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Notes on the nomenclature of species of Agave L. (Agavaceae / ) grown in and described from European gardens and nurseries Gideon F. Smith 1 and Estrela Figueiredo 2 1Office of the Chief Director: Biosystematics Research and Biodiversity Collections, South African National Biodiversity Institute, Private Bag X101, Pretoria 0001 South Africa / Acocks Chair, Schwe - ickerdt Herbarium, Department of Botany, University of Pretoria, Pretoria, 0002 South Africa / Cen - tre for Functional Ecology, Departamento de Ciências da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal (email: [email protected]). 2Department of Botany, P.O.Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth, 6031 South Africa / Centre for Functional Ecology, Departamento de Ciências da Vida, Universi - dade de Coimbra, 3001-455 Coimbra, Portugal (email: [email protected]).

Summary : Several species of Agave L. (Agavaceae Anwendung der bekannten Namen Agave ap - / Asparagaceae) have intricate nomenclatural his - planata Hort. ex K.Koch, A. verschaffeltii Lem. ex tories, given that material imported into Europe Jacobi und A. ghiesbreghtii Hort. ex Jacobi. Zusät - from their New World habitats took several years, zlich machen wir Bemerkungen über 12 weitere if not decades to flower, resulting in hasty and Artnamen, die von K. Koch verwendet wurden, in cryptic descriptions often being based on vegeta - einigen Fällen mit beschreibenden Bemerkungen tive material only. Furthermore, multiple authors in einem oft übersehenen Beitrag, den er 1862 sometimes, but not always, used the same bino - veröffentlichte. mials to refer to known in cultivation, mak - ing it important to clarify and establish correct The intricate nomenclature of Agave taxa author citations to determine which species are Many species of Agave L. have intricate referred to. In this paper we clarify the use of the nomenclatural histories (see Gentry, 1982 for sev - well-known names Agave applanata Hort. ex eral examples). Since the first representatives of K.Koch, A. verschaffeltii Lem. ex Jacobi and A. this New World genus were introduced to Europe, ghiesbreghtii Hort. ex Jacobi. In addition, notes these plants have been the subject of huge horti - are provided on 12 further species names that K. cultural interest (Ullrich, 1993). Because most Koch established and, where appropriate, fur - Agaves take several years to flower, and often will nished by him with descriptive statements in an not do so under the conditions in which they were often overlooked paper that he published in 1862. grown in Europe, numerous species were de - scribed based on vegetative material only. Most of Zusammenfassung : Mehrere Arten der Gattung these descriptions were devoid of other original Agave L. (Agavaceae / Asparagaceae) haben eine material (specimens or illustrations), which com - komplizierte nomenklatorische Vergangenheit, plicates their interpretation. Many of these names u.a. weil das aus den neuweltlichen Habitaten are now widely regarded as synonyms, while oth - nach Europa importierte Material bis zur Blüte ers are of unresolved application. For example, mehrere Jahre oder sogar Jahrzehnte benötigte, The List (www.theplantlist.org) records 246 weshalb oft übereilte und kryptische Beschrei - accepted names, 618 synonyms and 89 unresolved bungen allein auf der Basis von vegetativem Ma - names in the genus Agave . terial publiziert wurden. Zusätzlich verwendeten Different authors sometimes, but not always, mehrere Autoren manchmal, aber nicht immer, used the same binomials to refer to plants known dieselben binomialen Namen für aus Kultur in cultivation, making it important to clarify and bekannte Pflanzen, weshalb es wichtig ist, die ko - determine correct author citations to decide which rrekten Autorzitate zu klären und festzuhalten species are referred to. Here we provide clarity on um Klarheit zu schaffen, welche Arten jeweils the author citations of the well-known Agave gemeint waren. In diesem Beitrag klären wir die names Agave applanata Hort. ex K.Koch, Agave

