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The genus Manfreda Salisb. (Asparagaceae; Agavoideae) comprises polycarpic perennial herbs, with an upright bulb and a globose to cylindrical corm with fleshy and contractile roots. The plants produce thin to succulent leaves, organized in a basal rosette. The inflorescences are more frequently spiciform with solitary and sessile flowers at the nodes, but it can also be racemose with paired and pedicellate flowers, in both cases the flowers are subtended by one bract and one or two bracteoles (Castro-Castro et al. 2018). Manfreda includes 37 species distributed from the southeastern United States, throughout Mexico, and in Central America north of Honduras and El Salvador (Verhoek-Williams 1975, Castro-Castro et al. 2017, 2018). All species, except M. fusca Ravenna, occur in Mexico, whereas Guerrero and Jalisco are the most species rich states, with 14 and 12 species respectively (Castro-Castro et al. 2017, 2018).
Manfreda inhabit moist to xeric habitats with cold to mild winters. Commonly, the plants grow in open sunny or partial to deep shade sites in pine-oak forest, grassland, and tropical dry forest from sea level to 3,250 m (Castro-Castro et al. 2017, 2018). Although the range of distribution and ecological preferences of Manfreda are wide, there is a concentration of species towards the Pacific Lowlands biogeographic province in Mexico, where 16 species are found.
Throughout their area of distribution, Manfreda species are colloquially known as akasman, American aloe, American amole, amole, amole cenizo, amole de río, amole pa’ chango, amoli, amolilla, apintli, aporaka, azucena sabanera, biaa’toba, calpaich, cebolla de cerro, chupac, c`ipal, huaco, coyamole, jabón de hormiga, lechuguilla de castilla, lengua de lagartija, mezcalito, nama choco, oreja de liebre, soap plant, sotol, spice lily, taiicholi, Virginian agave, or wild tuberose (Verhoek 1978, Castillejos-Cruz 2009). In some regions of Mexico, their bulbs, corms, and fleshy roots are macerated, dissolved in water, and used as a soap substitute to wash clothes and hair, as poison for artisanal fishing, and as a remedy against snake bite, stomach colic and headache (Verhoek 1978). In addition, the plants have a high ornamental potential (Rodríguez & Castro-Castro 2007), and the scape of inflorescences have been employed to make arrows for hunting (Verhoek 1978).
Given the existence of the high diversity of Manfreda between the political and natural boundaries in Guerrero, in this document we decided to present an analysis of the richness and geographical distribution of the genus in the state. Furthermore, during the revision of herbaria and data collection for the biogeographic analyses, we noted some specimens from the north of Guerrero in the Balsas Basin biogeographic province that did not match with any known species of Manfreda due to the size of different vegetative structures of individuals, the quantity of flowers in the inflorescence, and the perigone shape. Hence, after an analysis of herbarium vouchers and live plants from a recent field collection, review of relevant literature, comparisons with the most related taxa in the genus, and critical comments from experts, we recognize this as a new species. The biogeographic analysis and the description of the new species are accompanied with a map, illustrations, and a key to Manfreda of Guerrero.
Materials and methods
Geographical data and richness analyses. Data were obtained from herbarium specimens in the following collections: CHAPA, CIIDIR, ENCB, FCME, FEZA, IBUG, IEB, and MEXU (acronyms according to Thiers 2019). We also examined the virtual herbaria of ARIZ, ASU, BH, F, GH, MICH, MO, NY, RSABG, US, and WIS; and searched for specimens in the Southwest Environmental Information Network (SEINet) from Symbiota content management system (Gries et al. 2014). In addition, taxonomic treatments, monographs, and floristic studies were reviewed for distributional data (Matuda 1975, Verhoek-Williams 1975, Piña 1985 a, b, c, Piña 1986, McVaugh 1989, Ayala-Hernández 2006, Rodríguez & Castro-Castro 2007, Castillejos-Cruz 2009, Rodríguez 2009, García-Mendoza 2011, Castro-Castro et al. 2017, 2018). When doubtful taxonomic identity records were detected, digitalized voucher specimens were analyzed, or images were requested from the staff of the corresponding herbaria.
