In disentangling two species limits of Hechtia (Bromeliaceae: Hechtioideae) from Sierra Madre Occidental, a new species is discovered from Durango, Mexico

Abstract

Epitypes are proposed for two species of Hechtia: H. subalata and H. jaliscana, both endemic to Western Mexico. The holotypes of both species consist of fruiting specimens. While the female flowers of both species are extremely similar, the staminate ones are noticeable different: we select staminate vouchers as epitypes to clearly circumscribe both taxa. As a result of the delimitation of the concept Hechtia subalata and its geographical distribution, we identify a new species from Durango, Mexico, previously misidentified as Hechtia subalata: Hechtia marthae. We provide images portraying plant parts of all species, as well as photographs in habitat. Finally, assessments of the conservation status of the three species sensu IUCN criteria are included.

Full text

Phytotaxa 523 (1): 055–072
https://www.mapress.com/j/pt/
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Eric Gouda: 28 Sept. 2021; published: 15 Oct. 2021
https://doi.org/10.11646/phytotaxa.523.1.3
55
In disentangling two species limits of Hechtia (Bromeliaceae: Hechtioideae) from
Sierra Madre Occidental, a new species is discovered from Durango, Mexico
IVÓN RAMÍREZ-MORILLO1,3,*, PABLO CARRILLO-REYES2,4, JOSÉ LUIS TAPIA-MUÑOZ1,5 & CLAUDIA J.
RAMÍREZ-DÍAZ1,6
1Centro de Investigación Científica de Yucatán, A. C., Unidad de Recursos Naturales-Herbario CICY, Calle 43 # 130 x 32 y 34. Colonia
Chuburná de Hidalgo, Mérida, Yucatán 97205, México.
2Universidad de Guadalajara, Departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias,
Camino Ramón Padilla Sánchez 2100, km. 15.5 carretera a Nogales, Predio Las Agujas, Zapopan, Jalisco 45110, México.
3
�
ramirez@cicy.mx; https://orcid.org/0000-0002-6288-7984
4
�
pablo.creyes@academicos.udg.mx; https://orcid.org/0000-0001-9278-0208
5
�
jltapia@cicy.mx; https://orcid.org/0000-0002-3281-9032
6
�
claujrd11@gmail.com; https://orcid.org/0000-0002-3593-4407
* Author for correspondence: Ivón Ramírez-Morillo, ramirez@cicy.mx
Abstract
Epitypes are proposed for two species of Hechtia: H. subalata and H. jaliscana, both endemic to Western Mexico. The
holotypes of both species consist of fruiting specimens. While the female flowers of both species are extremely similar, the
staminate ones are noticeable different: we select staminate vouchers as epitypes to clearly circumscribe both taxa. As a
result of the delimitation of the concept Hechtia subalata and its geographical distribution, we identify a new species from
Durango, Mexico, previously misidentified as Hechtia subalata: Hechtia marthae. We provide images portraying plant parts
of all species, as well as photographs in habitat. Finally, assessments of the conservation status of the three species sensu
IUCN criteria are included.
Keywords: Durango, epitype, Flora Novo-Galiciana, IUCN, Jalisco, Nayarit, Zacatecas
Resumen
Se proponen epitipos para dos especies de Hechtia: H. subalata y H. jaliscana, ambas endémicas del oeste de México. Los
holotipos de ambas especies consisten en ejemplares con frutos. Mientras que las flores pistiladas de ambas especies son
extremadamente similares, las flores estaminadas difieren considerablemente, por lo que se seleccionan ejemplares estami-
nados como epitipos que permiten circunscribir ambos taxones. Como resultado de la delimitación del concepto Hechtia
subalata y de su distribución geográfica, identificamos una nueva especie de Durango, México, previamente identificada
erróneamente como Hechtia subalata: Hechtia marthae. Proporcionamos imágenes de las especies, mostrando caracteres
relevantes y su hábitat. Finalmente, se incluyen evaluaciones del estado de conservación de las tres especies según criterios
de la IUCN.
Palabras clave: Durango, epitipo, Flora Novo-Galiciana, IUCN, Jalisco, Nayarit, Zacatecas
Introduction
Hechtioideae Givnish (2007: 23), which currently includes three genera, Bakerantha L.B. Smith (1934: 72),
Mesoamerantha I. Ramírez & K. Romero (2018b: 308), and Hechtia Klotzsch (1835: 401) (Ramírez-Morillo et al.
2018b), have at least 84 species (Gouda et al. cont. updated), a figure that has notably increased in the last decade,
with a description rate of 2–5 species a year (for example, just in 2020, five species were described, i.e.; H. elegans
Siekkinen et al. (2020: 466), H. ensifolia Hern.-Cárdenas et al. (2020: 469), H. medusae Hern.-Cárdenas et al. (2020:
469), H. platyphylla Hern.-Cárdenas et al. (2020: 471), and H. pycnostachya Hern.-Cárdenas et al. (2020: 473), the
result of new explorations in Mexico and a meticulous taxonomic work in the subfamily by several plant biologists.

RAMÍREZ-MORILLO ET AL.
56 • Phytotaxa 523 (1) © 2021 Magnolia Press
Identification of species in Hechtioideae, a group of herbaceous, rosetophilous, dioecious species, is not an easy task
due to: (a) sexual dimorphism, where male and female structures (inflorescences and flowers, not the rosette) differ
in many characteristics, although closely related species show no differences in flower features on one sex but differ
when flowers of the opposite sex are compared; (b) scantiness of information provided by type specimens and/or
original descriptions, which hinder the correct application of names; and (c) inaccurate provenance data (some referred
as “Mexico” only), which also makes identifications difficult. Thus, to understand species limits and correctly apply
names, information on all plant parts and geographical distribution is needed. Here is where epitypification becomes
useful, resolving the problems posed by the scantiness of type specimen and incomplete descriptions and avoiding the
proliferation of names by proposing new species and/or declaring names invalid and/or confusing.
Hechtia subalata was described based on two herbarium specimens: the holotype, which bears the apical part of
an infructescence with unopened fruits (J. N. Rose 3467, US!; photograph and fruits, GH!), collected in Durango in
Sierra Madre region, a vague locality since Sierra Madre could be referred to Sierra Madre Oriental that occupies ca.
5.22% of the state of Durango, to the Northeast, or Sierra Madre Occidental that occupies ca. 71.16 % of the state to
the West; INEGI (2001). Rose’s fieldnotes do not cite a specific locality in the state (https://www.biodiversitylibrary.
org/item/175144#page/163/mode/1up). Smith also cited a paratype (J. N. Rose 3556, GH [sub G]!) consisting of a
leaf (without sheath) and fruit fragments (in an envelope), collected in the Mexican State of Zacatecas, in San Juan
Capistrano, Sierra Madre Occidental area. A duplicate specimen of the paratype at NY, not cited in the protologue,
shows leaf features (abaxial and adaxial surface) and two inflorescence branches with opened fruits. Subsequent
publications (Smith 1938; Smith & Downs 1974) did not add more information on this species. McVaugh (1989)
added data on floral features, particularly those of male flowers, habitat, altitude, flowering, and fruiting periods, and
indicated the geographical distribution of the species from the Mexican States of Durango and Zacatecas to Jalisco and
Nayarit. Furthermore, the same author cited the type locality (originally as Durango, Sierra Madre) as “Huasemote”
[most likely Huazamota, Mezquital Municipality]. Recent reports of H. subalata by Villaseñor (2016) for the Mexican
States of Chihuahua and Sinaloa have to be taken with caution since no vouchers were provided in this publication and
this geographical extension of the species range is highly improbable since most species in Hechtioideae are narrowly
endemic, probably related to low seed vagility and/or high substrate affinity. Our observations, based on revision
of herbarium specimens and field work, are in agreement with McVaugh (1989) in that H. subalata is found in four
Mexican States: Durango (based on holotype and McVaugh, 1989), Jalisco, Nayarit, and Zacatecas. The northernmost
populations from Durango (in Nombre de Dios, Topia and northern portion of Mezquital Municipalities) previously
identified as H. subalata, differ from this species, among other features, by the presence of large primary bracts (on
both sexes specimens) and condensed pistillate branches, which is here assigned to a new taxon, Hechtia marthae,
described herein.
The obvious and pertinent question at this point is: if McVaugh completed the morphological characteristics
missing in the protologue and added geographical provenance to the type specimen of Hechtia subalata as he
understood it, what is the purpose of designating an epitype? Article 9.9 of the International Code of Nomenclature
for algae, fungi, and plants (ICBN [Shenzhen Code]; Turland et al. 2018) indicates that “an epitype is a specimen or
illustration selected to serve as an interpretative type when the holotype, lectotype, or previously designated neotype,
or all original material associated with a validly published name, is demonstrably ambiguous and cannot be critically
identified for purposes of the precise application of the name to a taxon”. When having an incomplete type, the
determination of a specimen will always be ambiguous, since many other vegetative and reproductive characters are
missing from the type specimen and not provided in the protologue, often leading to misidentifications, especially in
dioecious, sexual dimorphic species. Epitypes became in use in 2005, 19 years after McVaugh’s publication, a tool that
makes species delimitation more precise and avoids misidentifications.
The second species addressed here is Hechtia jaliscana, described from a specimen bearing an infructescence
with mature fruits, showing its characteristic sinuose to fractiflex floral axis and channeled rachis of the branches (in
both staminate and pistillate inflorescences). It was collected in Jalisco, below Presa de Santa Rosa, barranca de Río
Grande de Santiago, north of Amatitán (McVaugh 18530, MICH!; photo US!). A detailed study of female flowers
shows that they are extremely similar to those of Hechtia subalata, but staminate inflorescence on both species differs
in several features. To provide all morphological and geographical information leading to an accurate identification of
Hechtia jaliscana, having either female or male material at hand, we propose an epitype portraying its male flowers.
In addition, we include complete morphological descriptions of the three species, citation of herbarium specimens, and
evaluations of their conservation status sensu IUCN (2019). Images of plants in habitat, illustrations of plant features,
and a distribution map are also included.

