A Reevaluation of the Agave Kerchoveicomplex

60 Haseltonia 27: 60–72. 2020 A reevaluation of the Agave kerchovei complex (Asparagaceae/Agavoideae) including expanded descriptions of three species Greg D. Starr, The Southwest Center College of Social and Behavioral Sciences, University of Arizona, Tucson, AZ 85719 USA. email [email protected] Manuscript received 20th November 2019

Abstract: The taxonomy of the Agave kerchovei complex in the states of Puebla and Oaxaca is revisited, ana- lyzed, and discussed, and the status of the names considered to be synonyms is explored and discussed. The status of names synonymized with A. kerchovei (A. convallis, A. dissimulans) is discussed considering new field data from the type localities, with the confirmation of A. convallis as a species as put forth by Garcia-Mendoza and reinstatement of A. dissimulans as a recognized species. Keywords: Agave convallis, Agave dissimulans, Agave kerchovei, Agavaceae, Heteracanthae, Puebla, Oaxaca, Tehuacán-Cuicatlán Valley

INTRODUCTION mine if they were indeed distinct or simply within Two nineteenth century European taxonomists, the range of variation for other species. That later au- Joseph Salm-Dyck and Georg Albano von Jacobi, de- thor appeared in the guise of Howard Scott Gentry. scribed several Agave species using cultivated plants After 30+ years of traveling throughout continen- sent from Mexico, many without specific locality tal North America, researching old names, tracking information (for example Agave attenuata Salm- down localities, and sorting out the application of Dyck, Agave xylonacantha Salm-Dyck, Agave asper- these names, Gentry (1982) provided his monograph rima Jacobi, Agave chiapensis Jacobi). John Gilbert of the genus and the application of many old names. Baker (Kew, UK), another important nineteenth In some instances, Gentry recognized the distinctive- century botanist, also described several Agave species ness of certain agaves and described several new spe- without known localities (for example Agave maxi- cies, while in others, he grouped two or more very miliana Baker, and Agave warelliana Baker). It was distinct taxa under one umbrella species name, thus Berger (1915) who first compiled and organized the considering that species to be highly variable. genus Agave into a complete monograph in which The latter scenario was the case for Agave kercho- he included 274 species divided into three subgen- vei, a species Gentry apparently felt was highly vari- era, Manfreda, Littaea, and Euagave which is now able based on his inclusion of five previously recog- recognized as an invalid name (Thiede et al. 2019). nized species as synonyms, creating what is referred to None of these taxonomists had visited the native here as the “Agave kerchovei complex”. Those species habitats of the species they described or wrote about, are: Agave convallis, Agave dissimulans, Agave inopina- and worked with extremely limited plant material, bilis (all described by Trelease in 1920), Agave expa- thus they did not consider the variation within a spe- triata (described by Rose in 1900), and Agave noli- cies; the result was describing many species based on tangere (described by Berger in 1915). After extensive minor variations in morphology. Essentially the first fieldwork, new data have been gathered that addresses field botanist to publish on the genus was William inconsistencies with the accepted definition of Agave Trelease (1920) when he wrote the Agaves of Mexico kerchovei and sheds light on the validity of two names for Standley’s ‘Trees and Shrubs of Mexico’. Trelease that have been sunk into synonymy in Gentry (1982). did field work prior to describing most species and understood many of the factors affecting diagnosable population parameters (e.g. morphological variation, THE HISTORY OF AGAVE distribution, and soil type). Although Trelease trav- KERCHOVEI LEMAIRE eled in Mexico and collected and described several 1864 — LEMAIRE: THE ORIGINAL species of Agave, his publication (Trelease 1920) did DESCRIPTION OF AGAVE KERCHOVEI not address earlier names and their application to de- (FIGURES 1–8) termine whether any should be relegated to synonymy or left on their own. Lemaire (1864) provided the first description It was left to later authors to research the many of Agave kerchovei using a juvenile plant imported names applied to earlier-named species and deter- directly from Mexico by Jean Verschaffelt of the re- Author Copy HASELTONIA 27. 2020 61

