Thiede, J. (2016): A review of Agave mitis (Asparagaceae/Agavaceae). Bradleya 34: 200-216.

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A review of Agave mitis (Asparagaceae/Agavaceae) Joachim Thiede Schenefelder Holt 3, 22589 Hamburg, Germany (email: [email protected]).

Summary: A broad review of the history, discovery, introduction, taxonomy, nomenclature, distribution, habitats, ecology, and characteristics of Agave mitis (syn. A. celsii) is provided. A revised taxonomic concept is introduced: The invalidly published A. mitis var. albidior shows broad overlap in its diagnostic features (leaf colour and length of the tepal lobes) with A. mitis var. mitis and is therefore referred to the synonymy; no infraspecific taxa are recognized. A. ehrenbergii and A. goeppertiana are established as new synonyms. A neotype is designated for A. albicans. A. mitis is more widespread than previously known and occurs in the Mexican states of Coahuila, Nuevo León, Tamaulipas, San Luis Potosí, Guanajuato, Querétaro, Hidalgo and Puebla; a distribution map is given.

Zusammenfassung: Es wird ein breitgefaßter Überblick über die Geschichte, Entdeckung, Einführung, Taxonomie, Nomenklatur, Verbreitung, Standorte, Ökologie und Merkmale von Agave mitis (syn. A. celsii) gegeben. Ein revidiertes taxonomisches Konzept wird eingeführt: Die ungültig veröffentlichte A. mitis var. albidior weist in ihren diagnostischen Merkmalen (Blattfarbe und Länge der Petalenzipfel) eine breite Überschneidung mit A. mitis var. mitis auf und wird deshalb in die Synonymie verwiesen; es werden keine infraspezifischen Taxa anerkannt. A. ehrenbergii und A. goeppertiana werden als neue Synonyme etabliert. Für A. albicans wird ein Neotypus ausgewählt. A. mitis ist weiter verbreitet als bisher bekannt und kommt in den mexikanischen Staaten Coahuila, Nuevo León, Tamaulipas, San Luis Potosí, Guanajuato, Querétaro, Hidalgo und Puebla vor; eine Verbreitungskarte wird gegeben.

Keywords: Agavaceae, Agave celsii, Agave mitis, Asparagaceae, Barranca de Metztitlán, Flora of Mexico, nomenclature, taxonomy.

Introduction Agave mitis Martius (1848) is a long, wellknown species of ser. Micracanthae (Salm-Dyck) Hochstätter (= Group Polycephalae of Gentry

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(1982)) within subg. Littaea (Tagliabue) Herbert. It is characterized by rosettes of few, broad and soft leaves often forming large clumps, leaf margins with small, closely spaced, often bicuspid teeth, and inflorescences formed terminally on small side rosettes below the terminal rosettes ('axillary') with densely aggregated flowers (Figures 2–8, 10, 11). It is distributed from the barrancas and mountains of the eastern Chihuahuan Desert up to the humid cloud forests of the Mexican Sierra Madre Oriental (Figure 9). Contemporary taxonomic concepts applied different species names [Gentry (1982): A. celsii Hooker / Ullrich (1993): A. mitis Martius] and distinguished plants with pale glaucous to whitish leaves and longer tepal lobes ("tepals") locally distributed in the Barranca de Metztitlán (Hidalgo) at varietal level [Gentry (1982): A. celsii var. albicans (Jacobi) Gentry / Ullrich (1993): A. mitis var. albidior (Salm-Dyck) B.Ullrich)] from the widespread typical variety with green leaves and shorter tepal lobes [Gentry (1982): A. celsii var. celsii / Ullrich (1993): A. mitis var. mitis]. Albeit known for 168 years, the history, discovery, introduction, taxonomy, nomenclature, distribution, habitats, ecology, and characteristics of A. mitis remained incompletely known. This paper aims to fill these gaps of knowledge.

Methods Specimen and Observation Data were provided by the "Arizona State University Hebarium (ASU), Desert Botanical Garden (DES), Royal Botanic Gardens, Kew (K), United States National Herbarium, Smithsonian Institution (US)" via JSTOR Global Plants (https://plants.jstor.org/) and the Global Biodiversity Information Facility (http://www.gbif.net) accessed through the BioCASE Data Portal (http://search.biocase.org) or other web resources, 6th August 2016. Results and discussion The novelties period: a plethora of names (1848-1876) Agave mitis was published by the German botanist, explorer and ethnologist Carl Friedrich

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Philipp von Martius (1794–1868) on 16th Nov. 1848 in the seed distribution list (Index Seminum) of the Botanical Garden Munich (Germany) with a short description of the leaf features of a cultivated plant originating from Mexico (Figure 1). This earliest date and place of valid publication was not established until Ullrich (1993: 29); all earlier authors (e.g. Jacobi, 1865: 256; Baker, 1888: 191; Berger, 1915: 62; Gentry, 1982: 220) ascribed A. mitis to Salm-Dyck in Bonplandia 7: 93, 1859 [Gentry (1982: 220) wrongly cited “1855”, see below]. When cultivated plants of A. mitis came into bloom, its inflorescence, flower, fruit and seed features were added by Jacobi (1871: 82–83). Twenty-five species and varietal names now placed in the synonymy of A. mitis were published between 1856 and 1914, seventeen of them between 1856 and 1866 (Table 1). Between 1848 and 1859, all new taxa except A. celsii were published from sterile, non-flowering rosettes. The inflorescence and flower characters of these species were partly published later when cultivated plants came into bloom, as mentioned above for A. mitis. Between 1861 and 1914, nearly a half of the new taxa were published from plants in bloom and included inflorescence and flower characters (Table 1). Seven of the new species were illustrated with colour plates (Table 1) in Curtis's Botanical Magazine (5×), Gartenflora (1×) and Todaro's Hortus Botanicus Panormitanus (1×). The plates are reproduced here as Figures 2–8; they finely illustrate the considerable variability of A. mitis in leaf and flower colour, size, and shape. Four further plates were published in black and white (Table 1). All new taxa were published from cultivated plants mostly of unknown origin (Table 1). Five species only were based on plants known to be introduced from Mexico, but all without precise locality: introduced by Karwinski (A. rupicola, A. densiflora) or Ehrenberg (A. ehrenbergii), sent by Botteri (A. botterii), or from an unknown Mexican source (A. mitis; Table 1). Ullrich (1993) assumed that A. mitis was likewise introduced by Karwinski, but clear evidence is lacking. The first distribution records as to Mexican state were published not earlier than in 1920 by Trelease (1920: 135): “San Luis Potosí” for A. celsii and “Hidalgo or Veracruz?” for A. micracantha. Astonishingly, for a species that long- and wellknown, the first specimen-based localities were not published earlier than in Gentry's revision (1982: 233) which cited nineteen field-collected specimens, the oldest from 1878 (“1875”; Parry & Palmer s.n. [MO]) and 1891 (Pringle 3739 [K, MEXU, MO, UC, US]).

