Bradleya 32/2014 pages 146–163

A review of Agave ellemeetiana K.Koch (Asparagaceae/Agavaceae) Joachim Thiede Schenefelder Holt 3, D-22589 Hamburg, (email: [email protected])

Summary : A broad overview of the history, publi - ever, besides the discovery of novelties, several cation, author, introduction, rediscovery, distri - long-known species of unknown origin in the wild bution, habitats, typification, and relationships of were rediscovered in habitat such as A. albomar - Agave ellemeetiana K.Koch is given. The species ginata Gentry (Magallán & Hernández, 2000) and was first published by K.Koch in 1865 from culti - A. warelliana Baker (Lott & Garcia-Mendoza, vated plants of unknown origin and was rediscov - 1994; Cházaro et al ., 2008). The most notable ered in habitat only in the last couple of years in among the rediscovered species is certainly A. elle - Central Veracruz (Sierra de Zongolica), northern meetiana K.Koch, a species of subg. Littaea with Oaxaca (Sierra Mazateca and near San Bartolomé nearly stemless rosettes of rather few, broad, flat, Ayautla), and in western Oaxaca (Putla/Tlaxiaco soft and unarmed shiny green leaves with a soft region). Plants from western Oaxaca differ con - terminal spine (Figures 1–4, 6–14). Since its pub - sistently in their finely denticulate leaf margins lication in 1865, A. ellemeetiana was known from and are distinguished as Agave ellemeetiana cultivated plants of unknown origin in the wild subsp. subdentata (Trelease) Thiede comb. nov. and was not rediscovered in habitat until the last from the plants in Veracruz and northern Oaxaca couple of years (Köhres, 2008; Cházaro et al ., with entire, smooth leaf margins (= Agave elle - 2012). Albeit published nearly 150 years ago, the meetiana subsp. ellemeetiana ). history, publication, author and typification of the species and its introduction, rediscovery, distri - Zusammenfassung : Es wird eine breitgefaßte bution, habitats and relationships remained in - Übersicht über die Geschichte, Veröffentlichung, completely known. This paper aims to fill these Autorschaft, Einführung, Wiederentdeckung, Ver - gaps of knowledge. breitung, Standorte, Typifikation und Ver - wandtschaft von Agave ellemeetiana K.Koch 1. History and publication gegeben. Die Art wurde zuerst von K.Koch 1865 The history of Agave ellemeetiana is connected nach kultivierten Pflanzen unbekannter Herkunft with three people: Professor Karl Heinrich Emil veröffentlicht und erst in den letzten Jahren an Koch (1809-1879; see Anon., 1879; Wimmer, Naturstandorten in Zentral-Veracruz (Sierra de 2004), horticultural botanist at the Botanical Gar - Zongolica), in Nord-Oaxaca (Sierra Mazateca und den Berlin and author of nearly 2100 new plant bei San Bartolomé Ayautla) und in West-Oaxaca names including some 20 new names in Agave , (in der Region Putla/Tlaxiaco) wieder entdeckt. the Prussian general Georg Albano von Jacobi Pflanzen aus West-Oaxaca unterscheiden sich (1805–1874, see Gutte, 2006), renowned author of konstant durch ihren fein gezähnelten Blattrand an 11-volume treatise on European field artillery, und werden als Agave ellemeetiana subsp. sub - owner of a large agave-collection, and author of dentata (Trelease) Thiede comb. nov. von den about 110 new names in Agave , and the eponymee Pflanzen in Veracruz und Nord-Oaxaca mit of the species, Jonkheer Willem Cornelis Mary de ganzrandigem, ungezähneltem Blattrand (= Agave Jonge van Ellemeet (1811-1888, see Koch, 1864b; ellemeetiana subsp. ellemeetiana ) unterschieden. Alsemgeest, 2004), a Dutch landowner, mayor and keeper of an important succulent collection on his Introduction estate “Overduin” near Oostkapelle (The Nether - The systematics of the genus Agave L. are in a lands) which in 1856 comprised 200 names continuous flux and receive attention by many au - with emphasis on Mammillaria Haw. (Janse, thors: since the taxonomic synopsis by Thiede in 1977), but later Ellemeet specialized on Agave and 2001, some 30 new species were published, in - its relatives. cluding the most remarkable Agave albopilosa I. The first appearance of A. ellemeetiana in Cabral, Villarreal & A.E. Estrada (2007). How - printed matter as a mere name without descrip -

146 Bradleya 32/2014 Figure 1. Agave ellemeetiana (subsp. ellemeetiana ). Part-coloured lithograph from Saunders' Refugium Botanicum (plate 163, Baker 1869) from a plant flowering in the greenhouse of W.W. Saunders, with an inflorescence 4.25m (14 ft.) in height. This plate represents the first illustration of a flowering Agave ellemeetiana and is reproduced here in colour for the first time (courtesy of G.D. Rowley). Photograph by Jonathan Clark