Bradleya 31/2013 53 verschaffeltii Lem. ex Jacobi [a synonym of Agave culture, from ‘Hortus’ or ‘Garden’) in their author potatorum Zucc.] and Agave ghiesbreghtii Hort. ex citation. Agave ellemeetiana Hort. ex Jacobi is one Jacobi, along with notes on 12 further species such example. For comparison, only seven names names that Koch (1862a) established in a little of Aloe are known to have that origin. Further - known publication. more, about 6% of the Agave names encountered in the literature were published in plant cata - Names without description logues. These catalogues were produced by nurs - In the genus Agave many names were pub - eries for trade purposes, and often included very lished without a description (a written statement short statements of plant features. According to of a feature or features of a taxon) or diagnosis (a Art. 30.6 of the ICN, names are accepted as written statement of that which distinguishes the validly published in trade catalogues up to 31 De - taxon from other taxa) of the plant referred to. cember 1952. Catalogues in which Agave names These are nomina nuda (nomen nudum = naked were published originated from often well-known name) and not validly published according to Art. and reputable horticultural companies or nurs - 38.1 of the International Code of Nomenclature erymen who typically operated from continental for algae, fungi, and plants (ICN) (McNeill et al ., Europe, and included Van Houtte, Verschaffelt, 2012). When they are later validated with a de - De Smet, Haage & Schmidt, Cels and Besaucèle. scription, this is indicated in the authority of the These catalogues are today extremely rare and as name by adding ‘ex’ between the name of the per - far as we could ascertain, not available online, son who first referred to it, and the name of the which further complicates the assessment of the author who validated it. Quite remarkably, at nomenclatural status of the names provided in least 10% of the legitimate, validly published them. nomenclatural novelties in Agave have been vali - dated by another, later author. For comparison, The case of Agave applanata Hort. ex the figure for the Old World rosulate leaf-succu - K.Koch lent genus Aloe L. is only 1.5% (Figueiredo & In Agave, many names were used in the horti - Smith, 2010). These figures are only estimates be - cultural trade long before they were validly pub - cause according to the ICN the name of the person lished. These names were usually adopted by the preceding ‘ex’ (i.e. the person who originally pro - authors who later described the plants. In some vided the name, but did not give a description) cases, their etymology remains unknown. An ex - and the word ‘ex’, can be omitted from the author ample is Agave warelliana Hort. ex Baker for citation. which the origin of the name has not been When a description of the plant is provided, recorded. Like these agave names, the name there are also requirements that will apply for its Agave applanata existed in the horticultural trade acceptance as valid. For instance, according to for several years before the name was validated. Art. 38.3 of the ICN, “...statements describing The name was used for one of the many Mexican properties such as purely aesthetic features, eco - agaves introduced to Europe through nurseries in nomic, medicinal or culinary usage, cultural sig - the 19th century. The authority of Agave ap - nificance, cultivation techniques, geographical planata has been variously attributed to ‘K.Koch’, origin, or geological age...” do not meet the re - ‘K.Koch ex Jacobi’ or ‘Lem. ex Jacobi’. quirements. This is particularly relevant for Karl Koch (1809–1879) appears to be the first Agave where many names have been accompanied author to have provided a description of Agave ap - with brief statements focusing on the beauty or planata (1862a: 198) in a paper where he de - oddity of the plants. The validity of these descrip - scribed several species from the garden of Tonel tive statements is debatable. In Art. 38.4, the ICN in Belgium. Jean Baptiste and Constant Tonel also addresses these situations, stating that had a well-known garden near Ghent, a centre for “When it is doubtful whether a descriptive state - Agave horticulture in the 19th century. Jean ment satisfies the requirement of Art. 38.1(a) for Tonel travelled to Mexico where he eventually ob - a “description or diagnosis”, a request for a deci - tained a wing of a Franciscan monastery at Mex - sion may be submitted to the General Commit - ico City that he used for growing material that he tee…”. imported from Belgium to Mexico (Loriaux, 1993). This included horticultural and floricultural ma - Names of horticultural origin terial popular at the time, such as roses. He regu - About 10% of the validly published nomen - larly returned to Ghent where he also had a clatural novelties in Agave are of horticultural ori - garden with Mexican plants, particularly succu - gin. Over 70 names of Agave have the word ‘Hort.’ lents. Koch’s description of Agave applanata from (an abbreviation that indicates their use in horti - Tonel’s garden was very short (Figure 1), but it

