Hechtia carrilloi I. Ramírez, 2023

Ramírez-Morillo, Ivón, Ramírez-Díaz, Claudia J., Brunel, Juan Pablo Ortiz & Romero-Soler, Katya J., 2023, Male flowers reveal the true identity of a new species of Hechtia (Bromeliaceae) from the Mexican state of Jalisco, Phytotaxa 626 (1), pp. 21-31 : 22-29

Hechtia carrilloi I. Ramírez , sp. nov. ( Figures 1–4 ).

TYPE:— MEXICO: Jalisco: Municipio Tolimán, Arroyo La Ciénega , ca. 1 km al N de Tolimán, 19°36’33.4’’N, 103°54’33’’W, 783 m, collected by C. Ramírez-Díaz, 13 January 2015, cultivated and flowered in Yucatán, March 6 2018, I. Ramírez-Morillo 2020 J (Holotype, IBUG!) .

Diagnosis:—This new species resembles Hechtia santanae , but differs in its smaller rosettes (10–12 cm height × 28–30 cm diameter vs. 20–30 × 20–40 cm), strict sympodial growth pattern (vs. sympodial with precocious-flowering pattern); adaxially sparsely lepidote to glabrous leaves (vs. densely silvery-lepidote), staminate inflorescence features: stipes ebracteate and terete (vs. bracteate and dorsally flattened in H. santanae ), peduncle internodes 0.5–1.2 cm long (vs. 1–2.5 cm), peduncle bracts smaller (0.8–2.5 × 0.3–0.4 mm vs. 20–90 × 7–9 mm), branches forming an 85–90° angle with the axis of fertile part (vs. 40–75°), shorter branches ((1.5–) 2–6 vs. (3.5–) 5.5–13.5 cm), flowers with tubular corolla (vs. obconical), flower size (4–5 mm long, 1–2 mm in diameter vs. 5–5.5 mm long, 3.8–4.2 mm in diameter), filaments longer than the petals (vs. equaling the petals), anthers maroon (vs. reddish).

Description:—Plant forming caespitose rosettes, 10–12 cm height × 28–30 cm in diameter, producing offshoots by means of stolons, sometimes at the base of the inflorescence. Leaves 15–20, central ones erect, basal ones erect to slightly reflexed; sheath oblong, 2–3.5 × 1.3–1.5 cm, pale yellow, basally light brown when dry, mostly glabrous and densely white lepidote only close to the blade, both in adaxial and abaxial sides, margins serrulate; blade narrowly triangular, attenuate, 11–17 × 1.6–1.8 cm, succulent, green, sparsely silvery lepidote adaxially, densely white lepidote abaxially, margins spiny; spines antrorse, triangular, 1–2 mm long, 1–3 cm apart, brownish, with a thin tuft of white trichomes at the axil of the basal spines. Inflorescence emerging from the center of a mature rosette (strict sympodial growth pattern SPP, sensu Ramírez et al., 2014).

Staminate inflorescence a 1-divided panicle, fertile part cylindrical in outline, erect, 90–110 cm long (including the peduncle); peduncle terete, 42–65 cm long, 0.4–0.6 cm in diameter, green with purple blots, surface sparsely whitepuberulent, three times longer than the rosette height; peduncle internodes varying in length, from 0.5–1.2 cm long and decreasing in length distally; peduncle bracts triangular, acute, 0.8–2.5 × 0.3–0.4 cm, brownish, multinerved, usually much shorter than internodes; main axis 42–80 cm long, 0.3–0.4 cm in diameter at the base, terete, green tinged purple, sparsely white-lepidote; internodes 0.4–1 cm long; primary bracts narrow-triangular, attenuate, acute, (5–) 8–9 × 1–2 mm, sparsely lepidote on both surfaces, papyraceous, longer than the stipe of the lower branches, almost equaling the length of that of the shorter upper branches, margins entire to erose, brownish, 5-nerved; branches 30–50 in number, in an angle of 85–90° with to the main axis, (1.5–) 2–6 cm long, 0.6–1.0 cm in diameter, each with (8–) 20–65 densely arranged flowers; stipe (0.3–) 0.6–1 cm long, ebracteate; rachis 0.5–1 mm in diameter, terete; floral bracts triangular, acuminate, ca. 1–1.1 mm × 0.4–0.5 mm, concave, light green, margins erose, lepidote on both surfaces, 1-nerved, as long as or shorter than the pedicel. Flowers pedicellate, patent, 4–5 mm long, 1–2 mm in diameter (including petals and stamens up to 5 mm in diameter), actinomorphic; pedicels terete, 0.8–1 mm long, 1–1.5 mm in diameter; sepals ovate, rounded, green with red spots in the middle area, 0.8–1 x 0.9–1 mm, sparsely lepidote abaxially, glabrous adaxially, margins erose, 3-nerved, shorter than the petals; petals broadly elliptic, rounded, 2.5–2.8 × 1.5–2 mm, glabrous on both surfaces, white, faintly 5-nerved; stamens with filaments triangular, flattened, ca. 3 mm long, white, exceeding the petals at anthesis; anther oblong, ca. 0.8–1 × 0.3–0.5 mm, dorsifixed, maroon; pistillode ca. 1 mm long, yellowish.