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 57
Material & methods
Herbarium revision: Vouchers from several herbaria, collected in the area northern of Mexican Volcanic Belt,
particularly in the Mexican States of Aguascalientes, Durango, Jalisco, and Zacatecas, were studied. Specimens
were all studied in the respective herbaria or from high-resolution images taking by us from the following herbaria
(acronyms according to Thiers, 2021): CICY, CIIDIR, GH, GUADA, HNT (only digital images), HUAA, HUH,
IBUG, IEB, MEXU, MICH, MO, NY, UAMIZ, UJAT, US, WU, and XAL. Material seen by authors are indicated
with the exclamation mark (!). Information on morphological features from specimens with flowers of both sexes and
fruits were recorded, as well as locality data to elaborate description and distribution maps, and to calculate the IUCN
conservation risk.
Field work: We planned different field trips to collect material of both sexes for all the species. In the case of
Hechtia subalata, we decided to go to the most precisely known locality included in the protologue: the surroundings
of San Juan Capistrano, Municipality of Valparaiso, Zacatecas, where paratypes came from. Plants were found in fruit
and with dry male inflorescences; rosettes were collected and after 2–3 years, they bloomed (both sexes) in cultivation.
We also examined the type specimen of H. subalata (holotype, J. N. Rose 3467, US!; and paratypes J. N. Rose 3556,
GH! NY!). In addition, information published by Smith (1937, 1938) and McVaugh (1989) was analyzed, except
Norris 13985 (MICH), a specimen collected in the Mexican State of Nayarit, that despite all effort, we could not
localize it. We also visited localities in the Mexican states of Durango, Jalisco, and Zacatecas.
In the case of Hechtia jaliscana, we studied the types and material deposited in different herbaria collected in
the Mexican States of Jalisco, and Zacatecas. Field work in the Mexican State of Jalisco was not an easy endeavor for
security reasons, nevertheless, we were able to collect material of the species in Jalisco (San Cristóbal de la Barranca
Municipality), where the species was identified in the field, based on features of the female infructescence; plants of
both sexes were cultivated to obtain fresh material of female and male flowers. These specimens were compared to
the type material (R. McVaugh 18530 fruits (holotype, MICH!; isotype, US!) and other specimens collected at the type
locality (L.M. Villarreal de Puga 3423♂ (GUADA!, IBUG).
Morphological descriptions are based on herbarium and living material for the three species included in this
article. Morphological descriptions are organized as follows: rosette, staminate inflorescence, pistillate inflorescence
and fruit features, as well as seed. Morphological terms are based on Scharf and Gouda (2008) and for growth patterns
Ramírez-Morillo et al. (2014).
In order to document the phenology of the species, when citing herbarium specimens, we follow this method
throughout the article: the symbol ♂ is used to indicate that the voucher has male flowers; and the symbol ♀ when the
specimen feature female flowers; finally, when the voucher is in fruit, we indicate it as such; here we omit the symbol
♀ which would be redundant for a dioecious species.
The conservation status of all three species was assessed using the IUCN Red List Criteria (B-set of criteria)
(IUCN 2019). Determination of the Extent of Occurrence (EOO) and Area of Occupancy (AOO) of the species were
calculated through the geospatial conservation assessment tool GeoCAT platform (Bachman et al. 2011). A distribution
map was produced in QGIS Desktop version 3.16.1 (QGIS 2019), and the limits of the biogeographical areas are based
on Morrone et al. (2017).
The assessment of the conservation status of the three species was based solely on geographical distribution
criteria (criteria B) since the evaluation of the population status is hard to assess in the field (criteria A), and there are
no quantitative analyses (criteria E) performed at this time. These evaluations would be the matter of separate projects,
perhaps to be conducted in the future by this research team or other colleagues. As many species in the genus, the
species treated here, have restricted geographical distributions and small-sized populations, most likely associated
with the low vagility of the seeds and/or high affinity for particular types of soils, factors that estimate an increase
in the vulnerability of the species of Hechtia. It must be remembered that conservation status assessments are just
hypotheses based upon data at hand and are likely to be revisited as more data becomes available. We think that in
these cases, it is much better to err on the safe side and assign the evaluated taxa the most threatened status recovered
from the analyses of whatever data we have, until abundant data convincingly demonstrate that the species is less
threatened. In the meanwhile, in lieu of better data and in the best scenario, a threatened status would probably secure
the conservation efforts required to preserve the known populations of the species.

