Research Article |
Corresponding author: Jorge Gutiérrez ( jgutierrezg@chapingo.mx ) Academic editor: Clifford Morden
© 2020 Jorge Gutiérrez, Teresa Terrazas.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gutiérrez J, Terrazas T (2020) Xochiquetzallia (Asparagaceae, Brodiaeoideae), a new genus segregated from the paraphyletic Dandya. PhytoKeys 139: 39-49. https://doi.org/10.3897/phytokeys.139.46890
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A new genus, Xochiquetzallia, for the Brodiaeoideae, Asparagaceae family is here proposed. A taxonomic analysis based on morphology highlights its synapomorphies. The characters that distinguish Xochiquetzallia are the absence of a pith in the gynophore and the presence of an entire stigma. The recognition of Dandya purpusii as a monotypic genus is supported by the development of a short floral tube (< 2 mm) and a pith in the gynophore, as well as a divided stigma shared with the other genera of the Milla clade, Bessera, Jaimehintonia, Petronymphe and Milla. A key to its taxonomic determination is given for both the Xochiquetzallia species and the Milla clade genera.
Asparagales, geophyte, gynophore, Mexico, Milla clade
Dandya H.E. Moore is a genus endemic to Mexico and one of the five genera of the Milla clade (
Plant morphology was analyzed from field collected and herbarium material (ARIZ, BH, CHAPA, F, FC, GH, IEB, JEPS, MEXU, NY, RSA, UAMIZ, US and XAL). We studied the floral morphology from organisms collected in the field, except for Milla mortoniana (material removed from herbarium, MEXU). Morphological characters, vegetative and reproductive, were observed and analyzed with the help of a microscope (Nikon SMZ-2T) and terminology follows various authors (
Based on the phylogenetic clades recovered by
Perennial herbs, geophytic; terete; subcampanulate flowers, tube 1.5–3.0 mm long, erect; gynophore with pith, stigma divided.
Dandya purpusii (Brandegee) H.E. Moore, Gent. Herb. 266–268 (1953).
Muilla purpusii Brandegee, Univ. Calif. Publ. Bot. iv, 177 (1911).
Bloomeria purpusii (Brandegee) Macbride, Contr. Gray Herb. Ser. 2. 56: 1-20 (1918).
Brodiaea purpusii (Brandegee) Ingram, Madroño xii, 27 (1953).
Perennial herbs, 28–47 cm tall, including corm and inflorescence. Fibrous roots. Corm subglobose compressed, fleshy, 1.0–1.7 cm in diameter; tunic formed by the wide bases of the leaves, brown or dark brown, covering up to 2.0 cm from the base of the scape. Leaves 2–3, 13–27 cm long, dark green, linear, subterete, with scabrous surface, hyaline prominences on the veins; base truncated, apex acute. Inflorescence in umbel; Scape of 27–42 cm long, usually longer than leaves, terete, surface smooth or with acute prominences. Floral bracts 2–3, linear-lanceolate, triangular, 4.0–8.5 mm long; bracteoles, one per flower. Flowers 6–11, pedicels 2.0–4.0 cm long, subcampanulate, erect, articulate, floral tube 1.5–3.0 mm long; tepals blue, 6 in 2 series, external tepals ovate-lanceolate, 6.0–10.0 × 1.5–3.0 mm, apex acute and papillose, base cuneate, entire margin; internal tepals ovate-lanceolate, 6.0–9.5 × 1.6–3.0 mm, apex obtuse and papillose, base cuneate, entire margin. Stamens 6; filaments free, adnate to the throat of the tube, widened toward the base, 4.0–6.0 mm; anthers oblongs, yellow, basifixed, 1.5–2.0 mm; gynophore 0.5–1.1 mm long, adnate to the floral perigone formed three cavities, present pith. Ovary cylindrical, 2.5–5.5 mm, fused at its base to the floral perigone; style filiform, 2.1–4.0 mm; stigma divided, papillose; capsule loculicidal, subcylindric, glabrous, brown, 10.0–12.0 mm long; seed oblong-falciform, compressed, black, bright, 3.0 × 5.0 mm.
MEXICO. Coahuila; Sierra de la Paila, October 1910, Purpus 4959 (holotype UC!; isotypes F!, GH!).
MEXICO: Coahuila, Municipality of Ramos Arispe, Valle de los Ángeles, Sierra de la Paila, 6 August 1975, M. F. Robert & J. Passini 4675 (ENCB); El Cidral, Sierra de la Paila, 20 August 1987, J. A. Villarreal 3980 (TEX); 5.9 miles east of the road between Yucatan and Mexico, highway 40, west of the mountains along a gravel road to 17.2 miles north of Mexico highway 40, 1650 m elevation, 19 September 1996, J. M. Porter 11308 & J. T. Columbus (RSA); 15 km north of Estación Marte, on secondary road, 1550 m elevation, 24 October 2011, J. Gutiérrez et al. 1225 (FEZA, CHAPA, MEXU, UAMIZ).