Figure 1. Dark green leaved Agave ker- Figure 2. Classic form of Agave kercho- Figure 3. Agave kerchovei growing in chovei lacking marginal teeth in the vei with its long triangular leaves as seen the semi-arid desert southwest of Tehu- upper one-third of the leaf. along Mexico Highway 125 between Te- acán, Puebla along Mexico Highway 125 huacán, Puebla and Huajuapan de León, showing the lack of teeth in the upper Oaxaca. one-half of the leaf. nowned Verschaffelt family of Belgian horticulturists. of A. kerchovei, establishing a clearer picture of the According to Baker (1877) and Thiede et al. (2020), taxon. He explained that his plant was not quite Lemaire chose to honor Baron Gustave-Phillipe fully grown and measured about 40 cm tall and 60 de Kerchove van Ousselghem by giving this plant cm across. He described the leaves as widest at the the specific epithet of kerchovei. In his treatment of base and tapering gradually to the tip, which fits the the genus Agave, Baker (1877) included a detailed typical look of plants in habitat (Figs. 1–8). Berger illustration while also noting that the plant was used an additional two specimens, one collected by widespread in collections with at least half a dozen H. Schenck (No. 211) from the Cerro de la Yerva at different forms in Mr. J. T. Peacock’s collection San Luis Tultitlanapam in the Sierra de Mixteca in (Hammersmith, UK) in 1872. Puebla and the other by C.A. Purpus (No. 415) at Tehuacán in Puebla, in his inflorescence and flower TAXA RECOGNIZED BY LEMAIRE description. In this monograph, Berger recognized Although Lemaire (1864) used a juvenile plant the close affinities of A. expatriata Rose and A. noli- for his description, there were other forms in cultiva- tangere A. Berger to A. kerchovei by including these tion and he noted three varieties based on minor dif- three taxa in his section Pericamptagave A. Berger. ferences in marginal teeth, leaf shape, and leaf mar- TAXA RECOGNIZED BY BERGER gin; these three varieties were listed by Berger (1915) but fell out of favor by the time Gentry published Berger did recognize the varieties of Lemaire and his monograph and are currently not recognized: included A. beaucarnei Lemaire as a synonym of A. kerchovei: • A. kerchovei var. macrodonta Lemaire — • Agave kerchovei Lemaire with much larger spines (teeth) rather point- Synonym: A. beaucarnei Lemaire ed up the middle to strongly curved. * A. kerchovei var. macrodonta Lemaire • A. kerchovei var. diplacantha Lemaire — * A. kerchovei var. diplacantha Lemaire with leaves elongate and much more pointed * A. kerchovei var. distans Lemaire (presumably he meant the apex was more at- • A. expatriata Rose tenuate). • A. noli-tangere A. Berger • A. kerchovei var. distans Lemaire — hav- 1920 — WILLIAM TRELEASE ing brown margins and the teeth 8–90 mm CONTRIBUTIONS ADD TO THE GROWING apart. COMPLEX 1915 — ALWIN BERGER EXPANDS THE The next major treatment of the genus Agave was DESCRIPTION undertaken by Trelease (1920) for Trees and Shrubs of Mexico compiled by Paul C. Standley. Although Berger (1915) used a larger plant than Lemaire Trelease was an astute student of the genus, spending had and provided a more detailed description time in the field studying the plants and their nuances, Author Copy 62 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Figure 5. A dark, olive green leaved Agave kerchovei lacking teeth in the upper one-half of the leaf as seen on the arid hills near Tehuacán, Puebla.

Figure 4. Agave kerchovei with leaves a bright yellow green on the upper surface and rust brown on the lower surface showing the typically widely spaced marginal teeth.one-half of the Figure 6. The long triangular leaves of Agave kerchovei as seen leaf. in the semi-arid desert near Zapotitlan, Puebla.

Figure 7. The tall inflorescence ofAgave kerchovei showing Figure 8. Leaf detail of Agave kerchovei showing the lack of the flowers flushed with red. teeth in the upper one-third of the blade. Author Copy HASELTONIA 27. 2020 63

Figure 9. Note the dense rosette of Figure 10. A green leaved plant and Figure 11. The attractive dual colors on Agave convallis at the type locality with two red leaved plants of Agave convallis Agave convallis near San Jerónimo south its lanceolate to narrowly oblanceolate as seen near San Jerónimo south of El of El Parián. leaves and teeth to nearly the tip. Parián.

he may have, on occasion, had too narrow a definition of what constituted a species. Might that have been the case when, in 1920, he described three new species that clustered near A. kerchovei (Figs. 1–8) in his key to species? Gentry must have considered this question when he relegated all three to synonymy under A. kerchovei. These three —A. inopinabilis, A. convallis (Figs. 9–18), and A. dissimulans (Figs. 19–23) — were all collected from near existing railroad stops along the route from Tehuacán to Oaxaca. Apparently Trelease did not keep a field notebook, but it is safe to conclude that he rode this line on at least two separate occasions (1903, 1905) based on his agave collections. TAXA RECOGNIZED BY TRELEASE Because Berger’s Die Agaven was published dur- ing World War I, Trelease was unaware of his description of A. noli-tangere, thus it was not included in his treatment of the genus. Trelease (1920) described three additional species in the A. kerchovei complex and recognized the following five species: • Agave kerchovei Lemaire Synonym: A. beaucarnei Lemaire • A. convallis Trelease • A. dissimulans Trelease • A. expatriata Rose • A. inopinabilis Trelease 1982 – HOWARD SCOTT GENTRY REVISES THE CONCEPT After 30+ years of field work involving numerous collections, Gentry (1982) provided a detailed description for his concept of A. kerchovei. In general, Gentry had a broad species concept, and he considered A. kerchovei to be an extremely variable species encompassing several morphological forms with a Figure 12. Flowering Agave convallis plants near San Jeróni- distribution ranging from central Hidalgo south to mo south of El Parián, showing affinities to A. kerchovei. Author Copy 64 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Figure 13. A stout specimen of Agave convallis in the moun- Figure 14. Agave convallis in the mountains near San Pedro tains near San Pedro Jaltepetongo above Tomellín Canyon Jaltepetongo above Tomellín Canyon proper showing a faint proper. central stripe on its leaves.

Figure 15. A very colorful Agave convallis near San Pedro Figure 16. Agave convallis with bicolored leaves showing a Jaltepetongo above Tomellín Canyon proper. distinct central stripe found southeast of Huajuapan de Léon.

