarum and cognate botanical works of Carl Linnaeus. In vol. 1 of 1957-1959 facsimile of Linnaeus (1753), q.v. Utech, F.H. (2002). Medeola Linnaeus. Pp. 150-151 in: Flora of North America Editorial Committee, eds. Flora of North America North of Mexico. Volume 26. Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford University Press, New York and Oxford.