Manfreda Occidentalis (Agavoideae, Asparagaceae) a New Species from Western Mexico

https://doi.org/10.11646/phytotaxa.321.1.2

Manfreda occidentalis (Agavoideae, Asparagaceae) a new species from western Mexico

ARTURO CASTRO-CASTRO1, GUADALUPE MUNGUÍA-LINO2, PABLO CARRILLO-REYES3,4 & AARÓN RODRIGUEZ3,4,* 1Cátedras CONACYT–Instituto Politécnico Nacional, Centro Interdisciplinario de Investigación para el Desarrollo Integral Regional, Unidad Durango (CIIDIR), Sigma #119, Fracc. 20 de Noviembre II, 34220, Durango, Mexico. 2Cátedras CONACYT–Universidad de Guadalajara, Laboratorio Nacional de Identificación y Caracterización Vegetal (LaniVeg), Camino Ramón Padilla Sánchez 2100, 45110, Nextipac, Zapopan, Jalisco, México. 3Laboratorio Nacional de Identificación y Caracterización Vegetal, Consejo Nacional de Ciencia y Tecnología (CONACyT), Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Camino Ramón Padilla Sánchez 2100, 45110 Nextipac, Zapopan, Jalisco, México. 4Herbario Luz María Villarreal de Puga, Instituto de Botánica, Departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Camino Ramón Padilla Sánchez 2100, 45110 Nextipac, Zapopan, Jalisco, México. e-mail: [email protected] *author for correspondence

Abstract

The genus Manfreda (Asparagaceae) contains 35 species. A species complex includes plants similar to M. gutttata, which are characterized by the protrusion of the ovary into the perigone tube. This feature was found in specimens recently collected in western Mexico. After a morphological analysis, we found that some of the plants differ from the other species in this complex. Hence, we describe a new species named M. occidentalis morphologically similar to M. planifolia but can be dis- tinguished from it by the oblong-ovoid corm, channeled leaves, which are papillate on both sides, perigone 1.8–2.5 cm long, funnel-shaped, yellowish and light purple striate, perigone tube 0.3–0.7 × 0.2–0.4 cm, filaments adnate to the perigone and arising 0.3–0.7 cm above the ovary apex, anthers 0.9–1.1 cm long, and style 2.1–3.5 cm long, exceeding the perigone tube by 1.8–3 cm at anthesis. Manfreda occidentalis also resembles M. chamelensis; however, phenologic asynchrony is a notable difference between them as well as the shape of the leaves and the length and shape of perigone tube and lobes. A distribution map, photographs, and a key to identify species of Manfreda which grow in western Mexico accompany the description.

Resumen

El género Manfreda (Asparagaceae) agrupa a 35 especies. Existe un complejo de especies que incluye plantas similares a M. guttata que se caracterizan por la protrusión del ovario en el interior del tubo del perigonio. Este carácter fue observado en algunos especímenes de Manfreda recientemente recolectados en el occidente de México. Después del análisis de estas plantas, notamos que no se asemejan a ninguna de las especies del complejo antes señalado. Por lo tanto, proponemos una especie nueva, M. occidentalis morfológicamente similar a M. planifolia, pero puede distinguirse de ella por su cormo oblongo-ovado, hojas acanaladas y papilosas en ambas caras, perigonio de 1.8–2.5 cm de longitud, infundibuliforme, amarillo y púrpura estriado, tubo del perigonio de 0.3–0.7 × 0.2–0.4 cm, filamentos adnados al perigonio y surgiendo 0.3–0.7 cm por arriba del ápice del ovario, anteras de 0.9–1.1 cm de longitud, y estilo de 2.1–3.5 cm de largo, excediendo el tubo del perigonio por 1.8–3 cm en antesis. Manfreda occidentalis también se asemeja a M. chamelensis, sin embargo, existen diferencias entre ambas especies en la forma de las hojas y en la longitud y forma del tubo y lóbulos del perigonio. Además, existe una notable asincronía fenológica entre ambas. La descripción de la especie nueva es acompañada de un mapa de distribución geográfica, fotografías y una clave para la identificación de las especies de Manfreda que habitan el occidente de México.

