Hechtia deceptrix I. Ramírez & Hornung, 2015

Ramírez-Morillo, Ivón, Hornung-Leoni, Claudia T., González-Ledesma, Manuel & Treviño-Carreón, Jacinto, 2015, A new species of Hechtia (Bromeliaceae: Hechtioideae) from Hidalgo (Mexico), Phytotaxa 221 (2), pp. 157-165 : 158-163

Hechtia deceptrix I. Ramírez & Hornung , spec. nov. ( Figs. 1 , 2 , 3 , Table 1)

This new species shares with Hechtia epigyna the epigynous flowers but it differs in its larger overall vegetative size, in its strict sympodium growth pattern with terminal inflorescence (vs. pseudomonopodial with lateral inflorescence), glabrous adaxial foliar surface (vs. white lepidote), green petals (vs. pink) in fresh flowers, and erect fruits (vs. pendulous).

TYPE:— MEXICO. Hidalgo: Municipio de Actopan, Puente de Dios, alrededores de Puente de Dios, 20°18’08’’N, 98°47’23’’W, matorral con Juniperus , Dasylirion y Brahea , sobre sustrato calizo, 1735 m elevation, 15 May 2008, López-Ferrari, Espejo & Zamudio 3309 ♂ (holotype UAMIZ!; isotype IEB!).

Plants lithophytic, rosettes caespitose, in general shape globose, 40–60 cm tall, 50–60 cm diameter, generally forming dense, small colonies of 2–3 rosettes, rarely clumps of 8–12 rosettes. Rhizomes large, 16–36 cm long, 22–34 mm diameter. Leaves 50–80 in number, flexible, central ones erect, basal ones slightly reflexed; sheaths broadly ovate, 4–6 × 4–5 cm, light brown, margins erose, densely white lepidote abaxially, lustrous and glabrous adaxially; blades narrowly triangular, attenuate, 26–40(–58) × 1–3(–4) cm, succulent, barely channeled in cross section, green, sometimes with purple spots at the apex or margins, densely white lepidote abaxially, white lepidote at base but glabrous and glossy adaxially, margin straight, armed; spines antrorse, triangular, 1.5–2.5 mm long, 4–10 mm apart, light green or occasionally purple, with a short tuff of white trichomes at the axile of basal spines. Inflorescence central, erect, emerging from a mature rosette (strict sympodium growth pattern, type SPP sensu Ramírez et al., 2014).

Staminate inflorescences usually a 1-divided panicle, sometimes the basal 2–4 branches with 1–2 secondary branches (then 2-divided), cylindrical, erect, 1.8–2.5 m long; peduncle terete, purple, pruinose, 0.8–1.2 m long, 10–16(–25) mm diameter at the base, much surpassing the rosette; internodes 4.1–8 cm long; peduncle bracts with triangular sheaths, 2.5–4 × 3–4 cm, light brown drying almost white, margins entire to erose, thin; blades triangular, long attenuate, slightly pungent, foliaceous, multinerved, 4–6 × 0.6–1.2 cm, spinose, green with purple spots, densely white lepidote abaxially, glabrate adaxially, exceeding internodes; main axis 1.8–2.2 m long, ca. 1 cm diameter, upwards to ca. 5 mm, terete, purple, internodes 3.5–6.5 cm long; primary bracts ovate-triangular, attenuate, 2.6–5.5 × 7–10 cm, spinose-dentate, green with purple spots, sparsely white-lepidote on both surfaces, multinerved, shorter than branches; branches ca. 30 in number, in an angle of 45° or less with the main axis, 4–17 cm long, 1.5–2 cm diameter, 28–38 flowers; rachis ca. 2 mm diameter, flattened at its base, drying light brown, glabrous, sterile basal portion 2.7–3 cm long, upward decreasing in length to be completely sessile; floral bracts ovate-elliptical, acute, 6–7 × 2.6–2.9 mm, membranous, green, brownish at the apex, margins erose and hyaline, glabrous, 1-nerved, falsely appearing, due to levels of insertion along the floral axis, to be shorter than sepals at anthesis. Flowers pedicellate, poystichous, divaricate, actinomorphic, erect; pedicels obconic, 4–5 mm long, ca. 1.5 mm diameter, slightly lepidote; sepals triangular, green, apically brownish, 3.5–3.8 × 1.8–2 mm, entire, glabrous, 3-nerved, shorter than petals; petals wide-elliptical, rounded, 4.2–4.5 × 3.3–3.5 mm, spreading in their apical half, light green, veins not visible; filaments triangular, flattened, 1.9–3.5 mm long, white; anthers oblong, ca. 0.9–1.5 mm long, dorsifixed, green, pollen yellow; pistillode reduced, greenish-white, stigmatic lobes much reduced.