Bradleya 32/2014 147 as ” Wochenschr .” in the references) do not men - tion any Agave. Thus, Koch may have seen the plant in Paris in 1864 together with Jacobi, but the first appearance of the name A. ellemeetiana in print was two years earlier in October 1862 by Koch, as mentioned above. During the 27th to 29th April 1864, the collec - tion of the Belgian agave-collector van der Vinnen was auctioned in Brussels, Belgium (Vermeulen, 1864; Koch, 1864a), and Ellemeet purchased an especially large specimen of A. ellemeetiana . Shortly thereafter, Ellemeet donated the plant to the Berlin for further study where it was kept from no later than 29th May 1864 onwards (Koch, 1864c). Based on the van der Vinnen plant at Berlin, Koch (1865a) validly pub - lished A. ellemeetiana as a new species on 1st April 1865 in his “ Wochenschrift ”. Just one week later, from 7th to 12th April 1865, Koch attended the ”Exposition Universelle d’Horticulture” at Amsterdam (NL) where he, as his report shows (Koch, 1865b), had intensive con - tacts with Ellemeet. Most certainly, Koch pre - sented a reprint of his publication of A. ellemeetiana to Ellemeet, albeit no written record could be found. Eduard August von Regel (1815- 1892), botanist and later director of the Imperial Botanical Garden of St. Petersburg (Russia), also attended the Amsterdam exposition and men - tioned in his report (Regel, 1865) that Ellemeet re - ceived an award for his agaves. The specimen of A. Figure 2. Agave ellemeetiana (subsp. ellemeetiana ). ellemeetiana exhibited at Amsterdam was most Hand-coloured copper-engraving from Curtis's Botani - probably Ellemeet’s plant, but doubtless not the cal Magazine (plate 7027, Baker 1888b) from a plant specimen from the van der Vinnen auction which flowering at Kew Gardens. was kept in Berlin from no later than 29th May (courtesy of The Biodiversity Heritage Fund). 1864 onwards (Koch, 1864c, see above). M. Witte (1865), chief gardener at BG Leiden (The Nether - tion was in October 1862 when Koch (1862) re - lands) published a report on a visit to Ellemeet's ported on a plant exhibition in June 1862 in Mid - estate where he saw an “admirable plant” of A. delburg (The Netherlands) close to Ellemeet’s ellemeetiana with a diameter of 2m (!) and a leaf estate where the latter exhibited his plants, in - size of 90 × 20cm “throning in the centre” of Elle - cluding “his” A. ellemeetiana . During that occa - meet’s agave collection. sion, Koch also visited Ellemeet’s collection with a Six month after Koch’s publication, Jacobi specimen of the ”absolutely rare A. ellemeetiana ” (1865) independently published A. ellemeetiana (Koch, 1864b). based on the same specimen in Berlin in the Oc - Jacobi (1865) mentioned that A. ellemeetiana tober-issue of the “ Hamburger Garten- und Blu - received its name in Paris and lists the 'author menzeitung ” with the 'author name' ”Hort. Paris” name' “Hort. Paris” for it. In 1873 (reproduced in and without the additive ' Nob .' (nobis, Latin = us) Alsemgeest, 2004), Ellemeet mentioned that Koch which he used for the species he intended to de - and Jacobi found the rare agave named after him scribe as new (Thiede, 2014: 296). Albeit Koch al - in Paris. Koch made his first visit to Paris not ready noted in 1871 that his publication in until August to September 1864 (Wimmer, 2004), “Wochenschr . 7: 103, 1865” antedates that of Ja - but his reports in the journal which he edited cobi, most later works wrongly cited Jacobi as au - (“ Wochenschrift des Vereines zur Beförderung des thor (e.g. Baker, 1888a; Berger, 1915; Gentry, Gartenbaues in den Königlich Preussischen 1982; Thiede, 2001; Govaerts, 2014); only a few Staaten für Gärtnerei und Pflanzenkunde ”; ab - works correctly cited Koch (e.g., IPNI, 2014). breviated as ” Wochenschrift ” in the following and Regel (1865), in his report on the Amsterdam ex -

148 Bradleya 32/2014 Figure 4. Agave ellemeetiana (subsp. ellemeetiana ) flow - ering in cultivation at Rotterdam Zoo in 1922, with an inflorescence more than 4m in height (from van den Houten 1930).

(Anon., 1873a, 1873b; Alsemgeest, 2004). The auc - Figure 3. Agave ellemeetiana (subsp. ellemeetiana ) flow - tion was completely unsuccessful, and Ellemeet ering in cultivation at Kew, showing an inflorescence presented his whole collection to the Zoological about 4.50m high and with about 3150 flowers (from Anonymous 1910; courtesy of The Biodiversity Heritage Gardens at Rotterdam (The Netherlands) in 1874 Fund). and 1875 (Krelage, 1885). In 1885, Ellemeet's agaves at Rotterdam still represented the best position, provided a short description of A. elle - agave collection in the Netherlands (Krelage, meetiana (“Grosse, hellgrüne, länglich-ovale, 1885). Several plants at Rotterdam Zoo possibly ganzrandige Blätter, die in einen Krautstachel going back to Ellemeet were depicted by van den ausgehen”) which must be considered to represent Houten in 1930, and some agaves going back to a further, albeit unintended, publication of that Ellemeet were still in cultivation in 2004 name unrecognized by previous authors. Regel’s (Alsemgeest, 2004), 130 years after Ellemeet’s col - publication appeared in the August issue of the lection was disintegrated. In 2011, no plant from Gartenflora 1865, thus antedating Jacobi’s publi - Ellemeet was still present at Rotterdam, but a few cation from October 1865. were still kept at Amsterdam (Alsemgeest, pers. In 1871, Ellemeet’s agave collection comprised comm . 2011). 137 different “names”, including 4 rather large specimens of “his” A. ellemeetiana measuring up 2. Introduction into cultivation to 1.55m in diameter (Ellemeet, 1871; reproduced Europe. by Morren, 1871; André, 1872). However, many of Agave ellemeetiana has most certainly been these species names are now placed in the syn - continuously in cultivation in Europe from not onymy of long-known variable species such as A. later than 1862 onwards, albeit it was (and still mitis Mart., A. americana L., A. salmiana L., A. is) uncommon. The first record from the UK dates angustifolia Haw. and A. lophantha Schiede ex from 1865 when Jacobi (1867) saw a living plant Kunth (Alsemgeest, 2004). About two years later, in the collection of the British insurance broker, Ellemeet’s whole collection of 323 specimens was entomologist and botanist William Wilson Saun - auctioned on his estate on 22nd. September 1873 ders (1809-1879); the plant came into flower two