Figure 18. A colorful young Agave convallis with orange-red Figure 17. A young Agave convallis with purplish red leaves leaves found above San Pedro Jaltepetongo above Tomellín seen southeast of Huajuapan de Léon. Canyon proper. Author Copy HASELTONIA 27. 2020 65

Figure 19. Several Agave dissimulans with their drooping, Figure 20. A drooping-leaved Agave dissimulans and one of blue glaucous leaves at the type locality of Cerro Campana. the rare plants with more erect leaves at the type locality of Cerro Campana. southern Oaxaca. He thus lumped all the previously TAXA RECOGNIZED BY GARCÍA-MENDOZA described taxa into his expanded concept of A. kerchovei. • Agave convallis Trelease Synonym: A. dissimulans Trelease TAXA RECOGNIZED BY GENTRY • Agave kerchovei Lemaire Gentry (1982) primarily used flower characteris- Synonyms: A. beaucarnei Lemaire, A. tics to cluster similar species into “groups” - an infor- expatriata Rose, A. inopinabilis Trelease, A. mal taxonomic designation for an amalgamation of noli-tangere A. Berger related species. In his group Marginatae (now section Heteracanthae Salm-Dyck; Thiede et al., 2019), Gen- try included 21 species, including A. kerchovei with A DETAILED ANALYSIS OF the following synonyms: SPECIES NAMES ASSOCIATED • Agave kerchovei Lemaire WITH AGAVE KERCHOVEI Synonyms: A. convallis Trelease, A. TRELEASE COLLECTIONS OF AGAVE dissimulans Trelease, A. expatriata Rose, A. CONVALLIS, AGAVE DISSIMULANS, AND inopinabilis Trelease, A. noli-tangere A. Berger AGAVE INOPINABILIS 2011 — ABISAÍ GARCÍA-MENDOZA REVISITS The William Trelease story is an essential center- AGAVE CONVALLIS piece to the A. kerchovei species complex. As noted earlier, Trelease traveled on the train between Te- In his work on the Agave family of the Tehuacán- huacán and Oaxaca at least twice, collecting agave Cuicatlán Valley of southern Puebla and northern specimens at three stops along the way. In August Oaxaca, García-Mendoza (2011) considered A. con- 1903, Trelease collected an agave from near Tehuacán, vallis to be distinct from A. kerchovei and restored which he initially called A. vanderdonckii hort. Ex it to species status. He listed several important and Baker but later described it as A. inopinabilis. consistent characteristics that differentiateA. con- Trelease separated A. inopinabilis with its leaves vallis from A. kerchovei. Among them, A. convallis spreading, oblong, thin, or else fleshy rather than has a greater number of leaves that are shorter and hard, from A. kerchovei with its leaves often falcate, broadly lanceolate, with a shorter length to width ascending, thick, and stiff. The area around Tehu- ratio, and teeth continuing closer to the tip. Agave acán has been quite popular with agave collectors as convallis also occurs at a higher elevation in oak for- it was accessible by train and very rich with arid veg- est. García-Mendoza noted that the two specimens etation, including a cluster of agaves in what is now of A. dissimulans cited as syntypes correspond more considered the section Heteracanthae. closely with A. convallis than with A. kerchovei. He, Trelease’s second journey on the train between Te- therefore, without visiting the type locality, moved A. huacán and Oaxaca occurred in February 1905. It is dissimulans into synonymy with A. convallis. unclear whether he made his collections while travel- Author Copy 66 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Figure 21. A hillside full of Agave dissimulans with drooping, Figure 22. Another Agave dissimulans with stiff, erect blue blue glaucous leaves on Cerro Campana. glaucous leaves. Note the teeth are present nearly to the leaf tip.

ing from Tehuacán to Oaxaca or from Oaxaca to Te- huacán as his collection number of 4 was made on February 12, 1905 and his two collections numbered 81 and 82 were dated from February 11, 1905. Near the El Parián train station (Fig. 24), Trelease collected a plant he described as A. convallis. He grouped it close to A. inopinabilis and A. expatriata, all three with thin, fleshy leaves, yet separate from the thick and stiff leaved A. kerchovei. In the key to species, Trelease separated A. convallis with “spike very com- pact” from A. expatriata with “spike rather loose”. Trelease collected the plant he later described as A. dissimulans from near the stop of Mexia north of El Parián and south of San Antonio Nanahuati- pam in northern Oaxaca. At the time Trelease rode the train, there were two stops: Ignacio Mejía and Mexia, within about 3 kilometers of each other. Al- though the train no longer runs, tracks are still vis- ible through the overgrown vegetation, and there Figure 23. A hillside full of flowering Agave dissimulans with are still signs of the station in Ignacio Mejía and the drooping leaves making the plants look like octopuses on the hill. station at the stop that would have been Mexia (Fig. 25). The tracks run alongside the Río Salado from San Antonio Nanahuatipam to near the switching station at Mexia where the Río Salado joins the Río Hondo near Cerro Campana, the hill where Trelease likely collected A. dissimulans. I can picture William Trelease sitting at a window seat scanning the hills as the train sped along the tracks, anxious to reach a stop, so he could get out and wander about look- ing for unusual agaves. February is the height of the dry season in that part of Oaxaca, and the tropical, deciduous forest lives up to its name, the bulk of the vegetation being devoid of leaves rendering the envi- ronment an ideal time for spotting agaves. Because Trelease collected specimens in habitat Figure 24. The now abandoned and unused train station at from known localities, those three taxa can be exam- El Parián. Author Copy HASELTONIA 27. 2020 67