Key words: Agavaceae, El Tuito, reproductive isolation, savannoid vegetation, Sierra del Cuale

60 Accepted by Mario Martínez-Azorín: 16 Jun. 2017; published: 12 Sept. 2017 Introduction

The genus Manfreda Salisbury (1866: 78) (Asparagaceae) includes 35 species of perennial herbs with mainly spicate inflorescences. The flowers are subtended by one bract and one or two bracteoles (Fig. 1; Table 1). However, two recently described species, M. paniculata Hernández-Sandoval et al. (2008: 170) and M. petskinil R.A.Orellana, L.Her. & Carnevali in Hernández-Sandoval et al. (2008: 173), develop paniculate inflorescences and flowers subtended by a single bracteole. Hernández-Sandoval et al. (2008) pointed out that these characters in the two species relate Manfreda to Agave Linnaeus (1753: 323). The natural geographic distribution of Manfreda extends from southeastern USA, throughout Mexico (except Baja California), and southwards into Honduras and El Salvador in Central America. All species, except M. fusca Ravenna (1987: 17), occur in Mexico. Western Mexico is one of the most species rich areas, with 13 species (Table 1, Fig. 2). Their habitats vary from moist to xeric and from cold to mild winters. The plants grow in open sun or partial to deep shade, often in oak-pine forest at an elevational range from sea level to 3,050 m.

TABLE 1. Diversity and natural distribution of the genus Manfreda (Asparagaceae, Agavoideae). Species Distribution 1. M. brunnea (S.Watson) Rose Mexico (Chihuahua, Coahuila, Durango, Hidalgo, Zacatecas) 2. M. bulbulifera Castillejos & E.Solano Mexico (Guerrero) 3. M. chamelensis E.J.Lott & Verh.-Will. Mexico (Jalisco) 4. M. elongata Rose Mexico (Durango, Nayarit) 5. M. fusca Ravenna Guatemala (Chimaltenango, Comalapa) 6. M. galvaniae A.Castañeda, S.Franco & Mexico (Estado de México) García-Mend. 7. M. guerrerensis Matuda Mexico (Guerrero) 8. M. guttata (Jacobi & Bouché) Rose Mexico (Aguascalientes, Chihuahua, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, Michoacán, Nayarit, Querétaro, San Luis Potosí, Sinaloa, Zacatecas) 9. M. hauniensis (J.B. Petersen) Verh.-Will. Mexico (Estado de México, Guerrero, Morelos, Oaxaca) 10. M. involuta McVaugh Mexico (Jalisco, Nayarit, Zacatecas) 11. M. jaliscana Rose Mexico (Durango, Jalisco, Michoacán, Nayarit, Sinaloa, Sonora) 12. M. justosierrana García-Mend., A.Castañeda Mexico (Guerrero) & S.Franco 13. M. littoralis García-Mend., A.Castañeda & Mexico (Guerrero, Oaxaca) S.Franco 14. M. longibracteata Verh.-Will. Mexico (Jalisco, Michoacán) 15. M. longiflora (Rose) Verh.-Will. USA (Texas); Mexico (Nuevo León, Tamaulipas) 16. M. maculata (C. Mart.) Rose Mexico (Estado de México, Guerrero, Oaxaca) 17. M. maculosa (Hook.) Rose USA (Texas); Mexico (Coahuila, Nuevo León, Tamaulipas) 18. M. nanchititlensis Matuda Mexico (Estado de México) 19. M. occidentalis Art.Castro & Aarón Rodr. Mexico (Jalisco, Nayarit) 20. M. paniculata L.Hernández, R.Orellana & Mexico (Campeche, Quintana Roo, Yucatán) G.Carnevali 21. M. parva Aarón Rodr. Mexico (Guerrero) 22. M. petskinil R.Orellana, L.Hernández & Mexico (Yucatán) G.Carnevali 23. M. planifolia (S.Watson) Rose Mexico (Chihuahua, Nayarit, Sonora) 24. M. potosina (B.L.Rob. & Greenm.) Rose Mexico (Coahuila, San Luis Potosí, Zacatecas) 25. M. pringlei Rose Mexico (Distrito Federal, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Oaxaca, Puebla, Tlaxcala, Veracruz) 26. M. pubescens (Regel & Ortgies) Verh.-Will. Mexico (Chiapas, Guerrero, Morelos, Oaxaca, Puebla) ex Pina 27. M. revoluta (Klotzsch) Rose Mexico (Estado de México) 28. M. rubescens Rose Mexico (Jalisco, Nayarit) 29. M. scabra (Ortega) McVaugh El Salvador (San Salvador, Santa Ana, Sonsonante); Guatemala (Chimaltenango, Chiquimula, Huhutenango, Jalapa, Jutiapa, Quiché, San Juan Sacatepequez, San Marcos, Santa Rosa, Solola); Honduras (El Paraíso, Tegucigalpa); Mexico (Aguascalientes, Chiapas, Colima, Distrito Federal, Durango, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosí, Veracruz, Zacatecas) ...continued on the next page