RAMÍREZ-MORILLO ET AL.
58 • Phytotaxa 523 (1) © 2021 Magnolia Press
Taxonomy
Hechtia jaliscana L.B. Sm., Phytologia 10(6): 482, t. 1, f. 10. 1964.
TYPE:—MEXICO: Jalisco: [Amatitán Municipality], below presa de Santa Rosa, in the barranca of the río Grande, north of Amatitán,
750–800 m, Sep 1, 1960, McVaugh 18530 fruits (holotype, MICH!; isotype, US!)).
EPITYPE (here designated):—MEXICO: Jalisco: Municipio San Cristóbal de la Barranca, ca. rancho Jacalitos, a 1 km al SE de Santa
Cruz de Atístique, bosque tropical caducifolio con Pilosocereus, Stenocereus queretaroensis, Ceiba sp., Bursera sp., Pithecellobium
dulce, 21°06’37.9’’N, 103°26´34.1’’W, 920 m, originally collected by I. Ramírez, P. Carrillo-Reyes, W. Cetzal & J.L. Tapia, May
2015; flowering in Mérida, Yucatán, May 2018, I. Ramírez 2432♂ (CICY!).
Plants lithophytic, 1.3–1.4 m height when blooming rosettes cespitose, generally circular, 30–40 cm tall, 60–70 cm in
diameter, forming dense small clusters of 2–5 rosettes. Leaves 25–34 in number, reflexed; sheath broadly ovate, 3.5–4
× 5–6.8 cm, light brown, margins entire at the base, distally serrate, lustrous and glabrous at the base, lepidote distally
on both surfaces; blade narrowly triangular, acuminate to long attenuate, 50–75 × 3–3.5 cm, green, reddish toward
the margin when sun exposed, glabrous and shiny above, and white lepidote below; spines antrorse or divaricate,
triangular, ca. 3 mm long, 8–30 mm apart, green to light brown. Inflorescence central, erect, emerging from a fully
grown rosette (strict sympodium growth pattern, type SPP sensu Ramírez-Morillo et al. 2014).
Staminate inflorescence a once-divided panicle, cylindrical in general shape, erect to slightly curved, 1.3–1.4
m long; peduncle terete, green, sparsely white lepidote, 40–50 cm long, 8–10 mm in diameter at the base, about
as long as rosette height but shorter than rachis of the inflorescence; internodes 2.5–3.0 cm long; peduncle bracts
triangular, 10–13 × 0.5–1.0 cm, margins of its sheath sparsely spiny, the blade long attenuate, acute, entire, multi-
nerved, reddish at the base, already brownish and dry at anthesis, densely white lepidote on both surfaces, usually
twice as long as internodes; rachis (main axis) in a zig-zag fashion, 90–100 cm long, 8–10 mm in diameter at the
base, terete, green with brown stripes, the surface sparsely white lepidote, the internodes 5–5.5 cm long; primary
bracts triangular, attenuate, apex ending in a spine, 2.5–3 × 0.6–0.7 cm, margins sparsely spiny, brownish, densely
white lepidote abaxially, almost glabrous adaxially, multi-nerved, much shorter than the branches; branches ca. 22 in
number, forming an angle of 70–90° relative to the rachis, 9–16 cm long, 2–2.5 cm in diameter, (37–)78–90 flowered;
rachis (branch) terete, longitudinally sulcate, 5–6 mm in diameter, densely white lepidote, stipe nearly none; floral
bracts triangular, short-acuminate, concave, 6–9 × 4 mm, green basally, apex light brown, sparsely white lepidote
only on the basal abaxial surface, margins hyaline, erose, 7-nerved, mid nerved and apiculus conspicuous on abaxial
surface, dark brown, equaling the sepals in anthesis. Flowers sessile, verticillate, ca. 1 cm long, 4–5 mm in diameter at
anthesis, fragrant (sweet smell) during the morning; pedicel (a constriction of the floral receptacle) obconic, 0.5 –1.0
mm long, 0.8–1 mm in diameter, light brown, sparsely lepidote; sepals oblong, acute, cucullate, 4–6 × 4 mm, green
at the base, brownish at the apex, entire or slightly serrate at the apex, glabrous on both sides, or with some sparse
trichomes on the basal abaxial surface, multi-nerved; petals oblong, rounded, cucullate, 5.5–6 × 4 mm, white, glabrous,
multi-nerved, petal apices convergent in anthesis and stamen filaments emerging between them; filaments triangular,
flattened, 6–9 mm long, 5 mm wide, white; anthers narrowly oblong, 1.5–3 mm long, dorsifixed, green, pollen yellow;
pistillode reduced, 1.5–2.5 mm long, ca. 2 mm in diameter, green, stigmatic lobes much reduced.
Pistillate inflorescences a once-divided panicle, in general shape cylindrical, erect, 1–1.4 m long; peduncle terete,
ca. 66 cm long, 1–1.4 cm in diameter at the base, longer than the rosette; internodes 2–3.5 cm long; peduncle bracts
triangular, long attenuate, acute, 10–13 × 0.5–1.5 cm, multi-nerved, reddish at the base, brownish and dry even when
inflorescence is in anthesis, densely white lepidote on both surfaces; margins of its sheath sparsely spiny, the blade
entire; rachis (main axis) ca. 70 cm long, ca. 0.8–1 cm in diameter at the base, terete; internodes 1.5–3.5 cm long;
primary bracts narrow-triangular, acuminate, 2.5–4 × 0.7–0.9 cm, brown-reddish when fresh, brown when dry, sparsely
lepidote abaxially, glabrous adaxially, densely serrate, multi-nerved, much shorter than the branches; branches 20–22
in number, in an angle of ca. 45° with the rachis, 13–25 cm long, 20–34 flowered; rachis (branch) with stipe 10–12
mm long, terete, channeled, glabrous; floral bracts lanceolate-ovate, acuminate, 7–9 mm long, 3.5–5.5 mm wide,
brown, glabrous except the sparsely white lepidote base adaxially, brown, margin minutely denticulate, about equaling
the sepals. Flowers sessile, fragrant (sweet smell) during the morning, 13–15 mm long, 4.5–5.5 mm in diameter at
anthesis; sepals broadly ovate, acute, 3.3–4 × 3.3–3.6 mm, glabrous on both sides, except some sparsely trichomes
on the basal abaxial surface, greenish basally, apically brown, multi-nerved; petals triangular to ovate, acute, 3.5–4.3
× 1.5–2.1 mm, glabrous on both surfaces, with a few minute trichomes on basal abaxial surface, white, multi-nerved,

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 59
during anthesis adnate to the ovary and apical portion erect; staminodes six in number, narrowly triangular, laminar,
2–2.4 mm long, ca. 0.5 mm at the base; ovary oblongoid, 3.9–4.9 mm long, 1.8–2 mm in diameter, glabrous, greenish,
stigmatic lobes recurved, 1.3–1.7 mm long, white. Fruits narrowly ovoid, 11–15 mm long, 6–8 mm in diameter, base
shrinks and resembles a pedicel, glabrous, brown when dry; seeds fusiform, 5.5–7.5 mm long, 0.8–1.4 mm in diameter,
brown, reticulate, caudate, wings 2–3.5 mm long, brown reddish.
Additional specimens examined:—MEXICO. Jalisco: Municipio Amatitán, barranca de Santa Rita, 1000 m,
July 13, 1969, L.M. Villarreal de Puga 3423♂ (GUADA!, IBUG!); Municipio San Cristóbal de la Barranca, 3 km al
sur de Techaluta, 780 m, July 25, 1995, P. Carrillo-Reyes & R. Ramírez-Delgadillo 22♂ (IBUG!, IEB!); arroyo El
Tamarindo, 3.5 km al NNO de San Cristóbal de la Barranca, por el Río Cuixtla, 21°04’00”N, 103°25’00”W, 880 m,
February 26, 2000, P. Carrillo-Reyes & R. Bello 1136♂ (IBUG!, UAMIZ!); Santa Cruz de Enmedio, 0.5 km al NNO de
Santa Cruz de Atístique y 8.5 km en línea recta al NNO de San Cristóbal de la Barranca, 21°07’08”N, 103°27’10”W,
940 m, March 25, 2013, P. Carrillo-Reyes & A. Castro-Castro 6937♂ (IBUG!); La Mesa de los Caballos, ca. 8 km al
NW de San Cristóbal de la Barranca, 1565 m, July 17, 2013, P. Carrillo-Reyes et al. 7001♀ (CICY!, IBUG!); 0.5 km
al N de Ixcamilpa por el camino a Sta. Cruz, 21°06’07”N, 103°26’27”W, 910 m, September 9, 2014, P. Carrillo-Reyes
et al. 7506♀ (CICY!, IBUG!); 1 km en línea recta al N de El Escalón, 21°00’37”N, 103°25’24”W, 1050 m, March 26,
2015, P. Carrillo-Reyes & C.J. Ramírez-Díaz 7714♀(IBUG!); 0.5 km en línea recta al N de El Escalón, km 30 carretera
Guadalajara-Colotlán, 1040 m, January 22, 2016, P. Carrillo-Reyes et al. 8221♀ (IBUG!); Municipio Zapopan, cerca
de la ranchería La Coronilla del Ocote, llegando por Tesistán y Santa Lucía, 20°56’07”N, 103°36’03”W, 1260 m,
June 19, 2000, J.A. Lomelí et al. 3265 fruits (IBUG!, UAMIZ!). Nayarit: Municipio Amatlán de Cañas, Cerca de Los
Barrenitos, ca. 2 km en línea recta al NE de Amatlán de Cañas, 20°49’29”N, 104°23’29”W, 805 m, March 28, 2018,
P. Carrillo-Reyes & D. Cabrera-Toledo 8880-A♀ (IBUG!); 8880-B♂ (IBUG!). Zacatecas: Municipio Trinidad García
de la Cadena, El Abra, 5 km en línea recta al SW de García de la Cadena, 21°10’03”N, 103°29’22”W, 1760 m, August
24, 2014, P. Carrillo-Reyes et al. 7470 fruits (CICY!, IBUG!).
Phenology:—Staminate plants have been collected in January, March and July; female flowers are documented
during January-March and July-September, while fruits have been collected during June-August.
Distribution and habitat:—Hechtia jaliscana has been collected in the municipality of Amatlán de Cañas, in
the State of Nayarit, municipalities of Amatitán, San Cristóbal de la Barranca, and Zapopan in the State of Jalisco,
and municipality of Trinidad García de la Cadena, in Zacatecas, Mexico. Most of the localities are in the basin of
the Santiago River, except for those of Amatlán de Cañas, Nayarit, which is in the Ameca river basin, all of them
are situated in the Pacific Lowlands biogeographic province according to Morrone et al. (2017), in the low elevation
valleys of the southern portion of the Sierra Madre Occidental province (Figure 1). Individuals of this species grow as
lithophytes, mostly in northern exposure rocky slopes, inside tropical deciduous forest vegetation (Rzedowski 1978),
at elevations between 750–1750 m, frequently accompanied by individuals of Bursera fagaroides (Kunth) Engler,
Casearia corymbosa Kunth, Ceiba aesculifolia (Kunth) Britten & Baker f., Celtis caudata Planch., Haematoxylum
brasiletto H. Karst., Opuntia fuliginosa Griffiths, Pachycereus pecten-aboriginum Britton & Rose, Pseudobombax
palmeri (S. Watson) Dugand, Stenocereus queretaroensis (F.A.C. Weber) Buxb. In some localities, H. jaliscana grows
in pine-oak forest dominated by Quercus resinosa Liebm., Q. praineana Trel., and Pinus oocarpa Scheid., inhabiting
rocky slopes alongside species as Agave guadalajarana Trel., A. rzedowskiana P. Carrillo, Vega & R. Delgad., A.
schidigera Lem., Mammillaria scrippsiana (Britton & Rose) Orcutt, species of the genera Muhlenbergia Schreb.,
Opuntia Mill., and Selaginella P. Beauv. (Figure 2).
Discussion:—Hechtia jaliscana (Figures 2 and 3) is recognized by the cespitose rosettes, thick succulent leaves,
glabrous and shiny above, and white lepidote below, armed with coarse spines. The inflorescence, a once-divided
panicle in both sexes (Figure 3A), erect to slightly pending, is produced by young, newly formed rosettes. Flowers are
fragrant during morning hours, with a sweet scent that attract bees (Figure 3B). The rachis of the pistillate inflorescence
is sinuose to almost fractiflex (Figure 2A) while it is slightly sinuose in staminate inflorescences (Figure 2B), with
branches in almost 90° angle with the rachis (main axis); flowers open, but the petal apices converge and form a closed
corolla with the stamens protruding in between (Figure 3D, F), while petals on female flowers remain adnate to the
ovary with the three stigmatic lobes exerted (Figure 3E). The rachis of pistillate branches is longitudinally sulcate
(Figure 3G) with nearly sessile flowers, with a pedicel (rather the floral receptacle that shrinks and looks like a pedicel)
when in fruit (Figure 3H), a common phenomenon in species of Hechtia.
IUCN Conservation assessment:—The EOO (1434 km2) and AOO (44 km2) calculated from 13 collections
corresponding to 12 localities with the GeoCAT (Bachman et al. 2011), suggest that H. jaliscana is an Endangered
(EN) species, according to the IUCN B set of criteria.

