A new species of Hechtia (Bromeliaceae: Hechtioideae) from Hidalgo (Mexico)

Abstract

We propose that two populations previously referred to Hechtia epigyna, from the Mexican state of Hidalgo, represent a new species. Plants from the Hidalgo populations share the inferior ovary with Hechtia epigyna, an unusual trait in the genus, but they differ in their growth pattern (central vs. lateral inflorescence), characters of the adaxial foliar surface, petal color, and fruit position during dehiscence. We also provide a clarification on the typification of Hechtia epigyna. An assessment of the conservation status of the new species, Hechtia deceptrix following IUCN criteria resulted as CR (Critically Endangered).

Full text

Phytotaxa 221 (2): 157–165
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Copyright © 2015 Magnolia Press
Article
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Eric Gouda: 8 Jul. 2015; published: 30 Jul. 2015
http://dx.doi.org/10.11646/phytotaxa.221.2.5
157
A new species of Hechtia (Bromeliaceae: Hechtioideae) from Hidalgo (Mexico)
IVÓN RAMÍREZ-MORILLO
1,4
, CLAUDIA T. HORNUNG-LEONI
2
, MANUEL GONZÁLEZ-LEDESMA
2
&
JACINTO TREVIÑO-CARREÓN
3
1
Centro de Investigación Científica de Yucatán, A. C., Unidad de Recursos Naturales-Herbario CICY, Calle 43 # 130. Colonia Chu-
burná de Hidalgo, Mérida, Yucatán 97200, México.
2
Universidad Autónoma del Estado de Hidalgo, Centro de Investigaciones Biológicas, Instituto de Ciencias Básicas e Ingeniería, Km
4.5 Carretera Pachuca-Tulancingo, Mineral de La Reforma, Hidalgo 42184, México.
3
Universidad Autónoma de Tamaulipas, Facultad de Ingeniería y Ciencias, Centro Universitario Adolfo López Mateos, Cd. Victoria,
Tamaulipas 87149, México.
4
E.-mail ramirez@cicy.mx (Author for correspondence)
Abstract
We propose that two populations previously referred to Hechtia epigyna, from the Mexican state of Hidalgo, represent a new
species. Plants from the Hidalgo populations share the inferior ovary with Hechtia epigyna, an unusual trait in the genus, but
they differ in their growth pattern (central vs. lateral inflorescence), characters of the adaxial foliar surface, petal color, and
fruit position during dehiscence. We also provide a clarification on the typification of Hechtia epigyna. An assessment of the
conservation status of the new species, Hechtia deceptrix following IUCN criteria resulted as CR (Critically Endangered).
Key words: Epigynous flowers, growth pattern, Hechtia epigyna, Hidalgo, IUCN, Tamaulipas
Introduction
Hechtia Klotzsch (1835: 401) is represented in the Mexican State of Hidalgo by three species (Espejo et al. 2004),
namely H. glomerata Zuccarini (1840: 240), H. lundelliorum Smith (1938: 97), and H. podantha Mez (1896: 549). A
fourth species, H. lepidophylla Ramírez (2008: 65) also has been reported for Hidalgo. Hornung-Leoni & Ramírez-
Morillo (in prep.) have records of three more species in this state and overall, here we report a total of eight Hechtia
species for Hidalgo, including the new one described herein. Plants of some of these species form large colonies: H.
podantha does around the capital city, Pachuca, and H. glomerata in areas such as the Biosphere Reserve Barranca de
Metztitlán in the central-north portion of the state of Hidalgo.
Materials and methods
Field work was carried out in the state of Hidalgo, Mexico, where we collected plants of the new species from two
populations identified as Hechtia epigyna Harms (1935: 532) by Espejo et al. (2010). We found pistillate plants in
bloom at the Municipality of Atotonilco El Grande and staminate plants in bloom at Municipality of Cardonal; fruit
features were studied on specimens deposited at Herbarium UAMIZ (López-Ferrari et al. 3311; Zamudio et al. 13866;
Zamudio & Zamudio 14085). We based the description of the new entity on Espejo et al. (2010), and on our collections
deposited at HGOM (Hornung et al. 1344, 1354). In order to circumscribe the new species, we carried out field work
in Tamaulipas, Municipality of Jaumave, at Nogales at the type locality of H. epigyna where we collected plants in fruit
and with old staminate inflorescences and live plants of both sexes to cultivate and obtain fresh flowering material;
vouchers of these collections are deposited at Herbarium CICY (Ramírez et al. 1721, 1723). We also studied high
resolution images of the two sheets labeled H. W. Viereck 81 collected in Jaumave, Tamaulipas on March 2, 1930,
both deposited at Herbarium B (Röpert (ed.) 2000+ [continuously updated]), and examined two collections of the
same species at GH. Members of the San Diego Bromeliad Society kindly provided images and flowers in spirit of H.
epigyna. We measured dimensions and compare qualitative features across all the available specimens.