54 Bradleya 31/2013 Figure 2. The statement on Agave ghiesbreghtii extracted from Koch (1862b).

Figure 1. The valid description of Agave ap - planata extracted from Koch (1862a). was accepted by some later authors as validating to A.Versch.). When Lemaire eventually described the name (Berger, 1915; Trelease, 1920). A. verschaffeltii (Lemaire, 1868: t. 564) it was too However, both Gentry (1982) and Thiede late as the name had already been validated by (2001) considered this description as insufficient Jacobi (1865: 122). Therefore, the correct author - and cited Jacobi as the validating author of the ity for the name is A.verschaffeltii Lem. ex Jacobi, name mentioned by Koch (as Agave applanata and not A.verschaffeltii Lem. K.Koch ex Jacobi). In our view the original de - In the case of A. ghiesbreghtii , no description scription provided by Koch is certainly sufficiently by Lemaire or Verschaffelt could be found in the diagnostic to regard Koch as the only validating literature. The name is recorded in IPNI as author of the name. validly published by Koch in “Wochenschr. v. In 1864 in his extensive work on the genus (1862) 83, 143”, and this publication was accepted Agave , Georg Albano von Jacobi (1805–1874) com - by some authors (Trelease, 1914, 1920; Berger, mented that he was adopting the name Agave ap - 1915; Breitung, 1968). We could find no reference planata already in use in horticulture, because he to the name on page 143 while on page 83 only a thought a new name would increase nomenclat - short statement refers to it (Figure 2). Recent au - ural complexity. However, he did not mention thors (Govaerts et al ., 2011) considered this state - Koch. Instead he cited Lemaire as the author of ment to be insufficient and listed the name as the name. For this reason, recent authors (e.g. Go - validated two years later by Jacobi (1864: 545), vaerts et al ., 2011) cite the name as Agave ap - who provided a more comprehensive description. planata Lem. ex Jacobi. Jacobi (1864) further Both Koch (1862b) and Jacobi (1864) spelled the noted that his species was the same as the one epithet as ‘ ghiesbrechtii ’, but the correct spelling grown in Tonel’s garden in Ghent (which was the is ‘ ghiesbreghtii ’, as it commemorates Auguste one described by Koch). The name should there - Boniface Ghiesbreght (1810–1893). Article 60.1 in fore be cited as Agave applanata Hort. ex K.Koch. the ICN stipulates that orthographical errors in names or epithets can be corrected. Ghiesbreght The cases of Agave verschaffeltii and A. was a plant collector who introduced many Mexi - ghiesbreghtii can agaves into Europe. He went to Mexico in Charles Antoine Lemaire (1800–1871) was the 1836 as a collector, and eventually settled there. originator of several Agave names that he neg - We conclude that the names should be cited as lected to describe and that were eventually vali - Agave verschaffeltii Lem. ex Jacobi and A. ghies - dated by other authors. In 1864 he commented breghtii Hort. ex Jacobi. that he had examined and named several species from the collections of Tonel, Ambroise Verschaf - The names of other Agaves in Tonel’s garden felt (1825–1886) and Jean Verschaffelt (?–1884), In addition to Agave applanata , Koch (1862a) the latter two being associated with the famous – an often overlooked paper – included descriptive Verschaffelt family nurseries. Plants with which statements for 12 further species that he obtained these names are associated were exhibited at hor - from Tonel’s garden, as follows: ticultural exhibitions in Belgium, especially at Ghent, and later more widespread, without hav - 1. Agave montezumae Hort. ex K.Koch in Wochen - ing been adequately described by him. For exam - schr. Vereines Beförd. Gartenbaues Königl. ple, in 1864 Lemaire mentioned two names to be Preuss. Staaten 5: 197 (1862). described soon (Lemaire, 1864: 66): Agave ver - Probably a synonym of Agave salmiana Otto schaffeltii and A. ghiesbreghtii (that he ascribed ex Salm-Dyck. Although ‘ Agave montezumae Hort.