Pistillate inflorescences a 1-divided panicle, cylindrical in outline, erect, 90–145 cm long (including peduncle); peduncle terete to slightly compressed at the base, 48–66 cm long, 0.8–1.3 cm in diameter at the base, much longer than the rosette height but shorter than the main axis, green tinged brown, surface smooth, glabrous to scatteredly lepidote; peduncle internodes 1.5–4.5 cm long; peduncle bracts foliaceous, triangular, long-attenuate, acute, (0.8–) 2.5–7.5 cm long, 0.5–1.2 cm wide at the base, brownish, margins hyaline at the base, entire to erose, sparsely spinulose toward the apex, multinerved, glabrous at the base to densely white lepidote towards the apex on both surfaces, usually longer than to rarely equaling the internodes; main axis 42–79 cm long, 0.4–0.8 cm in diameter at the base, terete, green to brownish, internodes 0.5–3 cm long; primary bracts triangular, attenuate, acute, 0.5–1.4 × 0.4–0.8 cm, entire to erose or sometimes spinulose, brownish, multinerved, glabrous to sparsely white-lepidote on both surfaces, usually not exceeding the stipe of the branches; branches 35–45 in number, in an angle of 45–60° with the main axis, (2–) 6–8 (–10) cm long, (6–) 28–42 flowered; stipe 0.4–1 (–2) cm long; rachis 1–2 mm in diameter, sulcate, green, glabrous; floral bracts deltoid, acuminate, 1–1.5 × 1–1.5 mm, sparsely lepidote abaxially, glabrous adaxially, margins erose, 1–3-nerved, adnate to the pedicel for about half its length. Flowers 6.5–8 mm long, 1.8–3 mm in diameter, pedicellate, divaricate, actinomorphic; pedicels terete, 1–5 mm long, 0.5–1 mm in diameter; sepals triangular, acute, 0.8–1.3 × 0.8–1.5 mm, white lepidote abaxially, green with beige apex, entire, 1–3-nerved, much shorter than the petals; petals lanceolate, acute, 2.5–3.5 × 1.2–1.8 mm, glabrous on both surfaces, entire, white with light brown; staminodes six, triangular, laminar, 2.2–2.5 mm long, white; ovary superior, ovoid to ellipsoid, 3–4 mm long, 1.5–1.8 mm in diameter, white to yellowish tinged reddish, glabrous, stigmatic lobes recurved, nearly 1 mm long. Fruits ovoid to narrowly ovoid, 4–7 (–9) mm long, (2–) 3–5 mm in diameter, glabrous, erect, brown when mature; seeds fusiform, brown to reddish brown, reticulate, 5–6 mm long, 0.4–0.7 mm in diameter, with a lateral hyaline wing 1–2 mm long.

