García-Marínez et al. 2017 Calochortus ownbeyi

Phytotaxa 314 (2): 241–250 http://www.mapress.com/j/pt/ Copyright © 2017 Magnolia Press

Article

ISSN 1179-3155 (print edition)

PHYTOTAXA

ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.314.2.5

Validation of Calochortus ownbeyi (Liliaceae), a new species from northwestern Mexico MIGUEL A. GARCÍA-MARTÍNEZ1, AARÓN RODRÍGUEZ2 & HUGH P. MCDONALD3

Maestría en Ciencias en Biosistemática y Manejo de Recursos Naturales y Agrícolas (BIMARENA), Centro Universitario de Ciencias Biológicas y Agropecuarias, Apartado postal 1–139, Zapopan 45101, Jalisco, Mexico 2 Herbario Luz María Villarreal de Puga del Instituto de Botánica de la Universidad de Guadalajara (IBUG), Departamento de Botánica y Zoología, Apartado postal 1–139, Zapopan 45101, Jalisco, Mexico; E-mail: aaron.rodriguez@cucba.udg.mx 3 Retired Professor, New York City College of Technology, City University of New York, 300 Jay St. Brooklyn, New York, USA 1

Abstract The name Calochortus ownbeyi has been extensively used even though it was not validly published. Accordingly, we are here validating and illustrating this species. Calochortus ownbeyi can be confused with C. venustulus, but there is morphological and molecular evidence that they are distinct species. A comparison between these two taxa and a distribution map are provided.

Resumen El nombre Calochortus ownbeyi ha sido utilizado extensamente a pesar de que no fue publicado válidamente. Por consiguiente, en este trabajo se valida y se ilustra la especie. Calochortus ownbeyi puede ser confundido con C. venustulus pero existe evidencia morfológica y molecular que apoya la distinción entre ambas. Se presenta una comparación morfológica entre estos dos taxones y un mapa de distribución.

Key words: Cyclobothra, geographic information system, Marion Ownbey, Sierra Madre Occidental, Transmexican Volcanic Belt

Introduction Calochortus Pursh (1814: 240) contains approximately 70 species of bulbous plants with scapiform or leafy stems and lanceolate leaves. Calochortus exhibits an extensive range of floral morphology; flowers vary in shape from subglobose to broadly campanulate, and the orientation can be erect, divergent or nutant. The perianth is composed of two series of three tepals each, the inner surfaces glabrous or pubescent. The tepals sometimes possess a nectary variable in size and outline. Perianth shape, distribution of trichomes and morphology of the nectary are the most critical characters for species identification (McVaugh 1989, Ownbey 1940, Fiedler & Zebell 2002).

Ownbey (1940) authored the most complete treatment of Calochortus. The latest taxonomic revision of C. section Cyclobothra Sweet (1828: t273) was McDonald (2000), in which he presented an informal description of Calochortus ownbeyi, but the epithet was not validly published. Since then, some confusion is found in herbaria collections with the use of this name, and similar plants have been identified as either Calochortus ownbeyi or C. venustulus Greene (1888: 158). Calochortus ownbeyi is clearly a distinguishable entity based on morphological (McDonald 2000) and molecular evidence (Patterson & Givinsh 2003). According to Ownbey’s classification, C. ownbeyi (McDonald 2000) belongs to Calochortus section Cyclobothra subsection Ghiesbreghtiani Ownbey (1940: 521).

Since we are currently involved in a taxonomic and biogeographical study of Calochortus section Cyclobothra, we decided it was necessary to validate the name. We provide a description, illustration, distribution map of Calochortus ownbeyi and comparison with C. venustulus. We examined the specimens of Calochortus ownbeyi and C. venustulus deposited in 18 herbaria collections (ANSM, CAS, CHAP, CHAPA, CIIDIR, ENCB, FCME, GUADA, HUAA, IBUG, Accepted by Mark Chase: 9 Jul. 2017; published: 25 Jul. 2017

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IEB, IZTA, MEXU, QMEX, SLPM, UAMIZ, XAL, ZEA; acronyms following Thiers, 2017). Then, we verified taxonomic determinations of the specimens. Fieldwork was carried out in the states of Chihuahua, Durango, Jalisco and Sonora in October and November 2016. We generated a database with 152 records (106 for Calochortus ownbeyi and 45 for C. venustulus) containing taxonomic, curatorial, ecological and geographical information. The database is available from the author. Georeferences for those specimens lacking coordinates or with wrong spatial data (79, 48%) were carried out using QGIS 2.10.1 (2015) following Hijmans et al. (1999). The data were projected into a map of states (INEGI 2016) and biogeographic provinces of Mexico (Morrone 2002). Additionally, the preliminary categories of conservation of both taxa were estimated following IUCN guidelines (2016) for distribution data (criterion B). The former required measurement of the extent of occurrence (EOO) and the area of occupancy (AOO) using GeoCAT (Bachman et al. 2011).

