Page  118 ï~~118 THE MICHIGAN BOTANIST Vol. 46 NOMENCLATURE OF GENTIANOPSIS CRINITA (GENTIANACEAE) James S. Pringle Royal Botanical Gardens P.O. Box 399, Hamilton, ON Canada L8N 3H8 ABSTRACT The name Gentiana crinita Froel. and its homotypic synonyms are neotypified by a specimen of the greater or wide-leaved fringed gentian of eastern North America, consistent with the long-established use of the specific epithet. The greater or wide-leaved fringed gentian, which is native from Georgia north to Maine and Manitoba, was known historically as Gentiana crinita Froel. and is now called Gentianopsis crinita (Froel.) Ma. From a study by Fernald (1923; recently noted by Jarvis 2007), however, it has sometimes been suspected that the epithet crinita might correctly be applicable, instead, to Victorin's gentian, which was originally described as Gentiana victorinii Fernald and is now called Gentianopsis virgata subsp. victorinii (Fernald) Lammers. The latter is endemic to the intertidal flats of the St. Lawrence estuary near Qudbec City. With research on typification having been stimulated by the Linnaean Plant Names Typification Project, the Flora of North America North of Mexico, and the compilation of databases on type specimens at major herbaria, uncertainty as to the correct name of G. crinita should be resolved, lest the nomenclature of this well-known and much-admired species be disrupted unnecessarily. Gentianopsis crinita, as Gentiana crinita, was the first North American species of the fringed gentians to be recognized taxonomically. Froelich (1796) distinguished this species from the European Gentiana ciliata L. [Gentianopsis ciliata (L.) Ma] by its greater plant size, basally clasping, lanceolate rather than linear leaves, larger and more numerous flowers, dimorphic calyx lobes, and corolla lobes fringed all around rather than only proximally. The description suffices to distinguish Gentianopsis crinita s. str. from all other North American taxa of Gentianopsis as well as from G. ciliata. Froelich doubtless based his description of Gentiana crinita on specimens, because no such detailed description is present in either of the two publications he cited. In the synonymy of G. crinita, he listed "Gentiana ciliata. LINN. syst. pl. 1. p. 645. n. 17. var. americana" [fonts and punctuation as in original]. If the name Gentiana ciliata var. americana L. had previously been validly published, Froelich's citation of it as part of the basis of his G. crinita would necessitate consideration the identity of the plant so named by Linnaeus. Froelich also cited "Gentiana auctumnalis [sic] secunda," number 50 in Colden's (1749-1751) list of native plants in the vicinity of Newburgh, New York, a work that antedates the

Page  119 ï~~2007 THE MICHIGAN BOTANIST 119 1753 starting-point for botanical nomenclature. Colden's description of Gentiana autumnalis secunda suffices only to identify the taxon as a species of Gentianopsis, but no species other than G. crinita s. str. occurs in the area where Colden botanized. Froelich's only reference to any publication by Linnaeus in connection with Gentiana crinita was in the citation quoted above. Systema Plantarum is a compendium of works by Linnaeus edited by Reichard (1779-1780). One of its components is Linnaeus's Species Plantarum, based on the second edition (Linnaeus 1761-1762) but with later additions. This is interpreted as the fourth edition of Species Plantarum, the third edition having been a "straightforward reprint" of the second, differing only in the insertion of errata (Stafleu and Cowan 1981, 1983). As the additions are not credited, it is not known which, if any, were written by Linnaeus while he was still active and which were written by Reichard or by others. In that work, on page 645, Gentiana ciliata is said to occur "in Helvetiae, Italiae, Germaniae, Austriae, Carnioliae, Canadae montibus" [italics in original]. A note on geographic variation, not present in the earlier editions of Species Plantarum, is added: "Haec corolla in America maxime ciliata, in Italia mediocriter; in Islandia & Norwegia tantum serrata" ("This [species] with the corolla ciliate to the highest degree in America, only moderately so in Italy, merely serrate in Iceland and Norway"). In the earlier editions no American or Canadian component of Gentiana ciliata sensu Linnaeus had been distinguished from the European, either morphologically or taxonomically. Notably in the present context, the phrase "maxime ciliata" is applicable to Gentianopsis crinita but not to G. virgata subsp. victorinii. This