Additional specimens examined (paratypes):— MEXICO. Jalisco: Municipio Tolimán, arroyo La Ciénega , ca. 1 km al N de Tolimán, 19°36’33’’N, 103°54’33’’W, 783 m, January 13 2015, P. Carrillo-Reyes et al. 7627 J(IBUG!); P. Carrillo-Reyes et al. 7632, fruits (CICY!, IBUG!); arroyo La Ciénega , ca. 1 km al NE de Tolimán, 19°36’N, 103°54’W, 760 m, December 20 2005, P. Carrillo-Reyes & J. A. Lomelí-Sención 5109, leaves only (IBUG!); Las Canoas , 19°32’53’’N, 103°53’36’’W, 690 m, November 19 2018, K. Romero-Soler et al. 1243 fruits (CICY!); ca. de 1 km de Tolimán , a las orillas del río, 19°36’27.8’’N, 103°54’33.1’’W, 768 m, 19 November 2018, K. Romero-Soler et al. 1242 fruits (CICY!); Paso de Cedros , ca. 15 km al E de Tolimán, parte alta de la Barranca de Huisichi , 1820 m [820 m], 11 October 1997, J. A. Lomelí-Sención et al. 2811 fruits (GUADA-27156!, XAL!); Puerto de Toxín , 5–6 km al SW de San Pedro Toxín, 19°33’56’’N, 103°59’37’’W, 1200 m, 17 July 1994, F. Santana Michel & B. Benz 6716 fruits (ZEA!); La Taza , San Pedro Toxín , 500 m, 14 November 1992, G. Chávez & J. López s.n. fruits (GUADA-24111!); sitio de muestreo 2, proyecto Yesos del Occidente , Paraje La Sierrilla , 19°35’20’’N, 103°53’35’’W, 870 m, 18 June 2022, Brunel 1520 J (IBUG!); same locality, 21 August 2022, flowered in culture in September 2022, Brunel & P. Díaz 1702 ♀ (IBUG!); Rancho Araiza, El Pandito, Cerro El Yeso, 19°38’26.8’’N, 103°57’19.9’’W, 860 m, 5 March 2023, P. Díaz & Brunel 244, leaves only (IBUG!); Rancho Alegre, 3 km en línea recta al NE de El Huisichi, 19°34’36’’N, 103°50’52’’W, 960 m, 28 May 2021, P. Carrillo-Reyes & E. Acosta-Pérez 9808 fruits (IBUG!). Municipio Zapotitlán de Vadillo , 0.1 km al NO de Taberna de Tajipo (2.1 km al SE de Las Canoas), 19°31’30’’N, 103°53’11’’W, 670 m, 19 November 2018, K. Romero-Soler et al. 1245 fruits (CICY!); 0.5 km en línea recta al NNE de Paso de Alseseca, 19°29’32’’N, 103°50’19’’W, 750 m, 3 June 2021, P. Carrillo-Reyes et al. 9813 fruits (IBUG!) .

Phenology:—Flowering plants have been collected in January, March, June, and September; fruiting specimens from October to January and from May to July, since the fruits have a longer duration on the plants. The fructification occurs immediately after blooming.

Distribution and habitat: — Hechtia carrilloi is known from 10 localities, eight of them in the municipality of Tolimán and two in nearby Zapotitlán de Vadillo ( Figure 1 ). This area corresponds to a very dry valley influenced by the rain shadow caused by the Sierra de Manantlán to the west, whereas it is bounded by the Nevado de Colima and Volcán de Fuego volcanic complex to the east. The valley is furrowed by deep ravines coming from the Nevado de Colima (INE, 2000). These ravines have eroded the deposits and uncovered the bedrock, which consists of volcanic sandstone, limestone, and gypsum outcrops ( Ortiz-Brunel et al., 2023) as well as soils derived from these rocks. Hechtia carrilloi habits these ravines and does not seem to show a clear preference for any of the three mentioned types of soil as it occurs in all of them, at 500–1200 m elevation.

The vegetation of all the localities correspond to tropical deciduous forest ( Figure 2A ) (sensu Rzedowski, 1978), where the new species grows along with Amphipterygium adstringens ( Schlechtendal, 1844: 636) Standley (1923: 673) , Bursera spp. , Fouquieria formosa Kunth (1823: 83) , Havardia acatlensis ( Bentham, 1875: 593) Britton & Rose (1928: 42) , Isolatocereus dumortieri ( Scheidweiler, 1837: 220) Backeberg (1942: 47) , Lysiloma sp. , Mimosa rosei Robinson (1898: 317) , Opuntia sp. , Pachycereus pecten-aboriginum Britton & Rose (1909: 422) , Plumeria rubra Linnaeus (1753: 209) , Prosopis laevigata (Humboldt & Bonpland ex Willdenow 1806: 1059) Johnston (1962: 78) , Sarcomphalus amole (Sessé & Mociño, 1888:38) Hauenschild (2016: 55) , and Stenocereus queretaroensis ( Weber 1891: 27) Buxbaum (1961: 101) . Hechtia carrilloi closely shares the habitat with other succulent plants, most of them with distribution restricted to the dry valleys and ravines in the surroundings of Tolimán, such as Acanthocereus paradoxus González-Zamora et al. (2020: 146) , Echeveria rulfiana Jimeno-Sevilla et al. (2015: 72) , Euphorbia diazlunana ( Lomelí-Sención & Sahagún-Godínez, 1993: 15) Steinmann (2003: 48) , Sedum chazaroi Carrillo & Lomelí (2008: 77) , Mammillaria beneckei Ehrenberg (1844: 833) , and M. scrippsiana ( Britton & Rose 1923: 84) Orcutt (1926: 8) .