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FIGURE 1. Geographical distribution of Hechtia jaliscana, H. marthae and H. subalata. Biogeographical provinces are depicted in
different colors; their limits are based on Morrone et al. (2017).

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FIGURE 2. Hechtia jaliscana in habitat. A. Structure of the rachis in the infructescence. B. Details of the structure of the staminate
inflorescence at anthesis. C. Rosettes with male flowers in habitat. D. Individuals in habitat of tropical dry forests. (Credits: A, C. Ivón
Ramírez-Morillo. B, D. Claudia Ramírez-Díaz).

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FIGURE 3. Hechtia jaliscana. A. General structure of a male individual in bloom. B. Bee visiting fragrant male flowers. C. Origin of
the inflorescence in a female plant. D. Details of male flowers. E. Details of female flowers. F. Branch with male flowers. G. Branch with
female flowers. H. Preserved ripe fruits. (Credits: A, D, F. Claudia Ramírez-Díaz. B, C. Ivón Ramírez-Morillo. E, G. Pablo Carrillo-Reyes.
H. Fruit features taken from P. Carrillo-Reyes & D. Cabrera-Toledo 8880-A♀ (IBUG)).

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 63
Hechtia marthae I. Ramírez, sp. nov. (Figures 4, 5).
TYPE:—MEXICO: Durango: Municipio Nombre de Dios, adelante de Nombre de Dios, camino a El Venado, 11 km del desvío, matorral
de encino-pino, sobre laderas en suelo volcánico, 23°47’55’’N, 104°17’40’’W, 1745 m, originally collected by I. Ramírez, A. Espejo,
A. López-Ferrari, A. Mendoza & J. Ceja, September 2005, flowering under cultivation, June 2009, I. Ramírez & G. Carnevali 1627♂
(Holotype, CICY!).
Diagnosis:—Hechtia marthae is similar to H. subalata but differs in its pistillate branches much shorter than the
primary bracts (vs. conspicuously larger than the primary bract), with shorter branches (2–5 vs. (2–)5–15 cm long),
floral bracts narrowly triangular, 2–5 × 1 mm (vs. broadly ovate, 3–4.5 × 3–4 mm), ovary 1/3 inferior (vs. completely
superior), with longer petals (5–7 vs. 3–5 cm long).
Plants lithophytic or terrestrial, cespitose, 1.1–1.75 m height when flowering; rosettes generally round, 30–40 cm
tall, 35–45 cm in diameter, generally forming dense, colonies of 5–21 rosettes. Leaves 20–30 in number, central ones
erect to slightly reflexed in basal ones; sheath ovate to broadly ovate, 2.8–6.5 × 2–6.3 cm, light brown, margins entire
and erose distally, lustrous and glabrous at the base, slightly lepidote distally; blade narrowly triangular, acuminate to
long attenuate, 20–33(–50) × 1.3–4 cm, green, reddish toward the apex and margin when sun exposed, densely white
lepidote abaxially, white lepidote adaxially; marginal spines antrorse and retrorse, triangular, 2–5 mm long, 6–20 mm
apart, brown, with a short tuff of white trichomes in the concave portion of the spine. Inflorescence central, erect,
emerging from a mature rosette (strict sympodium growth pattern, type SPP sensu Ramírez-Morillo et al. 2014).
Staminate inflorescences a once-divided panicle, cylindrical in general shape, erect, 0.7–1.35 m long; peduncle
terete, green to light brown, slightly white lepidote, 33–79 cm long, 8–17 mm in diameter at the base, longer than
the rosette; internodes 0.8–2.5 cm long; peduncle bracts the basal ones, with ovate sheath, 2–3 × 1.5–2 cm, the
blades narrow triangular, 9–24 × 0.9–1.3 cm, the distal bracts with a sheath rectangular to triangular, 4–7 × 0.8–1.4
cm, margins entire then erose and spiny toward the blade, this long attenuate to acuminate, multi-nerved, brownish,
densely white lepidote abaxially, glabrate or slightly lepidote adaxially, longer than internodes, clasping the peduncle;
rachis (main axis) 33–62 cm long, 3–9 mm in diameter at the base, terete, green to brownish; internodes 0.8–3.6 cm
long, white lepidote; primary bracts rectangular to triangular, abruptly acuminate, 4–5 × 1.2–1.4 cm, entire at the base
to erose and spiny toward the apex, brown when dry, sparsely white-lepidote abaxially, glabrous to slightly lepidote
adaxially, multi-nerved, about the length of the branches; branches 18–27 in number, forming an angle of 15° or less
with the rachis, (1.8–) 4–7.5 cm long, ca. 5 mm in diameter, 16–29 flowered; rachis (branch) flattened at its base,
0.2–1 cm long, glabrous, stipe; floral bracts inconspicuous, narrowly-triangular, acuminate, 1.5–2(–4) × 0.1 mm,
green, brownish at the apex, slightly lepidote, inconspicuously nerved, equaling or slightly exceeding the pedicel.
Flowers pedicellate, polystichous, 7–9 mm long, 3–5 mm in diameter, fragrant during the morning (sweet smell);
pedicel obconic, 2–3 (–4) mm long, ca. 1 mm in diameter, light green, slightly lepidote; sepals ovate to triangular,
acute, 3–4 × 1.8–3 mm, entire, green at the base, apically brownish, slightly lepidote at the base, 3-nerved, shorter than
petals; petals elliptic, acute, 5–6 × 2–3.5 mm, green with brownish spots, multi-nerved; filaments triangular, terete,
4–5 mm long, ca. 1 mm in diameter, white to greenish; anthers oblong, 2–3 mm long, dorsifixed, green, pollen yellow;
pistillode reduced, 0.5–0.8 mm long × 1.5 mm in diameter, light green, stigmatic lobes much reduced.
Pistillate inflorescences a once-divided panicle, in general shape cylindrical, erect, 0.83–1.12 m long; peduncle
terete, 40–55 cm long, 0.8–1.9 cm in diameter at the base, longer than the rosette; internodes 1–2.5 cm long; peduncle
bracts without clear distinction between sheath and blade, basal ones with ovate sheath, blade triangular and long
acuminate, 5–8.5 × 1.8–2.6 cm, margins entire at the base to erose and spiny distally, brownish, densely white lepidote
abaxially, glabrate or slightly lepidote adaxially, multi-nerved; longer than internodes, clasping the peduncle; rachis
(main axis) 30–57 cm long, ca. 0.7–1.1 cm in diameter at the base, terete, green, internodes 1–3 cm long; primary
bracts rectangular to ovate-triangular, acuminate, 2–4 × 0.8–2 cm, brown when dry, sparsely white-lepidote abaxially,
glabrous to slightly lepidote adaxially, entire at the base to erose and spiny toward the apex, multi-nerved, as long or
shorter than the branches; branches 15–25 in number, in an angle of ca. 45° with the rachis, 2–5 cm long, ca. 1.3–1.8
cm in diameter (up to 4 cm when in fruit) 5 mm in diameter, 10–20 flowered; rachis (branch) terete, white lepidote,
glabrous in the infructescence, stipe nearly none; floral bracts inconspicuous, narrowly-triangular, acuminate, 2–5
× 1 mm, green, white lepidote, margins serrate. Flowers pedicellate, verticillate, fragrant during the morning (sweet
smell), 10–12 mm long, ca. 5–7 mm in diameter; pedicels which are a constriction of the base of the floral receptacle
resembling a pedicel, obconic, 2–3 mm long, 1–2 mm in diameter, tomentose; sepals ovate to triangular, acute, 4–5
× 3–4 mm, green, slightly lepidote abaxially, entire, 3-nerved; petals elliptic, acute, 5–7 × 3–4 mm, green, glabrous,
entire, multi-nerved; staminodes six in number, narrowly triangular, laminar, 2.5–2.9 mm long, ca. 1 mm at the base;
ovary 1/3 inferior, oblongoid, 7–9 mm long, 3–4 mm in diameter, glabrous, green, stigmatic lobes recurved, ca. 1 mm