Occurrence data was cleaned by eliminating repeated records and ambiguous locations and then visualized using the program ArcView GIS ver. 10.3 (ESRI 2014). The number of Manfreda species in Guerrero was quantified by municipality, vegetation type (Rzedowski 1990), biogeographic province (Morrone et al. 2017), and a standardized grid cell. The grid cell size was chosen according to the criteria of Willis et al. (2003), with the modification of Suárez-Mota & Villaseñor (2011). Based on this method, the maximum distance of each taxon was averaged, and the value was divided by the number of species with maximum distance values greater than zero, for which 13 species reached the value in our analysis. The quotient was used to define the length of the cell. The grid analysis was performed with DIVA-GIS 4.2 (Hijmans et al. 2004). In this way, the state of Guerrero was divided into 162 cells of 144 km2.
Taxonomic treatment. The morphological description and illustration of the new species are based on traditional taxonomic methods (Simpson 2006). Descriptions and measurements were carried out under a Zeiss Stemi 305 stereomicroscope. The line illustration of the new species was prepared from herbarium specimens, alcohol-preserved material and digital photos. The terminology used follows Verhoek-Williams (1975), Castillejos-Cruz (2009), and Castro-Castro et al. (2017, 2018). The morphological data obtained from the revised herbaria specimens were summarized to produce the descriptions, which served to prepare the identification key and the diagnosis of the new species. The conservation status of the new species was determined based on field observations and applying the IUCN red list criteria (IUCN 2019). We used GeoCAT program (Bachman et al. 2011) to estimate the extent of occurrence (EOO) and area of occupancy (AOO) indices for evaluating the geographic range.
Results
Geographical data and richness analyses. Our database included 86 different records, with 114 specimens, for 14 species of Manfreda in Guerrero (Appendix 1). We corroborated the taxonomic identifications and georeferenced 52 % of all records that lacked geographic coordinates. Manfreda collections in Guerrero are limited to 33 of the 81 municipalities in the state. The records and richness by municipalities are concentrated in Chilpancingo de Los Bravo (18 records / 5 spp.), Taxco de Alarcón (8 records / 3 spp.), Chilapa de Álvarez (7 records / 4 spp.), and Leonardo Bravo (7 records / 3 spp.). On the other hand, in Arcelia, Chilapa de Álvarez, Colotlipa, Carrizal de Bravo, Guayameo, Leonardo Bravo, Taxco de Alarcón, Técpan de Galeana, Tzitzio, and Zirándaro are restricted five endemic species to Guerrero (M. arceliensis Art.Castro, J.G. Zavala & Cruz Durán, M. bulbulifera Castillejos & E. Solano, M. guerrerensis Matuda, M. justosierrana García-Mend., and M. parva Aaron Rodr.; Table 1).
Table 1 Diversity and distribution of the genus Manfreda in Guerrero, Mexico.
| Species | Distribution in Mexico | Municipalities and records | Elevation range | Vegetation types | |
|---|---|---|---|---|---|
| 1. | M. arceliensis Art.Castro, J.G. Zavala & Cruz Durán | Guerrero | Arcelia (2) | 1,248-1,393 | Oak forest |
| 2. | M. bulbulifera Castillejos & E.Solano | Guerrero | Chilapa de Álvarez (1), Colotlipa (3) | 1,200-1,283 | Pine-oak forest |
| 3. | M. chamelensis E.J.Lott & Verh.-Will. | Guerrero, Jalisco | José Azueta (2), Técpan de Galeana (2) | 15-860 | Tropical deciduous forest, pine forest |
| 4. | M. guerrerensis Matuda | Guerrero | Carrizal de Bravo (1), Guayameo (1), Zirándaro (1) | 950-2,300 | Oak forest |
| 5. | M. hauniensis (J.B. Petersen) Verh.-Will. | Estado de México, Guerrero, Morelos, Oaxaca | Alpoyeca (1) Atlixtac (1), Atoyac de Álvarez (2), Chilpancingo de los Bravo (9), Eduardo Neri (1), Malinaltepec (1), Metlatonoc (1), San Miguel Totolapan (1), Xochihuehuetlán (1) | 545-2,628 | Oak-pine forest, tropical deciduous and sub deciduous forest |