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Agave mitis Mart. foliis laete viridibus, patulis, carnosis, e lineari-ovatis acuminatis planis aut antice concaviusculis, acuminatis, spina terminalis molliuscula, margine spinoso-denticulatis, dentibus frequentibus erectis aut reversis curvis molliusculis albis dein fuscis, infra spinam terminalem integerrimis. - Inter A. viviparam et atrovirentem quasi intermedia, sed minor et magis delicata. Folia minus firma quam in reliquis speciebus majoribus, facilius franguntur. Crescit in Mexico in regione calida, ideo cultura interior.

Figure 1. Agave mitis (“var. mitis”). Text of the protologue of Agave mitis Martius (reproduction of the text from “Delectus Seminum in Horto R. Botanico Monacensi collectorum anno 1848”, p. 4, published 16th Nov. 1848).

The synoptic period: first name reductions (1877–1981) Several early authors started to refer redundant names into synonymy. For example, the Kew botanist John Gilbert Baker (1834-1920), keeper of the herbarium 1890–1899 and authority on the systematics of Agave and many other monocotyledons, placed A. ousselghemiana, A. concinna, and A. micracantha var. albidior in the synonymy of A. albicans (Baker, 1877: 717; 1888: 191; 1891). Further name reductions were carried out in synoptic treatments by Baker (1877, 1888), by Alwin Berger (1871–1931), curator at Hanbury Gardens (Italy) 1897–1914 and Agave monographer (Berger, 1915), and by US botanist William Trelease (1857–1945), director of Missouri Botanical Garden 1889–1912 and eminent Agavaceae researcher (Trelease, 1920). However, Baker's synopses (1877, 1888) and Berger's monograph (1915) still recognized most of the names listed in Table 1 as distinct and placed only a few of them in the synonymy of others. An important step forward was provided by Trelease in his Agave treatment for P.C. Standley's Trees and Shrubs of Mexico. Trelease (1920: 135) was the first to carry out a drastic name reduction and recognized two species only (A. celsii and A. micracantha) and listed many further names as differing little from or being evidently allied to these two species or to A. polyacantha. Hermann Jacobsen's Handbuch der sukkulenten Pflanzen (1954) followed Berger's (1915) monograph and correspondingly upheld many redundant names. The synopsis of the US Agave collector August J. Breitung (1959) omitted most of the names, and Jacobsen's later Sukkulentenlexikon (1981) largely followed Trelease's synopsis and recognized the same two species.

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Figure 2. Agave mitis (“var. mitis”). Colour plate (hand-coloured copper engraving) from the protologue of the synonymous A. celsii, showing a plant flowering at Kew [Curtis's Botanical Magazine plate 4934; Hooker (1856)]. The leaves, given as “of a pale very glaucous green colour” in the description, are miscoloured on the plate showing whitish-grey leaves matching those of “var. albidior” (cf. Figure 6). Courtesy of The Biodiversity Heritage Fund.

A modern lumping concept: Howard Scott Gentry (1982) The extensive Agave research in the field for over 25 years and studies of specimens and literature by Howard Scott Gentry (1903–1993) culminated in his monumental monograph Agaves of Continental North America in which he recognized only one broadly defined species, distributed in the Mexican states of Tamaulipas, San Luis Potosí and Hidalgo (Gentry, 1982: 220–224; Table 3): Agave celsii Hooker (1856) with two varieties: The widely distributed A. celsii Hooker var. celsii (Fig-

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Figure 3. Agave mitis (“var. mitis”). Part-coloured lithograph from the protologue of the synonymous A. densiflora Regel (cited as “Hook.”), showing a plant flowering in the Imperial Botanical Garden St. Petersburg, Russia [Gartenflora pl. 410, Regel (1863)] Courtesy of The Biodiversity Heritage Fund.

ure 2) covers the range of the species (Figure 9) and is characterized by green to light grey glaucous leaves (Figure 12–17) and tepal lobes 12–18 mm long, and the newly combined A. celsii var. albicans (Jacobi) Gentry (Figure 6), a local endemic distributed on the eastern rim of the Barranca de Metztitlán (Hidalgo), distinguished by its pale glaucous leaves (Figures 10, 18–20) and lobes 20– 27 mm long. Gentry listed five species in the synonymy of A. celsii, v. a. A. mitis ”H. Monac. ex Salm” (1855), and referred to the synopsis of Trelease (1920) for further synonyms not listed by him.

Shortcomings - Nomenclature: Gentry applied the name A. celsii Hooker (1856), placing “Agave micracantha Salm, Bonplandia 7, 93, 1855” and “Agave mitis H. Monac. ex Salm, Ibid.” in its syn-

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Figure 4. Agave mitis (“var. mitis”). Colour plate (hand-coloured copper engraving) from the protologue of the synonymous A. botterii, showing a plant flowering in the collection of J. T. Peacock at Sudbury House, Hammersmith, England [Curtis's Botanical Magazine plate 6248, Baker (1876)]. Courtesy of The Biodiversity Heritage Fund.

onymy which, based on Gentry's citations, inverted the priority of names with the same rank (Art. 11.3 of the International Code of Nomenclature for algae, fungi, and plants [ICN]; McNeill et al., 2012). However, Bonplandia volume 7 was in fact published in 1859, so that Gentry's concept could be upheld. That A. mitis was in fact published much earlier in 1848 thus antedating A. celsii was not established until Ullrich (1993), as mentioned above. Furthermore, Gentry's new combination A. celsii var. albicans (Jacobi) Gentry (1982), based on A. albicans Jacobi (1865), did not consider the earlier name A. micracantha Salm-Dyck var. albidior Salm-Dyck (1859) as basionym; both names are identical as outlined earlier by Baker (1891), mentioned above.