ther south along Mexico Federal Highway 125, but they are consistent in critical characteristics and do not encompass more than the one taxon (Figs. 1–5). There are five sheets of A. inopinabilis housed at the Missouri Botanical Garden: four with leaves collected by Trelease and one with a photograph of a potted plant. None of the specimens have Trelease collection numbers; however, all except the sheet ac- cessioned as number 3346890 are classic A. kerchovei. It is unfortunate that García-Mendoza (2011) selected this outlier specimen when he lectotypified A. inopinabilis. This specimen was collected north of Tehuacán and has a wider than normal margin with heavier teeth and interstitial teeth not usually found on typical A. kerchovei leaving some wiggle room for the interpretation of the definition for A. inopinabilis. However, the other four specimens fit neatly into A. kerchovei, and A. inopinabilis is best left in synonymy there until plants north of Tehuacán can be examined more thoroughly. Figure 25. The author at the remains of the train station at Mejia Viejo near Cerro Campana and the type locality for Agave dissimulans. AGAVE CONVALLIS ined with a more discerning eye by visiting the type TRELEASE (FIGURES 9–11) localities, which enables us to understand Trelease’s Although not recognized as a species by Gentry reasoning for describing them. Through the University (1982), Agave convallis was recognized as distinct by of Arizona herbarium, a loan of the Trelease specimens García-Mendoza (2011). García-Mendoza collected of A. convallis, A. dissimulans, and A. inopinabilis was Agave convallis west of the train station at El Parián, requested from the Missouri Botanical Gardens for and amplified Trelease’s original description. Figures firsthand examination. Although pressed specimens 9–12 illustrate that the plants in the area around El of agaves are notoriously vague at times, viewing the Parián have solitary, dense rosettes of numerous, lan- actual specimens is far more informative than viewing ceolate to broadly lanceolate, medium green leaves scanned images on-line and is another integral meth- that are frequently flushed red or nearly all red, and od of determining the parameters of taxa. with marginal teeth occurring nearly to the apex. While, as seen in Figs. 1–8, Agave kerchovei has open FOLLOWING IN THE FOOTSTEPS OF TRELEASE rosettes of few, long triangular, dark olive-green, or The question surrounding the status of A. con- rarely lighter yellow-green leaves lacking marginal vallis, A. dissimulans, and A. inopinabilis is one that teeth in the upper one-third of the blade. persists because of the presumed variability in Agave In his section on Agave convallis, García-Mendoza kerchovei as defined by Gentry’s concept of the spe- (2011) cited Gentry specimens (Gentry 20278 and cies (Gentry 1982). García-Mendoza (2011) par- Gentry 22400 [both MEXU, US]) that were collect- tially resolved the issue by recognizing A. convallis ed about 14 miles southeast of Huajuapan de Léon as a legitimate species but included A. dissimulans as along Highway 190 and identified by Gentry as A. a synonym of A. convallis and not of A. kerchovei as kerchovei. Figures 16 & 17 show plants at the same Gentry had. Recently, Smith, et. al. (2018) clarified locality and confirm their similarity to those seen at the typification of the names Agave convallis, Agave and near the type locality of El Parián. dissimulans, and Agave kerchovei. To determine the The train station at El Parián is in the Tomellín validity of each name and their proper assignation, Canyon and plants of Agave convallis are found on it was necessary to visit the localities where Trelease the cliffs and canyon walls both north and south of El traveled and collected, which is what I accomplished Parián and its neighbor Faustino G. Olivera. Figures on several trips to the critical areas from 2015–2018. 10–12 show plants of Agave convallis that occur north of San Jerónimo, which is 4 kilometers by air south- AGAVE INOPINABILIS TRELEASE west of El Parián. Note the plants with red leaves or those flushed with red. This is a characteristic com- Trelease described A. inopinabilis from the area monly seen on this species but not on Agave kerchovei. around Tehuacán, and Gentry listed it as a synonym Another locality for Agave convallis documented of A. kerchovei, so it was imperative to visit the semi- by García-Mendoza (2011) is north of San Gabriel arid hills around Tehuacán to study the Heteracan- Almoloyas which is north of El Parián on the west thae agaves found there. Agave kerchovei is uncom- side of Tomellín Canyon. Figures 13–15, 18 show mon in the immediate vicinity of Tehuacán, and it plants in the vicinity of Jaltepetongo which is locat- is doubtful that more than one look-alike species ed just north of San Gabriel Almoloyas and about 21 occurs there. It does become more common far- kilometers by air from El Parián. Author Copy 68 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Plants at all these localities are very consistent in pressed name), therefore eliminating any potential their numerous, lanceolate to broadly lanceolate, me- confusion and the use of the name. dium green leaves that are frequently flushed red or Berger described Agave