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FIGURE 4. Hechtia marthae. A. Pistillate plant in bloom. B. Staminate plant in bloom. C. Detail of inflorescence with young fruits. D.
Female flowers at anthesis. E. Male flowers at anthesis. Illustration by Alberto Guerra based on photographs by A, D. Ricardo Quirino-Olvera.
B. Gustavo Romero-González. C. Adolfo Espejo-Serna from the specimen A. R. López-Ferrari et al. 2989 (UAMIZ). E and staminate rosette
based of photographs by Ivón Ramírez-Morillo. D and pistillate rosette in based of photographs by Ricardo Quirino-Olvera.

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 65
long, pale green. Fruits pedicellate (rather the base of the floral receptacle shrinks and resembles a pedicel), ellipsoid,
10–14 mm long, 5–8 mm in diameter, glabrous, green when mature, brown when dry; seeds fusiform, 3.5–5 mm long,
1–1.5 mm in diameter, brown, reticulate, caudate, wings ca. 1 mm long, brownish.
Additional specimens examined:—MEXICO. Durango: Municipio Mezquital, Temoaya, a 4.5 km por el camino
a Agua Zarca, 23°20’N, 104°30’W, matorral subtropical, 1650 m, August 18, 1983, M. González E. s.n. fruits (CIIDIR!,
HUAA!, UAMIZ!, UJAT!); Temoaya, 2.5 km al noreste por la carretera a Mezquital, 23°18’59’’N, 104°29’19’’W,
matorral subtropical, 1704 m, May 25, 2020 (cultivated), R. Quirino-Olvera 143♀ (CICY!); Municipio Nombre de
Dios, Chachacuaxtle, 33 km al SSW de Nombre de Dios, 22°40’N, 104°16’W, matorral subtropical, 1740 m, March
21, 1988, S. González E. 4086 fruits (CIIDIR!, HUAA!, UAMIZ!, UJAT!); ca. 10 km sobre la brecha a El Venado,
a partir de la carretera Nombre de Dios-Durango, 23°47’55’’N, 104°17’40’’W, matorral espinoso, 1739 m, July 25,
2003, A.R. López-Ferrari et al. 2989 fruits (UAMIZ!, UJAT!); 2993♂ (UAMIZ!, UJAT!); ca. 10 km en la brecha que
va de la carretera Durango-Nombre de Dios hacia El Venado, 23°47’54.3’’N, 104°17’38.8’’W, cañada con matorral
xerófilo, 1751 m, June 18, 2009, I. Ramírez 1627♂ (CICY!, WU!); same locality, June 3, 2013, I. Ramírez 2205♂
(CICY!); same locality, October 31, 2019, I. Ramírez et al. 2442 fruits (CICY!); Municipio Topia, canyon below Topia,
just downstream from Los Molinos, on cliff, 3800 feet alt., June 1978, M. Kimnach & H. Sánchez-Mejorada 1777♂
(HNT!, US!).
Phenology:—Staminate plants have been collected in June, pistillate plants only reported for May, while fruiting
specimens have been collected from March to October.
FIGURE 5. Hechtia marthae. A. Plants in habitat. B. Infructescence with open fruits in habitat. C. Seeds. (Credits A-C. Ivón Ramírez-
Morillo).
Distribution and habitat:—Populations of Hechtia marthae has been found thus far only on four localities in
the western slope of the Sierra Madre Occidental, in the state of Durango, in Mezquital, Nombre de Dios, and Topia

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municipalities. Its distribution matches partially with the extensions of the Pacific Lowlands province that penetrate
the lowlands of the Sierra Madre Occidental province (Morrone et al. 2017). It grows at elevation between 1158–1751
m, on xerophytic shrublands and subtropical scrub along with species of Agave L., Dasylirion Zucc., Fouquieria
Kunth, Opuntia, Prosopis L., and Vachellia Wight & Arn. (Figure 5).
Eponomy:—The species is named after Martha González-Elizondo in recognition for her outstanding contribution
to the knowledge on the Flora of Durango and for collecting material of the species.
IUCN Conservation assessment:—The GeoCAT (Bachman et al. 2011) indicates that, based upon the EOO
(7953 km2) and AOO (28 km2), H. marthae is classified as Vulnerable (VU) and Endangered (EN), respectively,
according to the IUCN B set of criteria. In order to better understand the conservation status of this species robustly, it
is necessary to evaluate demographic aspects of its populations and possible threats.
Discussion:—Populations included here in Hechtia marthae were misidentified previously as Hechtia subalata
or H. podantha Mez (1896: 549), due to a holotype that only represents an apical portion of an immature infructescence
(see Table 1 for comparisons of main differences among the three species). As many species of Hechtia, blooming
periods are very short and flowers in an inflorescence are open for 1–3 days, making difficult to document floral
structures which show more accurately differences among species (such as colors, shapes, fragrances, etc.). This
species shows wide and relatively long primary bracts, lepidote, with condense pistillate branches resembling species
of the Hechtia podantha complex (Martínez-Correa 2008), probably a case of convergent evolution grouping species
from different lineages (Ramírez-Morillo et al. 2018a) invading high elevation dry habitats occasionally exposed to
frosts.
TABLE 1. Main differences among Hechtia marthae, H. podantha and H. subalata. *Characters for Hechtia podantha were
mostly taken from Espejo-Serna et al. (2010) and completed with information from herbarium specimens; seed features were
from Fatzer (1996) and Gorocica (2017).
Characters Hechtia marthae Hechtia podantha* Hechtia subalata
Leaf blades: length x
width (cm)
20–33(–50) × 1.3–4 19–44.7 × 0.5–1.5 20–50 × 2.5–4.8
Leaf blade surfaces Densely white lepidote
abaxially, white lepidote
adaxially; rarely developing a
red color on the blade
Minutely white lepidote adaxially,
rather glaucous, white lepidote
abaxially; rarely developing red
longitudinal lines at the margin of
the blades
Densely white lepidote abaxially,
white lepidote at base but soon
glabrous and glossy adaxially;
sometimes developing a red spot at
the base of the spine
Spines length (mm);
separation
2–5; 6–20 apart, with white
tuff of hairs on concave portion
of spine
3.1–7 (–8); 6.4–32 apart 1.5–2.5; 8–25 apart
♂Inflorescence type 1-branched 3-branched 1–branched
♂Peduncle diameter
mm; color; surface
8–17; green to light brown,
slightly white lepidote
4.6–11(–15), dark green to purple,
sparsely white lepidote
5–14; green to brownish, densely
white lepidote
♂Primary bracts length
and width (mm); shape
40–50 × 12–14; rectangular
to triangular and abruptly
acuminate
22–48(–60) × 7–12(–25); widely
ovate
12–40 × 8–14; narrowly ovate-
triangular
♂Branch length (cm) (1.8–)4–7.5 1.6–6.6 2.5–10(–15)
♂Floral bracts (mm);
shape
2–4 × 1; linear-triangular,
acuminate
1.2–3.1 × 0.2–1; linear to triangular,
acute
4.5–6 × 2.5–4.5; broadly ovate,
acute
♂Pedicels length (mm) 2–3(–4) 1.4–2.7 0 (sessile flowers)
♀Inflorescence length
(cm)
Once-branched; 83–112 3-branched, 35–127 Once-branched; 120–170
♀Peduncle diameter
mm; color; surface
8–19; light green; densely
white lepidote
7.3–18; dark green to purple;
sparsely white lepidote
8–20 mm; green to brownish;
sparsely lepidote
♀Branch length (cm) 2–5 2.5–9 (2–)5–15
♀Primary bracts (mm) 20–40 × 8–20; rectangular to
ovate triangular
24–60 ×18–20; ovate triangular 15–25 × 8–15; triangular
♀Floral bracts (mm) 2–5 × 1; narrowly triangular,
acuminate
1.1–5.3 × 0.1–0.6; triangular, acute 3–4.5 × 3–4, broadly-ovate, acute
Fruit length; diameter
(mm), shape
10–14; 5–8, ellipsoid 7.1–12; 4.1–6.5, ovoid 9–15; 6–7; ellipsoid
Seeds length x diameter
(mm); shape
3.5–5 × 1–1.5; fusiform 2–52; 7.2–21; ellipsoid 35–40 × 1; triangular
Distribution (Mexican
States)
Durango Hidalgo, Aguascalientes,
Guanajuato, Querétaro
Durango, Jalisco, Nayarit, Zacatecas