Bradleya 32/2014 149 years later (see below). The British agave collector Breitung (1961) obtained plants from the Jardin W. B. Kellock of Stamford Hill exhibited a speci - Botanique “Les Cèdres” (France), wrongly stating men in May 1872 (Anon., 1872). At Kew Gardens, that this appeared to be a new introduction to the nine plants came into flower between 1874 and Southwestern USA. All these records invalidate 1949 (see below; Figure 2 & 3). The species was Gentry’s (1982: 97) statement ”that there is no seen at Kew by Howard S. Gentry (1903-1993, see record of A. ellemeetiana in North American gar - Ullrich 1993), the intrepid explorer and monogra - dens”. At present, A. ellemeetiana is rare in culti - pher of the genus in Continental North America, vation and is kept mainly in specialist collections in 1969 (Gentry, 1982: 96) and in 1975 by Janse and botanical gardens in the USA and Europe; (1977) and is still in cultivation there (Royal seeds or plants are commercially offered by sev - Botanic Gardens Kew, 2014), possibly uninter - eral nurseries. ruptedly from not later than 1874 onwards. At Hanbury Gardens in La Mórtola (; see Asia . Russo, 2001; Smith & Figueiredo, 2014a), A. elle - Agave ellemeetiana is (or was) cultivated at meetiana was cultivated no later than 1889 Bogor (Indonesia; Raja, 1963) and on the Philip - (Cronemeyer, 1889) and was still seen there in pines (Madulid, 1995). 1993 (M. Greulich, pers. comm .). John Gilbert Baker (1834-1920), botanist at Kew, keeper of the 3. Flowering specimens, plates and photo - herbarium, and monographer of agave and other graphs monocotyledons, studied a plant at Hanbury in The first flowering specimen: In June 1867, 1891 (Baker, 1892). Alwin Berger (1871-1931, see Jacobi when just in the field on a military cam - Schröder, 2011), curator at Hanbury from 1897 to paign, received a message from W. W. Saunders 1914, listed A. ellemeetiana in his “ Hortus Mor - that the latter's A. ellemeetiana was in flower with tolensis ” (Berger, 1912: 11) and depicted a flower - an inflorescence of nearly 4m (13 ft.) in height and ing plant at Hanbury in his agave monograph the upper 2.75m (9 ft.) densely beset with flowers (Berger, 1915: 125). Gentry (1982: 97) did not find (Jacobi, 1867). Jacobi saw that plant himself in A. ellemeetiana at Hanbury during his visit in autumn 1865 and received pressed flowers from 1955. At the Jardin Botanique “Les Cèdres” at St. Saunders and from which he provided the first, Jean-Cap-Ferrat (France), the species was culti - very detailed description of the flowers of A. elle - vated from no later than 1961 (Breitung, 1961, see meetiana (Jacobi, 1867). Saunders had received below) and was still present in 1990 (Kolendo, the plant from the Royal Horticultural Society 1990). According to SYSTAX (2014), the species is (RHS) at Chiswick. Baker (1869) published a at present cultivated in Germany in the botanical partly coloured plate of the same plant in Saun - gardens at Kiel and Ulm, but it was also seen by ders’ “ Refugium Botanicum ” showing the inflo - the author at BG Bochum in 2012. rescence of 4.25m (14 ft.) in height. This plate, reproduced here as Figure 1, represents the very USA. first illustration of a flowering specimen of A. elle - The earliest record for the cultivation of A. elle - meetiana . Eight years later, Baker (1877) pro - meetiana in the USA is found in a letter from ca. vided the first complete description of A. February 1885 by James Gurney (1831-1920), ellemeetiana including the inflorescence and flow - head gardener at Missouri Botanical Garden (St. ers accompanied by a black and white woodcut of Louis) since 1867 (Knotts & Pagels, 2014). Gur - a non-flowering plant (the plate was reproduced ney (1885) lists the agaves cultivated at Missouri, by Alsemgeest, 2004). among them A. ellemeetiana . In 1912, A. ellemee - Kew Gardens: At Kew, A. ellemeetiana flow - tiana was still kept at Missouri, as is indicated by ered in May 1874 (Anon. 1874), in February to a herbarium specimen made in spring 1912 by March 1877 (Anon., 1877a, 1877b) and in June Charles Henry Thompson (1870-1931), Assistant 1883 (Anon., 1883a, 1883b). From the specimen Botanist and keeper of the succulent collection flowering in 1877, four herbarium sheets were (Tropicos, 2014). The US Department of Agricul - prepared, three of which were much later selected ture imported A. ellemeetiana seeds from the as neotype by Gentry (1982: 96; see below). A fur - nursery Vilmorin-Andrieux & Co. (Paris, France) ther plant came into flower in 1888 at Kew and in 1923 (USDA, 1925) and in 1930 (USDA, 1930). was illustrated with a fine hand-colored copper In 1931, A. ellemeetiana was commercially offered engraving in “ Curtis’ Botanical Magazine ” (repro - by the nursery of Edward O. Orpet at Santa Bar - duced here as Figure 2; Baker, 1888b), noting that bara, California (Manning, 1931), and in 1933 it the species was introduced about 1864. This date was cultivated at Balboa Park in San Diego was repeated by later authors, but, as noted (Greer, 1933). The US agave collector August J. above, A. ellemeetiana is known for sure from no

150 Bradleya 32/2014 Figure 6. Agave ellemeetiana subsp. ellemeetiana in habitat on calcareous rock faces in the Sierra de Zon - golica near Totomachapa (central Veracruz). The larger rosettes with pendent dried leaves appear to be the bromeliad Tillandsia grandis which co-occurs with Figure 5. Distribution of Agave ellemeetiana in the Agave ellemeetiana in the Sierra de Zongolica (Wrinkle Mexican states of Veracruz and Oaxaca, and towns re - 2011) and also at Ayautla (Figure 10) ferred to in the text. Subsp. ellemeetiana : triangles, 15th January 2013, photo: Michael Greulich. subsp. subdentata : circles; filled symbols = herbarium specimens, open symbols = literature and photographic flowering at Kew (Figure 3) and at Rotterdam records. Symbols may represent more than one record. (Figure 4) both represent the most impressive (map: Wikimedia Commons 2014). photographs of A. ellemeetiana thus far published. later than 1862. At Kew, A. ellemeetiana flowered 4. Rediscovery, distribution and habitats again in 1910 (inflorescence about 4.5m high and All early agave authors (Koch, Jacobi, Baker, with about 3150 flowers; Anon., 1910 with a black Trelease) up to modern monographers (Berger, and white photograph reproduced here as Figure Gentry) knew A. ellemeetiana from cultivated 3), in 1918 (with black and white photograph, plants which may all only go back to the first in - Anon., 1918), in 1926 (with several lateral inflo - troduction no later than 1862. During the last cou - rescences below the lowest leaves which attained ple of years, localities in central Veracruz and in a height of nearly one meter and bore numerous three different areas in Oaxaca became known perfect flowers; Wright, 1926), in 1927 (2 plants, (see the map in Figure 5): Anon., 1928), and in 1949 (Anon., 1950). Central Veracruz . William Trelease (1857- Other gardens: A plant from Ellemeet’s collec - 1945), botanist, director of Missouri Botanical tion flowered at Rotterdam Zoo (The Netherlands) Garden, eminent researcher on the Agavaceae, in 1875 (Krelage, 1885). At Hanbury (Italy), A. and the most prolific author of names in Agave ellemeetiana flowered repeatedly before 1915, and (192; Smith & Figueiredo 2014b), was the first to in 1913, even with 3 specimens (Berger, 1915: 125, mention a possible wild locality for A. ellemeetiana with black and white photograph), at the Pal - from around Xalapa (Jalapa) in central Veracruz mengarten (D) in 1918 (Krauss, 1918, (Trelease, 1920). The source of this information with black and white photograph), the Botanical was not given, but the record was repeated by Garden, Berlin in 1918 or 1919, as is indicated by many later authors (e.g., Meyrán García, 1987; the distribution of seeds via the Index Seminum Sosa & Gómez-Pompa, 1994). Castillo-Campos et in 1919 and 1920 (Anon., 1919, 1920), Rotterdam al . (2005) and Espejo-Serna (2012) list A. ellemee - Zoo in 1921 (van den Houten, 1922), and the tiana as endemic to Veracruz and known from one Botanical Garden, Berlin in 2002 (Alseemgeest, herbarium specimen only (Castillo-Campos et al ., 2004; with the first colour photographs of a flow - 2005). ering plant by I. Richter which are also repro - Cházaro et al . (2012) provide the first printed duced in Richter, 2011: 62). The plant flowering habitat report of A. ellemeetiana from Veracruz: at Rotterdam in 1921 possibly goes back to Elle - They show photographs of flowering plants in meet and was depicted with a black and white habitat, cite two specimens from Totomochapa in photograph by van den Houten (1930; reproduced Mpio. de Tequila and from Joya Chica in Mpio. de here as Figure 4). The illustrations of the plants Zongolica (listed below), and list further sight