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Figure 5. Agave mitis (“var. albidior”). Colour plate (coloured lithograph) from the protologue of the synonymous A. macrantha Todaro, showing a plant flowering in BG Palermo [Todaro 1879: t. xxvii] Courtesy of The Biodiversity Heritage Fund.

Shortcomings - Typification: Since A. celsii lacked typification, Gentry (1982: 221, 233) designated the specimen “Hort. Kew June 1856, K” imprecisely as “type”. This specimen (= K000524810) was prepared in June 1856 from the living plant figured in plate 4934 in Curtis's Botanical Magazine (here reproduced as Figure 2) as part of the protologue published on 1st Aug. 1856 and consequently constitutes original material (ICN Art. 9.3.; McNeill et al., 2012) eligible as lectotype (ICN Art. 9.2.; McNeill et al., 2012); Gentry's “type” thus represents a lectotype. The protologue of A. albicans Jacobi in Hamburger Garten- und Blumenzeitung 21: 256 (1865) did not indicate a type. Gentry (1992) designated on p. 224 a specimen as well as a colour plate as “Neotype: Hort. Kew Gard., 22 May 1891. Curtis Bot. Mag. Tab. 7207)”, reproducing the plate on p. 224 as “Neotypic”, and cited on p. 233 the specimen as “Hort. K Type. Cult. Kew Bot. Gard., June 1856”. This is contrary to ICN Art. 8.1 (McNeill et

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Figure 6. Agave mitis (“var. albidior”). Colour plate (hand-coloured copper engraving) of the synonymous A. albicans, showing a plant flowering at Kew [Curtis's Botanical Magazine plate 7207, Baker (1891)]; This plate is designated in this paper as neotype for A. albicans Jacobi. Courtesy of The Biodiversity Heritage Fund.

al., 2012) which states that the type of a species name or infraspecific taxon is either a single specimen or an illustration (i.e., not both). Here, the plate in Curtis's Botanical Magazine is designated as neotype which adequately shows the characteristic leaf colouration of the plant (see below).

Shortcomings - Characters: Gentry took the measurements of the tepal lobe lengths of var. albicans (20–27mm) from a well-developed clump cultivated in fertile soil at Huntington BG (California; Gentry, 1982: Fig. 9.6. on p. 223). Measurements for cultivated plants given by various authors up to Berger (1915; Table 2), i.e., before the publication of Gentry's (1982) monograph, show a lobe length of 13–19 mm for var. albicans only which falls within the range of var. celsii (12– 22mm; Table 2); the length of the tepal lobes can thus not be used to distinguish varieties within A. celsii. The highest values for the lobe lengths of 20–27mm (Gentry l.c.) and of 22–25mm (Smith & Steyn, 2002; cf. Table 1) were apparently both ob-

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Figure 7. Agave mitis (“var. mitis”). Colour plate (hand-coloured copper engraving) of the synonymous A. haseloffii, showing a plant flowering at Kew [Curtis's Botanical Magazine plate 7527, Baker (1897b)] Courtesy of The Biodiversity Heritage Fund.

tained from especially well-nourished cultivated plants "having grown right out of their descriptions" (Lyons, 2002: 162) - a warning not to consider data from such “hyperdeveloped” plants for species descriptions or taxonomic decisions.

Gentry's concept reassessed: Bernd Ullrich (1993) Bernd Ullrich, a German agavophile who published extensively on Agave and its relatives between 1989 and 1996, in 1993 principally retained Gentry's varietal concept, but corrected Gentry's nomenclatural errors (cf. Table 3): Ullrich (1993) established A. mitis Martius (1848) as the earliest species name and placed A. celsii Hooker (1856) in its synonymy, and replaced Gentry's name A. celsii var. albicans by the new combination A. mitis var. albidior (Salm-Dyck) Ullrich (1993) based on A. micracantha var. albidior Salm-Dyck (1859), the earliest name at varietal rank. Subsequent authors followed Ullrich's concept (e.g., Thiede, 2001; Smith & Steyn, 2002), but some (e.g., Irish & Irish, 2000; Scheinvar, 2008) retained Gentry's nomenclaturally incorrect concept.

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ured on plate 7207 in Curtis's Botanical Magazine (here reproduced as Figure 6); it is the same specimen Gentry designated invalidly as neotype for A. albicans (see above). Ullrich (1993) added as new Mexican state records localities in Querétaro (SW of Jalpan) and in Nuevo León (E Zaragoza), a photographic record of A.B. Lau [Ullrich overlooked the earlier record of A. mitis from Nuevo León by Synnott (1989, as A. celsii)].

Shortcomings - Nomenclature: For his new combination A. mitis var. albidior (Salm-Dyck) Ullrich, Ullrich (1993: 32) wrongly cited as basionym reference for A. micracantha [misspelt “Agave micrantha”] var. albidior the whole page range of Salm-Dyck's paper [“Bonplandia 7(7): 87–93, 15. IV. 1859”; correct is: pp. 85–96], but not the exact page only (p. 93) on which the name was published as is required under ICN Art. 41.5 & Ex. 12. (McNeill et al., 2012). Ullrich's new combination was thus invalidly published.

Figure 8. Agave mitis (“var. mitis”). Colour plate (hand-coloured copper-engraving) of the synonymous A. bouchei, showing a plant flowering at Kew [Curtis's Botanical Magazine plate 7558, Baker (1897a)] Courtesy of The Biodiversity Heritage Fund.