noli-tangere as being cul- nearly all red, and easily distinguishable from A. ker- tivated in La Mortola (Italy) for many years as Agave chovei with fewer leaves that are thick and stiff, and horrida. However, he stated that it differed from lack marginal teeth in the upper third. Agave horrida by the narrow and long-pointed leaves that lack teeth in the upper part, which is indicative of Agave kerchovei. Without live specimens or origi- AGAVE DISSIMULANS nal localities and based solely on the descriptions and TRELEASE (FIGURES 19–23) vouchers, Agave noli-tangere is best left as a synonym Agave dissimulans is another species in the Agave of Agave kerchovei. kerchovei complex described by Trelease that was collected from near the train station at Mexia, which is near the station at Ignacio Mejía. The website EXPANDED DESCRIPTIONS http://www.railwaystationlists.co.uk/ lists both Igna- OF RECOGNIZED SPECIES cio Mejía and Mejía Viejo (Mexia) as separate stops Considering recent field work, with visits to the along the rail line. Ignacio Mejía was visited in Feb- known type localities and other documented localities ruary 2017 and a guide hired to visit the spot where for the species names included by Gentry as synonyms the station of Mexia would be. Our guide took us of Agave kerchovei, it is apparent that there are three to a hill he called Cerro Campana (The Bell), with closely related species in north-central Oaxaca and a dense population of agaves with long, drooping, southern Puebla: Agave convallis Trelease, Agave dis- blue leaves (Figs. 19–21, 23). Herbarium vouchers simulans Trelease, and Agave kerchovei Lemaire. were collected and deposited at ARIZ. More specimens were collected on a second visit to Cerro Cam- Agave convallis Trelease, Contributions from pana and are deposited at ARIZ also. The abandoned the U.S. National Herbarium 23(1): 138. 1920. Lec- train stop at Mexia appeared to be little more than a totype: Mexico. Oaxaca: El Parián, Tomellín Can- switching yard with no evidence of an actual depot yon, W. Trelease 4, 5 February 1905 (MO), (MO or civilization. However, Cerro Campana was vis- barcode: MO-1128755; MO specimen number ible from the stop, and Trelease must have seen the 3346866, online image at: http://www.tropicos. plants and decided to disembark at that point to org/Image/26009!); designated by García-Mendoza collect specimens. Trelease (1920) described them as (2011: 22) (Figs. 9–18). having “leaves widely and flaccidly spreading or re- ADDITIONAL SPECIMENS EXAMINED curved”, which was a bit of an understatement. Most of the plants had leaves that could best be described Mexico: Oaxaca: 2 miles past San Jerónimo So- as drooping (Figs. 19–21), while a small percentage sola, road to El Parián, 6250 feet elevation. Starr had leaves that could be described as widely spread- 2017-004, 11 February 2017 (ARIZ). Oaxaca: 2 ing (Fig. 22). Additional data were collected and miles past San Jerónimo Sosola, road to El Parián, have been incorporated into an expanded descrip- 6250 feet elevation. Starr 2018-028, 20 January tion for Agave dissimulans, which is highly distinct 2018 (ARIZ). Oaxaca: 2 miles past San Jerónimo from both A. convallis and A. kerchovei. Based on Sosola, road to El Parián, 6250 feet elevation. Starr this new information, this distinctive taxon is taken 2018-029, 20 January 2018 (ARIZ). Oaxaca: 2 out of synonymy with both A. kerchovei (Gentry miles past San Jerónimo Sosola, road to El Parián, 1982) and A. convallis (García-Mendoza 2011) and 6250 feet elevation. Starr 2018-030, 20 January returned to species status based on distribution and 2018 (ARIZ). Oaxaca: 2 miles past San Jerónimo leaf morphology as illustrated in the key to species. Sosola, road to El Parián, 6300 feet elevation. Starr See Smith et. al. (2018) for a discussion of the lecto- 2018-031-01, Starr 2018-031-02, Starr 2018-031-03, typification of Agave dissimulans. 20 January 2018 (ARIZ). Oaxaca: 0.6 mile north of El Parián, Tomellín Canyon, 4820 feet elevation. AGAVE EXPATRIATA ROSE AND AGAVE NOLI- Starr 2018-032, 20 January 2018 (ARIZ). Oaxaca: 4 TANGERE A. BERGER airmiles south of San Pedro Jaltepetongo, 7025 feet elevation. Starr 2018-034, 21 January 2018 (ARIZ). Gentry (1982) also included Agave expatriata Oaxaca: 4 airmiles south of San Pedro Jaltepetongo, Rose and Agave noli-tangere A. Berger as synonyms 7050 feet elevation. Starr 2018-035, 21 January of Agave kerchovei. Rose (1900) described Agave ex- 2018 (ARIZ). Oaxaca: 4 airmiles south of San Pedro patriata based on a plant of unknown origin being Jaltepetongo, 7050 feet elevation. Starr 2018-036, 21 cultivated as Agave brauniana (a synonym of A. ob- January 2018 (ARIZ). Oaxaca: 14 miles SE of Hua- scura Schiede, = A. polyacantha Haw.) at the Botani- juapan along the highway, elevation about 6000 feet. cal Garden in Washington. Because Agave expatriata Gentry 22400, 4 November 1967 (MEXU, US). was based on one specimen of unknown origin with a description that could fit any number of species in DESCRIPTION Heteracanthae, Smith & Starr (2019) proposed re- Plants: generally solitary or rarely cespitose and jecting the name as nomina utique rejicienda (sup- forming small colonies; medium size, acaulescent, Author Copy HASELTONIA 27. 2020 69