Bradleya 32/2014 151 records from near Naranjal, Córdoba, Tlilapan, Oaxaca, Mpio. de Huautla de Jiménez, vereda Orizaba, and near Texhuacán. Surprisingly, A. de Agua de Fierro, (3.5km del Puente de Fierro, ellemeetiana is rather abundant on the slopes and por la terracería a Sta. María Chilchotla), a la ter - limestone cliffs in the mountain regions of the racería de Sta. Cruz-Aguacatitla, 1460m, D. X. Sierra de Zongolica between San Andrés Tenejapa Munn Estrada & F. Mendoza 1387A, 26.09.2001. and the town of Zongolica, but it remained unno - 18°10'11"N 096°50'37"W. Ladera con exposición ticed by botanists for a long time (Cházaro et al ., W. Vegetación secundaria derivada de Bosque 2012). An A. ellemeetiana habitat near To - Mesófilo de Montaña. Peñas de roca en la ladera tomachapa (between Naranjal and the town of del cerro. Infructescencia de 1.90m de alto; frutos Zongolica) was also visited by the German agave secos. Abundante. MEXU (AGA1108992), MO. specialist Michael Greulich in January 2013 (Fig - (Instituto de Biología, 2010b; Tropicos, 2014). ures 6–8). In the “Reserva del Bicentenario”, a pri - Oaxaca, Mpio. de Huautla de Jiménez, aprox. vate nature reserve in the Sierra de Zongolica, A. 1.5km de Aguacatitla, por la terracería de Agua - ellemeetiana occurs on rocks in cloud forest catitla a Sta. Cruz de Juárez, 1589m, D. X. Munn (Reserva del Bicentenario, 2014). The US nurs - Estrada & T. Kasey 1181, 12.05.2002. 18°09'54"N eryman Guy Wrinkle (2011) offered young plants 096°50'43"W. Paredes de roca expuesta, a la orilla of an ” Agave species Zongolica“ which apparently de la terracería. Inflorescencia de 80cm; flor represent young A. ellemeetiana plants originat - blanca(!). Abundante. MEXU (AGA1109024), MO. ing from the Sierra de Zongolica where the plant (Instituto de Biología, 2010c; Tropicos, 2014). is “growing on vertical cliffs along with [the One of the two latter specimens was appar - bromeliad] Tillandsia grandis Schltdl”. ently used for the molecular study by Gil-Vega et Specimen data: Veracruz, Mpio. de Tequila, al . (2007; see below). Totomochapa, cantil rocoso “El Popocatl”, Y. A. Northern Oaxaca – San Bartolomé Ayautla . Gheno H . 100, 14.02.2011 (flowering). CORU, The first who verifiably rediscovered Agave elle - XAL (Cházaro et al ., 2012). meetiana in habitat was the plant explorer Alfred Veracruz, Mpio. de Zongolica, paraje Joya Bernhard Lau (1928–2007, see Staples, 2007). In Chica, junto al Instituto Tecnológico Superior de 1987, Lau (1993a, 1993b) travelled along the new Zongolica, 5 kilómetros al oeste de la ciudad de road MEX 182 winding from San Felipe Jalapa de Zongolica, 1300m, M. Cházaro B. & L. Escandon Díaz up to Huautla de Jiménez along the Rio 9805, 27.04.2010 (flowering). CORU (Cházaro et Santo Domingo gorge (Figure 9). Lau stopped in al ., 2012). between at San Bartolomé Ayautla to see the local Northern Oaxaca – Sierra Mazateca . The Mex - endemic cycad Dioon rzedowskii De Luca, ican botanist and Agavaceae specialist A. García- A.Moretti, Sabato & Vázq.Torres (1980) (Figure Mendoza (2002) briefly mentioned the rediscovery 12). On the almost vertical karstic cliffs of Cerro of A. ellemeetiana in Oaxaca, but without indicat - Rabón above Ayautla, Lau discovered an espe - ing a locality. Two years later, García-Mendoza cially large species of butterwort later published (2004) mentioned A. ellemeetiana to occur be - as Pinguicula gigantea Luhrs (1995) growing to - tween 400–1200m in the distritos Teotitlán and gether with a “large population of a form of Agave Tuxtepec, i.e., roughly in the Sierra de Mazateca attenuata Salm-Dyck in an area of 3000mm rain - (= Sierra de Huautla) in Northern Oaxaca and fall annually” (Lau, 1993a, 1993b, 1998). The ac - just southeast of the Sierra de Zongolica (Figure companying photographs show Pinguicula 5). The records by García-Mendoza were most gigantea and agave leaf tips (Lau, 1993a, 1993b). probably based on three A. ellemeetiana speci - Ullrich (2007: Figure 6 & p. 28) reproduces a pho - mens in the National Herbarium of Mexico tograph made by Lau in Oaxaca most probably at (MEXU) which were collected in 2001 in the Cerro Rabón (Ullrich l.c.: “near Jalapa de Diaz” course of a floristic study of the montane cloud which is ca. 10 road km E of Ayautla) and com - forests in the Sierra Mazateca (Lorea & Munn, ments that the plant cannot be determined with 2005). certainty, but may be A. ellemeetiana . Partrat Specimen data: Oaxaca, Mpio. de Eloxochitlán (2014) reproduces Lau’s habitat photograph from de Flores Magón, brecha de San Antonio Eloxo - Cerro Rabón showing Pinguicula gigantea and chitlán a San José Buenavista, adelante del se - agave leaf tips, and Rivadavia (2014) provides ad - gundo puente, 1320m, D. X. Munn Estrada & ditional photographs of both species which make F.Mendoza 1442, 1.10.2001. Bosque de niebla. the agave clearly identifiable as the stemless, Creciendo sobre roca en un ladera del cerro. In - green-leaved and toothless A. ellemeetiana . Lau’s fructescencia de 1.30m de alto, frutos secos. Abun - misidentification of A. ellemeetiana with the sim - dante. MEXU (AGA1108894), XAL. (Instituto de ilar A. attenuata ssp. dentata (J.Verschaff.) B.Ull - Biología, 2010a; Cházaro et al ., 2012). rich (= A. pedunculifera Trel.) whose eastern-most