| 6. | M. aff. jaliscana Rose | Durango, Guerrero, Jalisco, Michoacán, Nayarit, Sinaloa, Sonora, Zacatecas | Chilpancigo de los Bravo (3), Eduardo Neri (1), Tixtla de Guerrero (1) | 650-1,600 | Oak forest, tropical deciduous forest |
| 7. | M. justosierrana García-Mend. | Guerrero | Leonardo Bravo (2), Técpan de Galeana (1) | 1,940-2,328 | Oak-pine forest |
| 8. | M. littoralis García-Mend., A.Castañeda & S.Franco | Guerrero, Oaxaca | Acapulco (3) | 220-406 | Tropical deciduous forest |
| 9. | M. maculata (C. Mart.) Rose | Estado de México, Guerrero, Oaxaca | Chilpancingo de los Bravo (1), Iguala de la Independencia (1), Taxco de Alarcón (1) | 1,000-1,850 | Pine-oak forest, tropical deciduous forest |
| 10. | M. parva Aarón Rodr. | Guerrero | Taxco de Alarcón (4) | 1,818-1,845 | Coniferous forest, pine-oak forest |
| 11. | M. pringlei Rose | Ciudad de México, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Oaxaca, Puebla, Tlaxcala, Veracruz | Chichihualco (3), Coahuayutla de José María Izazaga (1), Leonardo Bravo (1), Mochitlán (1) | 850-2,450 | Cloud forest, oak-pine forest, tropical deciduous forest |
| 12. | M. pubescens (Regel & Ortgies) Verh.-Will. ex Piña | Chiapas, Guerrero, Morelos, Oaxaca, Puebla | Buenavista de Cuéllar (4), Chilapa de Álvarez (1), Chilpancingo de los Bravo (1), Iguala de la Independencia (1), Leonardo Bravo (1), Pilcaya (1), Taxco de Alarcón (3), Tlapa de Comonfort (1) | 800-2,420 | Oak-pine forest, palmar, savannoid vegetation, tropical deciduous forest |
| 13. | M. scabra (Ortega) McVaugh | Aguascalientes, Chiapas, Colima, Distrito Federal, Durango, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosí, Veracruz, Zacatecas | Ajuchitlán del Progreso (2), Arcelia (1), Azoyú (1), Chichihualco (1), Chilapa de Álvarez (2), Chilpancingo de los Bravo (4), Coyuca de Benítez (2), Eduardo Neri (1), Iguala de la Independencia (2), Mártir de Cuilapan (1) | 362-2,010 | Oak-pine forest, tropical deciduous forest |
| 14. | M. umbrophila García-Mend. | Guerrero, Oaxaca | Chilapa de Álvarez (3) | 1,860-1,950 | Oak-pine forest |
Manfreda species in Guerrero have been collected mostly in temperate and cold conditions with coniferous, oak-pine, oak, and pine forest, where 13 species are located: M. arceliensis, M. bulbulifera, M. chamelensis E.J. Lott & Verh.-Will., M. guerrerensis, M. hauniensis (J.B. Peterson) Verh.-Will., M. aff. jaliscana Rose, M. justosierrana, M. maculata (C. Mart.) Rose, M. parva, M. pringlei Rose, M. pubescens (Regel & Ortgies) Verh.-Will. ex Piña, M. scabra (Ort.) McVaugh, and M. umbrophila García-Mend. Meanwhile, in warm climatic conditions with tropical deciduous forest, as in subdeciduous tropical forest, savannoid vegetation, and palmar, it is possible to locate: M. chamelensis, M. hauniensis, M. littoralis García-Mend., A. Castañeda & S. Franco, M. maculata, M. pubescens, and M. scabra (Table 1).
The elevation range in which Manfreda has been collected in Guerrero is wide, from 15-450 m in Acapulco, Ajuchitlán del Progreso, José Azueta, and Técpan de Galeana [M. chamelensis, M. littoralis and M. scabra]; to 1,900-2,628 m in Alpoyeca, Atilxtac, Atoyac de Alvarez, Carrizal de Bravo, Chichihualco, Chilapa de Álvarez, Eduardo Neri, Leonardo Bravo, Metlaltónoc, Taxco de Alarcón, and Técpan de Galeana [M. guerrerensis, M. hauniensis, M. justosierrana, M. pringlei, M. scabra, M. pubescens, and M. umbrophila].
According to Morrone et al. (2017) biogeographic provinces in Guerrero, Manfreda has been collected in the Sierra Madre del Sur (37 records / 10 spp.), Balsas Basin (26 records / 9 spp.), Transmexican Volcanic Belt (8 records / 4 spp.), and Pacific Lowlands (8 records / 3 spp.) (Figure 1).
Figure 1 Species richness of Manfreda in Guerrero, Mexico. In the cutout of the lower right corner, the two cells with the greatest richness are indicated. The grid comprises cells with an area of 144 km2. Biogeographical regionalization according to Morrone et al. (2017).