descriptive wording indicates that in this work, when the American component was distinguished morphologically for the first time, the distinction was based on G. crinita s. str. The treatment of Gentiana ciliata in Systema Plantarum does not include the adjective americana, either as the epithet of a variety or otherwise, nor was any name given to the American variant. Therefore it seems likely that Froelich intended "var. americana," which he placed after rather than before the bibliographic citation, as a phrase denoting in Latin "the American variant," rather than as a name (italics not used here in accord with this interpretation). The similarity in format of "var. americana" to a present-day varietal name appears to be coincidental. This interpretation is supported not only by the sequence of wording but also by the absence of a Greek letter. When Froelich recognized varieties taxonomically, as he did with several European species, he designated each variety with a Greek letter, followed by a phrase name in italics rather than a oneword varietal epithet. Don (1837) cited "Gentiana ciliata f3, Americana Lin. syst. 1 p. 645" in the synonymy of Gentianella crinita (Froel.) G. Don, but the addition of "f3" was Don's own. Grisebach (1845) listed Gentiana ciliata var. americana L. as a name in the synonymy of G. crinita, and MacMillan (1892) cited it as the basionym of the combination Gentiana americana (L.) MacMill. Both Grisebach and MacMillan attributed the authorship to Linnaeus, with the citation abridged to "Linn. syst. 1. p. 645." Fernald (1923) did not mention the supposed name Gentiana ciliata var. americana L. The only publication by Linnaeus cited by Fernald in this context was Species Plantarum, ed. 2 (Linnaeus 1761-1762). In that work, as in ed. 1

Page  120 ï~~120 THE MICHIGAN BOTANIST Vol. 46 (Linnaeus 1753), Linnaeus merely gave the range of G. ciliata as "in Helvetiae, Italiae, Canadae montibus" [italics in original]. He neither distinguished the Canadian plants by name nor contrasted them morphologically. Fernald concluded that Linnaeus's inclusion of Canada in the range given for Gentiana ciliata had been based on the taxon now called Gentianopsis virgata subsp. victorinii, of which he assumed Linnaeus had possessed a specimen, not now extant, collected by Pehr Kalm near Qudbec City. Linnaeus had received Kalm's collections in 1751, prior to the publication of the first edition of Species Plantarum (Steamrn 1957). Fernald's conclusion was based on the similarity of the Qudbec endemic to the European Gentiana (now Gentianopsis) ciliata. Specimens from L'Islet, Qudbec (Marie-Victorin 3183, GH), "laid upon sheets of G. ciliata" were, according to Fernald, "quite indistinguishable from it until the technical details [were] examined." He noted that the leaves of both G. ciliata and the Qudbec endemic are linear-lanceolate, whereas G. crinita "has the upper leaves ovate or ovate-lanceolate and broadly rounded or subcordate at base." Linnaeus, in Fernald's opinion, "would hardly have identified it with the narrowleaved G. ciliata." Actually, Linnaeus, as discussed below, appears to have done exactly that, as did, even more definitely, Reichard or some other author of addenda to Species Plantarum. The author of the addendum quoted above had observed a specimen or specimens of Gentianopsis crinita s. str., as indicated by his description of the distinctive corolla fringing, and nevertheless chose to retain the North American plants he thus described in Gentiana ciliata. With so few specimens having been availabile at the time, the taxonomic significance of the difference in leaf shape in this case likely remained unrecognized. Linnaeus's (1761) inclusion of "Canada" in the range given for Gentiana ciliata was another point raised by Fernald. He identified "Canada" of Linnaeus's time, at least in this context, as "presumably Quebec." As noted by Steamrn (1957, p. 144), however, "'Canada' in the Linnaean sense does not correspond to [present-day] Canada but to a region... partly in Canada, mostly in the United States, where Kalm did much collecting, i.e. roughly from Philadelphia and New York northward, by way of Albany, to Montreal and Quebec, and from Albany westward along the Mohawk River to Lake Ontario and Niagara Falls." Perhaps because of the emphasis on "Canada" in Kalm's mandate for his explorations (Jarrell 1979), Linnaeus seems sometimes uncritically to have attributed a Canadian provenance to specimens received from Kalm. For example, although he correctly stated that Kalm had collected