152 Bradleya 32/2014 Figure 8. Rosette of Agave ellemeetiana subsp. ellemee - tiana on a vertical calcareous rock face in the Sierra de Figure 7. Plants of Agave ellemeetiana subsp. ellemee - Zongolica near Totomachapa (central Veracruz). The in - tiana on calcareous rock faces in the Sierra de Zongolica sert shows the entire leaf margin without denticles. near Totomachapa (central Veracruz) 15th January 2013, photos: Michael Greulich. 15th January 2013, photo: Michael Greulich.

Figure 9. Mountains with cloud forest/tropical rainfor - est near San Bartolomé Ayautla (Northern Oaxaca), habitat region of Agave ellemeetiana subsp. ellemee - tiana . San Bartolomé Ayautla is situated at the Rio Santo Domingo which is visible just in the midst behind the right banana plant. October 2008, photo: Art Vogel. localities lie about 300km westwards (Ullrich, 2007: 25) precluded the report of its rediscovery. The Dutch horticulturist Art Vogel ( pers. comm .) observed A. ellemeetiana growing on near vertical calcareous rock faces along MEX 182 in the Ayautla area (Figures 9–12). Vogel even found one plant growing epiphytically on the stem of a cycad (most probably Dioon rzedowskii , Figure 12); his photograph is the first record of A. ellemeetiana Figure 10. Vertical calcareous rock cliff in cloud for - est/tropical semideciduous forest near San Bartolomé growing as an epiphyte. Ayautla (Northern Oaxaca) with Agave ellemeetiana Western Oaxaca – Putla/Tlaxiaco region . In subsp. ellemeetiana and the bromeliad Tillandsia gran - May 2008, the German nurseryman Gerhard dis (rosettes with pendent old leaves), the largest species Köhres (2008) reported A. ellemeetiana from vol - in its genus occuring largely lithophytic. canic rocks at about 2300m northeast of Putla October 2008, photo: Art Vogel. near Santiago Nuyoó in Western Oaxaca. Köhres (2008) published habitat photographs of A. elle - meetiana , also including an inflorescence, and pro -

Bradleya 32/2014 153 Figure 11. Agave ellemeetiana subsp. ellemee - Figure 12. As a rare exception, Agave ellemee - tiana on vertical calcareous rock cliffs in cloud for - tiana subsp. ellemeetiana grows as epiphyte on est/tropical semideciduous forest near San the stem of a cycad, most probably the local en - Bartolomé Ayautla (Northern Oaxaca). demic Dioon rzedowskii in cloud forest/tropical October 2008; photo: Art Vogel. semideciduous forest near San Bartolomé Ayautla (Northern Oaxaca). October 2008, photo: Art Vogel. vided the first record of the species from the Sierra Madre del Sur, i.e., the southern mountain Dougall (1895-1973), intrepid botanist-explorer chain exposed to the Pacific Ocean (Figure 5). The who spent 42 winters until his death collecting UK garden designer Paul Spracklin found A. elle - plants and animals in Oaxaca and Chiapas meetiana in March 2010 and depicts it from about (Phillips, 2006), mentioned in his field notes that 20km east of Putla (which is roughly around San - on 19.11.1949 he saw “a large Agave sp. - maguey tiago Nuyoó) where it occurs around 2600m along pelon -, the leaf margins are spineless”. The agave with a large population of A. atrovirens Karw. ex was observed on an abandoned ranch at Salm-Dyck on a rock face (Spracklin, 2010; Fig - Zarzamora in Tlaxiaco Mpio. (MacDougall, 1971), ures 15 & 16). The US nurseryman Jeremy Spath which is west of Tlaxiaco and south of San Andrés and Kelly Griffin (2011) found A. ellemeetiana a Chicahuaxtla and thus just within the area men - couple of years before and depict the species from tioned above where A. ellemeetiana is now known between Putla and Tlaxiaco at about 2440m (8000 to occur. Since no other spineless agave is known ft.; Griffin, 2011). The Köhres locality was re-vis - from that area (cf. Lott & Garcia-Mendoza, 1994), ited by the Dutch agave specialists Bertus Spee MacDougall most probably already rediscovered and Wim Alsemgeest in January 2014 (Figures 17 A. ellemeetiana back in 1949(!), but without noting & 18). Plants of the Köhres' collection are culti - the importance of his find. vated in Germany (Figure 19), and plants of the Spath & Griffin collection were offered commer - 5. Habitats and cultivation cially (Figure 20). In Oaxaca, A. ellemeetiana subsp. ellemeetiana Most interestingly, Thomas (Tom) Baillie Mac - occurs between 400 and 1460m on rocks in mon -