Taxonomy Calochortus ownbeyi García-Mart., Aarón Rodr. & H.P.McDonald, sp. nov. (Figs. 1–3) Type:—MEXICO. Sonora: Yécora, Los Pilares, 28º23’38.88’’N, 108º47’24.3’’W, 1315 m, 7 August 2016 (fl), García-Martínez et al. 274 (holotype: IBUG; isotypes: CIIDIR, MEXU, CAS).

Calochortus ownbeyi is characterised by the following: flowers campanulate, erect, perianth white; external tepals glabrous, nectary absent; internal tepals densely hairy extending half the surface; nectary linear, often obscured by the trichomes.

Bulbous perennial herbs, 25–55(–65) cm. Bulb ovoid, 1.3–2.5 × 1.2–2.5 cm, with fibrous-reticulate coats. Stem erect, branching, (1–)3– 5(–9) ramifications, bulbils frequently present in most leaf axils, branches widely variable in length, lowest often as long as basal leaf, upper much shorter. Basal leaf flat (15.0–)25.0–45.0 × 0.3–0.9 cm, linear; cauline leaves lanceolate to linear, 21.0–46.0 × 0.2–0.5 cm. Flowers with a pair of bracts, 1.5–3.0 × 0.2–0.5 cm, opposite, narrow-lanceolate to linear. Flowers campanulate, erect to somewhat spreading; external tepals elliptic to lanceolate, 1.0–1.5 × 0.3–0.5 cm, apiculate, glabrous or with a few trichomes, white or more often green to pale green or reddish, nectaries absent; internal tepals obovate, 1.3–2.0 × 0.6–1.5 cm, white, inner face with trichomes from near base to 1/2 or 2/3 length, trichomes mostly white or yellow at base, denser around nectary; nectary linear, 4–5 × 1–2 mm, often obscured by the trichomes. Filaments basally dilated, 4.0–5.0 × 0.8–1.0 mm; anthers oblong, 2.0–4.0 × 0.5–1.0 mm, acute to apiculate, basifixed, lavender or white. Ovary fusiform, 4–7 × 1–2 mm, winged; style with a persistent trilobed stigma.

Capsule erect, 1.5–2.5 × 0.5–1.0 cm, ellipsoid; seeds compressed, 2.00–3.00 × 0.08–1.2 mm, surface rough (Fig. 1).

Etymology:—The specific epithet honors Francis Marion Ownbey (1910–1974).

Distribution, habitat and phenology:—Calochortus ownbeyi is exclusive to Mexico. It grows from southeastern Sonora to northern Colima. It inhabits open pine and oak forest along the biogeographic province of Sierra Madre Occidental and the western portion of the Transmexican Volcanic Belt at 1,000–2,900 m elevation (Fig. 3). It flowers in early August and fruits in October. The species covers a broad geographical range and grows in eight Mexican states, Chihuahua, Colima, Durango, Jalisco, Nayarit, Sinaloa, Sonora and Zacatecas. Although the populations are dense, they are rare, growing only in particular places with poor and well-drained soils. Using GeoCAT (Bachman et al. 2011) the EOO sums 174,000 km2, and the size of the AOO is 356 km2, based on cells of 2 × 2 km. Following the IUCN (2016) criteria, the EOO classifies it as least concern (LC), which means it is a widespread species. In contrast, the AOO value considered it endangered (EN), which means that C. ownbeyi can face a high risk of extinction in the wild. These analyses indicate that C. ownbeyi is a rare but widespread species.

Notes:—Calochortus ownbeyi has been confused with C. venustulus. In September 1935, H.S. Gentry collected and described a Calochortus species with white flowers and yellow hairs in the centre from Sonora: Álamos, Sierra Charuco, Río Fuerte, 27º35’34’’N, 108º45’21.2’’W, 1240 m, 13 September 1935 (fl, fr), Gentry 1819 (MEXU!). In 1936, he found another species with a different flower colour, this time describing it as yellow from Chihuahua: Uriachi, Los Cascarones, Río Mayo, 28º0’14.43’’N, 108º10’2.41’’W, 2420 m, 11 September 1936 (fl), Gentry 2670 (MEXU!). These two specimens were determined as C. venustulus by Ownbey (1940), and he did not notice the differences between them. Fishbein (1998) contributed additional information on Calochortaceae in Gentry’s Río Mayo Plants. He analysed the specimens collected by H.S. Gentry, P.S. Martin, G. Starr, J. Laferrièr and his own collections of Calochortus. As a conclusion, he wrote “yellow-flowered and white-flowered forms may represent distinct taxa”.

After examining these two specimens, we concluded that the first corresponds to C. ownbeyi and the second to C. venustulus.

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FIGURE 1. Calochortus ownbeyi. A. Habit. B. Flower front view. C. Flower posterior view. D. Internal tepal. E. External tepal. Illustrated

by C. Rodríguez based on type material (García-Martínez et al. 274).