Ullrich (1993) noted that the leaf colour of A. mitis var. mitis may vary from dark and pale green to yellow green (“vert foncé, vert pâle et aussi vert jaune”) and that the whitish colour of var. albidior also varies gradually, and adds that the varietal differences in the length of the tepal lobes given by Gentry (21–27 mm in var. albidior [as var. albicans] vs. 12–18mm in var. mitis [as var. celsii]) are not consistent, since already Baker (1891) indicated a lobe length of 13mm for var. albidior [as A. albicans]. Consequently, Ullrich doubted the taxonomic value of var. albidior. In addition, Ullrich (1993) designated neotypes for both names accepted by him: For A. mitis, he designated the specimen Gentry 20077 at MEXU, and for A. mitis var. albidior the specimen “n° 109 (22. V. 1891)” at Kew. The latter was prepared from the living plant of A. albicans fig-

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Additional data from recent studies Taxonomy: Hochstätter (2015: I, 18) recently elevated var. albidior to the rank of subspecies as A. mitis ssp. albidior (Salm-Dyck) Hochstätter. Characters: Recent field studies by Michael Greulich (Germany) and Paul Spracklin (England) showed that the leaf colour of both var. mitis and var. albidior varies considerably and overlaps: The leaf colour of var. mitis plants near Rio Verde ranges from green (Figure 12) to glaucous (Figure 13) plants at Tula de Allende have whitish-glaucous leaves (Figure 14), and plants with light green to glaucous leaves grow intermixed on Cerro la Laja (Figure 15) and in the Parque Nacional Los Marmoles (Figure 16). Vice versa, in the Barranca de Metzitilán, the classical and sole site of var. albidior, the leaf colour varies from alabaster-white (Figure 18), glaucous (Figure 20) and light green (Figure 19) up to a darker green [Figure on p. 49 in Scheinvar (2008, as A. celsii var. albicans)]. The glaucous plants within var. mitis (Figures 13–16) match those found within var. albidior (Figures 19–20), and, vice versa, the green plants within var. albidior (Figure 19) match those found within var. mitis (Figures 15–17), even if some inaccuracy of the leaf colours on the photographic reproductions must be taken into account. Distribution: New state records for A. mitis var. mitis were provided for Guanajuato (Zamudio & Galván, 2011) and for Coahuila and Puebla (Instituto de Biología 2016a, 2016b, 2016c; cf. map in Figure 9). Scheinvar (2008: 50) provided a dot distribution map for A. mitis var. albidior (as A. cel-

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bidior (as A. celsii var. albicans) and one population of A. mitis var. mitis (as A. celsii) from San Luis Potosí; the results of both studies were summarized by Scheinvar (2008) and Eguiarte et al. (2013): The probability that a flower develops into a fruit is 43.9%, and the probability that at least one ovule of a flower develops into a seed is 35%. On average, each inflorescence develops 51 fruits with 49 viable seeds each. Main floral visitors are bats and also bees, bumblebees, sphingids, and hummingbirds. The populations in the Barranca de Metztitlán exhibit a very low genetic distance, while the distance of the San Luis Potosí population is clearly larger (Scheinvar, 2008; Eguiarte et al. 2013). The notion of Eguiarte et al. (2013) that the status of subspecies for the Metztitlán population is adequate due to the expressed genetic distance, and may even be considered a separate species given the morphological differences described by Gentry (1982), is rejected here (see above and below).

Figure 9. Distribution of Agave mitis in Mexico. Black dots: localities for “var. mitis” given by Gentry (1982; as var. celsii), yellow dot: locality for “var. albidior” given by Gentry (1982; as var. albicans), grey dots: additional localities recorded since Gentry (1982), yellow dots with grey margins: localities for “var. mitis” with leaf colourations approaching or matching that of “var. albidior”. Dots may represent more than one record. Coa = Coahuila, NL = Nuevo León, Tam = Tamaulipas, SLP = San Luis Potosí, Gto = Guanajuato, Qro = Querétaro, Hid = Hidalgo, Pue = Puebla. Map: Wikimedia Commons 2016.

sii var. albicans) in the Barranca de Metztitlán with 20 localities, showing that the plant is more common than previously thought and also occurs at several spots on the western rim of the barranca from where it has not been reported hitherto. Ecology: Rocha et al. (2005) studied the reproductive ecology of A. mitis var. albidior (as A. celsii var. albicans), and Rocha (2006) the population genetics of three populations of A. mitis var. al-

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Summary: a revised concept for Agave mitis Gentry (1982) distinguished two varieties within A. celsii (see Table 3): var. celsii with green to light grey glaucous leaves and lobes 12–18mm long, and var. albicans with pale glaucous leaves and lobes 20–27mm long. This concept was questioned by Ullrich (1993; Table 3), noting that the leaf colour of A. mitis var. mitis (= var. celsii of Gentry) varies from dark to pale and yellow green and that the white colour of var. albidior (= var. albicans of Gentry) also varies gradually, and that the varietal differences in the length of the tepal lobes given by Gentry are not consistent, since already Baker (1891) indicated a lobe length of 13mm for var. albidior. In this paper, data are presented showing that the two varieties recognized by Gentry and Ullrich show broad overlap in both distinguishing characters (Table 3): A. mitis “var. mitis” exhibits dark green or light green to glaucous leaves and tepal lobes 12–22mm long, and “var. albidior” alabaster-white or glaucous to light green leaves and tepal lobes 13–19(–27)mm long. Since “var. albidior” is neither separable by distinct diagnostic characters nor geographically (glaucous plants are also found within the range of “var. mitis” as shown above), it is referred to the synonymy of A. mitis here, and a uniform concept for the species is introduced here without recognition of infraspecific taxa. In order to retain a name widely established in literature and horticulture, “var. albidior” could be downgraded to the rank of a forma and re-circumscribed as to include plants with alabasterwhite leaves only. This action is not followed here

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Figure 10. Agave mitis (“var. albidior”). Cultivated plant in bloom. The leaf colour of the plant, labelled A. celsii, would place it in A. mitis “var. albidior” [BG Berlin (D), 15.06.1992]. Photograph: Joachim Thiede.

Figure 12. Agave mitis (“var. mitis”). Plants growing in the shade with green leaves [San Luis Potosí, near Rio Verde, 29.10.2010]. Photograph: Michael Greulich.

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Figure 11. Agave mitis (“var. albidior”). Upper part of the inflorescence with open flowers of the same plant as in Figure 10 [BG Berlin (D), 15.06.1992]. Photograph: Joachim Thiede.

Figure 13. Agave mitis (“var. mitis”). Plants at the same locality growing in the open show some variability in leaf colouration, including plants with green leaves (below) and glaucous leaves (above) [San Luis Potosí, near Rio Verde, 29.10.2010]. Photograph: Michael Greulich.