Map 1. The distribution of the three species in the Agave kerchovei complex: Agave convallis A. dissimulans & A. kerchovei . moderately densely leaved, to 45–70 cm tall and 75– turning gray or white with age. Inflorescence: the 120 cm across. Leaves: 30–80 per rosette, uniformly whole flower stalk is 3–5 m tall with the fertile sec- spreading, light green to dark green, sometimes with tion 1.5–3.6 m long. Flowers: in pairs, tepals are yel- red, purple-red, or brownish red on the edges or low to greenish yellow, sometimes green and flushed tips, occasionally entirely red or purplish red, usually with red or purplish red; overall length is 35–50 with a prominent, light green or yellow-green cen- mm, ovary 12–20 mm long, neck is 3–8 mm long, tral stripe; 30–60 cm long by 6–10 cm wide; broad- tube is 2–4 mm long by 5–8 mm wide, tepals are est from base to near the middle or narrowed at the 15–24 mm long by 3–8 mm wide, filaments are yel- base and broadest near the middle, linear-triangular- low or reddish and showy. lanceolate, lanceolate to broadly lanceolate; rigid, thick and succulent at the base; flat to concave to- DISTRIBUTION wards the apex above, convex below; margin straight, (MAP 1) corneous, white or grayish. Teeth: small to medium, Agave convallis was first collected in 1905 by Wil- 2–10(–20) mm long, 10–60 mm apart, straight, an- liam Trelease from Tomellín Canyon, near the train gled back toward base, or variously flexed or curved, station at El Parián in northern Oaxaca. In 2011, chocolate brown to reddish brown when young, be- Abisaí García-Mendoza listed specimens from near coming white or grayish with age, present nearly to San Gabriel Almoloyas in the mountains north of El tip. Terminal spine: awl-like, shallowly to deeply Parián and along Mexico Federal Highway 190 be- grooved above, rounded to ridged below, 2–3 cm tween Asunción Nochixtlán and Huajuapán de León. long, chocolate brown or reddish brown when young, Agave convallis occurs in the districts of Cuicatlán, Author Copy 70 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Huajuapán, Nochixtlán, and Teposcolula in Oaxaca, the old train station at Mexia, 2400 feet elevation, and is reported from Cerro de la Yerba at San Luis in Starr 2018-043, 23 January 2018 (ARIZ). Oaxaca: Puebla, which is the locality listed by Berger for the Cerro Campana at Cañon Oscuro, 2.1 airmiles south Agave kerchovei specimen collected by H. Schenck. of Ignacio Mejia, near the old train station at Mexia, Having visited several of the localities listed by 2400 feet elevation, Starr 2018-044, 23 January García-Mendoza, the plants were found to be dis- 2018 (ARIZ). Oaxaca: Cerro Campana at Cañon tinct from Agave kerchovei. While Agave kerchovei has Oscuro, 2.1 airmiles south of Ignacio Mejia, near the open rosettes of dark green leaves that are broadest old train station at Mexia, 2420 feet elevation, Starr near the base and long tapering to the tip, Agave con- 2018-045, 23 January 2018 (ARIZ). vallis has denser rosettes of medium to bright green leaves, frequently flushed with red or purplish red, or DESCRIPTION even all red, and the same width from the base to the Plants: medium to large, solitary, acaules- middle or above and then quickly tapering to the tip. cent, sparsely leaved rosettes; 50–120 cm tall by They appear to favor steep, rocky, shaded cliff faces 100–150 cm across. Leaves: 15–40 per rosette; although they occasionally can be found growing on blue-glaucous, rarely with a faint light yellow-green open, exposed slopes and in steep, narrow drainages. mid-stripe; smooth to very lightly asperous; mostly They can be found growing among thorn scrub and flaccid, drooping, recurved, sinuous, rarely straight in pine-oak forests 1420–2250 meters (4920–7375 and widely spreading; 65–110 cm long by 4–9 cm feet). wide; broadly linear to long linear-triangular; thick and succulent at the base; flat to concave above, con- DISCUSSION vex below; margin straight, horny, white or grayish. Agave convallis was described by Trelease in 1920 Teeth: small to medium, 1–13 mm long, 10–60 mm and later included in with Agave kerchovei by Gentry apart; chocolate brown when young, becoming white (1982). However, it is distinctive and easily separable or grayish with age. Terminal spine: awl-like; 2–4.5 by the rosette density, leaf shape, and leaf color as cm long; chocolate brown when young turning gray noted above. Generally, the two are not found grow- or white with age. Inflorescence: total flower stalk ing together; however, I collected a leaf voucher from length is 3–4.5 m, the fertile section 1.3–2.4 m one sterile plant that looked much like Agave conval- long. Flowers: campanulate; tepals are green on the lis in a side canyon near Puente Santa Lucia along outside, sometimes flushed with a light blush of red- Mexico Federal Highway 135D. There were a few dish purple, the inside of the tepals are light green at plants of Agave kerchovei and Agave titanota growing the base and reddish purple toward the tip; overall in this side canyon as well. I believe Agave convallis length is 37–44 mm, ovary 15–18 mm long, neck accounts for much of the diversity of Agave kerchovei is 4–5 mm long, tube is 1–2 mm long by 5–8 mm as defined by Gentry. wide, tepals are 16–23 mm long by 7–9 mm wide, filaments are yellowish green to dark reddish purple Agave dissimulans Trelease: Contributions and are very showy even from a distance. from the U.S. National Herbarium 23(1): 138. 1920. Lectotype: Mexico. Oaxaca: [Ignacio Mejía] Mexia DISTRIBUTION (MAP 1) (toward Tomellín), Trelease 81, 11 February 1905 (MO), (MO barcode: MO-1128757; MO specimen Agave dissimulans has a restricted distribution number 3346870) [online image at: http://www. and is known only from the Cerro Campana and tropicos. org/Image/26018!]; designated by García- adjacent hills in the Cañon Oscuro south of Ignacio Mendoza (2011: 22). Isolectotype: Mexico. Oaxaca: Mejía along the Rio Salado and near the old train Mexia, Trelease 81, 11 February 1905 (MO), (MO tracks. William Trelease first collected this in Febru- barcode: MO-1128758; MO specimen number ary 1905 with no other specimens being vouchered 3346869) [online image at: http://www.tropicos.org/ until the Starr collections in February 2017 and Image/26019!; designated by García-Mendoza (2011: January 2018 from the same locality (see the above 22). Syntype: Mexico. Mexia, Trelease 82, 11 Febru- section on additional specimens examined). Plants ary 1905 (MO), (MO barcode: MO-1256273; MO are found mostly on the rocky north and east slopes specimen number 3346871) [online image at http:// above the river bottom with some found on a south- www.tropicos.org/Image/26020]. (Figs. 19–23) southeast slope in a side drainage, all occurring between 2200–3000 feet elevation. ADDITIONAL SPECIMENS EXAMINED DISCUSSION Mexico: Oaxaca: Cerro Campana at Cañon Oscuro, 2.1 airmiles south of Ignacio Mejia, near Agave dissimulans has an interesting past, first the old train station at Mexia, 2275 feet elevation, being described as a new species, then included in Starr 2017-008, 15 February 2017 (ARIZ). Oaxaca: Agave kerchovei, to which it is certainly related, then Cerro Campana at Cañon Oscuro, 2.1 airmiles south removed from there and included in the concept of of Ignacio Mejia, near the old train station at Mexia, Agave convallis, another species to which it is cer- 2350 feet elevation, Starr 2018-042, 23 January tainly related, and now finally coming full circle and 2018 (ARIZ). Oaxaca: Cerro Campana at Cañon here being recognized again as a valid species. The Oscuro, 2.1 airmiles south of Ignacio Mejia, near saga of Agave dissimulans is a reminder that closely Author Copy HASELTONIA 27. 2020 71