Validation of Calochortus ownbeyi (Liliaceae)

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Calochortus ownbeyi and C. venustulus exhibit highly divergent floral syndromes. Calochortus ownbeyi is described as the “cat’s ear” syndrome, whereas C. venustulus has the characteristics of the “star tulip” syndrome (Patterson & Givinsh 2003). The syndromes are associated with specific groups of pollinators that could represent a strong selective force (Dilley et al. 2000). Calochortus ownbeyi is most easily distinguished by its white and obovate internal tepals, whereas C. venustulus has yellow and lanceolate to obovate internal tepals (Table 1). Also, the external tepals of C. ownbeyi can be white, pale green or reddish, but in C. venustulus they are always yellow. Trichomes on the tepals, which are generally white, extend halfway or higher up the tepals in C. ownbeyi, whereas in C. venustulus they are yellow and limited to the nectary area. In general, C. ownbeyi is taller than C. venustulus and has consistently branched stem with bulbils in the leaf and bract axils, whereas in C. venustulus bulbils are always absent. The leaves are similar and variable in the two species, but floral bracts in C. ownbeyi are shorter.

TABLE 1. Morphological comparison of Calochortus ownbeyi and C. venustulus.

Character Plant size Number of ramifications Basal leaf size Cauline leaves size Floral bracts size Bulbils External tepals colour, shape and size Internal tepal colour, shape and size Distribution of trichomes on external tepals Distribution of trichomes on internal tepals Nectary shape and size Filaments colour and size Anthers colour and size Ovary colour and size Capsule shape and size Distribution

C. ownbeyi 25–55(–65) cm (1–)3–5(–9) (15.0–)25.0–45.0 × 0.3–0.9 cm 21.00–46.00 × 0.20–0.50 cm 1.5–3.0 × 0.2–0.5 cm present often green to pale green or reddish, lanceolate, 1.0–1.5 × 0.3–0.5 cm white with yellow at base or completely white, obovate, 1.3–2.0 × 0.6–1.5 cm absent or slightly hairy (3–10 trichomes) densely hairy in nectary area extending over half the surface linear, 4.0–5.0 × 1.0–2.0 mm yellow, cream or green, 4.00–5.00 × 0.80–1.00 mm white, cream or lavender, 2.0–4.0 × 0.5–1.0 mm green-yellow-reddish, 4–7 × 1–2 mm erect, lanceolate, apiculate, 1.5–2.5 × 0.5–1.0 cm western Mexico: Chihuahua, Colima, Durango, Jalisco, Nayarit, Sinaloa, Sonora and Zacatecas

C. venustulus (15–)25–30(–45) 1–3(–6) 15.0–35.0 × 0.2–0.7 14.00–29.00 × 0.15–0.50 2.0–5.0(–6.0) × 0.2–0.3 absent completely yellow, lanceolate, 1.0–1.8 × 0.4–0.9 cm yellow, elliptical to obovate, 1.2–2.0 × 0.4–1.9 cm absent or slightly hairy (3–10 trichomes) densely hairy only in nectary area elliptical, 1.0–3.0 × 0.9–1.5 mm yellow, 3.00–4.00 × 0.05–1.00 mm yellow, 2.0–3.0 × 0.9–1.5 mm green-reddish, 4–6 × 1–25 mm erect, lanceolate, apiculate, 2.0–3.5 × 1.0–1.5 cm northern Mexico: Chihuahua and Durango

Calochortus ownbeyi and C. venustulus have been found in similar environments along the Sierra Madre Occidental biogeographical province (Fig. 3), although they grow on different slopes. Calochortus ownbeyi can be found in the western (Pacific) slope, whereas C. venustulus is found more consistently on the eastern slopes of the same mountain range. The two species come into contact in the southern portion of the province in Coyotes and La Michilía, Durango.

Calochortus ownbeyi continues south over the western end of the Transmexican Volcanic Belt. Calochortus ownbeyi is found at 1,000–2,900 m elevation, whereas C. venustulus reaches higher elevations, 1,500–3,300 m. In accord with the estimations of EOO and AOO, C. ownbeyi is a rare and widespread species. The same analysis for C. venustulus generated an EOO value of 130,000 km2 and 164 km2 for AOO based on cells of 2 × 2 km. Following the IUCN (2016) criteria, C. venustulus is also classified into LC for the EOO and EN for the AOO values. Thus, C. venustulus and C. ownbeyi are both rare but widespread species.

Phylogenetic analyses, based on plastid DNA sequence evidence (Patterson & Givinsh 2003), placed C. ownbeyi with other Mexican taxa but far away from C. venustulus. The samples of C. ownbeyi used in the analysis came from collections made in coastal (Cabo Corrientes, 20º21’28’’N, 105º15’16’’W, 1030 m, Patterson 1044, WIS) and inland Jalisco (Cuautla, 20º14’37’’N, 104º27’50’’W, 1900 m, Patterson 1045, WIS). These coastal populations differ only in having lavender anthers and trichomes (Fig. 2B), whereas in the inland populations the anthers are white with yellow trichomes (Fig. 2C).