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Figure 14. Agave mitis (“var. mitis”). This plant, growing in full sun, has whitish-glaucous leaves approaching those of “var. albidior” in colouration (cf. Figure 18) [Hidalgo, near Tula de Allende, 15.11.2010] Photograph: Michael Greulich. since it would merely represent a concession to horticultural needs contrasting established taxonomic praxis. Instead, especially attractive cultivated clones could be named as cultivars.

Agave mitis Martius, Index Seminum (M, Monacensis) 1848: 4 (1848). Type (neotype, designated by Ullrich (1993: 29)): Mexico, Tamaulipas, Rancho Cielo, Sierra de las Cucharas, 8 miles SW of Gómez Farías, 19 June 1963, elev. 3500–4000 ft., limestone outcrop over forest (Gentry 20077 [MEXU [neo], DES, US [isoneo]]).

≡ Agave micracantha Salm-Dyck var. mitis (Martius) A.Terracciano, Primo Contr. Monogr. Agave: 25 (1885). = Agave celsii Hooker, Bot. Mag. 82: t. 4934 (1856) ≡ Agave bollii var. celsii (Hooker) A.Terracciano, Prim. Contr. Monogr. Agave: 26 (1885) (nom. illeg., Art. 52.1, 25.1). Type (lectotype, designated by Gentry, 1982: 221): “Hort. Kew, June 1856” (K K000524810; digital image!). = Agave rupicola Regel, Gartenflora 7: 312 (1858). = Agave rupicola var. brevifolia Regel, Gartenflora

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7: 312 (1858). = Agave rupicola var. longifolia Regel, Gartenflora 7: 312 (1858). = Agave rupicola var. rubridentata Regel, Gartenflora 7: 312 (1858). = Agave micracantha Salm-Dyck, Bonplandia 7: 93 (1859). = Agave micracantha var. albidior Salm-Dyck, Bonplandia 7: 93 (1859) = Agave mitis var. albidior (Salm-Dyck) B.Ullrich, Succulentes 16: 32 (1993) (nom. inval., Art. 41.5.) = Agave mitis ssp. albidior (Salm-Dyck) Hochstätter, Agave I: 18 (2015). Type (neotype, designated by Ullrich, 1992: 32)]: ex cult., Anonymus, "n° 109 (22. V. 1891) in Herb. K". The neotype was prepared from the living plant figured on plate 7207 in Curt. Bot. Mag. (here reproduced as Figure 6) syn. nov. = Agave celsiana Cels, Rev. Hort. 1861: 335 (1861) syn. nov. (see notes below). = Agave concinna Hort. Tonel ex K.Koch, Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 5: 198 (1862) (nom. illeg., Art. 53.1, due to Agave concinna Lemaire, Hort.

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Figure 15. A large, high altitude population at about 3000m, growing in an open and exposed location as opposed to their usual shady and wet rock cliff habitats, together with Nolina parviflora. Besides the typical and common green-leaved plants, many plants with glaucous leaves occur. Agave mitis hybridises with A. montana which grows on the other side of the canyon [Querétaro, Cerro la Laja NE of Zimapán, 12.11.2014] Photograph: Paul Spracklin.

Vanhoutt. 1(2): 23 (1846)) = Agave macroacantha (as "macracantha") Zuccarini var. concinna (Hort. Tonel ex K.Koch) A.Terracciano, Prim. Contr. Monogr. Agave: 29 (1885) [(see notes below). = Agave oblongata Hort. Tonel ex K.Koch, Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 5: 198 (1862) ≡ Agave sartorii var. oblongata (Hort. Tonel ex K. Koch) A.Terracciano, Prim. Contr. Monogr. Agave: 23 (1885). = Agave densiflora Regel, Gartenflora 12: 274, t. 410 (1863) (nom. Illeg., Art. 53.1, due to Agave densiflora Hooker, Bot. Mag. 83: t. 5006 (1857)). = Agave albicans Jacobi, Hamburger Garten- Blumenzeitung 21: 256 (1865) ≡ Agave micracantha Salm-Dyck var. albicans (Jacobi) A.Terracciano, Primo Contr. Monogr. Agave: 25 (1885) ≡ Agave celsii var. albicans (Jacobi) Gentry, Agaves Cont. N. Amer.: 223 (1982). Type (neotype, here

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designated): Curt. Bot. Mag. t. 7207 (1891) (here reproduced as Figure 6). = Agave bouchei Jacobi, Hamburger Garten- Blumenzeitung 21: 217 (1865) ≡ Agave rupicola var. bouchei (Jacobi) A.Terracciano, Prim. Contr. Monogr. Agave: 24 (1885). = Agave celsiana Jacobi, Hamburger Garten- Blumenzeitung 21: 446 (1865) (nom. illeg., Art. 53.1, due to Agave celsiana Cels, Rev. Hort. 1861: 335 (1861)) [see notes below]. = Agave ehrenbergii Jacobi, Hamburger GartenBlumenzeitung 21: 255 (1865) syn. nov. (see notes below). = Agave goeppertiana Jacobi, Hamburger GartenBlumenzeitung 22: 219 (1866) syn. nov. (see notes below). = Agave haseloffii Jacobi, Hamburger GartenBlumenzeitung 22: 220 (1866). = Agave ousselghemiana Jacobi, Nachtr. 41 (1868).

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Figure 16. Agave mitis (“var. mitis”). On this rock face, green (above), green-glaucous (mid left) and whitish-glaucous forms (below) co-occur [Hidalgo, Parque Nacional Los Marmoles around 25km S of Jacala along MEX 85, 12.11.2014] Photograph: Paul Spracklin.

Figure 18. Agave mitis (“var. albidior”). Plants with narrow, alabaster-white leaves on the classical site in the Barranca de Metztitlán, accompanied by Agave xylonacantha (lower left) [Hidalgo, Barranca de Metztitlán ca. 6km S of Metztitlán, 11.01.2013] Photograph: Michael Greulich.

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Figure 17. Agave mitis (“var. mitis”) growing epiphytically in a cloud forest [Tamaulipas, at MEX 86 between Tula and Ocampo, 7.03.2013] Photograph: Bertus Spee.