MANFREDA OCCIDENTALIS, A NEW SPECIES FROM MEXICO Phytotaxa 321 (1) © 2017 Magnolia Press • 61 TABLE 1. (Continued) Species Distribution 30. M. sileri Verh.-Will. USA (Texas); Mexico (Tamaulipas) 31. M. singuliflora (S.Watson) Rose Mexico (Chihuahua, Durango, Jalisco, Sinaloa, Zacatecas) 32. M. umbrophila García-Mend. Mexico (Guerrero, Oaxaca) 33. M. variegata (Jacobi) Rose USA (Texas); Mexico (Hidalgo, Nuevo León, Puebla, San Luis Potosí, Tamaulipas, Veracruz) 34. M. verhoekiae García-Mend. Mexico (Oaxaca) 35. M. virginica (L.) Salisb. ex Rose USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Kentucky, Louisiana, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia, West Virginia); Mexico (Nuevo León, Tamaulipas)

FIGURE 1. Floral and fruit diversity in Manfreda from western Mexico. A–B. M. occidentalis (A. Rodríguez et al. 5924, CIIDIR, IBUG, IEB, MEXU). C–D. M. chamelensis (A. Rodríguez et al. 4830, CIIDIR, IBUG, IEB, MEXU). E–F. M. elongata (A. Rodríguez et al. 4836, IBUG). G–H. M. guttata (A. Rodríguez et al. 4489, IBUG). I–J. M. involuta (A. Rodríguez et al. 4852, IBUG). K–L. M. jaliscana (A. Rodríguez et al. 4838, IBUG). M–N. M. longibracteata (A. Rodríguez et al. 4161, IBUG). O–P. M. rubescens (A. Rodríguez et al. 5787, IBUG). Q. M. planifollia (A. Castro-Castro et al. 2942a, IBUG). R. M. scabra (A. Rodríguez et al. 4485, IBUG). S–T. M. singuliflora (A. Rodríguez et al. 4527, IBUG). All pictures by Aarón Rodríguez.

62 • Phytotaxa 321 (1) © 2017 Magnolia Press CASTRO-CASTRO ET AL. FIGURE 2. Distribution of Manfreda species that occur in western Mexico. A. Western Mexico amplified highlighting the municipalities of Cabo Corrientes (Jalisco) and Compostela (Nayarit).

In Mexico, some species of Manfreda are traditionally useful. Both, the leafy base of the stem, known as bulb, and the structure below the bulb with attached roots, referred as corm, are macerated and employed as a vermifuge and antidote against snake venom. Commonly, they are used as “barbasco” (fish-poison) and soap substitute (Verhoek 1978, Hernández-Sandoval et al. 2008). Also, several species are appreciated for their ornamental value (Hannon 2002, Rodríguez & Castro-Castro 2007). The phylogenetic position of Manfreda has been addressed several times. Bogler & Simpson (1996), based on phylogenetic analyses of nuclear ribosomal DNA sequences, showed the close relationship between Manfreda, Polianthes Linnaeus (1753: 316) and Prochnyanthes Watson (1887: 457). In contrast, Rocha et al. (2006) produced a phylogenetic hypothesis based on different sources of evidence (Álvarez de Zayas 1987, Hernández-Sandoval 1995, Bogler & Simpson 1996, Bogler et al. 1995, 2006, Eguiarte et al. 2000), and stressed that the basic relationships