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Hechtia marthae, H. podantha and H. subalata are very similar in several features: they share the central
inflorescence that emerges from a fully formed rosette, inflorescences in both sexes have their primary branches
subtended by a conspicuous primary bract, branches are usually condensed, and flowers are usually green, as well as
the fact that inflorescences are in bloom for 2–3 days, a common feature of Hechtioideae but with few exceptions.
Main differences among the three species are depicted on Table 1.
Hechtia subalata L.B. Sm., Contribution Gray Herbarium 117: 15–16, t. 1, f. 29. 1937. TYPE:—MEXICO: Durango,
Sierra Madre, August 13, 1897, J. N. Rose 3467 fruits (holotype, US!; photograph and fruits, GH!).
EPITYPE (designated here):—MEXICO: Zacatecas: Municipio Valparaíso, 1 km SE del Puente Tepetatita, 26 km W de Huejuquilla el
Alto rumbo a San Juan Capistrano, 22°40’36.1’’N, 104°03’03.3’’W, 1078 m, matorral xerófilo con Prosopis, Jatropha, Croton,
Myrtillocactus, originally collected May 27, 2015 by P. Carrillo-Reyes, J.L. Tapia, C.J. Ramírez-Díaz and I. Ramírez; flowering in
cultivation in 2019, I. Ramírez et al. 2430♂ (epitype CICY!; isoepitypes, IBUG!, MEXU!).
Plants lithophytic, cespitose, 1.4–1.8 m when flowering; rosettes in general shape globose, 40–60 cm tall, 50–60 cm
in diameter, generally forming dense, small colonies of 2–6 rosettes, rarely clumps of 8–12 rosettes. Leaves 20–35
in number, central ones erect, basal ones slightly reflexed; sheath broadly ovate to widely elliptical, 4–5.2 × 4.5–7
cm, margins entire to erose toward the apex, lustrous and glabrous at the base adaxially and slightly lepidote at the
apex, white lepidote abaxially; blade narrowly triangular, acuminate to long attenuate, 20–50 × 2.5–4.8 cm, green,
sometimes with red spots at the apex and or margins, at the base of every spine, densely white lepidote abaxially, white
lepidote at base but soon glabrous and glossy adaxially; marginal spines retrorse rarely antrorse, triangular, 1.5–2.5
mm long, 0.8–2.5 cm apart, light brown or occasionally with red spots close to the spines. Inflorescence central, erect,
emerging from a fully grown rosette (strict sympodium growth pattern, type SPP sensu Ramírez-Morillo et al. 2014).
Staminate inflorescences a once-divided panicle, cylindrical in general shape, erect, 0.65–1.8 m long; peduncle
terete, green to brownish, sparsely lepidote, 20–56 cm long, 5–14 mm in diameter at the base, surpassing the rosette;
internodes (0.8–) 1.1–4.8 cm long; peduncle bracts the basal ones with wide-ovate sheath, 1–2.2 × 1.1–7 cm, the blade
narrow triangular, long attenuate, slightly pungent, (3.5–)8.7–17 × 0.3–0.7 cm, green to brownish, the apical ones
without a clear distinction between sheath and blade, ovate to triangular, long attenuate to acuminate, 2.2–4.5 × 0.8–1.1
cm, margins entire to erose and spiny toward the blade multi-nerved, brownish when dry, white lepidote abaxially,
glabrate adaxially, longer than internodes; rachis (main axis) 0.44–1.32 m long, 5–10 mm in diameter at the base,
terete, brownish; internodes 1–3.4 cm long; primary bracts ovate-triangular, acuminate, 1.2–4 × 0.8–1.4 cm, entire
to dentate, light brown, densely lepidote abaxially, slightly lepidote adaxially, multi-nerved, shorter than branches;
branches 16–59 in number, forming an angle of 45° or less with the rachis, 2.5–10 (–15) cm long, ca. 1.5 cm in
diameter, 28–72 flowered; rachis (branch) terete, light brown, 7–8 cm long, 0.8–1.2 mm in diameter, densely lepidote,
stipe nearly none; floral bracts broadly ovate, acute, 4.5–6 × 2.5–4.5 mm, brown, margins erose to dentate, glabrous
on both surfaces, multi-nerved, sometimes with a conspicuous central nerve, equaling or slightly surpassing the sepals.
Flowers sessile, polystichous, divaricate, fragrant during the morning (sweet smell); sepals ovate to triangular, acute,
cucullate, 3–4 × 2–2.5 mm, entire to slightly erose, brownish, glabrous on both surfaces, multi-nerved; petals elliptical,
acute to rounded, cucullate at the apex, 4–5 × 2–3 mm, white with brown spots, glabrous on both surfaces, multi-
nerved; stamens exserted, ca. 4.5 mm long; filaments triangular, flattened, ca. 4 mm long, white; anthers oblong, 2–3
mm long, dorsifixed, yellow, pollen yellow; pistillode reduced, 1.5 × 2 mm, stigmatic lobes much reduced.
Pistillate inflorescences a once-divided panicle, cylindrical in general shape, erect, 1.2–1.7 m long; peduncle
terete, 0.35–1.18 m long, 8–20 mm in diameter at the base, surpassing the rosette; internodes (1–)2–4 cm long; peduncle
bracts the basal ones with triangular sheath, 1.5–25 × 1.4–1.8 cm, the blades narrow triangular, long attenuate, slightly
pungent, 10–27 × 0.5–0.7 cm, foliaceous, green to brownish, the apical ones without a clear distinction between
sheaths and blades, ovate to triangular, 2–3.5 × 0.9–1.2 cm, margins entire to erose and spiny toward the blade, long
attenuate to acuminate, brownish when dry, white lepidote abaxially, glabrate adaxially, multi-nerved, longer than
internodes; rachis (main axis) 63–73 cm long, 6–9(–14) mm in diameter at the base, terete, green; internodes (1–)2–3.5
cm long; primary bracts triangular, acuminate, 1.5–2.5 × 0.8–1.5 cm, light brown, glabrous on both surfaces, entire
to dentate, multi-nerved, shorter than branches; branches 22–32 in number, forming an angle of 45° or less with the
rachis, (2–)5–15 cm long, 4–8 mm in diameter, with 10–44 flowered, flowers occasionally emerging from the base of
the branch; rachis (branch) terete, green, 0.4–0.8 cm long, 2–4 mm in diameter, glabrescent, stipe nearly none; floral
bracts broadly ovate, acute, 3–4.5 × 3–4 mm, without covering or exceeding the sepals, margins erose to dentate,
glabrous on both surfaces, multi-nerved, sometimes with a conspicuous central nerve, brown, membranous. Flowers
sessile, polystichous, divaricate, fragrant (sweet smell during the morning); sepals ovate, acute, 2–3.5 mm long, ca.