Perennial herbs, geophytic; terete or flattened leaves; subcampanulate or hypocrateriform flowers, tube 1.0–25.0 mm long, erect or reclined; gynophore without pith, stigma entire.
Perennial herbs, 20–60 cm tall, including corm and inflorescence. Fibrous roots, some fleshy. Corm subglobose compressed, fleshy, 1.0–2.5 cm in diameter; tunic formed by the wide bases of the leaves, brown or dark brown, covering up to 2.0 cm from the base of the scape. Leaves 5–9, 20–49 cm long, dark green, linear, flattened or terete, with glabrous or scabrous surface, hyaline prominences on the veins; base truncated, apex acute. Inflorescence in umbel; Scape of (16–) 20–50 cm long, usually shorter than leaves, terete, surface smooth or with acute prominences. Floral bracts 2–3, linear-lanceolate, triangular, 3.0–9.0 mm long; bracteoles, one per flower. Flowers 4–20, pedicels 0.8–3.5 cm long, subcampanulate or hypocrateriform, erect or decumbent-descending, articulate, floral tube 1–25 mm long; tepals white or blue, 6 in 2 series, external tepals elliptic, 8.0–16.0 × (1.5–) 2.0–7.0 mm, 1–3 veins, apex acute and papillose, base cuneate, entire margin; internal tepals elliptical to broadly elliptical, (6.5–) 8.0–16.0 × (2.0–) 3.0–11.0 mm, apex obtuse and papillose, base cuneate, entire margin. Stamens 6; filaments free, adnate to the throat of the tube, widened toward the base or columnar, 2.0–5.0 (–7.0) mm; anthers linear, lanceolate-deltoid, yellow, basifixed, 1.0–2.5 mm; gynophore 0.8–1.6 mm long, adnate to the floral perigone formed three cavities. Ovary cylindrical, 1.0–5.0 mm, fused at its base to the floral perigone; style filiform, 1.8–7.0 mm; stigma entire, papillose; capsule loculicidal, subglobose or subcylindric, glabrous, brown, 6.0–13.0 mm long; seed oblong-falciform, compressed, black, bright, seed coat papillose, 4.0 × 1.5 mm.
Xochiquetzallia mortoniana (H.E. Moore) J.Gut.
This genus is named in honor of the goddess of Aztec flowers, in Nahuatl "Xōchiquetzalli" (beautiful flower) "xṓchitl" (flower), "quétzalli" (beautiful). The Aztecs developed majestic architectural works, had extensive knowledge of astronomy and great respect for nature, particularly plants.
1 | Stigma entire | Xochiquetzallia |
– | Stigma dissected | 2 |
2 | Connate stamens, dorsifixed anthers | Bessera |
– | Free stamens, dorsifixed or subdorsifixed anthers | 3 |
3 | Subcampanulate flowers, gynophore < 2 mm | Dandya |
– | Tubular flowers, gynophore > 2 mm | 4 |
4 | Flowers green-yellow, orange-red; tepals ascending or erect | Petronymphe |
– | Flowers white, purple or pink; tepals divaricate | 5 |
5 | Diffuse-ascendant tepals; filaments 7-8 mm; floral tube < 15 mm long | Jaimehintonia |
– | Divaricated tepals, occasionally reflexed; filaments < 7 mm; floral tube >15 mm long | Milla |
≡ Dandya balsensis López-Ferr. & Espejo in Act. Bot. Mex. 18: 11–15, f. 1–2. (1992), basionym Type:– MEXICO. Morelos: Municipality Talquiltenango, road between Valle de Vázquez and Chimalacatlán, 1200 m elevation, 25 June 1989, A. Flores Castorena 1075 & D. Martínez Alvarado (holotype, UAMIZ!) (Figs
MEXICO, Morelos: Municipality Talquiltenango, road between Valle de Vázquez and Chimalacatlán, 1200 m elevation, 25 June 1989, A. Flores Castorena 1075 & D. Martínez Alvarado (isotypes, ENCB! IEB!); Ibid, 06 July 2006, J. Gutiérrez 797 (CHAPA, FEZA, UAMIZ, MEXU); 9 June 2007, J. Gutiérrez 839 (CHAPA, FEZA, UAMIZ, MEXU).
≡ Dandya hannibalii L.W. Lenz in Aliso 7(3): 316, f.2. (1971), basionym Type:– MEXICO, Michoacán: about 10 miles south of Cuatro Caminos, at km 165, Mexico 37, on the northwestern slopes of Sierra Madre, 31 July 1967, T. M. Howard 67-64 (holotype, RSA 190791!) (Figs
MEXICO, Michoacán: Municipality of Huetamo, Balsas road km 72, near Las Cruces bridge, 9 July 2006, J. Gutiérrez 805 (CHAPA, FEZA, UAMIZ, MEXU); ibid., 2 September 2006, J. Gutiérrez 813 (CHAPA, FEZA, UAMIZ, MEXU).