Gentiana quinquefolia L. [Gentianella quinquefolia (L.) Small] in Pennsylvania, he said that Kalm had collected Chironia campanulata L. [Sabatia campanulata (L.) Torr.] in Canada. The latter species does not occur in Canada even as defined at that time, but could have been found by Kalm in New Jersey or along the Delaware River in easternmost Pennsylvania. Conversely, Linnaeus said that his specimen of Potentilla pensylvanica L., which is not native to Pennsylvania, was from Canada, but since he had already used the name P canadensis, it seems that when he had another species from "Canada" he chose an epithet that to him was essentially synonymous. Gentianopsis crinita is known from upland habitats in Pennsylvania and

Page  121 ï~~2007 THE MICHIGAN BOTANIST 121 New York, i.e. within "Canada" sensu Linnaeus, and was, as noted below, found in Pennsylvania by Kalm. Linnaeus, toward the end of his life, is known to have discarded or given away specimens from his herbarium that he considered superfluous, so the possibility that he once possessed a specimen of the taxon Gentianopsis virgata subsp. victorinii cannot be ruled out. Kalm was in the vicinity of Qudbec at the right season and mentioned seeing estuarine species such as Elymus arenarius L. [Leymus mollis (Trin.) Pilg.] and Plantago maritima L. (Kalm 1753-1761). He did not mention gentians in that area, although he did note having found "some species of Gentiana" in Pennsylvania. Whether Kalm attributed any fringed gentian to "Canada" in his manuscript flora of Canada is not known, as that manuscript, which was seen by Linnaeus, was subsequently destroyed in a fire (Benson 1964, introduction to translation of Kalm 1753-1761). The major component of Linnaeus's herbarium, now at LINN, contains only one specimen of a North American fringed gentian. (None are at S-LINN.) That specimen, LINN 328.38, represents Gentianopsis crinita s. str., and, according to its label, was collected by Kalm in Pennsylvania. It is likely that it was the basis for the contrast between the European and American representatives of Gentiana ciliata in Systema Plantarum by Reichard or whoever wrote that addition. According to Fernald (1923), "in his herbarium, as shown by memoranda made at various times by Asa Gray, Linnaeus had G[entiana] crinita set apart as a distinct species though under an unpublished name." It is highly unlikely that any specimens from Linnaeus's herbarium were removed after Gray's time, and the set of microfiches of its contents the images now available on the Internet (<>, accessioned 6 December 2007) are thorough, showing all of each sheet, yet the only specimen of Gentianopsis crinita therein is the one cited above. It may be that the abbreviation "pensylv." in an annotation by Linnaeus, interpreted by curators at LINN as part of "K[alm] sylvae pensylv.," designating the provenance, with "silvae" partly covered, was interpeted by Gray as a specific epithet that remained unpublished. In this interpretation, no unpublished name appears on that sheet. Where Froelich saw specimens of Gentianopsis crinita and by whom they had been collected is not known. He did not travel to North America, and his own herbarium is not extant. He cited no specimens of this species, and stated only that he had seen non-European species "in variis plantarum exsiccatarum collectionibus." His attribution of Gentiana crinita not only to New York and Pennsylvania but also to Carolina suggests that he saw specimens collected by John Bartram, who botanized in all of those regions and sent specimens to several botanists in Europe. In North Carolina, Gentianopsis crinita is restricted to the mountains in the westernmost part of the state, which had been visited by few botanical explorers other than Bartram prior to 1796 (Core 1970). (Andrd Michaux, who had explored that region, mentioned no such gentian.) Linnaeus's inclusion of "Canada" in the range given for Gentiana ciliata and Froelich's inclusion of Pennsylvania in the range given for G. crinita may have been based on the same specimen, the one cited above. In Froelich's time the main component of Linnaeus's herbarium was in England, in the care of J.E. Smith, and was