= Agave micracantha var. picta J.Croucher ex R.Hogg, Florist & Pomol. 27: 135 (1874). = Agave botterii Baker, Bot. Mag. 102: t. 6248 (1876). = Agave macrantha Todaro, Hort. Bot. Panorm. 2: 11 (1879) ≡ Agave macracantha Zuccarini var. macrantha (Todaro) A.Terracciano, Prim. Contr. Monogr. Agave: 29 (1885). = Agave albicans var. variegata Hort. Berlin, Notizbl. Königl. Bot. Gart. Berlin 2(18): 335 (1899) (nom. inval., Art. 38.1) syn. nov. = Agave albicans var. ctenophora Trelease in L.H.BAILEy, Stand. Cycl. Hort. 1: 236 (1914). = Agave albicans var. mediopicta Trelease in L.H.BAILEy, Stand. Cycl. Hort. 1: 236 (1914). (The published rankless names for variegated cultivars listed by Trelease (1908) are omitted here). Illustrations (photographs and drawings, for plates, see Table 1; bwph = black & white photograph, colph = color photograph, hab = habitat; listing does not indicate exhaustiveness): Berger (1915: 62 [bwph], 63 [flower transection]; as A. mitis); van den Houten (1930: Figs. 2, 6, 7 [bwph; as A. haseloffii], Fig. 12 [bwph], Fig. 13 [bwph; as A. bouchei]; these plants cultivated at Rotterdam

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Figure 19. Agave mitis (“var. albidior”). These plants in the Barranca de Metztitlán show the variability in leaf colouration, ranging from glaucous (below) to light green (above), the latter matching typical plants of “var. mitis” (cf. Figures 15–17). Associated succulents: Mammillaria geminispina subsp. geminispina (left and lower right), Hechtia glomerata (above, narrow leaves), Agave xylonacantha (lower left), and Pachyphytum bracteosum (below centre) [Hidalgo, Barranca de Metztitlán ca. 1km S of Metztitlán, 11.01.2013]. Photograph: Michael Greulich.

Zoo may go back to the collection of the Dutch Agave collector van Ellemeet (1811–1888) which was dispersed 1874–75; see Thiede, 2014: 149); Breitung (1959: 46, as A. micracantha; 48: as A. micracantha & A. albicans; 90: as A. micracantha [all bwph]; Gentry (1982: 217 [flower transection], 220 [map], 222 [bwph hab], 223 [bwph]; as A. celsii); Ullrich (1993: 27, 30 [bwph]; as A. mitis var. albidior); Irish & Irish (2000: pl. 9 [colph]; as A. celsii); Smith (2002: 223 [colph]; as A. celsii var. albicans); Smith & Steyn (2002: 398 [drawings of habit, inflorescence, flowers, and fruit], 399 [bwph inflorescence, map]; as A. celsii var. albicans); Heller (2006: 108, 158, 160 [colph]); partly as A. mitis var. albidior); Greulich (2007: 32–36 [colph]; as A. ehrenbergii); Spracklin (2007: 148 [colph hab]); Scheinvar (2008: 48, 49 [colph hab], 50 [map]; as A. celsii var. albicans); Richter (2011: 36, 65, 122 [colph]); partly as A. mitis var. albidior); Starr (2012: 136–139 [colph hab]); Pilbeam (2013: 133–134 [colph hab]; partly as A. mitis var. albidior); Greulich (2014: 10 [colph hab epiphytic]); Hochstätter (2015: IX: 7 [colph hab]; as A. mitis ssp. albidior). Description: Rosettes branching axillary, forming large, long-lived, dense clumps; leaves ovate, oblong, or spatulate, ascending to outcurving, thickly soft-fleshy, short-acuminate, sometimes

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Figure 20 Agave mitis (“var. albidior”). Plants with broader, glaucous leaves in the Barranca de Metztitlán, accompanied by Ferocactus glaucescens (upper right) [Hidalgo, Barranca de Metztitlán ca. 6km S of Metztitlán, 11.01.2013] Photograph: Michael Greulich.

undulate, convex below, guttered or concave above, 30–60(–70) × 7–13cm, green to light greyglaucous, margins straight to sometimes +- wavy; marginal teeth small, frequently 2-tipped, sometimes with ciliate crests, 1–3mm, whitish to reddish-brown, closely spaced; terminal spine acicular, weak, 10–20mm, brownish, decurrent along the leaf tip for 1–6cm and more; inflorescence 1.5–2.5m, `spicate', (always?) from axillary branches below the terminal rosettes, peduncle with dense, chartaceous, deltoid, long-caudate, persistent bracts 1.5–7cm long, fertile part dense, becoming lax at fruiting time, partial inflorescences with geminate flowers; flowers fleshy, 40– 60 mm; tepals green outside, yellow to reddish or lavender to purplish within, tube funnel-shaped, thick-walled, 3-angled, 6-furrowed, 10–17 × 10– 12mm (at the apex), lobes linear-lanceolate, thickfleshy, ascending to recurving, shortly acuminate, cucullate, glandular-floccose at the tip, unequal, 12–18 × 5–8mm, inner with a broad fleshy keel; filaments 40–60mm, whitish with fine pale purple stripes, yellowish, brownish, or brownish purple, inserted at the tube apex; anthers mostly excen-

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Table 1. Agave mitis and its synonyms, arranged according to the year of publication. For each taxon, the varietal designation, the origins of the original material, the publication based on either either sterile rosettes (ros.) or flowering rosettes (flow.), and published plates are indicated. Year

Taxon / author

Variety

Origins

Ros./ flow.

Mexico, in warm regions (“in regione calida”)

ros.

A. rupicola Regel with var. brevifolia Regel, var. rubriden- mitis 1858 tata Regel & var. longifolia Regel

Mexico, introduced by Karwinski

ros.

1859 A. micracantha Salm-Dyck

mitis

ex. cult. Salm-Dyck, Dyck

ros.

A. micracantha var. albidior 1859 Salm-Dyck

albidior ex. cult. Salm-Dyck, Dyck

ros.