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FIGURE 2. Calochortus ownbeyi and C. venustulus flower differences. A. Calochortus ownbeyi type collection (García-Martínez et al.

274). B. Calochortus ownbeyi, coastal population (Rodríguez et al. 5935). C. Calochortus ownbeyi, inland population (Rodríguez et al.

5914). D. Calochortus venustulus habit (García-Martínez et al. 281). E. Calochortus venustulus (Rodríguez et al. 6156). F. Calochortus venustulus (Rodríguez & Castro-Castro 4567). Pictures: M.A. García-Martínez (A, D); A. Rodríguez (B, C, E, F).

Validation of Calochortus ownbeyi (Liliaceae)

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FIGURE 3. Distribution of Calochortus ownbeyi and C. venustulus.

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Specimens Examined:—MEXICO. Chihuahua: Guadalupe y Calvo, north side of Cerro Mohinora, 13 miles southwest of Guadalupe y Calvo, 25º57’N, 107º03’W, 2950 m, 20 August 1988 (fl), Nesom & McDonald 6466 (CAS, MEXU); Temósachic, Nabogame, 28º30’N, 108º30’W, 1800 m, 28 August 1987 (fl), Laferrière 996 (ANSM, CHAPA).

Colima: Minatitlán, Ejido Agua Fría, parte superior del arroyo Agua Fría, 19º19’32.95’’N, 104º1’29.4’’W, 1904 m, 4 October 2004 (fl, fr), Padilla-Velarde 1519 (CHAPA, UAMIZ). Durango: Durango, 52.1 miles SW of Durango, 23º25’30.91’’N, 105º0’11.04’’W, 2347 m, 1 September 1967 (fl), Oliver 678 (MEXU); 5.5 miles E of El Salto, 2 miles W of El Coyotes, 23º48’52.74’’N, 105º18’36.43’’W, 2470 m, 21 September 1974 (fl), Rollins & Kathryn 7433 (MEXU); 65–75 km SW of Durango City on road to La Flor, 23º27’15.9’’N, 104º44’41.48’’W, 2600 m, 17 September 1979 (fl), Breedlove 44172 (ENCB); between Coyotes and Llano Grande, along highway 40 between Mazatlán and Durango, 23º51’46.32’’N, 105º14’45.62’’W, 2450 m, 24 August 1986 (fl), Breedlove 63081 (CAS, MEXU); Ejido Las Navajas, bajío frente a Navajas, 23º52’N, 105º09’W, 2400 m, 28 August 2001 (fl), Carrillo 200 (CIIDIR, IBUG, MEXU); Ejido Las Navajas, al W de Los Artículos, frente a Navajas, 23º54’27’’N, 105º9’50’’W, 2400 m, 29 August 2001 (fl), Carrillo 261 (CIIDIR, IBUG, MEXU); km 70 carretera México 40 entre Durango y Mazatlán, 1 km al E de Las Navajas, 23º52’37.5’’N, 105º7’56.3’’W, 2538 m, 6 September 2010 (fl, fr), Rodríguez et al. 6120 (IBUG, IEB, UAMIZ); Mezquital, 19 km de Los Charcos, por el camino a Santa María Ocotán, 22º57’48.78’’N, 104º29’51.93’’W, 2400 m, 4 November 1983 (fl), González & González 2562 (ANSM, CIIDIR, ENCB, HUAA, IEB, MEXU, SLPM); 19 km al SW de Los Charcos, camino a Santa María Ocotán, 22º58’19.49’’N, 104º29’28.46’’W, 2250 m, 5 November 1983 (fl), Fernández 1821 (ANSM, CIIDIR, ENCB, IBUG, IEB, MEXU); W de Santa María de Ocotán, a lo largo de márgenes de arroyo, 22º54’45.62’’N, 104º37’14.91’’W, 1970 m, 16 October 1984 (fl), González & González 1546 (CIIDIR, HUUA); entronque a Laguna del Chivo, por el camino Charcos-La Escondida, en los alrededores del rancho, 23º7’N, 104º17’W, 2640 m, 24 September 1985 (fl), González 1858 (CIIDIR, IBUG, MEXU, UAMIZ); Sombrero Quemado, 2 km rumbo a La Guajolota, 22º58’54’’N, 104º42’4’’W, 2580 m, 15 August 1985 (fl), Solís 198 (CIIDIR, IBUG, MEXU, UAMIZ); Plátano Tatemado, alrededores, 12 km de La Guajolota, 22º58’N, 104º40’W, 2640 m, 12 September 1985 (fl), Solís 288 (CIIDIR, IBUG, IEB, MEXU, UAMIZ); Los Chapotes, al S de Santa María de