Eponymy: —The specific epithet honors Pablo Carrillo-Reyes, a researcher at the Universidad de Guadalajara, in recognition of his extraordinary contribution to the study of the flora of Jalisco and his remarkable work in the training of the new generation of taxonomists in the country.

Common name: —“Gatitos” (Carrillo-Reyes & Acosta-Pérez 9808, IBUG).

IUCN Conservation assessment: —The estimated EOO is 134.17 km 2 and the AOO is 40 km 2, both within the threshold for “Endangered” status. Following the IUCN (2022) criteria B1 (extent of occurrence) and B2 (area of occupancy), Hechtia carrilloi should be considered as Endangered (EN).

It is very possible that the species occurs in many more ravines and lowlands of the region of the Zapotitlán- Tolimán Valley. However, several areas of the region have been widely disturbed due to intensive cattle ranching and the cultivation of grapes and Agave spp . Derived from the AOO and EOO results and our observations, we suggest a preliminary risk category of EN.

Discussion: — Hechtia carrilloi has been collected in the biogeographical region of the Pacific Lowlands (sensu Morrone et al., 2017), a narrow strip that extends from Mexico (Sinaloa) to Costa Rica. This region is one of the thirteen biogeographical provinces within the Megamexico III region (sensu Rzedowski, 1991) and houses the third largest number of Hechtia species (24) after the Sierra Madre del Sur (25) and the Balsas Basin (40) ( Rivera-Martínez et al., 2022).

Hechtia carrilloi is characterized by its relatively small rosette size not exceeding 30 cm in diameter and 12 cm in height ( Figure 2B ), with succulent, rigid, light green leaves sparsely covered by trichomes ( Figures 2D, E ). Rosettes form colonies of 8–12 individuals, propagating by means of stolons ( Figure 2C ), but sometimes rosettes produce offshoots or pups at the base of the old inflorescence, giving the appearance of having lateral inflorescences, which is not the case. Inflorescences emerge from the center of a mature rosette and have a strict sympodial growth pattern (SSP sensu Ramírez et al., 2014; Figure 3F ).

Several remarkable morphological characters distinguish H.carrilloi from H. santanae (see Table 1 for comparison, and figures 2B, D, E). One of the main differences is the smaller overall size of the new species in terms of rosette, inflorescence, and flower compared to H. santanae . Furthermore, the stipe of the branches of H. santanae are covered by bracts giving it a scaly appearance, while in H. carrilloi the stipes are ebracteate. The third most important difference among these species is the male flower size and shape. In H. santanae , the shape of corolla is obconical and larger (5.0– 5.5 mm long, 3.8–4.2 mm in diameter), while in H. carrilloi it is tubular and smaller (4–5 mm long, ca. 2 mm in diameter) ( Figures 3A–B ). Finally, the color of the anthers also differs between the two species, they are reddish in H. santanae , while those of H. carrilloi are maroon.

Figures

FIGURE 1. Geographical distribution of Hechtia carrilloi (black triangles), and H. santanae (white dots).

FIGURE 2. A. Habitat of Hechtia carrilloi in Municipality of Tolimán, Arroyo La Ciénega. B. Rosettes of H. carrilloi (left) and H. santanae (right). C. Production of new rosettes by stolons in H. carrilloi. D. Adaxial (two upper ones) and abaxial (two lower ones) foliar surfaces; largest leaf H. santanae, shorter H. carrilloi. E. Leaf cross section of H. santanae (above) and H. carrilloi (below). (Credits: A. Katya Romero-Soler. B. Ivón Ramírez-Morillo. C. Pablo Carrillo-Reyes. D, E. Claudia Ramírez-Díaz).

FIGURE 3. Hechtia carrilloi. A. Staminate inflorescence. B. Staminate flowers. C. Pistillate flowers. D. Line drawing of staminate (above) and pistillate (below) flowers. E. Fruits. F. Rosette showing a central infructescence with strict sympodial growth pattern.(Credits: A, B. Ivón Ramírez-Morillo. C. Juan Pablo Ortiz-Brunel. D. Alberto Guerra. E, F. Katya Romero-Soler).

FIGURE 4. Illustration of Hechtia carrilloi. A. Branch with pistillate flowers. B. Female plant with inflorescence. C. Male plant with inflorescence. D. Branch with staminate flowers. D. Detail of staminate flowers. E. Branch with immature fruits. Illustration by Alberto Guerra.