A. concinna Hort. Tonel ex 1862 K.Koch nom. illeg.

mitis

1848 A. mitis Martius

mitis

1856 A. celsii Hooker

mitis

1861 A. celsiana Cels

A. oblongata Hort. Tonel ex 1862 K.Koch A. densiflora Regel 1863 nom. illeg.

ex cult. Kew, obtained from flow. Cels, Paris

mitis

ex. cult. Cels, Paris

mitis

ex. cult. Tonel, Ghent

mitis

ex. cult. Tonel, Ghent

Mexico, introduced by Karwinski

1865 A. albicans Jacobi

albidior ex. cult. Salm-Dyck, Dyck

1865 A. bouchei Jacobi

mitis

1865 A. celsiana Jacobi nom. illeg.

mitis

1866 A. goeppertiana Jacobi

mitis

Plates

Baker (1877: 717 black & white woodcut)

Hooker (1856: Bot Mag. t. 4934, as part of the protologue hand-coloured copper engraving) (reproduced here as Figure 2)

[Baker's (1873) part-coloured lithograph of a flowering rosette labelled “Agave micracantha, Salm-Dyck” represents A. micracantha Baker ≡ A. flaccifolia A.Berger, synonyms of A. polyacantha Haworth]

flow.

Cels (1861: Fig. 32 & 33, as part of the protologue black & white lithograph)

ros.

(see also Smith & Figueiredo 2013: 56)

flow.

flow. ros.

Regel (1863: t. 410, as part of the protologue, as “Agave densiflora Hook.” part-coloured lithograph) (reproduced here as Figure 3)

Baker (1877: 717 black & white woodcut ros.) Baker (1891: Bot Mag. t. 7207, hand-coloured copper engraving flow.) (reproduced here as Figure 6)

Baker (1897a: Bot Mag. t. 7558, hand-coloured copper engraving) (reproduced here as Figure 8)

ex cult. BG Berlin, obtained from Sanssouci

flow.

mitis

Mexico, introduced by Ehrenberg

ros.

1866 A. haseloffii Jacobi

mitis

ex. cult. Haseloff, Berlin

ros.

Baker (1897b: Bot Mag. t. 7527, hand-coloured copper engraving) (reproduced here as Figure 7)

A. micracantha var. picta 1874 J.Croucher ex R.Hogg

mitis

ex. cult. Peacock, Hammersmith

ros.

(variegation with broad white median stripe)

1865 A. ehrenbergii Jacobi

1868 A. ousselghemiana Jacobi

protologue Hooker (1856) & ex. cult. ex cult. BG Breslau & ex cult. Saunders, Reigate

albidior ex. cult. Tonel, Ghent

Mexico, introduced by M. Botteri, sent to W.W. Saunders, Reigate

1876 A. botterii Baker

mitis

1879 A. macrantha Todaro

albidior ex cult. BG Palermo

A. albicans var. variegata 1899 Hort. Berlin nom. inval.

A. albicans var. ctenophora 1914 Trelease in L.H.Bailey

A. albicans var. mediopicta 1914 Trelease in L.H.Bailey

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mitis

ex cult. BG Berlin

albidior ex cult. albidior ex cult

flow. ros.

flow.

flow. flow. ros. ros. ros.

Baker (1876: Bot Mag. t. 6248, as part of the protologue hand-coloured copper-engraving) (reproduced here as Figure 4) Baker (1877: 265 black & white woodcut, ros.) Todaro (1879: t. xxvii, as part of the protologue coloured lithograph) (reproduced here as Figure 5)

(variegation with broad white median stripe) (variegation with broad white median stripe)

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Table 2. Length of the tepal lobes given for the two Agave mitis varieties in the literature, supplemented by measurements taken from digital images (di) of specimens available from the web. Name in original publication

Length of tepal lobes

Agave mitis Martius “var. mitis“

Reference

A. celsiana Cels

12mm

Cels (1861 protologue)

A. oblongata K.Koch

19mm (“9 L. [Linien] lang”)

Jacobi (1868: 40)

A. haseloffii Jacobi

A. ehrenbergii Jacobi A. mitis Salm-Dyck A. botterii Baker

A. mitis Salm-Dyck A. haseloffii Jacobi

A. micracantha Salm-Dyck A. rupicola Regel A. rupicola Regel

A. bouchei Jacobi

A. mitis Salm-Dyck

A. micracantha Salm-Dyck A. celsii Hooker A. celsii Hooker A. ousselghemiana Jacobi A. albicans Jacobi A. albicans Jacobi A. albicans Jacobi

15mm (“7 L. [Linien] lang”) 15mm (“7 L. [Linien] lang”) 22mm

Jacobi (1868: 44) Jacobi (1871: 82)

< 25mm (“nearly an inch long”) Baker (1876 protologue) < 25mm (“under 1 inch”)

Baker (1877: 717)

18–20mm

# Berger s.n. (US 1023670 di!)

13mm (“half an inch long”)

Baker (1897b)

# Berger s.n. (US 1023665 di!)

16–17mm

# Berger s.n. (US 1023666 di!)

15–16mm 16mm

Berger (1915: 58)

17mm

Berger (1915: 63)

20mm

Berger (1915: 66)

12–18mm

Gentry (1982: 218, 220)

13–16mm

# Garcia Mendoza 4127 (ASU 245167 di!)

Agave mitis “var. albidior (Salm-Dyck) Ullrich” 17mm (“8 L. [Linien] lang”)

Jacobi (1869: 41 protologue)

13mm (“half an inch long”)

Baker (1891)

19mm (“¾ inch”) c. 13mm

A. celsii var. albicans (Jacobi) Gentry 21–27mm

A. celsii var. albicans (Jacobi) Gentry 15–17mm A. celsii var. albicans (Jacobi) Gentry 16–17mm A. celsii var. albicans (Jacobi) Gentry 22–25mm

tric, 17–22mm, yellow, green(ish), red or brown, before opening yellowish, brown, purplish-brown or dark purple; ovary 3-angled, cylindrical, 6grooved, truncate at base and apex, 13–20mm, without neck; style stout, shorter, as long as or longer than Fil, fade and dotted with purple, reddened towards tip, purple, brown; stigma weakly 3-lobate; fruits angularly ovoid, thick-walled, 18– 25 × 9–12mm, rough, dark brown, beaked; seeds hemispherical, 3–4 × 2–3mm, black, with well- developed complete marginal wing. – Cytology: 2n = 60 (Cave 1964). Note: The considerable variability in the coloura-