MANFREDA OCCIDENTALIS, A NEW SPECIES FROM MEXICO Phytotaxa 321 (1) © 2017 Magnolia Press • 63 among the main groups are congruent in the majority of the studies and suggested a broad concept of Agave including Manfreda, Polianthes and Prochnyanthes. Recently, a molecular phylogeny for Agavaceae (Good-Ávila et al. 2006), showed that these genera are closely related but relationships are unclear. All authors admitted the difficulty in solving the relationships within Agave s.l. (including Agave s.str., Manfreda, Polianthes, and Prochnyanthes), due to little variation in molecular markers and insufficient time for lineage sorting resulting in conflicts and uncertainties in Agave s.l. classification. It seems preferable to continue recognizing traditional circumscriptions of the genera until a well supported and stable phylogenetic hypothesis is produced. Manfreda was divided into four non-exclusive groups (Verhoek-Williams 1975), based on the ovary protrusion into the perigone tube, succulence of the leaves, geographic distribution, and flowering time. One of these groups embraces a set of species, allied to M. guttata (Jacobi & Bouché 1865: 561) Rose (1903: 21), having a 3–5 mm long protrusion of the ovary into the perigone tube (Fig. 1B, H and T). In all other species, the tube is inserted either near the base of the style or on a neck formed by a narrowed extension of the fused ovary and perigone tube without developing the protrusion (Fig. 1D, F, J, L, N and P). A recent field exploration, in western Mexico, resulted in the discovery of some plants having this ovary protrusion, which did not match any known species of the M. guttata alliance or other known species. Hence, it is proposed as a new taxon, named Manfreda occidentalis. Photographs, illustrations and a distribution map are included as well as a key for the identification of Manfreda in western Mexico.

Description

Manfreda occidentalis Art.Castro & Aarón Rodr. sp. nov. (Fig. 1A–B and Figs. 2–5) Species nova Manfreda planifolia magis proxima, a qua praecipue differt cormis oblongo-ovatis (contra globosis ad ovatis); foliis canaliculatis (contra applanatis), supra et subtus in facie papillatis (contra non papillatis); perigonio 1.8–2.5 cm longo (contra 2.8– 3.5 cm longo), infundibuliformi (contra hypocrateriformi), flavido et fasciato cum fasceis subpurpureis angustis (contra uniformiter flavido), perigonii tubo 0.3–0.7 × 0.2–0.4 cm (contra 0.9–1.1 × 0.1–0.3 cm); filamentis insertis ad 0.3–0.7 cm supra ovarii apice (contra insertis 0.9–1 cm supra ovarii apice); anthera usque ad 0.9–1.1 cm longa (contra 1.2–1.5 cm longa); et stylo 2.1–3.5 mm longo (contra 4.5–5 cm longo), exserto 1.8–3 cm ab perigonii tubo per anthesin (contra exserto 3.5–4.2 cm). Type:—MEXICO. Jalisco: Cabo Corrientes, km 176 de la carr. Mex. 200, entre Puerto Vallarta y Puerto Melaque, 753 m elevation, 20º22’1.4’’N, 105º18’52.8’’W, 6 September 2009 (fl, fr), A. Rodríguez 5924 et al. (holotype IBUG!, isotypes CIIDIR!, SLPM!, IEB!, MEXU!).

Perennial herb. Corm oblong-ovoid, (2.9–)4.5–5.6(–13.5) × 1.4–2.4(–5) cm, with growth buds and thickened contractile and fleshy roots. Roots cuneiform, 7–17 cm long. Bulb oblong-ovoid, 2–2.6 × 1.5–1.9 cm, tunicate. Leaves 4–7(–10) in a basal rosette, erect, alternate, lanceolate to oblong-elliptic, 48–83 × 1.8–4.5 cm, widest at the middle and tapering at both ends, channeled, acute, apex hardened, not pungent, and formed by the rolling of the lamina, margin hyaline, finely denticulate-erose under magnification, thin to leathery, dull green, concolorous, upper and lower surface papillate on the veins, the papilla dome shaped, 0.1–0.2 mm in diameter. Inflorescence spicate, (60–)90–190(–220) cm long, erect, internodes decreasing in size towards the apex; peduncle (53–)80–130(–170) cm long, bracts 6–10 along the peduncle, two basal and sterile bracts similar to the leaves, linear-lanceolate, (5.5)30–40(–60) × 0.4–2 cm, acute, truncate, thin to leathery, dull green, concolorous, upper and lower surface papillate on the veins, the papilla dome shaped, margin finely dentate; rachis dense, 7–28 cm long, with 15–30 floral nodes, primary floral bracts triangular to subulate, 0.7–1.8 × 0.2–0.6 cm, acute, truncate, smooth, soon drying, scariose, frequently deciduous. Bracteoles ensiform to acicular, 1–3 × 1 mm, entire. Flowers solitary at the inflorescence nodes, sessile [but pedicels 0.2–0.5 mm long in A. Rodríguez et al. 5924 (CIIDIR)], 1.8–2.5 cm long in anthesis, funnel-shaped, ascending to diffuse, yellowish and light purple striate; perigone tube 0.3–0.5 cm long, 0.2–0.4 cm wide at the base of the lobes, tubular, straight; lobes 1–1.5 × 0.2–0.4 mm, lanceolate, reflexed, barely hyaline, apices papillate. Stamens epitepalous, antitepalous, phanerantherous; filaments subulate, 1.4–2.5 cm long from their insertion point into the perigone tube, inserted 0.3–0.7 cm above the ovary apex, yellow; anthers linear, 0.9–1.1 × 0.1 cm, green and red striate, helicoidal after dehiscence. Ovary 0.5–0.7 × 0.3–0.4 cm, oblong-ovate, inferior, protruding 1–2 mm into the perigone tube; style filiform, 2.1–3.5 cm long, exerted 1.8–3 cm from the perigone tube at anthesis, dextrorse towards the stigma; stigma trilobed, papillose, clavate. Fruit a trigonous loculicidal capsule, 1.5–2 × 0.7–1.3 cm, globose to ovoid, crowned by the remains of the perigone and an acute ovarian protrusion. Seeds lunulate to ovate, 4–6 × 3–4 mm, flat, shiny black. Etymology:—The specific epithet refers to its restricted geographic distribution in western Mexico. Distribution, habitat and phenology:—Manfreda occidentalis is endemic to two mountain systems in western Mexico. Seven populations were found in close proximity to the Sierra El Cuale (Cabo Corrientes, Jalisco). It is part