Ocotán, 22º55’48.39’’N, 104º34’23.63’’W, 2200 m, 8 September 1986 (fl), Mendía 133 (CIIDIR, IBUG, MEXU, UAMIZ); 5 km al W de Santa María de Ocotán, 22º56’18.18’’N, 104º38’30.27’’W, 2120 m, 11 September 1986 (fl), Acevedo 304 (ANSM, CHAPA, CIIDIR, IEB, MEXU); La Candelaria, 23º4’N, 104º32’W, 1900 m, 15 September 1987 (fl), González 2359–O (CIIDIR, IBUG); 20 km carretera Mezquital, 23º25’21’’N, 104º21’1’’W, 2260 m, 12 October 1991 (fl), García 358A (HUAA, IEB); km 82.6 carretera Huejuquilla El Alto-Santa Lucía de la Sierra, 5 km al S de las Canoas (Durango) y 2 km al N de Ciénega de Mirasoles, 22º34’28.7’’N, 104º18’8.9’’W, 2257 m, 11 October 2009 (fl), Castro-Castro et al. 1928 (IBUG); Pueblo Nuevo, alrededores de Coyotes, 23º51’39.06’’N, 105º18’33.25’’W, 2450 m, 21 August 1981 (fl), Herrera & Rzedowski 176 (CIIDIR); Estación Coyotes, cerca de El Salto, 23º51’13.68’’N, 105º18’26.58’’W, 2450 m, 21 August 1981 (fl), Rzedowski 37516 (CIIDIR, ENCB, IEB, MEXU); Mesa de San Francisco, 24 km al SE de El Salto, 23º24’11.67’’N, 105º23’2.91’’W, 2430 m, 10 September 1983 (fl), Tenorio 4304 (MEXU); 41.3 miles W-SW of Ciudad Durango, along highway 40, toward Sinaloa, just to the E of Las Adjuntas, 23º44’12.36’’N, 105º27’47.98’’W, 2680 m, 27 September 1984 (fl), Lavin & Sundberg 5020 (QMEX, MEXU); Las Higueras, San Bernardino de Milpillas, 23º23’12.28’’N, 105º7’26.9’’W, 2280 m, 26 September 1981 (fl), Pérez 818 (ENCB); Santiago Papasquiaro, 30 km por el camino a Canelas, 25º6’N, 105º35’W, 2450 m, 27 August 1991 (fl), González & Acevedo 4928 (CIIDIR, IBUG); Súchil, La Mesa del Burro, 6 km al SW de Piedra Herrada, 23º24’38’’N, 104º16’24.26’’W, 2650 m, 16 September 1982 (fl), Fernández 1225 (ENCB); Potreros Ajolotes, San Juan de Michis, 23º24’6.2’’N, 104º10’31.77’’W, 2377 m, 18 October 1985 (fl), Alvarado 382 (CHAP, CIIDIR, IBUG, IEB, MEXU); carretera a Charcos, entre Michis y Piedra Herrada, 23º23’28’’N, 104º11’32.87’’W, 2410 m, 9 October 1985 (fr), Alvarado 225 (CHAPA, CIIDIR, IEB, MEXU); Cerro de las Iglesias, a 6 km de Alemán, 23º21’32.16’’N, 104º6’36.32’’W, 2770 m, 1 September 1986 (fl), Acevedo 199 (ANSM, CHAP, CIIDIR, IEB, MEXU); Reserva de la Biosfera La Michilía, El Alemán, entronque camino viejo a Puerta Roja, 23º20’N, 104º10’W, 2250 m, 19 October 1992 (fl), García & Medina 1727 (CIIDIR, IBUG, MEXU, UAMIZ); Reserva de la Biosfera La Michilía, Laguna de Ajolotes, 120 m al E de la laguna, camino a El Alemán, 23º24’N, 104º10’W, 2470 m, 23 September 1993 (fl), García & Medina 1849 (CIIDIR, IBUG, MEXU); 5.8 km al SO de San Juan Mochis camino a Temascal, 23º23’9.5’’N, 104º11’0.23’’W, 2377 m, 16 September 2005 (fl), Rodríguez & Castro-Castro 4526 (IBUG). Jalisco: Atenguillo, Sierra de la Campana, along road to Mascota, 7–8 miles NW of Los Volcanes, 20º24’11’’N, 104º34’10.21’’W, 1900 m, 23–25 October 1952 (fl), McVaugh & Scooby 13719 (MEXU); W slopes of Sierra de la Campana, 7–8 miles west of Los Volcanes, steep mountainside and nearby valley, 1800 m, 20º24’13’’N, 104º33’52.14’’W, 10 October 1960 (fl), McVaugh et al. 20024 (CHAPA, IEB, MEXU); 14 km E of Los Volcanes, on road from Ayutla to Talpa de Allende, 20º15’11’’N, 104º29’13.08’’W, 1830 m, 27 November 1983 (fl, fr), Breedlove 60582 (CAS); km 82–83 carretera México 70 justo en la Puerto La Campana, 20º22’24’’N, 104º35’69.7’’W, Validation of Calochortus ownbeyi (Liliaceae)