154 Bradleya 32/2014 Figure 13. Old cultivated plant of Agave ellemee - tiana subsp. ellemeetiana as it is known in culti - vation in Europe for nearly 150 years. Berkenhof Tropical Zoo, Kwadendamme, The Netherlands. 8th September 2009; photo: Bertus Spee. tane cloud forests (bosque mesófilo de montaña) and tropical semideciduous forests (bosque tropi - cal subperennifolio; Garcia-Mendoza, 2004), i.e., on edaphically arid microsites within humid habi - tats with fog during most of the year. Garciá-Men - doza (2002) mentions four Agave species that occur in cloud forests: A. ellemeetiana as well as A. atrovirens , A. attenuata , and A. warelliana . Agave gomezpompae Cházaro & Jimeno-Sevilla (Cházaro et al ., 2010), A. gracielae Galván & Za - mudio (2013) and A. mitis (Gentry, 1982: 222; Al - Figure 14. Agave ellemeetiana subsp. ellemee - cántara & Luna, 2001; Cartujano et al ., 2002; all tiana . Pure green leaf of a cultivated plant show - as A. celsii Hook.) which occur, among other veg - ing the entire leaf margin and the soft-herbaceous etation types, also in cloud forests can be added, terminal spine. as well as as five species of the former genus Man - 14th May 2014; photo: Michael Greulich. freda Salisb. and two species of the former genus Polianthes L., both now placed in Agave subg. in a greenhouse for bromeliads or orchids which Manfreda (Salisbury) Baker (Thiede, 2001): Agave are frequent companions on its habitats, than in a debilis A.Berger (as Manfreda pringlei Rose), A. hot desert greenhouse. Agave ellemeetiana can pubescens Regel & Ortgies [as Manfreda pubes - rapidly grow into large specimens and does not cens (Regel & Ortgies) Espejo & López-Ferrari], necessarily need a pronounced resting period in A. scabra Ortega (as Manfreda scabra (Ortega) winter; with sufficient light, it can grow almost McVaugh; all three in Castillejos Cruz 2009), A. continuously. justosierrana (García-Mendoza) Thiede (as Man - freda justosierrana García-Mendoza (2011)), and 6. Structure of the leaf margin A. umbrophila (García-Mendoza) Thiede (as Man - Early authors (Koch, 1865a; Jacobi, 1865; freda umbrophila García- Mendoza (2011)), as Baker, 1869, 1877, 1888a, 1888b) described the leaf well as A. coetocapnia (M.Roemer) Govaerts & margin of A. ellemeetiana as white, parchmenty Thiede (López-Pérez & al. 2011; as Polianthes and completely toothless. Later authors (Berger, geminiflora (Lexarza) Rose)), and the recently 1915; Gentry, 1982; Cházaro et al ., 2012) added published Polianthes alboaustralis Solano & Ríos- that the margin may sometimes be reddish and Gómez (2014) for which a binomial in Agave is yet finely serrulate towards the leaf apex. Cultivated wanting. Thus, with fourteen species, agaves are plants (Figure 14) as well as plants in habitat in much more common in cloud forests than previo - Veracruz (Figure 8) and Northern Oaxaca match suly thought. this description, but since the plants from both The habitats give a clue on the cultivation re - areas cannot be distinguished, it is not possible to quirements: A. ellemeetiana should tolerate a prove that Ellemeet’s original plants came from Ve - higher air and soil humidity and would do better racruz as (indirectly) stated by Trelease (1920).

Bradleya 32/2014 155 Figure 15. Agave ellemeetiana subsp. subdentata Figure 16. Agave ellemeetiana subsp. subdentata on a rock face in temporarily dry semideciduous in habitat 20km east of Putla (Western Oaxaca); a forest 20km east of Putla (Western Oaxaca), young plant with pure green leaves. The insert around 2600m. The leaves of old plants are glau - shows the margin of the plants densely beset with cous. 5th March 2010, photo: Paul Spracklin. small denticles. 5. March 2010, photos: Paul Spracklin.

Figure 18. Agave ellemeetiana subsp. subdentata on rocks northeast of Putla (Western Oaxaca); old plant with glaucous leaves. The inserts show the leaf margins of adult leaves (left) and of young, in - folded leaves (right), both densely beset with small denticles. 28th January 2014, photos: Bertus Spee.

Plants from Western Oaxaca differ in having whitish to reddish leaf margins which are finely denticulate throughout (Köhres, 2008; Spracklin, 2010; Figures 16, 18, 19 & 20). For the latter, the name Agave ellemeetiana var. subdentata Tre - lease in L.H.Bailey with ”leaves very minutely denticulate” (Trelease, 1914: 236) is re-established here at subspecific rank.

7. Nomenclature, typification and synonymy

Figure 17. Agave ellemeetiana subsp. subdentata Agave ellemeetiana K.Koch, Wochenschr . 8: on volcanic rocks northeast of Putla (Western 103, 1.April 1865. Holotype : sterile plant culti - Oaxaca); one specimen showing an inflorescence vated at BG Berlin (not known to be preserved). – with wilted flowers. Neotype (designated here): “ Hort. Kew . March 28th January 2014, photo: Bertus Spee. 10, 1877, sheet 1, 2, 3” (= K000524804,