64 • Phytotaxa 321 (1) © 2017 Magnolia Press CASTRO-CASTRO ET AL. of the more extensive Sierra Madre del Sur. In contrast, only one population was encountered in the Sierra de Mazatán (Compostela, Nayarit) in the western extreme of the Transmexican Volcanic Belt (Fig. 2A). It inhabits the ecotone between the Quercus Linnaeus (1753: 994) and Pinus Linnaeus (1753: 1000) forest, subdeciduous tropical forest, and the savannoid vegetation. Manfreda occidentalis is associated with Aristolochia emiliae Santana-Michel & Solís- Magallanes (2008: 8), Bessera tuitensis Ramírez (1992: 131), Bursera simaruba (Linnaeus 1753: 1026) Sargent (1890: 260), B. vazquezyanesii Rzedowski & Calderón de Rzedowski (2000: 55), Byrsonima crassifolia (Linnaeus 1753: 426) Kunth (1821: 149), Clethra Linnaeus (1753: 396), Echeandia coalcomanensis Cruden (1987: 130), Manfreda involuta McVaugh (1989: 231), Muhlenbergia cualensis Herrera & Peterson (1992: 114), Pinus jaliscana Pérez de la Rosa (1983: 290), P. oocarpa Schiede ex Schlechtendal (1838: 491), Prochnyanthes mexicana (Zuccarini 1837: 319) Rose (1903: 14), Quercus aristata Hooker & Arnott (1841: 444), Q. cualensis González-Villarreal (2003: 49), Q. resinosa Liebmann (1854: 182), Salvia aequidistans Fernald (1900: 512), S. cualensis González in González-Gallegos & Castro-Castro (2012: 50), Schoenocaulon jaliscense Greenman (1907: 20), and Verbesina Linnaeus (1753: 901). The plants are found in areas with deep and acidic soils between 700 to 1000 m elevation. Manfreda occidentalis flowers in August and fruits in September. Additional specimens examined (paratypes):—MEXICO. Jalisco: Cabo Corrientes, km 12.5 de la brecha El Tuito-Zimapán, 1074 m elevation, 20º21’20.5’’N, 105º15’1.8’’W, 12 August 2011 (fl, fr), A. Castro-Castro et al. 2511 (IBUG!); Km 5 de la brecha El Tuito-Zimapán, 2.5 km al E de La Provincia, 937 m elevation, 20º20’56’’N, 105º17’36.4’’W, 12 August 2011 (fl, fr), A. Castro-Castro et al. 2522 (IBUG!, MEXU!); Km 175 de la carr. Méx. 200, justo en el cruce con la brecha que conduce a El Cuale, 707 m elevation, 20º21’9.7’’N, 105º19’1.9’’W, 8 August 2011 (fr), A. Rodríguez et al. 6298 (IBUG!); Rancho Las Primaveras, km 186 de la carretera Méx. 200 Melaque- Puerto Vallarta, 394 m elevation, 20º26’23.9’’N, 105º17’51’’W, 4 August 2011 (sterile), A. Rodríguez et al. 6288 (IBUG!); Km 10.5 del camino entre El Tuito y Tehuamixtle, 500 m al N de Zicatán, 556 m elevation, 20º18’50.9’’N, 105º24’13.5’’W, 4 August 2011 (fr), A. Rodríguez et al. 6307 (IBUG!); Km 175 carretera Méx. 200 entre Puerto Melaque y Puerto Vallarta, 39 km al S de Puerto Vallarta, entrada a la mina de Zimapán, Puente La Hacienda, Rancho Maguey, 729 m elevation, 20º21’17.4’’N, 105º19’0.4’’W, 13 September 2015 (fr), A. Rodríguez & A. Castro-Castro 7603 (IBUG!). Nayarit: Compostela: Km 48 carr. Compostela-Puerto Vallarta, 6 km antes de