Bradleya 31/2013 55 ex Jacobi (1864)’ was cited by Thiede (2001), this 9. Agave coccinea Roezl ex K.Koch in Wochenschr. citation apparently reflects the listing of A. mon - Vereines Beförd. Gartenbaues Königl. Preuss. tezumae as a possible synonym of A. jacobiana by Staaten 5: 198 (1862). Jacobi (1864: 557). There was no validation of the Probably a synonym of Agave atrovirens Karw. name by Jacobi (1864). ex Salm-Dyck. This publication yields as illegiti - mate Agave coccinea Roezl ex Jacobi (1865). Ja - 2. Agave ferdinand-cortez Hort. ex K.Koch in cobi’s name is considered a synonym of Agave Wochenschr. Vereines Beförd. Gartenbaues atrovirens Karw. ex Salm-Dyck. Königl. Preuss. Staaten 5: 198 (1862). Probably a synonym of Agave salmiana Otto 10. Agave serrata Hort. ex K.Koch in Wochenschr. ex Salm-Dyck. Jacobi also saw plants under this Vereines Beförd. Gartenbaues Königl. Preuss. name at Tonel’s garden, and listed it as a possible Staaten 5: 198 (1862). synonym of A. jacobiana Salm-Dyck (Jacobi, 1864: A name of unresolved application. 557). 11. Agave taeniata Hort. ex K.Koch in Wochen - [3. Agave concinna Lem.] schr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 5: 199 (1862). [4. Agave vivipara L.] A name of unresolved application. Probably the same as ‘ Agave toeniata Hort.’ that was cited 5. Agave oblongata Hort. ex K.Koch in Wochen - by Berger (1915: 94) as used by Besaucèle, with schr. Vereines Beförd. Gartenbaues Königl. Thiede (2001) considering it as of unresolved ap - Preuss. Staaten 5: 198 (1862). plication. Probably a synonym of Agave mitis Mart. This publication yields as illegitimate the later valida - 12. Agave noacksii Hort. ex K.Koch in Wochen - tion of the name for the plants from Tonel’s gar - schr. Vereines Beförd. Gartenbaues Königl. den by Jacobi, as Agave oblongata Hort. ex Jacobi Preuss. Staaten 5: 199 (1862). (1869). Jacobi’s name is considered a synonym of Probably a synonym of Agave pendula Agave mitis Mart. Schnittsp. It could be an orthographical variant of A. noackii Jacobi (1865), which is considered a 6. Agave toneliana Hort. ex K.Koch in Wochen - synonym of Agave pendula , making it illegitimate. schr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 5: 198 (1862). [13. Agave schidigera Lem.] A name of unresolved application. This publi - cation yields as illegitimate Agave toneliana Hort. Conclusion ex Baker. Baker (1881) noted that the name was The common 19th century practice of describ - in use but he did not know if it had been published ing nomenclatural novelties in nursery catalogues before. Baker’s name is of unresolved application. and popular journals with brief statements (that are often acceptable for valid publication of plant 7. Agave applanata Hort. ex K.Koch in Wochen - names) has contributed to the confusion of names schr. Vereines Beförd. Gartenbaues Königl. in some taxa, including Agave . As we argue above, Preuss. Staaten 5: 198 (1862). the names established by Koch in his 1862 paper An accepted name. This publication yields as on Tonel’s plants are validly published and their illegitimate Agave applanata Lem. ex Jacobi authorities should be cited as indicated above. (1865) which was described from the same plants from Tonel’s garden. References BAKER , J.G. (1881). New garden plants. Gard. 8. Agave malinczii Hort. ex K.Koch in Wochen - Chron.,n.s. 15 : 362. schr. Vereines Beförd. Gartenbaues Königl. BERGER , A. (1915, reprinted 1988). Die Agaven. Preuss. Staaten 5: 198 (1862). Beiträge zu einer Monographie. Gustav Fis - A name of unresolved application. It is listed cher Verlag, Jena. as valid in IPNI (misspelled ‘ malinezii ’). Curi - BREITUNG , A.J. (1968). The Agaves . In: GLASS , C. & ously, it also has a very short description, but in FOSTER , R.A. ( EDS .) . The Cactus and Succulent the recent literature (Govaerts et al ., 2011), A. ma - Journal 1968 Yearbook. Abbey Garden Press, linczii is listed as a validly published name and a Reseda, California. synonym of Agave rudis Lem. ex Jacobi, itself an unresolved name.