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2.5 mm wide at the base, entire, green to brownish, glabrous on both surfaces, multi-nerved; petals triangular, acute,
cucullate at apex, 3–4.5 mm long, ca. 2 mm wide at the base, entire, white with brown spots, multi-nerved; staminodes
six in number, triangular, laminar, 1.5–2 mm long, ca. 0.5 mm wide, inserted in the corolla; ovary superior, oblongoid
to ellipsoid, ca. 7 mm long, 3 mm in diameter, light green, glabrous, stigmatic lobes recurved, ca. 1 mm long, light
green, placentation axillary. Fruits ellipsoid, 9–15 mm long, 6–7 mm in diameter, glabrous, erect, brown when dry;
seeds narrowly triangular, brown, 3.5–4 mm long, 1 mm wide, caudate, apical wing ca. 2 mm long, basal wing reduced,
brown.
Additional specimens examined:—MEXICO. Jalisco: Municipio Huejuquilla el Alto, 3.6 km al NW del
poblado Los Arroyos del Agua, km 17.5 de la carretera Huejuquilla el Alto-Ruiz, 22º40’30.7’’ N, 103º59’21.2’’ W,
1260 m, October 6, 2018, D. Figueroa et al. 235♂ (IBUG), 236♀ (IBUG!); Municipio Bolaños, Bolaños, orilla
del pueblo, 21°49’48.0”N, 103°46’48.0”W, 885 m, November 7, 2018, K. Romero-Soler et al. 1239 fruits (CICY!,
IBUG!), 1240♂ (CICY!); Municipio Chimaltitán, foothills 8–12 km SW of Bolaños, R. McVaugh 25929♀ (MICH!);
Municipio Mezquitic, 3 km al E de Tetakararo, bajando la barranca, 800 m, November 10, 1985, C. Chávez-Reyes D.
169♀ (IBUG!); Municipio San Martín de Bolaños, 1 km al S del Rancho La Joya, brecha San Martín al Platanito, 900
m, May 18, 1989, A. Flores et al. 1583 fruits (XAL!); Municipio Villa Guerrero, 11.2 km al oeste de Villa Guerrero
camino a Bolaños, 21°58’03”N, 103°40’36”W, 1780 m, November 6, 2018, K. Romero-Soler et al. 1238 fruits (CICY!);
11.2 km al WSW de Villa Guerrero por la carretera a Bolaños (7.7 km en línea recta), 21°58’03”N, 103°40’36”W, 1780
m, April 4, 2017, P. Carrillo-Reyes et al. 8520 fruits (IBUG!). Nayarit: Municipio El Nayar, arroyo El Fraile, al SE de
la Mesa del Nayar, 22°09’N, 104°33’W, 520 m, October 23, 1989, P. Tenorio & G. Flores 16620♀ (MEXU!, MO!);
Vereda de la Mesa del Nayar al Cangrejo, que cruza por la barranca, 900–1400 m, August 5, 1990, R. Ramírez et al.
551♀♂ (MEXU!). Zacatecas: Municipio Valparaíso, 1 km SE del Puente Tepetatita, 26 km W de Huejuquilla el Alto
rumbo a San Juan Capistrano, 22°40’36.1’’N, 104°03’03.3’’W, 1078 m, May 27, 2015; flowering under cultivation in
2018, I. Ramírez et al. 1989 fruits (CICY!); I. Ramírez et al. 1989a♂ (CICY!, IBUG!, MEXU!); San Juan Capistrano,
August 23, 1883, J. Rose 3556 fruits (HUH!, NY!); 16.8 km al W de Huejuquilla rumbo a San Juan Capistrano,
alrededores del río Atengo o Chapalagana, 1019 m, March 24, 2010, A. Castro-Castro 2027♀ (IBUG!); Valley of río
Atenco (Chapalagana), 8–15 km northeast of San Juan Capistrano, road to Huejuquilla el Alto, sheltered rocky canyon,
and adjoining arid flats, 1000–1200 m, January 14, 1975, R. McVaugh 25795♀ (MEXU!); 6 miles before San Juan
Capistrano on road to Jesus María, 1036 m, January 22, 1976, M. Kimnach 1855♀ (MEXU!); aprox. 2 km antes de
llegar a Chimaltitán, después de Florencia, 790 m, May 20, 1985, J.A. Lomelí-Sención 85-00190♂ (GUADA!).
Phenology:—Staminate plants have been collected in January, March, August, October and November; female
flowers are documented during May, August, October, and November; while fruits have been collected during April,
May, August, and November.
Distribution and habitat:—Hechtia subalata has been collected in the Mexican States of Durango (Municipality
of Mezquital), Jalisco (Bolaños, Chimaltitán, Huejuquilla el Alto, San Martín de Bolaños, and Mezquitic and Villa
Guerrero Municipalities), Nayarit (Del Nayar Municipality) and Zacatecas (Valparaíso Municipality). According to the
biogeographical provinces of Morrone et al. (2017), this taxon occurs in the Pacific Lowlands province, particularly in
canyons that penetrate the Sierra Madre Occidental province (Figure 1), at 800–1300 m of elevation, growing on rocky
slopes, within tropical dry forest, xerophytic shrubland and ecotones between those vegetation types with species of
Agave, Bursera bipinnata (DC.) Engl., B. fagaroides, Coryphantha (Engelm.) Lem., Croton L., Dodonaea viscosa (L.)
Jacq., Echinocereus pamanesiorum Lau, Euphorbia schlechtendalii Boiss., Forestiera phillyreoides (Benth.) Torr.,
Heliocarpus terebinthinaceus (DC.) Hochr., Ipomoea murucoides Roem. & Schult., Leucaena Benth., Myrtillocactus
geometrizans (Mart. Ex Pfeiff.) Console, Opuntia, Prosopis laevigata (Humb. & Bonpl. ex Willd.) M.C. Johnst.,
Stenocereus queretaroensis, and Vachellia pennatula (Schltdl. & Cham.) Seigler & Ebinger (Figure 6).
IUCN Conservation assessment:—Hechtia subalata is known from 17 collections (ten localities); most of them
by roadsides, and most likely occurs as isolated populations on appropriate microniches, rocky slopes in rugged canyons.
It has values of EOO de 6592.7 km² and an AOO of 44 km². According to the IUCN, therefore this species should be
considered as Vulnerable (VU) to Endangered (EN); nevertheless, because of the lack of population information, we
rely mostly upon distributional data, namely, the set of B criteria, geographical distribution.
Discussion:—Species of Hechtioideae, in general and those of Hechtia in particular, are still poorly represented
in herbaria: nothing is more discouraging for the collector than spiny, usually large rosettes with large inflorescences.
Most collectors usually find species in fruit because flowers last only one day and even inflorescences with tens or
hundreds of flowers, last but a few days. It is also known that populations of some species do not bloom every year,
especially those with strict monocarpic growth pattern (SMP, sensu Ramírez-Morillo et al. 2014) where long-lived,
well-developed rosettes take years to produce an inflorescence (for example Hechtia zamudioi Espejo, López-Ferr.

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 69
FIGURE 6. Hechtia subalata: A. Detail of the branching with young fruits. B. Detail of a dry staminate inflorescence. C. Rosette with
infructescence. D. Plants in habitat, showing the red coloration on leaves. (Credits: A, C. Sabina Ascencio. B. Ivón Ramírez-Morillo. D.
Pablo Carrillo-Reyes).

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70 • Phytotaxa 523 (1) © 2021 Magnolia Press
FIGURE 7. Hechtia subalata portraying a rosette in bloom, note the red coloration on the leaves. A. Pistillate inflorescence. B. Female
flowers at anthesis. C. Branch with open fruits. D. Staminate inflorescence. E. Details of the male flowers with open exposing the pollen.
F. Open flowers before anthesis. (Illustration by Alberto Guerra based of photographs by A, B. Omar Góngora. C-F, Ivón Ramírez-
Morillo).