Mesillas, 736 m elevation, 21º12’56.8’’N, 104º58’28.1’’W, 15 August 2013 (fl, fr), A. Castro-Castro & J.G. González-Gallegos 3275 (IBUG!, IEB!, MEXU!). IUCN Conservation assessment:—Manfreda occidentalis is known from eight locations (10 to 50 individuals for each one) within a radius of 65 km. Using GeoCAT (Bachman et al. 2011) the Extent of Occurrence (EOO) was of 699.9 km2, and the size of the Area of Occupancy is 24 km2, based on cells of 2 x 2 km. A preliminary category of endangered (EN/B2a) is proposed following the IUCN (2012) criteria. Taxonomic relationships:—Manfreda occidentalis is similar to the M. guttata group. The protrusion of the ovary into the perigone tube separates M. elongata Rose (1903: 21), M. guttata, M. planifolia (Watson 1887: 479) Rose (1903: 22), M. pringlei Rose (1903: 19), M. rubescens Rose (1903: 22) and M. singuliflora (Watson 1887: 479) Rose (1903: 16) from the rest of the Manfreda species (Verhoek-Williams 1975). Within the complex, the length of the ovary protrusion, the leaf margin, and the aggregation of the flowers along the inflorescence rachis differentiate each species (Fig. 1A, B, G, H, Q and T). Morphologically, M. planifolia is the most similar species to M. occidentalis (Fig. 1A, Q). However, M. occidentalis develops an oblong-ovoid corm as opposed to the globose to ovate corm in M. planifolia. The leaves are channeled with the upper and lower surfaces papillate (vs. flat and not papillate). In M. occidentalis, the perigone is shorter (1.8–2.5 vs. 2.8–3.5 cm), funnel-shaped (vs. salverform), and yellowish with light purple stripes (vs. yellowish). Likewise, the perigone tube is also shorter (0.3–0.7 × 0.2–0.4 vs. 0.9–1.1 × 0.1–0.3 cm). In M. occidentalis the filaments are inserted 0.3–0.7 cm above the ovary tip (vs. 0.9–1 cm) and the anthers reaches 0.9–1.1 cm (vs. 1.2–1.5 cm). A notable difference is the style length which is shorter in M. occidentalis than in M. planifolia (2.1–3.5 vs. 4.5–5 cm). Lastly, the style exceeds 1.8–3 cm the perigone tube in anthesis (vs. 3.5–4.2 cm; Table 2; Fig. 4). Manfreda occidentalis might also be confused with a sympatric species, M. chamelensis Lott & Verhoek-Williams (1991: 366). However, Manfreda occidentalis flowers from early August to mid-September, during the rainy season whereas M. chamelensis blooms from December to February, in the dry season. The morphological differences are highlighted in the Table 2. Lastly, Manfreda occidentalis inhabits western Mexico with 12 congeneric species [M. brunnea (Watson 1891: 156) Rose (1903: 19), M. chamelensis, M. elongata, M. guttata, M. involuta, M. jaliscana Rose (1903: 22), M. longibracteata Verhoek-Williams (1978: 166), M. planifolia, M. pringlei, M. rubescens, M. scabra (Ortega 1797: 13) McVaugh (1989: 234), and M. singuliflora]. Taxa treated in this study, can be identified with the following key.