Page  122 ï~~122 THE MICHIGAN BOTANIST Vol. 46 available for study by visiting botanists (Stafleu 1971), so it is possible that Froelich saw the specimen collected by Kalm. From this study I conclude that Froelich based the name Gentiana crinita entirely and exclusively on the species now called Gentianopsis crinita. No evidence suggests that the name was based even in part on any other taxon, either through the specimens he examined or through the literature he cited. To prevent instability either in the use of the epithet crinita in Gentianopsis or in the specific epithet applied to the species now called Gentianopsis crinita, a type specimen should be designated. It is not known with certainty whether the extant Kalm specimen was part of the "original material" studied by Froelich. However, if that specimen is selected as the neotype, any future uncertainty will be limited to whether it should be considered a neotype or a lectotype; there will be no question as to what specimen can legitimately be designated the nomenclatural type. Fortunately, this specimen is in good condition, and it well and clearly represents G. crinita s. str. Gentiana crinita Froel., Gentiana 112. 1796. Gentianella crinita (Froel.) G.Don, Gen. Syst. 4: 179. "1838" [1837]; Gentianopsis crinita (Froel.) Ma, Acta Phytotax. Sin. 1: 15. 1951. Neotype, designated here: Specimen collected by P. Kalm in Pennsylvania, LINN 328.38. ACKNOWLEDGMENTS I am grateful to the library staff of the Missouri Botanical Garden for access to historic publications. This paper is Contribution No. 180 from the Royal Botanical Gardens, Hamilton, Ontario, Canada. LITERATURE CITED Colden, C. (1749-1751). Plantae Coldenghamiae in provincia Noveboracensi Americes sponte crescentes, quas ad methodum Cl. Linnaei Sexualem, anno 1742, &c. Acta Soc. Reg. Soc. Sci. Upsaliensis 4: 81-136; 5: 47-82. Core, E.L. (1970). The botanical exploration of the southern Appalachians. 1970. Virginia Polytechn. Inst. State Univ. Res. Div. Monogr. 2: 1-65. Don, G. ("1838" [1837-1838]). A General History of the Dichlamydeous Plants, Comprising Complete Descriptions of the Different Orders; Together with the Characters of the Genera and Species, and an Enumeration of the Cultivated Varieties, their Places of Growth, Time of Flowering, Mode of Culture, and Uses in Medicine and Domestic Economy, the Scientific Names Accentuated, their Etymologies Explained, and the Classes and Orders Illustrated by Engravings, and Preceded by Introductions to the Linnaean and Natural Systems, and a Glossary of the Terms Used. The Whole Arranged According to the Natural System. Vol. IV-Corolliflorae. J.G. and F. Rivington, London, U.K. [Gentianeae in part 1. 1837.] Fernald, M.L. (1923). The gentian of the tidal shores of the St. Lawrence. Rhodora 25: 85-89, plate 139. Froelich, J.A. (1796). De Gentiana Libellus Sistems Specierum Cognitarum Descriptiones cum Observationibus. W. Walther, Erlangen. Jarrell, R.A. (1979). Kalm, Pehr (baptised Petter). In: Dictionary of Canadian Biography. University of Toronto Press, Toronto. 4: 406-408. Jarvis, C. (2007). Order out of Chaos: Linnaean Plant Names and their Types. The Linnean Society of London, London, U.K. Kalm, P. (1753-1761). En Resa til Norra America [original title]. English translation of portions

Page  123 ï~~2007 THE MICHIGAN BOTANIST 123 dealing with North America published 1937 as Peter Kalm's Travels in North America: The English Version of 1770 Revised from the Original Swedish and Edited by Adolph B. Benson... with a Translation of New Material from Kalm's Diary Notes. Wilson-Erickson Inc., New York. Republished 1964. Dover Publications, New York. 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. Facsimile published 1957-1959 as Ray Soc. Publ. 140 and 142. The Ray Society, London. Linnaeus, C. (1761-1762). Species Plantarum Exhibitentes Plantas Rite Cognitas ad Genera Relatas, cum Differentiis Specificis, Nominibus Trivialibus, Synonymis Selectis, et Locis Natalibus, Secundum Systema Sexuale Digestas, ed. 2. Laurentius Salvius, Stockholm. Macmillan, C. (1892). Metaspermae of the Minnesota Valley: A List of the Higher Seed-Producing Plants of the Drainage Basin of the Minnesota River. Rep. Geol. Nat. Hist. Surv. Minnesota, Bot. Ser. 1. Reichard, J.J., ed. (1779-1780). Caroli Linnaei...Systema Plantarum Secundum Classes, Ordines, Genera, Species: Cum Characteribus, Differentiis, Nominibus Trivialibus, Synonymis Selectis, et Locis Natalibus, Editio Novissima. Varrentrapp Filius et Wenner, Frankfurt am Main. Stafleu, F.A. (1971). Linnaeus and the Linnaeans: The Spreading of their Ideas in Systematic Botany 1735-1789. Regnum Veg. 79. Stafleu, F.A., and R.S. Cowan (1981, 1983). Taxonomic Literature: A Selective Guide to Botanical Publications and Collections with Dates, Commentaries and Types, ed. 2. Vol. III: Lh-O. Regnum Veg. 105; Vol. IV: P-Sak. Regnum Veg. 110. 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.