Pistillate inflorescences a 1-divided panicle, sometimes 1–2 of them with 1 secondary branch (then 2-divided), cylindrical, erect, 1.7–2 m long; peduncle terete, 85–100 cm long, 1.6–2 cm diameter at the base, much higher than rosette and as long as the main axis, purple, pruinose, internodes 5–7.5 cm long; peduncle bracts with triangular sheath, 2.5–4 × 3–4 cm, green with purple spots, drying light brown, margins entire to erose, thin; blade triangular, long attenuate, slightly pungent, foliaceous, multinerved, 7–9 × 0.6–1 cm, spinose, green with purple spots, densely white lepidote abaxially, glabrate adaxially, longer than internodes; main axis 85–100 cm long, ca. 1 cm diameter at the base, terete, purple, internodes 5–6.5 cm long; primary bracts narrowly ovate - triangular, attenuate, (1–)2.6–5.5 × 0.5– 1 cm, as long or shorter than the sterile portion of the branch, spinose-dentate, green with purple spots, apex brownish, margin erose, hyaline; branches 7–16 in number, stipitate, forming an angle of ca. 45° with the main axis, 5–14 cm long, (2–) 4–5 cm diameter cylindrical, with 18–40 flowers; basal sterile portion ca. 1.5 cm long decreasing in length to be completely lacking at distal end of inflorescence, dorsiventrally flattened; rachis ribbed, glabrous, green with purple hues; floral bracts ovate-elliptical, acute, ca. 8 × 4–5 mm, margin erose, hyaline, green, as long as the ovary. Flowers almost sessile, erect; pedicel ribbed, less than 1 mm long, ca. 0.5 mm diameter; sepals triangular, acute, 4–5 × 2–2.5 mm, green, entire, glabrous, 1-nerved; petals elliptic, rounded, cucullate at apex, 4–5.2 × 2–2.8 mm, entire, green; staminodes six, triangular, laminar, 1.6–1.9 mm long, white; ovary inferior, oblongoid to ellipsoid, 5–7 mm long, 2–3 mm diameter, green, slightly lepidote, stigmatic lobes recurved, 2–3.5 mm long, adnate at their bases, white, falsely appearing, due to level of insertion along the floral axis, to be as long as the petals at anthesis; placentation central, ovules white-greenish. Fruits ellipsoid, sessile, 10–14 mm long, 4–4.5 mm diameter, glabrous, erect, and brown when mature; seeds fusiform, brown to reddish brown, reticulate, ca. 3 mm long, 1 mm diameter, with a lateral wing ending in two caudae, these hyaline.

Habitat & Distribution:— Hechtia deceptrix occurs in Rio Amajac basin in the Municipality of Atotonilco El Grande, limiting with the municipalities of Actopan and Cardonal, in the central portion of Hidalgo. Plants of the new species form small (2–3 rosettes) to rarely large (8–12 rosettes, Figure 2A ) colonies on limestone cliffs at 1700–1800 m of elevation. These localities are part of the strip of submontane scrub that extends from Jacala and Pacula in the northwestern portion of Hidalgo, to the southeast into the region of Santa María Amajac, in the physiographic subprovince of Carso Huasteco in the Sierra Madre Oriental Province ( Anonymous, 1992). Locally, H. deceptrix is associated with other rosetofilous plants such as Agave celsii Hooker (1856 : t4934), Agave striata Zuccarini (1833: 678) , A. xylonacantha Salm-Reifferscheid-Dyck (1859: 92), Dasylirion longissimum Lemaire (1856: 91) , H. glomerata , Tillandsia albida Mez & Purpus , in Mez (1916: 248), and T. grandis Schlechtendal (1845: 424) , among other typical elements of these particular dry forests. The population at Puente de Dios is located at the limit of the Faja Volcánica Transmexicana Province in a transition to a forest of Juniperus flaccida Schlechtendal (1838: 495) , together with Brahea berlandieri Bartlett (1935: 31) .