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1947 m, 2 September 2006 (fl, fr), Rodríguez 4899 (IBUG); km 82–83 carretera México 70 justo en la Puerto La Campana, 20º22’24’’N, 104º35’69.7’’W, 1947 m, 17 September 2006 (fl, fr), Rodríguez et al. 4948 (IBUG); km 82–83 al SW de Ameca, Puerto La Campana por la carretera México 70 rumbo a Mascota, 20º22’24’’N, 104º35’69.7’’W, 1947 m, 28 September 2005 (fl, fr), Rodríguez 4620a (IBUG, IEB, MEXU); Puerto la Campana, km 82–83 carretera México 70 Ameca-Mascota, 20º22’15.95’’N, 104º35’37.36’’W, 1940 m, 7 September 2016 (fl), García-Martínez & Anguiano 290 (IBUG, MEXU, ZEA); Autlán de Navarro, Los Cuartones entre Corralitos y la Estación Científica Las Joyas, Sierra de Manantlán, 19º36’18’’N, 104º18’3’’W, 1945 m, 13 August 2002 (fl), Santana-Michel et al. 11108 (ZEA); Ayutla, km 30 camino a San Miguel de la Sierra, 20º7’57’’N, 104º34’0.2’’W, 2100 m, 13 August 2007 (fl, fr), Rodríguez et al. 5360 (IBUG); km 32 camino a San Miguel de la Sierra, 20º7’22’’N, 104º34’52.8’’W, 2100 m, 13 August 2007 (fl, fr), Rodríguez et al. 5365 (IBUG); km 31.2 del camino entre Ayutla y San Miguel de la Sierra, 20º7’33’’N, 104º34’32.2’’W, 1870 m, 23 October 2007 (fl, fr), Rodríguez & Harker 5354 (IBUG, IEB); 41 km al O de Ayutla y 1 km al E de San Miguel de la Sierra, 20º6’19’’N, 104º35’33.6’’W, 2112 m, 23 October 2007 (fl), Rodríguez 5359 (IBUG, IEB); km 36.6 en la camino de Ayutla a San Miguel de la Sierra, 20º7’17’’N, 104º35’8.11’’W, 2107 m, 11 November 2008 (fl, fr), Harker & Rodríguez 3921 (IBUG); km 9.6 del camino a San Miguel de la Sierra, a partir del entronque con la carretera Ayutla-Cuautla, 20º7’3.4’’N, 104º28’42.2’’W, 1845 m, 10 October 2010 (fl), CastroCastro & Villa-Vázquez 2278 (IBUG, IEB); Cabo Corrientes, on dirt road between km 11 and 12, 6.5 mi from highway 200, 1.7 miles beyond a settlement, 20º21.44’N, 105º15.2’W, 1220 m, 9 October 1996, McDonald s.n. (CAS); up forest logging road, off road 200, S of Puerto Vallarta, 20º21’28’’N, 105º15’16’’W, 1030 m, 9 October 1996, Patterson 1044 (WIS); Ejido la Provincia, brecha a Mina Zimapan-El Cuale, 20º13’33’’N, 105º15’18.06’’W, 1131 m, 6 September 2008 (fl), Ramírez-Delgadillo & Frías-Castro 7672 (IBUG); km 10–11 de la brecha a El Cuale; a partir del km 174.5 de la carretera México 200 entre Puerto Melaque y Puerto Vallarta, 3 km al E del ejido La Provincia, 20º21’33’’N, 105º15’26.8’’W, 1156 m, 6 September 2009 (fl), Rodríguez et al. 5935 (IBUG); Cuautitlán de García Barragán, Sierra de Manantlán, 1.5 km al SE de Corralitos, 19º36’9’’N, 104º17’53’’W, 1936 m, 9 October 2012 (fl), Carrillo-Reyes et al. 6823B (IBUG); Cuautla, road to Mascota, 6 mi NW of Cuautla, open oak forest, 20º14’18’’N, 104º27’27.79’’W, 1850 m, 21 October 1952 (fl), McVaugh & Scooby 13675 (IEB, MEXU); 5–6 mi W Cuautla, on road to Talpa, 20º14’28’’N, 104º27’44.9’’W, 1900 m, 10 October 1996 (fl), McDonald s.n. (CAS); 5–6 miles W of Cuautla, on road to Talpa 20º14’37’’N, 104º27’50’’W, 1900 m, 10 October 1996, Patterson 1045 (WIS); km 41–42 carretera San Vicente-Volcanes, 9.5 km al SE de Volcanes, 20º15’43’’N, 104º29’54.5’’W, 1687 m, 23 October 2007 (fl), Rodríguez & Harker 5348 (IBUG); 10 km al SE de Volcanes, Rancho Fresno Hueco, 20º15’23’’N, 104º29’45.2’’W, 1750 m, 23 October 2007 (fl, fr), Rodríguez & Harker 5351 (IBUG, IEB); Ejutla, cima del cerro localizado al SW de Ejutla, junto al Cerro El Narigón carretera Ejutla-El Grullo, 19º52’8.1’’N, 104º11’00.3’’W, 1365 m, 24 August 2004 (fl, fr), Rodríguez & Castro-Castro 3997 (IBUG, IEB, MEXU, UAMIZ); Mascota, 49 km W of Ayutla on road to Talpa, 20º23’49.2’’N, 104º37’15.41’’W, 1850 m, 21 September 1983 (fl), Anderson et al. 12746 (CAS, CHAPA, IEB, MEXU); 6 km después de Mascota, rumbo a Ameca, 20º29’49’’N, 104º45’47.66’’W, 1380 m, 15 August 1990 (fl), Espejo et al.