In the grid analysis, 34 cells had at least one record, that is, ±7 % of the analyzed area. The number of species per cell varied from one to four, but just two cells contained four species. In the first of these cells inhabit Manfreda aff. jaliscana, M. justosierrana, M. maculata, and M. pringlei. While M. guerrerensis, M. justosierrana, M. pringlei, and M. pubescens were found in the second cell. In general, Manfreda richness is concentrated in 8 cells located in the municipalities Chilpancingo de los Bravo, Leonardo Bravo and Taxco de Alarcón, which are in the Balsas Basin, Sierra Madre del Sur, and Transmexican Volcanic Belt biographical provinces, and bring together nine species (Figure 1).
Discussion
As might be expected, the Manfreda species richness distribution patterns in Guerrero are similar to those patterns found in other angiosperm groups. For example, there is a geographical correspondence between Manfreda richness areas in Guerrero and the diversity and endemism patterns for another 507 species of Mesoamerican geophyte monocots (Sosa & Loera 2017). Moreover, the same areas identified here are also associated with a high degree of richness, endemism and conservation priority for other plant groups; such as the perennial herbs species of Cosmos section Discopoda (DC.) Sherff (Vargas-Amado et al. 2013, 2019), bulbous herbaceous of Iridaceae tribe Tigridieae (Munguía-Lino et al. 2016), 79 species of the genus Salvia L. (Martínez-Gordillo et al. 2016) and the areas of endemism found for 32 taxa restricted to the Sierra Madre del Sur (Santiago-Alvarado et al. 2016).
The greatest species richness of Manfreda in Mexico is found in Guerrero, wherein 14 species have been registered and five are endemic (Castro-Castro et al. 2017, 2018; Table 1). The relatively low number of collections for the genus, which have been carried out in a period of just less than 100 years (1926-2014), is striking. Manfreda collections in Guerrero are limited to 33 of the 81 municipalities in the state, 40.7 % of the political territory of the state. Given the high diversity of Manfreda in Guerrero, the elevated endemism and the low coverage of botanical explorations focus on this group in the state, current estimates of species diversity are likely under-estimates and new species are likely to be found. For example, although in Guerrero the greatest diversity of Manfreda has been found in temperate and cold environments, it is advisable to focus on the collection efforts towards the tropical areas of Pacific Lowlands, since according to Castro-Castro et al. (2017, 2018), in this province is where the genus Manfreda has its highest species richness in Mexico. A final recommendation in this regard, is the detailed review of the plants included as M. aff. jaliscana. The six specimens analyzed show a homogeneous morphology among them, but contrasting in the sizes of the leaves, flowers and filaments with respect to other specimens collected in western and northwestern Mexico, the area where the species was described (Rose 1903), and it has been reported more frequently (Castro-Castro et al. 2017). In addition, the phenology of the populations of Guerrero differs in what is known for M. jaliscana, which has been reported in bloom mostly during the dry period (see Appendix 1 and Rodríguez & Castro-Castro 2007).
During the present investigation it was possible to detect another set of specimens that could not be assigned to any of the known species in Manfreda. The plants were collected in the municipality of Arcelia, an area where only two specimens of the genus had been previously collected before 2006 (Appendix 1). Given the general morphology, these plants could be related to M. maculata and M. parva. However, there are multiple differences that allow them to be recognized as an undescribed species. These dissimilarities and similitudes are detailed in the following lines together with data on the ecological preferences, taxonomic notes of the new species and a key for the identification of Manfreda in Guerrero.
Taxonomic treatment. Manfreda arceliensis Art.Castro, J.G. Zavala & Cruz Durán sp. nov. (Figure 2).
Type: Mexico. Guerrero: Arcelia municipality, Filo del Cerro el Tepehuaje, Campo Morado, 1393 m, 19 July 2006 (fl), R. Cruz-Durán et al. 6428 (Holotype: FCME; Isotype: CIIDIR).
Figure 2 Manfreda arceliensis Art.Castro, J.G. Zavala & Cruz Durán. A. Habit. B. Lateral view of flower. C. Perigone dissection and ovary longitudinal section. D. Detail of the tuft of short hairs on the top of perigone lobes. E. Floral bract and bracteoles. Illustrated by Lesslie Martínez Rubí.