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Jacobi (1866: 221 protologue)

Baker (1877: 717) Berger (1915: 65)

Gentry (1982: 218, 223)

# Garcia Mendoza 4740 (ASU 244548 di!) # Garcia Mendoza 5448 (ASU 245170 di!) Smith & Steyn (2002)

tion of tepals, filaments, anthers and styles is remarkable and may occur within the same population from plant to plant (Gentry, 1982: 223). Distribution (cf. map in Figure 9): Mexico (SE Coahuila, S Nuevo León, W & SW Tamaulipas, SW & S San Luis Potosí, NE Guanajuato, N & NE Querétaro, N & C Hidalgo, N Puebla); mostly on limestone outcrops, also in streamside thickets, in cloud forests, rarely epiphytic, 460–2440m; flowers March to July. Agave mitis ranges from arid desert scrub environments (Figures 18–20) and open mountain scrub habitats (Figures 15 & 16) up to shady and

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Table 3. Overview of the taxonomic and nomenclatural concepts for Agave mitis applied by Gentry (1982) and Ullrich (1993) and introduced in this paper. A. celsii var. celsii

A. celsii var. albicans

A. mitis var. mitis

A. mitis var. albidior

Leaves green to light grey glaucous Leaves pale glaucous Tepal lobes 12–18mm long Tepal lobes 20–27mm long Tamaulipas, San Luis Potosí, Hidalgo Barranca de Metztitlán Leaves dark to pale or yellow green Tepal lobes 12–18mm long Nuevo León, Tamaulipas, San Luis Potosí, Querétaro, Hidalgo “var. mitis”

Whitish leaf colour varies gradually Tepal lobes 13–27mm long Barranca de Metztitlán

A. mitis

Leaves dark green or light green to glaucous Tepal lobes 12–22mm long Coahuila, Nuevo León, Tamaulipas, San Luis Potosí, Guanajuato, Querétaro, Hidalgo, Puebla

“var. albidior”

Leaves alabaster-white or glaucous to light green Tepal lobes 13–19(–27)mm long Barranca de Metztitlán

humid rock cliffs (Figure 12). As mentioned by Thiede (2014: 155), A. mitis is not uncommon in cloud forests (Gentry, 1982: 222; Alcántara & Luna, 2001; Cartujano et al., 2002; all as A. celsii Hook.) where it may even occur as an epiphyte (Greulich, 2014: 10; Figure 17).

Taxonomic notes Agave celsiana Jacobi (1865: 446–448) is an illegitimate younger homonym of A. celsiana Cels (1861), a name apparently unknown to Jacobi. Jacobi's description is based on Hooker's (1856) protologue of A. celsii and thus a redescription of the latter, adding some data from young cultivated plants. A. celsiana Cels (1861) differs clearly from A. celsii Hooker (1856) / A. celsiana Jacobi (1865) in having lanceolate leaves with young teeth green and the oldest black, and flowers wine-reddish above and greenish below, and represents another variant of A. mitis. Agave concinna is ascribed to Baker in Gard. Chron., n.s., 7: 137 (1877) and represents an invalid later homonym of the earlier A. concinna Lemaire in Hort. Vanhoutt. 1(2): 23 (1846) (IPNI 2016; Govaerts 2016). Baker (l.c.) lists the name in the synonymy of A. albicans, thus rendering it invalid (ICN Art. 36.1.c; McNeill et al., 2012). However, A. concinna was published earlier with a short description by Koch (1862: 198) who considers it as an “A. celsiana Hook.”, a synonym of A. mitis. Agave ehrenbergii Jacobi (1865: 255–256) was published based on sterile plants with weak fleshy leaves which were introduced from Mexico by Ehrenberg and cultivated at Berlin. Jacobi com-

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Gentry (1982): new concept established; one species with two varieties Ullrich (1993): one species with two varieties upheld, but its status questioned

This paper: one species without varietal differentiation recognized

pared it with A. micracantha Salm-Dyck, a synonym of A. mitis. The inflorescence and flower descriptions for A. ehrenbergii were published six years later (Jacobi, 1871: 82–83). Thiede (2001) placed it among the unresolved names. Govaerts (2014) re-established it as earliest name for the Cuban A. legrelliana Jacobi (1866) of subg. Agave, apparently based on a note by Baker (1888: 188) that he (Baker) cannot separate it from the latter. Later, Govaerts (2016) noted his error and placed A. ehrenbergii among the unplaced names. Koch (1865: 187) mentioned that A. ehrenbergii belongs to A. mitis. Berger (1915: 41, 54) placed A. ehrenbergii between species most of which are now placed in the synonymy of A. mitis. Plants cultivated at BG Berlin labelled A. ehrenbergii of unknown origin but possibly dating back to original material belong to A. mitis (Greulich, 2007). Based on Jacobi’s and Berger's descriptions and further evidence, A. ehrenbergii is referred to the synonymy of A. mitis here. Agave goeppertiana Jacobi (1866: 219–220) was published based on sterile plants with soft, grooved leaves cultivated at the Botanic Garden Breslau and in Saunders' collection. In the following year, Jacobi (1867: 355–360) described the inflorescence and flowers, adding that it was first thought to represent A. bouchei and that the seeds are most similar to A. haseloffii, two names now placed in the synonymy of A. mitis. Berger (1915: 41, 54–56) published a photograph of a flowering plant at Breslau and placed the name among species now mostly referred to the synonymy of A. mitis. Thiede (2001: 12) and Govaerts (2016) placed A. goeppertiana among the unresolved

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names. The descriptions of Jacobi and Berger match A. mitis, and Berger's photograph matches those of A. mitis and its synonyms published by van den Houten (1930, see above) which partly show similar grooves on the leaves. Based on the above evidence, A. goeppertiana is referred to the synonymy of A. mitis here.

Acknowledgements The author is indebted to Michael Greulich (D), Paul Spracklin (UK) and Bertus Spee (NL) for providing photographs and additional information, to an anonymous reviewer for his comments, and to Graham Charles (UK) for his editorial work.

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