4181 (CIIDIR, IBUG, IEB, UAMIZ); km 17.3 camino a Mascota-Juanacatlán, 11.5 km al ENE de Mascota, 20º33’45’’N, 104º40’45’’W, 1890 m, 12 August 2001 (fl), Carrillo-Reyes 2508 (GUADA, IBUG, UAMIZ); alrededores del Puente Piedras Chinas, 1 km al NW del puerto La Campana, km 81 carretera Ameca-Mascota, 20º22’12’’N, 104º36’9.6’’W, 1925 m, 4 September 2009 (fl), Rodríguez et al. 5914 (IBUG); km 19.5 carretera Mascota-Puerto Vallarta, 20º39’39’’N, 104º51’56.9’’W, 1619 m, 4 September 2009 (fl), Rodríguez et al. 5918 (IBUG); km 22 de la carretera Mascota-Puerto Vallarta, 20º40’8.6’’N, 104º53’3.7’’W, 1531 m, 5 September 2009 (fl), Rodríguez et al. 5920 (IBUG); 10 km después de Mascota, rumbo a Ameca, 20º28’47’’N, 104º53’31’’W, 1516 m, 15 September 2009 (fl), Ceja et al. 2068 (IBUG, UAMIZ); Rancho Tlalpuyec, paraje La Berenjena, 1 km al SSE de Los Sauces de Yerbabuena y 3.7 km al SO de Tecoani en línea recta, 20º31’6’’N, 104º51’6’’W, 1370 m, 27 September 2014 (fl, fr), Carrillo-Reyes et al. 7525 (IBUG); Mezquitic, 3 km al norte del campamento Rivera Aceves, 22º5’29’’N, 103º53’5.23’’W, 2700 m, 3 October 1994 (fl), Cházaro-Bazañez & García-Guerrero 7435 (IBUG); San Sebastián del Oeste, brecha Mascota-San Sebastián del Oeste, 20º39’2’’N, 104º51’39.65’’W, 1390 m, 24 August 1985 (fl), González-Villarreal 12774 (IBUG); brecha saliendo a la carretera Mascota-San Sebastián del Oeste en dirección a la Bufa, empezando donde está localizada La Virgencita, 20º41’37’’N, 104º52’29’’W, 1815 m, 3 September 2006 (fl), Harker et al. 3157 (IBUG); Talpa de Allende, entre El Cuale y Las Minas de Oro, O de Los Lobos, 20º24’49’’N, 104º48’7.2’’W, 1500 m, 8 September 1971 (fl), González-Tamayo 374 (ENCB); entre La Crucecita, y La Cumbre del Tejamanil, 20º22’34’’N, 104º58’40.64’’W, 2080 m, 22 October 1971 (fl), González-Tamayo 530 (ENCB, MEXU); La Mina, Sierra El Caule, 20º22’18’’N, 105º5’10.55’’W, 1875 m, 7 November 1978 (fl, fr), Magallanes 1218 (MEXU); 20 km al S de Talpa de Allende, 20º13’38’’N, 104º46’24’’W, 1500 m, 4 October 1982 (fr), Hernández-Magaña et al. 9081 (MEXU); E of Talpa de Allende in Sierra de Cuale, 20º23’4.2’’N, 105º3’48.34’’W, 1585 m, 10 September 1986 (fl), Breedlove 64017 (CAS, MEXU); km 83–84 248 • Phytotaxa 314 (2) © 2017 Magnolia Press