66 • Phytotaxa 321 (1) © 2017 Magnolia Press CASTRO-CASTRO ET AL. FIGURE 4. Manfreda occidentalis Art.Castro & Aarón Rodr. A. Inflorescence apex. B. Flower dissection. C. Infrutescence ápex. D. Seeds. Illustrations by Lesslie Martínez Rubí based on type material.

FIGURE 5. Detail of papillae on upper (above) and underside (below) surfaces of leaves in Manfreda occidentalis Art.Castro & Aarón Rodr.

MANFREDA OCCIDENTALIS, A NEW SPECIES FROM MEXICO Phytotaxa 321 (1) © 2017 Magnolia Press • 67 TABLE 2. Morphological comparison of Manfreda occidentalis and related species. Character M. occidentalis M. planifolia M. chamelensis Corm oblong-ovoid, (2.9–)4.5–5.6(–13.5) × globose to ovoid, 3–4 × 3–3.8 cm cylindrical, 3–15 × 2–3 cm 1.4–2.4(–5) cm Bulb oblong-ovoid, 2–2.6 × 1.5–1.9 cm oblong-ovoid, 3–5 × 7–9 cm ovoid, 2–3.5 × 1.5–3 cm Leaves 4–7(–10), lanceolate to oblong- 6–8, lanceolate to oblong-elliptic, 50– 9–12, linear lanceolate, 40–75 × elliptic, 48–83 × 1.8–4.5 cm, erect, 75 × 2.6–4.5 cm, erect, flat, margins 1–3 cm, extended to prostrate, channeled, margins finely denticulate- finely denticulate-erose, yellow-green conduplicate, margins finely erose, dull green, concolorous, upper concolorous, upper and lower surface denticulate-erose, variegate, upper and and lower surface papillate not papillate lower surface papillate Inflorescence (60–)90–190(–220) cm long, dull 120–160 cm long, dull green 75–120 cm long, glaucous green green Peduncle 6–10 7–8 10–12 bracts number Rachis dense, 7–28 cm long, with 15–30 lax, 10–20 cm long, with (15–)20–36 dense, 10–20 cm long, with 10–35 floral nodes floral nodes floral nodes Perigone 1.8–2.5 cm long; funnel-shaped, pale 2.8–3.5 cm long; salverform, pale 2–3 cm long; funnel-shaped, yellow- yellow with light purple stripes yellow purplish Perigone tube 0.3–0.5 × 0.2–0.4 cm 0.9–1.1 × 0.1–0.3 cm 0.6–1.3 × 0.3–0.5 cm Perigone ascending to diffuse ascending erect to ascending orientation during anthesis Perigone 1–1.5 × 0.2–0.4 mm, lanceolate, 1.4–1.7 × 0.2–0.4 mm, lanceolate, 0.8–1.1 × 0.4–0.5 mm, oblong, apices lobes apices papillate apices with a tuft of white hairs with a tuft of white hairs Filament 1.4–2.5 cm long; inserted 0.3–0.7 cm 2.3–2.6 cm long; inserted 0.9–1 cm 2.1–3.2 cm long; inserted 0.3–0.7 cm above the ovary tip, yellow above the ovary tip, yellow above the ovary tip, variegate Anthers 0.9–1.1 cm long 1.2–1.5 cm long 1.3–1.5 cm long Style 2.1–3.5 cm long, exceeding 1.8–3 cm 4.5–5 cm long, exceeding 3.5–4.2 cm 4.2–6 cm long, exceeding 1.8–3 cm the perigone tube at anthesis the perigone tube at anthesis the perigone tube at anthesis Fruit 1.5–2 × 0.7–1.3 cm, globose to 1.3–2 × 1.3–1.5 cm, globose, crowned 1.2–1.6 × 1–1.5 cm, globose, not cylindrical, crowned by the remains by the remains of the perigone and an crowned by the remains of the of the perigone and an acute ovarian acute ovarian protrusion perigone and without an acute ovarian protrusion protrusion Seeds lunulate lunulate deltate-triangular Phenology flowering and fruiting from August to flowering and fruiting from September flowering and fruiting from December September to October to February Distribution western Jalisco and southwestern from central Chihuahua and Sonora to western Jalisco in Mexico Nayarit central Nayarit Elevation 400–1070 m 400–1000 m 10–570 m range