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158 • Phytotaxa 221 (2) © 2015 Magnolia Press
Hechtia epigyna was described in 1935 by Hermann August Theodor Harms, a German botanist associated with
the Botanical Museum in Berlin. The original description includes characters of both staminate and pistillate flowers
based on H. W. Viereck 81 (B!) from Jaumave, Tamaulipas, which consists of two sheets numbered I and II (Röpert
(ed.) 2000+ [continuously updated]). The first sheet (B1003990477, marked I) has two mounted leaves and only
(three) staminate inflorescences whereas the other (B1003990476, marked II) has also two leaves but both a staminate
and a pistillate inflorescence. A fragment consisting of a few staminate and pistillate flowers was sent to Lyman
Smith by Harms (Smith, 1937) and deposited at Herbarium GH (GH00275616). Here we reject the non-published
lectotypification and isolectotypification by K. Burt-Utley as indicated on the sheets and agree with Espejo et al.
(2010), who indicated that the type of H. epigyna (H. W. Viereck 81) consists of two sheets.
Taxonomy
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.

A NEW SPECIES OF HECHTIA FROM MEXICO
Phytotaxa 221 (2) © 2015 Magnolia Press • 159
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.

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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”.

A NEW SPECIES OF HECHTIA FROM MEXICO
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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).
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.

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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 Ivón Ramírez-Morillo; C by Jacinto
Treviño-Carreón; D by Michael Wisnev; E by Carlos Jiménez).

A NEW SPECIES OF HECHTIA FROM MEXICO
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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.
TABLE 1. Comparative features of Hechtia deceptrix and H. epigyna.
Characters H. deceptrix H. epigyna
Growth pattern Strict sympodium with central (terminal; Figures
1A, 2C) inflorescence
Pseudomonopodial, with lateral
inflorescence (Figure 3C)
Leaf adaxial surface Glabrous (Figure 3A) Finely white lepidote (Figure 3B)
Petal color at anthesis; Greenish white to cream-white (Figures 2D–F) Pink to white or white pink (Figure 3D)
Fruit orientation during dehiscence Erect or ascending (Figure 3E) Pendulous (Figure 3F)
Locality and elevation Hidalgo, 1700–1800 m Tamaulipas, 585–612 m
Physiographic subprovince on
the Sierra Madre Oriental Province
Carso Huasteco Gran Sierra Plegada
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.
Acknowledgments
We are indebted to Evelyn Rios and Laura Rodríguez for field assistance when collecting in Tamaulipas, and to Gil
Mendoza as well as Paula Moreno when collecting in Hidalgo. We thank the curators of the following herbaria B,
GH, IEB, MEXU, MICH, MO, OAX, UAMIZ, UC, US, WU, and XAL for sending Hechtia material on loan. The
first author is indebted to the Elizabeth Bascom Fellowship and the Missouri Botanical Garden, the DAAD-ANUIES,
and the Klarff foundation for financial support to study the Bromeliaceae collection in the herbaria B, BM, K, MO,
OXF, W, and WU. We are indebted to CONACyT for funding the project “Phylogeny, evolution and biogeography of
Hechtia Klotzsch (Hechtioideae: Bromeliaceae)” (number 183281) granted to the IMRM. We would also like to thank
PROMEP for the financial support to the project: “Distribución de las Bromeliaceae del Estado de Hidalgo” and to

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164 • Phytotaxa 221 (2) © 2015 Magnolia Press
FOMIX-CONACYT for funding the project “Diversidad del Estado de Hidalgo” second phase (95828) and third phase
(191908) both granted to the CTHL. We also acknowledge the Universidad Autónoma de Tamaulipas for the financial
support to the project: “Diversidad del género Hechtia (Bromeliaceae) en el Altiplano de Tamaulipas y Regiones
Adyacentes” (PFI2014-16), granted to JTC. We also thank Silvia Hernández-Aguilar for handling the herbarium
material and loans, José Luis Tapia Muñoz for compiling and maintaining a database of Mexican Bromeliaceae, and
to Gustavo A. Romero-González (AMES), who supplied literature on the genus, facilitated the study of herbarium
material at the Harvard University Herbaria, and reviewed the English version of this paper. Germán Carnevali (CICY)
reviewed a late draft of the manuscript. Robert Kopfstein, Andy Siekkenen, Dan Kinnard, Eloise Lau, and Pamela
Koide-Hyatt, members of the San Diego Bromeliad and Saddleback Valley Bromeliad Societies, kindly showed the
first author the true identity of Hechtia epigyna. We are grateful to Michael Wisnev for sending pictures of Hechtia
epigyna in bloom, cultivated at the Huntington Botanical Garden, to Carlos Jiménez Nah for the photographs of the
specimens of the new species deposited at Herbarium UAMIZ, to Jorge A. González Martínez for the illustration of
the new species, and, finally, to Eric Gouda for editorial comments and to two anonymous reviewers for observations
and suggestions that greatly improved the text.
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