DISTRIBUTION related agaves can be difficult to separate based only (MAP 1) on herbarium specimens, and diligent field work is needed to determine whether a population or se- Agave kerchovei is found in open, exposed loca- ries of populations warrant recognition as a species tions in the semi-arid desert of northern Oaxaca and separate from other, morphologically similar plants. southern Puebla from Tehuacán south-southwest to- Agave dissimulans is recognized as distinct from its wards Villa de Tamazulapam del Progresso and south two nearest relatives by its geographic isolation, dis- and southeast towards Tepelmeme Villa de Morelos tinctly blue-glaucous leaf color, and propensity to and Quiotepec. They occur in xerophytic scrub com- have drooping leaves. munities on soil of limestone origin between 1,950– 7,000 feet elevation. Lemaire: L’Illustration Agave kerchovei DISCUSSION Horticole 11: 64. 1864. Neotype: Jacobi’s sketch as traced by Berger in U.S. National Herbarium. (Figs. The name Agave kerchovei has a long history be- 1–8) ginning with Charles Lemaire’s 1864 original de- Additional specimens examined scription of the species using a young plant growing Mexico: Oaxaca: Puente Santa Lucia on Mex- in cultivation. In 1915, continuing through Alwin ico Highway 135D, about 16.5 miles north of Te- Berger expanded description by using a larger plant pelmeme, 5,760 feet elevation, Starr 2018-002, 15 but still not one from a known locality. In 1982, January 2018 (ARIZ). Oaxaca: Side canyon of the Howard Scott Gentry added more to the description Rio Hondo, 0.25 airmiles south of the junction but included two closely related, yet distinct, spe- where Mexico Highway 135D crosses the Oaxaca- cies that William Trelease described in 1920. García- Puebla border, 3,540 feet elevation, Starr 2018-012, Mendoza (2011) then resurrected Agave convallis, 15 January 2018 (ARIZ). Oaxaca: Along the trail one of Trelease’s species from 1920, but placed Agave to La Huerta, about 13.5 airmiles northeast of Te- dissimulans as a synonym under Agave convallis in- pelmeme, 5,300 feet elevation, Starr 2018-026, 18 stead of Agave kerchovei. Finally, to complete the saga, January 2018 (ARIZ). Oaxaca: Rancho Tambor, after examining the plants at Cerro Campana, it is about 7.5 airmiles west-southwest of San Antonio apparent that Agave dissimulans is distinct from both Nanahuatipam, 3700 feet elevation, Starr 2018-051, Agave convallis and Agave kerchovei. 24 January 2018 (ARIZ). Puebla: 6 miles south- When scrutinizing Table 1, it is apparent that west of Tehuacán, Gentry 22418, 12 November 1967 the three species are most likely very closely related, (DES, MEXU, US). Puebla: Tehuacán, Trelease s.n., and if morphometrics are the only consideration, 8 December 1903 (MO) (Type of Agave inopinabilis). they could all three be considered synonymous as is evident by Gentry’s subsuming both Agave convallis DESCRIPTION and A. dissimulans into A. kerchovei. However, when Plants: medium to large, solitary, acaulescent, field work, along with nuanced, and not so nuanced open and sparsely leaved rosettes; 60–100 cm tall by differences are included in the analysis, the case for 100–150 cm across. Leaves: 30–50 per rosette; me- separating A. convallis and A. dissimulans from A. dium green to dark green, rarely with a faint, light kerchovei becomes quite strong. How then can these green mid-stripe; smooth to very lightly asperous; three species be distinguished in the field? The fol- widely spreading; 60–125 cm long by 5–12 cm wide; lowing key can be useful for field identification. widest from the base to near mid-blade, long linear- triangular to triangular-lanceolate; thick and succu- ACKNOWLEDGEMENTS lent at the base; flat to concave above, convex below; margin straight, horny, white or grayish; the long- First and foremost, I would like to thank Carol for acuminate apex lacking teeth. Teeth: few and re- her never-ending support for allowing me the free- motely spaced, small to medium, straight to curved, dom to go chasing after answers to the many agave 6–15 mm long, 20–70 mm apart; chocolate brown questions. My three trips to Oaxaca allowing me to when young, becoming white or grayish with age, research this intriguing cluster of agaves were funded absent in roughly upper one-third or more of leaf. by the Tucson Cactus and Succulent Society, the Terminal spine: awl-like; deep groove that is broad Cactus and Succulent Society of America, the Cen- or narrow, rounded below; 3–6 cm long; chocolate tral Arizona Cactus and Succulent Society, and the brown when young turning gray or white with age. Tim Gregory Agave Research Project. I would like to Inflorescence: total flower stalk length is 2.5–5 m, thank Tristan Davis for physical and moral support the fertile section 1.2–3 m long. Flowers: greenish in the field and his valuable insights and suggestions or yellowish, sometimes flushed with red on the tepal for the manuscript; Jeff Chemnick for his traveling tips, filaments and anthers yellow or reddish; overall companionship and linguistic talents to secure per- length is 38–54 mm, ovary 18–21 mm long, neck mission from various towns to allow the collection of is 3–6 mm long, tube is 2–6 mm long by 7–10 mm material from their property; Silvia Salas of SERBO wide, tepals are 15–21 mm long by 7–9 mm wide, for securing proper permits (SEMARNAT-03-058-A) filaments are yellowish green to dark reddish purple that allowed me to collect in the Tehuacán-Cuicatlán and quite showy. Biosphere Reserve; Brian Kemble and Walker Young for their help in the field. Finally, many thanks to the Author Copy 72 STARR — REEVALUATION OF AGAVE KERCHOVEI COMPLEX