IN DISENTANGLING TWO SPECIES LIMITS OF HECHTIA Phytotaxa 523 (1) © 2021 Magnolia Press • 71
& I. Ramírez (2008: 55)). Species more frequently found in bloom, year after year, are those with sympodial growth,
with precocious-flowering pattern (i.e. Hechtia rosea E. Morren ex Baker (1889: 140)). We still do not know what
triggers plants to produce inflorescences. In the case of Hechtia subalata, as for ca. 26% of the described species, type
specimens only have fruits, and few of them included additional information on flowers or rosettes in their protologues.
For H. subalata, few features are discernable in the holotype (fruit size, inflorescence branching, branch length, etc.)
or the paratype (foliar blade, particularly the margins: sinuose-concave between spines, and spines alternating on both
margins). These data, combined with its geographical distribution, allow us to finally complete our understanding of
this concept and assign a more precise (other than what is found in type specimens) geographical distribution. It is
important to mention that the female flowers of H. subalata (Figure 7B) are practically identical to those of H. jaliscana
(Figure 3E), but their arrangement on the branches (agglomerated in the first and lax in the second) differentiates these
two taxa. It is important to emphasize that this close similarity of female flowers between both species is another
reason to propose epitypites based on a staminate specimen.
Acknowledgments
We thank the curators of the following herbaria: CICY, CIIDIR, IBUG, GH, GUADA, HUAA, HUH, IEB, MEXU,
MICH, UAMIZ, UJAT, US, and ZEA for allowing the study of Hechtia material. We are indebted to CONACyT for
funding the projects “Phylogeny, evolution and biogeography of Hechtia Klotzsch (Hechtioideae: Bromeliaceae)”
(number 183281) and “Diversification of Hechtia Klotzsch (Bromeliaceae) and its colonization of Megamexico:
phylogenetic and phylogeographical evidence and the exploration of patterns in the geographical, ecological and
morphological space” (number 283357) granted to the first author. A. Espejo, A. López-Ferrari, A. Mendoza, J. Ceja,
CICY, as well as academic staff of Herbarium CIIDIR, by financial support for field work in Durango. Thanks also to
Silvia Hernández-Aguilar and Carlos Jiménez-Nah for handling the herbarium material and loans. Thanks to Gustavo
Romero-González from Herbarium AMES, who supplied literature on the genus and facilitated the study of herbarium
material at HUH Herbaria. Thanks to Katya J. Romero-Soler, Sabina Ascencio Ramírez, and Dante S. Figueroa
Martínez for collecting and documenting the species in Jalisco, Mexico, and to Abraham Torres Soto, Arturo Castro
Castro, Jorge Noriega Villa, Karen Rostro del Muro, Néstor Raigoza, and Ricardo Quirino Olvera for documenting
H. marthae in the field. To the Roger Orellana Regional Botanical Garden, for cultivating the specimens. To Germán
Carnevali, Gustavo Romero-González, Ricardo Quirino Olvera, and Omar Góngora for photographs that were used to
elaborate the plates of H. subalata and H. marthae. To Alberto Guerra and Ixel Osorio Herrera for the illustrations of H.
subalata and H. marthae respectively. To Ricardo Rivera for allowing us to use preliminary data on the distribution of
the species. To Katya J. Romero-Soler for editing the manuscript and photographs, and for her comments on a previous
draft of this manuscript. Thanks to two anonymous reviewers for their advice, comments, and meticulous editing,
specially to Eric Gouda, for which we are grateful.
References
Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial
conservation assessment tool. ZooKeys 150: 117–126.
https://doi.org/10.3897/zookeys.150.2109
Baker, J.G. (1889) Handbook of the Bromeliaceae. 243 pp. [https://www.biodiversitylibrary.org/page/35547198#page/156/mode/1up]
Espejo-Serna, A., López-Ferrari, A.R. & Ramírez-Morillo, I. (2008) Dos nuevas especies de Hechtia (Bromeliaceae; Pitcairnioideae) del
centro de México. Acta Botanica Mexicana 83: 49–61.
https://doi.org/10.21829/abm83.2008.1060
Espejo-Serna, A., Martínez-Correa, N. & López-Ferrari, A.R. (2010) Neotypification of Hechtia podantha (Bromeliaceae). Journal of the
Botanical Research Institute of Texas 4: 221–223.
Fatzer, D.M. (1996) An analysis of seed morphology and systematic relationships in Hechtia (Bromeliaceae). M. S. Thesis. University of
New Orleans. USA.
Givnish, T.J., Millam, K.C., Berry, P.E. & Systma, K.J. (2007) Phylogeny, adaptive radiation, and historical biogeography of Bromeliaceae
inferred from ndhF sequence data. Aliso 23: 3‒26.
https://doi.org/10.3732/ajb.1000059
Gorocica, E.A. (2017) Morfología de semillas en Hechtia Klotzsch (Hechtioideae: Bromeliaceae). Bachelor Thesis. Instituto Tecnológico

RAMÍREZ-MORILLO ET AL.
72 • Phytotaxa 523 (1) © 2021 Magnolia Press
de Conkal, Mérida, Yucatán, 75 pp.
Gouda, E.J., Butcher, D. & Gouda, C.S. (2021 [cont. updated]) Encyclopaedia of Bromeliads, Version 4. Available from: http://bromeliad.
nl/encyclopedia/ University Botanic Gardens, Utrecht (accessed 1 April 2021)
Hernández-Cárdenas, R.A., Siekkinen, A., López-Ferrari, A.R. & Espejo-Serna, A. (2020) Five New Species of Hechtia (Bromeliaceae;
Hechtioideae) from Guerrero, Mexico. Systematic Botany 45: 466–477.
https://doi.org/10.1600/036364420X15935294613608
INEGI (Instituto nacional de estadística, geografía e informática) (2001) Conjunto de Datos Vectoriales Fisiográficos. Continuo Nacional.
Escala 1:1’000,000. Serie I. Instituto Nacional de Estadística y Geografía, Aguascalientes. Available from: http://www.inegi.gob.mx
(accessed 14 October 2021)
IUCN Standards and Petitions Committee (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Prepared
by the Standards and Petitions Committee. Available from: http://www.iucnredlist.org/documents/RedListGuidelines.pdf (accessed
3 August 2019)
Klotzsch, J.F. (1835) Hechtia, eine neue Gattung der Bromeliaceen. Allgemeine Gartenzeitung 3: 401–403.
Martínez-Correa, N. (2008) Sistemática del complejo de especies de Hechtia podantha Mez (Pitcairnioideae, Bromeliaceae). M. S. Thesis.
México City, México: Universidad Autónoma Metropolitana.
McVaugh, R. (1989) Bromeliaceae to Dioscoreaceae, Flora Novo-Galiciana 15. University of Michigan Press. Ann Harbor, 398 pp.
Mez, C. (1896) Bromeliaceae. In: De Candolle, A.L.P.P. & De Candolle, A.C.P. (Eds.) Monographiae Phanerogamarum Prodromi nunc
continuatio, nunc revisio, vol. 9. Sumptibus G. Masson, Paris, 990 pp.
Morrone, J.J., Escalante, T. & Rodríguez-Tapia, G. (2017) Mexican biogeographic provinces: Map and shapefiles. Zootaxa 4277: 277–
279.
https://doi.org/10.11646/zootaxa.4277.2.8
QGIS Development Team (2019) QGIS Geographic Information System. Open Source Geospatial Foundation Project. Available from:
http://qgis.osgeo.org (accessed 14 October 2021)
Ramírez-Morillo, I., Jiménez, C.F., Carnevali Fernández-Concha, G. & Pinzón, J.P. (2014) Three new species and growth patterns in
Hechtia (Bromeliaceae: Hechtioideae). Phytotaxa 178: 113–127.
https://doi.org/10.11646/phytotaxa.178.2.3
Ramírez-Morillo, I., Carnevali, G., Pinzón, J.P., Romero-Soler, K., Raigoza, N., Hornung-Leoni, C., Duno, R., Tapia-Muñoz, J.L. &
Echevarría, I. (2018a) Phylogenetic relationships of Hechtia (Hechtioideae; Bromeliaceae). Phytotaxa 376: 227–253.
https://doi.org/10.11646/phytotaxa.376.6.1
Ramírez-Morillo, I., Carnevali, G., Pinzón, J.P., Romero-Soler, K., Raigoza, N., Hornung-Leoni, C., Duno, R. & Tapia-Muñoz, J.L.
(2018b) The reestablishment of Bakerantha, and a new genus in Hechtioideae (Bromeliaceae) in Megamexico, Mesoamerantha.
Harvard Papers in Botany 23: 301–312.
https://doi.org/10.3100/hpib.v23iss2.2018.n15
Rzedowski, J. (1978) Vegetación de México. Limusa México, D.F. 432 pp. Available from: https://www.biodiversidad.gob.mx/
publicaciones/librosDig/pdf/VegetacionMx_Cont.pdf (accessed 14 October 2021)
Scharf, U. & Gouda, E.J. (2008) Bringing Bromeliaceae back to homeland botany. Journal of Bromeliad Society 58: 123–129.
Smith, L.B. (1934) Studies in Bromeliaceae –V. Contributions from the Gray Herbarium of Harvard University 104: 71–83.
Smith, L.B. (1937) Studies in the Bromeliaceae – VII. Contributions from the Gray Herbarium of Harvard University 117: 3–33.
Smith, L.B. (1938) Bromeliaceae, in Britton. North American Flora 19: 97–145.
Smith, L.B. (1964) Notes on Bromeliaceae–XII. Phytologia 10: 454–488. [https://www.biodiversitylibrary.org/page/13190376#page/9/
mode/1up]
Smith, L.B. & Downs, R.J. (1974) Pitcairnioideae (Bromeliaceae). In: Flora Neotropica Monograph 14, Part 3. Haffner Press, New York,
658 pp.
Thiers, B. (2021 [continuously updated]) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical
Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 9 January 2020)
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold,
K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (eds.) (2018) International Code of Nomenclature for
algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017.
Regnum Vegetabile 159. Glashütten: Koeltz Botanical Books.
https://doi.org/10.12705/Code.2018
Villaseñor, J.L. (2016) Checklist of the native vascular plants of México. Revista Mexicana de Biodiversidad 87: 559–902.
https://doi.org/10.1016/j.rmb.2016.06.017