Key to the species of the genus Manfreda from western Mexico 1. Leaves succulent, 4–8 mm wide, with deltoid teeth on the margins ...... M. brunnea 1. Leaves chartaceous, 0.5–2 mm wide, with erose margins...... 2 2. Fruit protruding into the perigone tube as a dome or cone up to 3–5 mm long ...... 3 2. Fruit not protruding into the perigone tube or hardly so as a cone, up to 1 mm long ...... 8 3. Corm 1.7–2 × 1–1.5 cm; perigone tube funnelform and curved 180° ...... M. singuliflora 3. Corm 2.9–13.5 × 1.4–5 cm; perigone tube cylindrical and straight or curved 20–45° ...... 4 4. Leaves lanceolate-elliptic, frequently variegate; inflorescence rachis 2.3–14 cm long ...... M. guttata 4. Leaves lanceolate, linear-lanceolate to oblong, concolorous; inflorescence rachis 10–33 cm long ...... 5 5. Perigone tube 0.3–1.1 cm long; leaves lanceolate to oblong ...... 6 5. Perigone tube 1.3–1.8 cm long; leaves linear-lanceolate ...... 7 6. Leaves papillate on both surfaces; perigone 1.8–2.5 cm long, funnel-shaped, yellowish and light purple striped; perigone tube 0.3–0.7 × 0.2–0.4 cm; filaments inserted 0.3–0.7 cm above the ovary tip; style 2.1–3.5 cm long, exceeding 1.8–3 cm the perigone tube at anthesis ...... M. occidentalis 6. Leaves smooth on both surfaces; perigone 2.8–3.5 cm long, salverform, yellowish; perigone tube 0.9–1.1 × 0.1–0.3 cm; filaments inserted 0.9–1 cm above the ovary tip; style 4.5–5 cm long, exceeding 3.5–4.2 cm the perigone tube at anthesis ...... M. planifolia 7. Leaves 24–45(–60) × 0.8–2.8 cm, flat; inflorescence rachis 10–25(–33) cm long ...... M. pringlei 7. Leaves (40–)50–95 × 2.5–5 cm, channeled; inflorescence rachis 20–40(–70) cm long ...... M. elongata 8. Perigone tube 15–25 mm long, about as long as or longer than the lobes; flowering from July to September, during the rainy season...... 9 8. Perigone tube 4–13 mm long, about as long as the lobes; flowering from January to May, during the dry season ...... 10 9. Floral bracts (1.7–)4.5–6.5 cm long; perigone (3–)4–7 cm long; perigone tube 2.7–3.3 cm long ...... M. longibracteata

68 • Phytotaxa 321 (1) © 2017 Magnolia Press CASTRO-CASTRO ET AL. 9. Floral bracts 1–3.2(–4) cm long; perigone 3–3.5 cm long; perigone tube 1.5–2 cm long ...... M. scabra 10. Leaves reclined on the floor; flowers yellow-green; filaments exceeding the perigone tube by 4–8 cm ...... 11 10. Leaves erect; flowers reddish-green; filaments exceeding the perigone tube by 1–4 cm ...... 12 11. Perigone tube 0.4–0.6(–1) cm long, shorter than the lobes; style 6–9 cm long ...... M. jaliscana 11. Perigone tube 0.6–1.3 cm long, as long as the lobes; style 4.2–6 cm long ...... M. chamelensis 12. Leaves 30–50 × 0.2–0.5 cm, involute and often appearing tubular, then 2–3 mm in diameter, wanting during the flowering time ...... M. involuta 12. Leaves 17–30(–38) × 0.6–1.4 cm, flat to channeled, present during the flowering time ...... M. rubescens

Acknowledgments

We are grateful to the Consejo Nacional de Ciencia y Tecnología (CONACyT) for the research grant # 80200 and doctoral scholarships to ACC (229039, 335340) and GML (251844). The Universidad de Guadalajara offered financial aid to ACC. Mollie Harker and Jorge A. Pérez de la Rosa and Jorge A. Tena Flores helped with manuscript revisions, discussions about plant speciation and image edition, respectively. Jesús González Gallegos helped with the Latin diagnosis and reviewed the manuscript. We appreciate the attention extended by the curators and staff of the following herbaria: CIIDIR, CIMI, CREG, GUADA, IBUG, IEB, MEXU, SLPM, XAL and ZEA. Last but not least, we appreciate the illustrations made by Lesslie Martínez Rubí.

References

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