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carretera México 70 justo en el puente La Piedra China, 2 km al NW del puerto La Campana, 20º22’18’’N, 104º36’17.2’’W, 1902 m, 17 September 2006 (fl), Rodríguez et al. 4950 (IBUG); 17 km al O de Talpa de Allende en línea recta, por la brecha rumbo a El Cuale, 3.2 km al O del cerro de La Tetilla de El Cuale, 20º22’32’’N, 104º59’14.1’’W, 2290 m, 26 September 2011 (fl), González-Gallegos & Guerrero 1103 (IBUG); 3–4 km al SO de El Cuale por la brecha hacia las Minas de Zimapán, 20º23’43’’N, 105º4’49.8’’W, 1630 m, 28 October 2011 (fl), González-Gallegos & Juárez 1160 (IBUG); Tecolotlán, Sierra de Quila, al lado del camino a 50 m antes de llegar a El Árbol de La Lira, 20º17’52’’N, 104º4’12.6’’W, 2082 m, 20 September 2008 (fl), Villa-Vázquez & Zamora-Tavarez 4 (IBUG); Sierra de Quila, La Mojonera, 3.7 km al NO de base La Ciénega, 20º18’45’’N, 104º2’35.5’’W, 2178 m, 26 September 2009 (fl), FloresArgüelles et al. 65 (IBUG); Sierra de Quila, en los alrededores de El Árbol de La Lira, 20º17’50’’N, 104º4’19.5’’W, 1947 m, 23 September 2011 (fl, fr), Paizanni & Paizanni 83 (IBUG); Tenamaxtlán, recorrido de Tenamaxtlán a Los Picachos, 20º14’18’’N, 104º26’8.58’’W, 1900 m, 20 August 1994 (fl), Machuca 7199 (IBUG, XAL); Los Picachos de Tenamaxtlán, 20º15’28’’N, 104º24’33.25’’W, 2000 m, 11 August 1996 (fl, fr), Machuca & Cházaro-Bazañez 7934 (IBUG); camino Tenamaxtlán-Los Picachos, Sierra de Quila, al E de promontorio rocoso, 20º14’80’’N, 104º9’9’’W, 1840 m, 28 September 1997 (fl), Machuca 8126 (IBUG); Villa Purificación, Villas de Cacoma, 19º48’20’’N, 104º32’35’’W, 1831 m, 28 October 2010 (fl), Rodríguez et al. 386 (ZEA). Nayarit: La Yesca, Sierra de las Palomas, Sierra de Alicia, beyond Huajimic, 21º38’14’’N, 104º25’37.93’’W, 1990 m, 9 October 1963 (fl), Schubert & Sousa 2033 (ENCB, MEXU); Rancho Los Planes, camino a San Pelayo, 21º22’7.8’’N, 104º3’35.24’’W, 2200 m, 16 October 1990 (fl), Flores-Macías & López 2246 (CHAPA, ENCB, IBUG, IEB). Sinaloa: Concordia, carretera MazatlánDurango, brecha Potrerillos-La Petaca, El Cuantatal, 4.5 km al SE de La Petaca, 23º23’52’N, 105º48’37’’W, 1700 m, 10 September 2000 (fl), Rito-Vega & Gutiérrez-García 10655 (MEXU). Sonora: Álamos, Sierra Charuco, Río Fuerte, 27º35’34’’N, 108º45’21.2’’W, 1240 m, 13 September 1935 (fl, fr), Gentry1819 (MEXU); 55.4 miles NE Álamos, 0.3 miles N Huicochi, 27º19’38’’N, 108º37’40.19’’W, 1555 m, 11 August 1980 (fl, fr), Lehto 24853 (ENCB); Sahuaripa, 8 km NW of Trigo on road from Yécora to Suarahipa, 28º37’29’’N, 109º3’24.02’’W, 1280 m, 25 August 1984 (fl, fr), Breedlove 61093 (MEXU); Yécora, field across from cementary, 28º22.5’N, 108º54.5’W, 1540 m, 17 August 1998 (fl), Fishbein et al. 3656 (MEXU); 15 km al O de Maycoba sobre la carretera Cuauhtémoc-Yécora, 28º23’49.3’’N, 108º44’3’’W, 1450 m, 6 August 2016 (fl), García-Martínez et al. 271 (CIIDIR, IBUG). Zacatecas: Jímenez del Teúl, Cerro de Urica, 23º18’46’’N, 104º5’6.26’’W, 2220 m, 17 September 1986 (fl), Acevedo 123 (ANSM, CHAPA, CIIDIR, IEB, MEXU); Valparaiso, km 58.6 carretera Huejuquilla El Alto-Santa Lucía de la Sierra, 3 km al W del entronque a Cuevitas, 22º37’3.6’’N, 104º11’58.5’’W, 2100 m, 10 October 2009 (fl), Castro-Castro et al. 1920 (IBUG); 500 m al W de Santa Lucia de la Sierra, rumbo a Santa Cruz de Tepetates, 22º28’24’’N, 104º13’38.6’’W, 2298 m, 11 October 2009 (fl, fr), Castro-Castro et al. 1947 (IBUG).

Acknowledgements The first author thanks the Consejo Nacional de Ciencia y Tecnología (CONAyT) for academic scholarship 581025.

Our thanks to curators of ANSM, CAS, CHAP, CHAPA, CIIDIR, ENCB, FCME, GUADA, HUAA, IBUG, IEB, IZTA, MEXU, QMEX, SLPM, UAMIZ, XAL and ZEA. A. Castro-Castro and J.G. González-Gallegos made important comments and suggestions, and C. Rodríguez created the illustration. HPM thanks F. Almeda and staff at the California Academy of Sciences for their assistance in making the material available. We appreciate the suggestions made by editor Mark Chase and an anonymous reviewer that improved the manuscript.

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