Table 1. Comparison of key characteristics for Agave convallis, A. dissimulans, and A. kerchovei. Agave convallis Agave dissimulans Agave kerchovei

Plant size (height × 45–70 cm × 75–120 cm 50–120 × 100–150 cm 60–100 cm × 100–150 cm width) Leaf number 30–80 15–40 30–50 Leaf size (length × 30–60 cm × 6–10 cm 65–110 cm × 4–9 cm 60–125 cm × 5–12 cm width) Leaf color Light green to dark green, Blue-glaucous, rarely with a Medium green to dark green, sometimes with red, purple- faint light yellow-green mid- rarely with a faint, light green red, or brownish red on the stripe. mid-stripe. edges or tips, occasionally entirely red or purplish red, usually with a prominent, light green or yellow-green central stripe. Inflorescence 3–5 m 3–4.5 m 2.5–5 m Peduncle 1.4–1.5 m 1.7–2.1 m 1.3–2 m Fertile section 1.5–3.6 m 1.3–2.4 m 1.2–3 m Flower color Yellow to greenish yellow, Green on the outside, Greenish or yellowish, sometimes green and flushed sometimes flushed with a light sometimes flushed with red on with red or purplish red. blush of reddish purple. the tips. Flower length 35–50 mm 37–44 mm 38–54 mm Ovary length 12–20 mm 15–18 mm 18–21 mm Neck length 3–8 mm 4–5 mm 3–6 mm Tube (l × w) 2–4 mm × 5–8 mm 1–2 mm × 5–8 mm 2–6 mm × 7–10 mm Tepal lobes (l × w) 15–24 mm × 3–8 mm 16–23 mm × 7–9 mm 15–21 mm × 7–9 mm

Key to the three species of Agave in the A. kerchovei complex. 1. Leaves blue-glaucous, flaccidly drooping, rarely stiff and wide spreading. - ...... A. dissimulans 1’. Leaves variously green or red to red-purple or flushed with red or red-purple...... 2. 2. Leaves lanceolate to broadly lanceolate, 30–60 cm long × 6–10 cm wide, marginal teeth present nearly to leaf tip...... A. convallis 2. Leaves long linear-triangular to triangular-lanceolate, 60–125 cm long by 5–12 cm wide, marginal teeth absent in roughly upper one-third or more of leaf...... A. kerchovei many communities, Tepelmeme Villa de Morelos, San Annual Report of the Missouri Botanical Garden. 11: Antonio Nanahuatipam, Ignacio Mejía, San Pedro 80–83, plates 7–10. Jaltepetongo, El Parián, and San Jerónimo Sosola and Smith, G., E. Figueiredo, and G. Starr. 2018. Notes on various local guides for their permission to collect the typification and nomenclature of Agave dissimulans from their territories and their assistance in the field. Trelease, Agave convallis Trelease, and Agave kerchovei Lem. (Asparagaceae: Agavoideae/Agavaceae). Haselto- LITERATURE CITED nia 24: 36–39. Smith, G.F. and G. Starr. 2019. Proposal to reject the Baker, J. G. 1877. The genus Agave. The Gardener’s Chron- name Agave expatriata (Asparagaceae: Agavoideae / icle & Agricultural Gazette 7(1): 527–528. Agavaceae) Taxon 68(3): 597–598. Berger, A. 1915 Die Agaven: Beiträge zu einer Monographie. Thiede, J., G. Starr, G. Smith. 2020. Gustave-Philippe de 288 pp. Kerchove d’Ousselghem, his Agave collection, and the García-Mendoza, A. 2011. Agavaceae. Flora del Valle de correct eponymy of Agave kerchovei Lem. (Asparagace- Tehuacán-Cuicatlán. Fascículo 88, Instituto de Bi- ae: Agavoideae/Agavaceae) Haseltonia, in preparation. ología Universidad Nacional Autónoma de México. 98 Thiede, J., G.F. Smith, and U. Eggli. 2019. Infrageneric pp. classification of Agave L. (Asparagaceae: Agavoideae / Gentry, H. S. 1982. Agaves of Continental North America. Agavaceae): a nomenclatural assessment and updated University of Arizona Press, Tucson, Arizona, USA. classification at the rank of section, with new combi- 670 p. nations. Bradleya 37:240–264. Lemaire, C. 1864. Observations sur le genre Agave et de- Trelease, W. 1920. Agave. Contributions to the U.S. Na- scription de quelques especes nouvelles. L’Illustration tional Herbarium. 23:107–142. Horticole 11: 64. Rose, J.N. 1900. Agave expatriata and other Agaves flowering in the Washington Botanic Garden in 1898.

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