Diagnosis. Manfreda arceliensis is morphologically similar to M. maculata and M. parva, but it can be distinguished from the former by the herbaceous leaves, broadly oblanceolate, flat, and glabrous (vs. coriaceous, lanceolate to oblanceolate, slightly channeled, undulate, and densely pubescent on both faces), inflorescence 41-52 cm long with 2-4 flowers [vs. (30-) 60-120 cm long with 6-22 flowers], first sterile bracts 1.4-3.3 cm long (vs. 4.5-8.5 cm long), perigone funnel-shaped (vs. tubular), and filaments inserted 5-5.2 mm below the sinuses of the lobes (vs. 6-10 mm). For its part, it distinguishes from M. parva for developing a bulb ovoid, 3-3.8 × 2.3-2.7 cm (vs. oblong, 2-2.5 × 1-2 cm), leaves 2-3, broadly oblanceolate, flat, green with densely purple spots [vs. (5-)11-24, linear, channeled, green concolorous], inflorescence with 3-4 flowers [vs. (4-)9-15], peduncle 38-48 cm long with 3-4 bracts (vs. peduncle 19-26 cm long with 4-7 bracts), and the first sterile bracts linear, 1.4-3.3 × 0.12-0.25 cm (vs. lanceolate, 6-8 × 0.7-1 cm).
Perennial herb, 42-52 cm tall. Corm ovoid 3-3.8 × 2.3-2.7 cm, with small buds and thickened, contractile and fleshy roots. Roots filiform, 3.5-6.5 cm long. Bulb ovoid, 3-3.8 × 2.3-2.7 cm, tunicate, straw. Leaves 2-3 in a basal rosette, alternate, broadly oblanceolate, 10-14.5 × 1.9-4 cm, erect, flat, thin to leathery, green with densely purple spots, upper and lower surface tiny papillate, acute and soft apex, narrowed into a basal petiolar portion, margin hyaline, smooth. Inflorescence spiciform, 41-52 cm long, erect, arcuate apex, internodes decreasing in length towards the apex; peduncle 38-48 cm long, bracts 3-4, first sterile bracts linear, 14-33 × 1.2-2.5 mm, the closest to the rachis 6.6-7 × 0.4-0.7 mm, both acute, truncate, clasping, thin, brownish green, concolorous, upper and lower surface smooth, hyaline, adpressed to divaricate; rachis lax, 1-2.6(-4.5) cm long, with 2-4 floral nodes, internodes 1.2-3 cm long, primary floral bracts linear, 1.6-4.4 × 0.5-0.6 mm, acute, truncate, smooth, persistent, yellow-brow, erect or reflexed. Bracteoles 1 at each floral node, ensiform to acicular, 0.8-1.1 × 0.1-0.2 mm, entire, yellow-brow. Flowers actinomorphic, funnel-shaped, ascending, yellowish green and with dense and fine points of reddish purple on the outside and yellow pale inside in anthesis; perigone tube 1.5-1.8 cm long, 0.6-0.9 cm wide at the base of the lobes, tubular but widen slightly towards its distal portion, slightly curved; lobes 6.5-7.5 × 3.6-4.4 mm, ovate-deltate, markedly divergent to reflexed in anthesis, apices with a tuft of short white hairs. Stamens epitepalous, antitepalous, phanerantherous; filaments subulate, 1.7-1.9 cm long from their insertion into the perigone tube, inserted 5-5.2 mm below the sinuses of the lobes, yellow brown to reddish; anthers elliptic, 8-9 × 1.5-1.8 mm, yellow with dense and fine points of reddish purple. Ovary 7.5-9.7 × 3.2-4 mm, ovoid to narrowly ovoid; style filiform, 2.8-3 cm long in anthesis, purple, exserted from the perigone tube by 1.3-1.5 cm in anthesis; stigma trigonous, trilobed, papillose, clavate. Fruit unknown.
Distribution, ecology, and phenology. Manfreda arceliensis is endemic to northern Guerrero in the transition zone between Balsas Basin and Sierra Madre del Sur biogeographic provinces. It inhabits oak forest with savannoid vegetation elements; in areas with deep and acid soils at an elevation of 1,248-1,393 m. Manfreda arceliensis is associated with Byrsonima crassifolia (L.) Kunth, Cardiostigma longispatha (Herb.) Baker, Clethra hirsutovillosa S. Valencia & Cruz Durán, Dodonaea viscosa Jacq., Psidium sp. and Quercus magnoliifolia Née. The species flowers in July.