In contrast, the only known population of Hechtia epigyna is located in the subprovince of the Gran Sierra Plegada in the state of Tamaulipas in the Sierra Madre Oriental Province, at lower elevation (585–612 m) in submontane scrub vegetation with a different floristic composition ( Puig, 1991). At the type locality, H. epigyna grows on vertical karstic walls with northern exposure, forming dense colonies of several rosettes (4–12) along with Brahea berlandieri , Stenocereus griseus ( Haworth 1812: 182) Buxbaum (1961: 100) , Myrtillocactus geometrizans Martius , in Pfeiffer (1837: 90) Console (1897: 10), and more frequently on less steep stone walls together with Pilosocereus chrysacanthus ( Weber 1897: 178) Byles & Rowley (1957: 66) , Agave lophantha Kunth (1850: 838) , and Dioon edule Lindley (1843: 59) , as well as spikemosses and Mexican snow balls ( Selaginellaceae ).

Etymology: —The specific Latin epithet, deceptrix means deceiver, after the fact that the new species was confused with Hechtia epigyna when first collected by A. Espejo and collaborators.

Additional specimens examined (paratypes):— MEXICO. Hidalgo: Municipio Actopan, Puente de Dios , 20°18’13’’ N, 98°47’20’’ W, 1740 m, 25 August 2007, Zamudio et al. 13866, fruits ( IEB, UAMIZ!) ; alrededores de Puente de Dios , municipio de Actopan, 20°18’08’’ N, 98°47’23’’ W, 1735 m, 15 May 2008, López-Ferrari et al. 3311 ♂ ( IEB, UAMIZ!) ; Municipio de Cardonal , barranca de Tolantongo, 20°37’52’’ N, 98°59’30’’ W, 1739 m, 16 May 2008, Espejo et al. 7150 ♀ ( IEB, UAMIZ!) ; 7151 ♂ ( IEB, UAMIZ) ; ejido San Cristóbal , 20º38’20’’ N, 98º59’30’’ W, 1800 m, 18 March 2008, Zamudio & Zamudio 14085 ♀ ( IEB, UAMIZ!) ; Municipio de Atotonilco el Grande, Puente de Dios , 20°18’08’’ N, 98°47’22’’ W, 1700 m, 02 April 2012, Hornung-Leoni et al. 1344 ♀ ( HGOM!) ; Municipio Cardonal , barranca de Tolantongo, 20°38’8’’ N, 98°59’27’’ W, 1791 m, 03 May 2012, Hornung-Leoni et al. 1354 ♂ ( HGOM!) .

Discussion: —We located two collections of Hechtia epigyna : the type, collected in Jaumave, Tamaulipas, and one deposited at GH, H. W. von Rozynski 741 (leaf and staminate inflorescence), collected in “ Tamaulipas, Jaumave near Nogales, II. 1933 ”.

The locality provided by von Rozynski 741 took us to Nogales in the Municipality of Jaumave, at the edge of “Reserva de la Biosfera El Cielo” in Tamaulipas. Our search was fruitful and we found large populations of Hechtia epigyna : round rosettes with old, lateral staminate inflorescences and infructescences.