156 Bradleya 32/2014 K000524803, K000524802, teste N.E.Brown; K sometimes reddish (Figures 8 & 14). digital images!) ≡ Agave ellemeetiana K.Koch, Distribution & habitats (Figure 5): Mexico: Wochenschr . 7: 164, 28. May 1864, nomen . Central Veracruz (Sierra Madre Oriental: Sierra = Agave ellemeetiana Regel, Gartenflora 14 : 266, de Zongolica; Figures 6–8), Northern Oaxaca Aug. 1865 (nom. illeg . Art. 53.1). Type: not indi - (Sierra Madre Oriental: Sierra Mazateca and near cated. Syn. nov . San Bartolomé Ayautla; Figures 9–12); on often = Agave ellemeetiana Jacobi, Hamburger Garten- near-vertical cliffs and rock-faces, in cloud or trop - & Blumenzeitung 21 : 457, Oct. 1865 ( nom. illeg . ical semideciduous forests, in the cloud belt, 400– Art. 53.1). Holotype: sterile plant cultivated at BG 1460m; flowers February to May. Berlin (not known to be preserved). – Neotype Illustrations: (hab. = habitat, inf. = inflores - (designated by Gentry, 1982: 96): “ Hort. Kew . cence, bwpl. = black & white plate, colpl. = colour March 10, 1877, sheet 1, 2, 3” (= K000524804, plate, bwph. = black & white photograph, colph. = K000524803, K000524802, teste N.E.Brown; K colour photograph): digital images!). Syn. nov . Baker (1869, inf. colpl. [reproduced as Figure 1]); Baker (1877, bwpl.); Baker (1888b, inf. colpl. Typification : Since no original material was [reproduced as Figure 2]); Berger (1915: 125–126, known to be preserved, Gentry (1982: 96) desig - inf. bwph.); Anonymous (1910, inf. bwph. [repro - nated three sheets (1, 2, 3) at Kew (K) as neotype duced as Figure 3]); Anonymous (1918, inf. for A. ellemeetiana Jacobi (Gentry omitted sheet bwph.); Krauss (1918, inf. bwph.); van den Houten 4). All three sheets bear the same handwritten (1930: 3, inf. bwph. [reproduced as Figure 4]); Bre - label by N. E. Brown and belong to the same spec - itung (1961: 22, bwph.); Janse (1977, bwph.); Gen - imen since “the parts are clearly labelled as being try (1982: 94–97, bwph.); Kolendo (1990, bwph.); part of that same specimen” (ICN Art. 8.3, Mc - Lau (1993a, hab. colph.); Lau (1993b, hab. colph.; Neill et al ., 2012). The same sheets are designated both as A. attenuata ); Alsemgeest (2004, inf. here as neotype for A. ellemeetiana K.Koch which colph.); Heller (2006: 82–83, colph.); Ullrich (2006, was based on the same cultivated plant as A. elle - hab. colph.; as A. attenuata (?)); Richter (2011: 62, meetiana Jacobi, as exemplified above. inf. colph.); Cházaro et al . (2012, hab. inf. colph.); Rivadavia (2014, hab. colph.); Reserva del Bicen - Etymology : Named after Jonkheer Willem Cor - tenario (2014, hab. colph.). nelis Mary de Jonge van Ellemeet (1811-1888, see Material studied: Ex cult., Anonymus s.n ., Koch, 1864b; Alsemgeest, 2004), a Dutch March 10, 1877 (sheets 1, 2, 3, 4 [= K000524804, landowner, mayor and keeper of an important K000524803, K000524802] K digital images!) (1, agave collection on his estate “Overduin” near 2, 3 = neotype); living plant at BG Bochum. Oostkapelle (The Netherlands). Agave ellemeetiana K.Koch subsp. subdentata Characters : For more detailed morphological de - (Trelease) Thiede, comb. nov . scriptions, see Jacobi (1865), Baker (1869), Baker = Agave ellemeetiana var. subdentata Trelease (1877), Baker (1888b), Berger (1915: 125–126), in L.H.Bailey, Stand. Cycl. Hort . 1: 236 (1914). Gentry (1982: 94, 96), Irish & Irish (2000: 118– Type : (not indicated). Neotype (designated here): 119), Thiede (2001: 30), and Cházaro et al . (2012). ex cult., J. Thiede s.n., 8th October 2014; from liv - Distribution (Figure 5): Mexico: Veracruz ing plant raised from seed collected in Mexico, (Sierra de Zongolica; Figures 6–8), Oaxaca (Sierra Oaxaca, between Putla and Tlaxiaco, ca. 2400m; Mazateca; near San Bartolomé Ayautla, Figures J. Spath & K. Griffin s.n. (ZSS!) (Figure 20) 9–12; Putla/Tlaxiaco region, Figures 15–18). Taxonomic rank: The morphologically as well as geographically discrete populations from the Key to the subspecies of Agave ellemeetiana : Putla/Tlaxiaco region are classified as subspecies - Leaf margin mostly without denticles, sometimes which necessitates a new combination based on finely denticulate towards the apex, whitish, some - times reddish ...... subsp. ellemeetiana var. subdentata Trelease.  Leaf margin with denticles throughout, whitish or Typification: In his short protologue (“With reddish ...... subsp. subdentata lvs. [= leaves] very minutely denticulate it be - comes var. subdentata , the distinction of which from A. pruinosa [= A. attenuata ssp. dentata Agave ellemeetiana K.Koch subsp. ellemee - (Roezl) B. Ullrich (2007)] is not clear.”), Trelease tiana (1914: 236) did not indicate a type. In an assess - Diagnostic characters: Leaf margin parch - ment of Trelease's online type material, Smith & menty and mostly completely toothless, some - Figueiredo (2014b) did not designate a type for the times finely serrulate towards the apex, white, name. Here, a neotype is designated from culti -

Bradleya 32/2014 157 Figure 19. Agave ellemeetiana subsp. subden - tata . A cultivated plant raised from seed from the Köhres collection near Santiago Nuyoó. The in - sert shows the leaf margin of young leaves densely beset with small denticles. 15th June 2014, photos: Michael Greulich. vated material of the Spath & Griffin collection (Figure 20). Diagnostic characters: Leaf margin whitish or reddish, finely, but clearly denticulate throughout (Figures 16, 18, 19 & 20). Leaves of adult plants have a distinct wax bloom (Figures 15 & 18) ab - sent from subsp. ellemeetiana (Figures 7, 8 &11) and appear to be at least sometimes broader (cf. Figure 15 with Figures 8, 11 & 13), but measure - ments are not available. Figure 20. Commercially offered young plant of Distribution & habitats (Figure 5): Mexico: Agave ellemeetiana subsp. subdentata , from seed Western Oaxaca (Sierra Madre del Sur: collected in the Putla/Tlaxiaco region (Spath & Putla/Tlaxiaco region (Figures 15–18); on volcanic Griffin s.n., = neotype collection). The insert rocks in cloud or tropical semideciduous forests, shows the leaf margin of young, infolded leaves 2300–2600m; flowers in January. densely beset with small denticles. Illustrations: Köhres (2008, hab. inf. colph.); 22nd August 2012, photos: Joachim Thiede. Spracklin (2010, hab. colph.); Griffin (2011, hab. colph.); Pilbeam (2013: 78 hab. colph.); all as A. Saunders (mentioned above), noted that its nearly ellemeetiana . tubeless flowers differ much from those of A. at - Material studied: Living plants raised from tenuata with a clearly developed tube, and con - seed collected between Putla and Tlaxiaco (Mex - cluded that both species cannot be placed ico); J. Spath & K. Griffin s.n. (= neotype collec - together. Nevertheless, one year later, Jacobi tion; Figure 20). (1868) continued to place A. ellemeetiana closest to A. attenuata , together with A. pruinosa Lem. ex 8. Placement and relationships – morpho - Jacobi and A. debaryana Jacobi (both synonyms logical data of A. attenuata subsp. dentata ; formerly known as Early agave authors (Koch, 1865a; Jacobi, A. pedunculifera ; Ullrich 2007). 1865; Baker, 1877, 1888a, 1888b) stressed the Berger (1915) combined floral features with similarity in habit and leaf structure between A. leaf characters and established a section Anopla - ellemeetiana and A. attenuata and placed both gave A.Berger for species with short or lacking species as close relatives, partly together with fur - tube and soft leaves subdivided into two subsec - ther unarmed and soft-leaved species such as A. tions: Dracontagave A.Berger for the stem-form - sartorii K.Koch (= A. pendula Schnittsp.) or even ing, short-tubed A. attenuata and taxa now placed with species of Agave subg. Manfreda (Salisb.) in its synonymy, and Leptagave A.Berger with the Baker. Jacobi (1867), when studying pressed flow - tubeless and stemless species A. ellemeetiana and ers of A. ellemeetiana he obtained from W.W. A. bracteosa S.Watson ex Engelm. Gentry (1982)