56 Bradleya 31/2013 FIGUEIREDO , E. & S MITH , G.F. (2010). What’s in a LEMAIRE , C. (1864). Observations sur le genre name: epithets in Aloe L. (Asphodelaceae) and Agave et description de quelques espèces nou - what to call the next new species. Bradleya 28 : velles. Illust. Hort . 11 : 59–66. 79–102. LEMAIRE , C. (1868). Agave verschaffeltii . Illust. GENTRY , H.S. (1982). Agaves of Continental North Hort . 15 : t. 564. America . The University of Arizona Press, LORIAUX , F. (1993). De Belgen en Mexico . Leuven Tucson. University Press, Leuven. GOVAERTS , R., Z ONNEVELD , B.J.M. & Z ONA , S.A. MCNEILL , J., B ARRIE , F.R., B UCK , W.R., D EMOULIN , (2011). World Checklist of Asparagaceae. Fa - V., G REUTER , W., H AWKSWORTH , D.L., H EREN - cilitated by the Royal Botanic Gardens, Kew. DEEN , P.S., K NAPP , S., M ARHOLD , K., P RADO , J., Published on the Internet; PRUD ’HOMME VAN REINE , W.F., S MITH , G.F., http://apps.kew.org/wcsp/ Accessed 22.1.2011. WIERSEMA , J.H. & T URLAND , N.J. ( EDS .) (2012). JACOBI , G.A. VON (1864). Bersuch zu einer sys - International Code of Nomenclature for algae, tematischen Ordnung der Agaveen. Hamb. fungi, and plants (Melbourne Code) adopted by Gart. Blumenz . 20 : 499–515. the Eighteenth International Botanical Con - JACOBI , G.A. VON (1865). Bersuch zu einer sys - gress Melbourne, Australia, July 2011 . Koeltz tematischen Ordnung der Agaveen. Hamb. Scientific Books, Königstein. [ Regnum Veg - Gart. Blumenz . 21 : 114–126. etabile 154 ]. JACOBI , G.A. VON (1869). Nachträge zu dem ver - THIEDE , J. (2001). Agave . In: EGGLI , U. ( ED .) Illus - such einer systematischen Ordnung der trated handbook of succulent plants. Mono - Agaveen. Abh. Schles. Ges. Vaterl. Cult., Abth. cotyledons. Springer-Verlag, Berlin, pp. 6–76. Naturw. Med . 1868: 138–176. TRELEASE , W. (1914). Agave . In: BAILEY , L.H. ( ED .) KOCH , K. (1862a). Einige Worte über der The standard cyclopedia of horticulture 1: Tonel’schen Agaveen in Gent. Wochenschr. 230–239. MacMillan, New York. Vereines Beförd. Gartenbaues Königl. Preuss. TRELEASE , W. (1920). Agave . In: STANDLEY , P.C. Staaten 5: 197–199. (ED .) Trees and Shrubs of Mexico . Contr. U. S. KOCH , K. (1862b). Die grosse Pflanzen- und Blu - Nat. Herb . 23 : 107–142. men-Ausstellung in Gent vom 2., 3. und 4. ULLRICH , B. (1993). Agaven. Illustrationen März. Wochenschr. Vereines Beförd . Garten - blühender Exemplare bis 1800. Palmengarten baues Königl. Preuss. Staaten 5: 81–87. 21 : 1–83.

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