The first author also had the opportunity to observe plants named Hechtia epigyna in San Diego, California, in March 2014, long in cultivation, one of which at the time was in bloom bearing a lateral inflorescence with pink petals ( Figure 3D ) and presenting other features (vegetative and floral ones) that match well the protologue of H. epigyna ; the available locality data for these cultivated plants indicated that they originally came from somewhere in Tamaulipas, Mexico. Overall, all the evidence indicates that H. epigyna is confined to Tamaulipas, and that both populations from Hidalgo represent the new species described in this article.

Hechtia deceptrix differs from H. epigyna in several important characters ( Table 1, Figure 3 ): the most important one is that the former has central or terminal inflorescences or strict sympodium pattern (SPP) growth as defined by Ramírez et al. (2014) while H. epigyna has lateral inflorescences and pseudomonopodial growth pattern (SMP, Figure 3C ). In this sense, H. epigyna forms part of the H. glomerata complex as defined by Jiménez (2011), whose members are distributed in several biotic provinces sensu Morrone (2014), namely: Mosquito ( Honduras), Península de Yucatán, Tierras Altas de Chiapas, Sierra Madre Oriental, Altiplano Mexicano, Provincia Veracruzana, and Provincia de Tamaulipas, all draining chiefly into the Atlantic watershed of Megamexico. All taxa in the H. glomerata complex share a pseudomonopodial growth pattern. In addition, H. deceptrix has greenish-white petals in flowers of both sexes at anthesis and erect fruits, whereas H. epigyna has flowers with pink petals in both sexes and pendent fruits ( Figure 3D, F ).

Epigyny, shared by Hechtia deceptrix and H. epigyna , probably evolved twice in the genus as suggested by preliminary results of a phylogenetic study based on cpDNA and nuclear DNA (I. Ramírez, in prep.), where the H. glomerata complex sensu Jiménez (2011) is monophyletic.

IUCN Conservation assessment:—The conservation status of Hechtia deceptrix was assessed using the IUCN Red List Criteria ( IUCN 2010). Because of the lack of hard population information, we relied mostly upon distributional data, namely, the set of B criteria, geographical distribution assessed both as B1 (extent of occurrence, EOO) or B2 (area of occupancy, AOO).To estimate distributional ranges, both extent of occurrence and area of occupancy, we used the GeoCat software (GeoCat 2014; Bachman et al. 2011). Using both B criteria the species ranks as CR (Critically Endangered), with an EOO of 20.127 km 2 and AOO of 8.000 km 2. The species is known from five collection points all near roadsides and most likely occurs as isolated populations on appropriate microniches in Hidalgo in the Sierra Madre Oriental Province and neighboring areas of the Faja Volcánica Transmexicana Province.

Figures

FIGURE 1. Hechtia deceptrix. Staminate plant (A, C). A. Plant in bloom. C. Flower at anthesis. Pistillate plant (B, D, E, F). B. Distal portion of the inflorescence. D. Flowers at anthesis. E. Flowers showing details of ovary surface and floral bract. F. Longitudinal section of the ovary showing placentation and ovules. (Based on Hornung et al. 1344♀, 1354♂, HGOM). Illustration by Jorge A. González Martínez.

FIGURE 2. Hechtia deceptrix. A. Plants in habitat. B. Comparative size of a staminate inflorescence and rosette. C. Central origin of the inflorescence according to the strict sympodium growth pattern. D. Details of staminate flowers at anthesis. E. Staminate inflorescence showing position of branches and flowers arrangement. F. Pistillate flowers at anthesis. (Photographs by Claudia T. Hornung-Leoni).

FIGURE 3. Hechtia deceptrix (A, E) A. Rosette showing the glabrous abaxial area and dark green color of leaves. E. Erect fruits (Based on Zamudio et al. 13866, UAMIZ). Hechtia epigyna (B, C, D, F). B. Rosette showing densely white lepidote adaxial surface and light green color of leaves. C. Lateral origin of inflorescence according to the pseudomonopodial growth pattern.D. Staminate flowers showing pink petals. F. Pendent fruits (Based on Ramírez et al. 1956, CICY). (Photographs: A, B, F by Ivon Ramírez-Morillo; C by Jacinto Treviño-Carreón; D by Michael Wisnev; E by Carlos Jiménez).