≡ Milla mortoniana H.E. Moore in Gentes Herbarum 8: 291 (1953), basionym Type: MEXICO, Guerrero: Distrito Mina, Tierras Blancas, 1400 m., 19 October 1936, Hinton 9725 (holotype GH!; isotypes NY!, US!) (Figs
MEXICO, Michoacán: Municipality of Aquila, Sánchez-Mejorada et al. 4301 (MEXU!).
≡ Dandya thadhowardii L.W. Lenz in Aliso 7(3): 314, f.1. (1971), basionym Type: MEXICO, Guerrero: About 25-30 miles south of Iguala, on hillsides in calcareous soil, at km 216 on Mexico 95, July 1964, 1965, 1966, Howard 64-74 (holotype!, RSA 100784) (Figs
MEXICO, Guerrero: Municipality of Cutzamala de Pinzón, 3 km north of Cutzamala river, road to Bejucos, 340 m elevation, 21 July 1986, A. Espejo 2481 & T. Chehaibar (UAMIZ); Municipality of Eduardo Neri, Barranca de Xococoapa, 1000 m elevation, 20 July 1991, S. Peralta et al. 231 (FCME); Municipality of Xochipala, Llano Delgado, 1035 m elevation, 21 July 1991, M. Gual 260 (FCME); km 62 highway Iguala-Chilpancingo, 910 m elevation, 4 July 1980, Campos & Castelo 56 (FCME); ibid., Campos & G. Velázquez 118 (FCME); 6 km east-northeast of Xochipala, 2 July 1980, J. Saldivar & Sánchez s. n. (FCME); ibid., Velázquez Toledo & Campos 63 (FCME); ibid. on the northwest hillside, 13 July 1991, M. Luna Flores 43 (FCME); Km 169 highway Iguala-Chilpancingo, 9 June 2007, J. Gutiérrez 840 (CHAPA, FEZA, UAMIZ, MEXU); Highway Iguala-Chilpancingo, 2 km towards Filo de Caballo, 9 June 2007, J. Gutiérrez 841 (CHAPA, FEZA, UAMIZ, MEXU); Municipality of Coyuca de Catalán, 3 km west of Coyuca de Catalán, 10 June 2007, J. Gutiérrez 844 (CHAPA, FEZA, UAMIZ, MEXU). Michoacán: Municipality of Huetamo de Nuñez, 500 m towards Petachícuaro, 430 m elevation, 20 July1986, A. Espejo 2467 T. Chehaibar (UAMIZ); Petachícuaro, 9 km north of Huetamo, 400 m elevation, 9 July 1982, José C. Soto & Esteban Martínez 4047 (ENCB, MEXU, UAMIZ); Municipality of San Lucas, highway Cd. Altamirano-San Lucas, km. 211, Rancho el Ovispo, 339 m elevation, 9 July 2006, J. Gutiérrez 805 (CHAPA, FEZA, UAMIZ, MEXU); highway Cd. Altamirano-San Lucas, km. 188, 352 m elevation, 9 julio 2006, J. Gutiérrez 806 (CHAPA, FEZA, UAMIZ, MEXU).
1 | Flowers white, divaricate-recline | 2 |
– | Flowers blue-violet, erect | 3 |
2 | Filaments ≤ 3 mm long; style ≤ 4 mm long; known from the State of Morelos | X. balsensis |
– | Filaments > 3.0 mm long; style 4-6 mm long; known from the States of Guerrero and Michoacán | X. thadhowardii |
3 | Perianth tube ≤ 2 mm long; tepals ≤ 1 cm long; filaments ≤ 4 mm long | X. hannibalii |
– | Perianth tube 2-2.5 cm long; tepals 1.5-1.6 cm long; filaments 2 mm long | X. mortoniana |
Xochiquetzallia balsensis (Dandya balsensis) and X. thadhowardii (D. thadhowardii) present morphological similarity and have a sympatric geographic distribution (Fig.
In Dandya and Xochiquetzallia, the connate stamens have been a recurrent character in the descriptions (
The existence of gynophore has also been a controversial feature. A gynophore was not described for the former species of Dandya now in Xochiquetzallia (X. balsensis, X. hannibalii and X. thadhowardii;
J. Gutiérrez thanks CONACYT for the grant awarded (160287) to perform doctoral studies. The authors also thank Ramiro Ríos Gómez and Eliseo Bravo Acevedo for their help in the field collection of the specimens studied and to Julio César Montero Rojas for artwork. The comments of a reviewer and the subject editor, Clifford Morden, are highly appreciated.