Conservation status. Manfreda arceliensis is represented by two collections in close proximity. The EOO was of 0.76 km2, and the size of the AOO is 8 km2, based on cells of 2 × 2 km. Adjusting to EOO and AOO areas results and following the IUCN (2019) criteria (extent of occurrence estimated to be less than 100 km2 and known to exist at only a single location), a preliminary category of Critically Endangered is proposed (CR/B1a and B2a). It is advisable to perform additional explorations and collections in the area where the species is found.
Etymology. The name of the new species refers to the municipality of Arcelia, Guerrero, Mexico.
Additional specimens examined. Mexico. Guerrero: Arcelia municipality, cerca de la cañada El Limón, Campo Morado, 1,248 m, 17 July 2006 (fl), R. Cruz-Durán et al. 6287 (CIIDIR, FCME).
Relationship and taxonomic position. Manfreda arceliensis is the smallest species among all the taxa of the genus present in Guerrero. Moreover, it is the species that develops fewer flowers in Manfreda. Morphologically, M. arceliensis is similar to M. maculata and M. parva, the differences and similarities in morphological traits among these three species are highlighted in the diagnosis and the key for Manfreda in Guerrero provided below.
Key for Manfreda in Guerrero, Mexico
1. Perigone tube 0.5-2.8 mm long ... 2
1. Perigone tube 6-33 mm long ... 3
2. Leaves 9-27 × 2-6 cm, elliptic to lanceolate-elliptic, flat, green concolorous; perigone lobes 2.5-3.5 cm long, linear, reflexed; filaments 1.3-2.1 cm long … M. littoralis
2. Leaves 12-25 × 0.3-0.6 cm, linear to lanceolate, channeled, green with purple macules; perigone lobes 0.8-1.1 cm long, linear-oblong, erect to reflexed; filaments 5-6.5 cm long … M. bulbulifera
3. Leaves pubescent … 4
3. Leaves glabrous … 6
4. Filaments 1.7-2.2 cm long … M. maculata
4. Filaments 3.5-5.2 cm long … 5
5. Leaves 2-3, 20-45 × 6-8 cm, markedly narrowed and pubescent toward the blade base … M. guerrerensis
5. Leaves 3-5(-9), 18-28(-70) × 2.2-4.2(-6) cm, not markedly narrowed toward the base and, densely pubescent on the entire surface … M. pubescens
6. Rachis dense, internodes 0-1 cm long … 7
6. Rachis lax, internodes 1.2-4(-6) cm long … 8
7. Corm 3-5 × 1.5-2.5 cm; leaves 15-56(-80) × 0.8-2.8 cm, dark green; filaments 2.6-4.5(-5.4) cm long … M. pringlei
7. Corm (4-)8-10 × (1.5-)2-3.5 cm; leaves (25-)35-45 × (5-)7-10 cm, light green, lustrous, with dark macules; filaments (4.5-)7-8.5 cm long … M. umbrophila
8. Plants flowering from January to May, during the dry season … M. chamelensis
8. Plants flowering from July to November, mainly during the rainy season … 9
9. Plants (1-)1.5-3.8 m tall … 10
9. Plants 0.4-0.6 m tall … 12
10. Leave margin irregularly denticulate to denticulate-erose, the teeth small and simple to large and bifid or trifid; inflorescence 2-3.8 m tall; filaments exceeding the tube by 6.3-10.5 cm … M. hauniensis
10. Leave margin whole to papillose; inflorescence 1.4-2.5 m tall; filaments exceeding the tube by 3-5.5 cm … 11
11. Leaves widely oblong to elliptical, coriaceous; basal peduncle bracts similar in shape and size to the leaves; flowers sessile to pedicellate; filaments inserted 1/3 to 1/5 from top of the tube or just in the tube mouth … M. scabra
11. Leaves linear-lanceolate, coriaceous to herbaceous; basal peduncle bracts clearly shorter than the leaves; flowers sessile; filaments inserted 1/4 from top of the tube … M. justosierrana
12. Bulb ovoid, 3-3.8 × 2.3-2.7 cm; leaves 2-3, broadly oblanceolate, flat, green with densely purple spots; inflorescence with 2-4 flowers; peduncle 38-48 cm long, sterile bracts 3-4, the first one linear, 1.4-3.3 × 0.12-0.25 cm … M. arceliensis
12. Bulb oblong, 2-2.5 × 1-2 cm; leaves (5-)11-24, linear, channeled, green concolorous; inflorescence with (4-)9-15 flowers; peduncle 19-26 cm long, sterile bracts 4-7, the first one lanceolate, 6-8 × 0.7-1 cm … M. parva