158 Bradleya 32/2014 followed a similar concept and established an in - ulations of A. ellemeetiana from the Sierra Madre formal Group Choritepalae for species with tube - del Sur in Western Oaxaca designated as subsp. less flowers which includes A. ellemeetiana , A. subdentata here need further study of their char - bracteosa and A. guiengola Gentry. I.e., over the acteristics, variability and differences from subsp. time, agave classifications based largely or exclu - ellemeetiana . sively on leaf-characters (Jacobi, Baker) were re - placed by systems combining floral features with Acknowledgements leaf characters (Berger, 1915; Gentry, 1982). The author is indebted to Michael Greulich The German agave specialist Bernd Ullrich (D), Paul Spracklin (UK), Bertus Spee (NL) and (1991) suggested that A. ellemeetiana might rep - Art Vogel (NL) for providing photographs and ad - resent a ”garden clone” of A. pedunculifera (= A. ditional information, to Wim J. Alsemgeest (NL) attenuata subsp. dentata ) only, an assumption for photographs, literature and additional infor - clearly rebutted by the rediscovery of the former mation, to Gordon D. Rowley (UK) for permission in habitat. Ullrich (1991) placed both species in a to photograph the plate from Baker (1869) in his re-established Group Attenuatae Baker, together library and to Jonathan Clark (UK) for taking with A. attenuata , A. vilmoriniana A.Berger, A. the picture of it reproduced here as Figure 1, to chrysoglossa I.M.Johnst. and A. ocahui Gentry. Graham Charles (UK) for his editorial work, and Data from leaf anatomy (Blunden et al ., 1973) to an anonymous reviewer for valuable com - and phytochemistry (steroidal sapogenins; Blun - ments. den et al ., 1978) are inconsistent with regard to the relationships of Agave ellemeetiana . Literature Cited ALCáNTARA AYALA , O. & L UNA VEGA , I. (2001). 9. Placement and relationships – molecular Análisis florístico de dos áreas con bosque data mesófilo de montaña en el estado de Hidalgo, Gil-Vega et al . (2007) provide a broad assess - México: Eloxochitlán y Tlahuelompa. Bol. Soc. ment of relationships in agave based on molecular Bot. Méx. 54 : 51–87. AFLP data for 85 accessions, including a specimen ALSEMGEEST , W. (2004). De geschiedenis rond of A. ellemeetiana (subsp. ellemeetiana ) from Agave ellemeetiana Jac., een plant genoemd Huautla, Oaxaca, most probably representing one naar een Nederlander. Succulenta (NL) 83 : of the two collections cited above, as well as a spec - 105–118. English version on the internet at imen named “ A. aff. pedunculifera JE s/n. Putla, http://www.agaves.nl/Articles/E_Agave%20ell Oaxaca”, which most probably represents A. elle - emeetiana.htm meetiana subsp. subdentata which occurs in just ANDRé , E. (1872). Collection d'Agaves de M. de that area (see above). In the resulting dendrogram, Jonge-Van Ellemeet. Ill. Hort . 19 : 148–150. the three species of Group Choritepalae sensu Gen - ANONYMOUS (1872). Second May Show. Gard. try, 1982 ( A. bracteosa , A. guiengola and A. elle - Chron ., ser. 3, 2: 672. meetiana ) are placed separate from each other. ANONYMOUS (1873a). Notes of the week [auction - Agave ellemeetiana subsp. ellemeetiana and A. elle - ing of Ellemeet's agave collection]. The Garden meetiana subsp. subdentata are placed separate, 4: 144. but not far from each other, between members of ANONYMOUS (1873b). Chronique Horticole [auc - Group Amolae (A. attenuata , A. nizandensis Cutak, tioning of Ellemeet's agave collection]. Rev. A. pedunculifera (= A. attenuata subsp. dentata ), A. Hort . 45 : 322. vilmoriniana , and A. yuccaefolia Redouté (= A. spi - ANONYMOUS (1874). Agave Ellemeetiana . The Gar - cata Cav.)) as well as A. filifera Salm-Dyck (Group den 5: 416. Striatae ) and A. garciae-mendozae Galván & ANONYMOUS (1877a). Agave ellemeetiana . Gard. L.Hern. (Group Marginatae ). The cladogram of Gil- Chron ., ser. 3, 7: 246. Vega et al . (2007) provides the first molecular evi - ANONYMOUS (1877b). Agave ellemeetii . Gard. dence that Gentry’s Group Choritepalae is artificial Chron ., ser. 3, 7: 278. and that A. ellemeetiana might best be grouped to - ANONYMOUS (1879). Professor Karl Koch. Gard. gether with other soft- and few-leaved species of Chron. , ser. nov. 12 : 329–331. Group Amolae , as suggested by Ullrich (1991). ANONYMOUS (1883a). Agave ellemeetiana. Gard. The finding that the two accessions of A. elle - Chron ., ser. nov. 19 : 698. meetiana (i.e., subsp. ellemeetiana and subsp. sub - ANONYMOUS (1883b). The evening meeting held in dentata ) studied by Gil-Vega et al . (2007) are the Royal Society. Gard. Chron ., ser. nov. 19 : placed somewhat separated from each other might 768. indicate that subsp. subdentata could possibly even ANONYMOUS (1910). Agave ellemeetiana. Bull. merit separate specific rank. In any case, the pop - Misc. Inf. Kew 1910 : 65.

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