Pinguicula zamudioana (Lentibulariaceae) a new species endemic to western Mexico

Abstract

Pinguicula zamudioana (Lentibulariaceae), a new species endemic to western Mexico, is described and illustrated. This new species belongs to Pinguicula section Orcheosanthus and is morphologically close to Pinguicula oblongiloba and P. michoacana, but differs in having a homophyllous rosette with a single type of leaves (summer leaves), light green leaves with glabrous petioles, short peduncles, a calyx with triangular-lanceolate to lanceolate lobes, a pink corolla with oblong to suborbiculate lobes and the fact that it only inhabits calcium concretions. Pinguicula zamudioana is easily distinguished from the other two taxa, due to the lack of a winter rosette, i.e. isomorphic leaves during the entire year, and to the fact that it is in constant growth and blooms all year round.

Full text

Phytotaxa 372 (4): 243–255
http://www.mapress.com/j/pt/
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Andreas Fleischmann: 22 Sept. 2018; published: 19 Oct. 2018
https://doi.org/10.11646/phytotaxa.372.4.1
243
Pinguicula zamudioana (Lentibulariaceae) a new species endemic to western
Mexico
HÉCTOR DAVID JUÁREZ-GUTIÉRREZ1, MIGUEL ÁNGEL MUÑIZ-CASTRO2* JULIÁN HERNÁNDEZ-
RENDÓN3 & ANA TERESA NUÑO-RUBIO3
1 Mareterra Consultores, Ottawa 1186A, Providencia Italia, Guadalajara, C.P. 44648, Jalisco, Mexico:
david.juarez@mareterraconsultores.com
2Herbario IBUG (“Luz María Villarreal de Puga”), Laboratorio de Ecosistemática, Instituto de Botánica, Departamento de Botánica y
Zoología, Universidad de Guadalajara, Camino Ramón Padilla 2100, Las Agujas, Nextipac, Zapopan, Jalisco, Mexico:
miguel.muniz@academicos.udg.mx
3Centro Universitario de Ciencias Biológicas y Agropecuarias (CUCBA), Universidad de Guadalajara, Camino Ramón Padilla 2100,
Las Agujas, Nextipac, Zapopan, Jalisco, Mexico.
*author for correspondence
Abstract
Pinguicula zamudioana (Lentibulariaceae), a new species endemic to western Mexico, is described and illustrated. This
new species belongs to Pinguicula section Orcheosanthus and is morphologically close to Pinguicula oblongiloba and P.
michoacana, but differs in having a homophyllous rosette with a single type of leaves (summer leaves), light green leaves
with glabrous petioles, short peduncles, a calyx with triangular-lanceolate to lanceolate lobes, a pink corolla with oblong
to suborbiculate lobes and the fact that it only inhabits calcium concretions. Pinguicula zamudioana is easily distinguished
from the other two taxa, due to the lack of a winter rosette, i.e. isomorphic leaves during the entire year, and to the fact that
it is in constant growth and blooms all year round.
Keywords: carnivorous plant, Jalisco, Nueva Galicia region
Introduction
The genus Pinguicula Linnaeus (1753a: 17) is one of the three genera in the carnivorous plant family Lentibulariaceae
(Lamiales), with ca. 98 species (Roccia et al. 2016, Rivadavia et al. 2017, Zamudio et al. 2018, Crespo et al. 2018,
Burelo et al. 2018) distributed in North America, Europe, northern and eastern Asia, Central America, Antilles and
South American Andes (Casper 1966; Zamudio 2005). America is the richest continent in Pinguicula species with
ca. 78 species (Ulloa–Ulloa et al. 2017, Rivadavia et al. 2017, Zamudio et al. 2018, Burelo et al. 2018), having its
main center of diversification in Mexico, with ca. 49 accepted species (Lampard et al. 2016, Rivadavia et al. 2017,
Fleischmann & Roccia 2018, Zamudio et al. 2018, Burelo et al. 2018). All taxa of the genus are herbaceous, rosette–
forming carnivorous plants with typical lamialean, bilabiate, tubular and spurred flowers (Fleischmann & Roccia
2018).
During the preparation of the treatment of Lentibulariaceae for the “Flora de Jalisco y áreas colindantes”
corresponding approximately to the Nueva Galicia region (sensu McVaugh 1961) in Mexico, the existence of a new
species of the genus Pinguicula was recognized. This taxon is described here as a species new to science based on
comparative herbarium studies and field observations.
Materials & Methods
A comprehensive literature review was conducted. The material of Pinguicula deposited in the herbaria CHAPA,
CIIDIR, EBUM, ENCB, FCME, HUAA, IBUG, IEB, MEXU, MICH, UAMIZ, UAT and XAL (herbarium acronyms

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244 • Phytotaxa 372 (4) © 2018 Magnolia Press
following Thiers 2018+) was reviewed and determined. Additionally, we studied types and digitalized images from the
following herbaria: CAS, COLO, DS, F, GH, K, LL, MO, NMC, NY, P, PH, POM, RSA, TEX, UMO, WIS and WU
(see Appendix 1). Exploration and field gathering of Pinguicula species were carried out mainly in the Mexican state
of Jalisco, but other adjacent areas corresponding to the Nueva Galicia region (sensu McVaugh 1961) where explored.
Voucher specimens collected from field surveys were made following Bridson & Forman’s method (1989) and were
deposited at IBUG. A distribution map was created in ArcMap 10.5 (Environmental Systems Research Institute 2014)
by using available free spatial geodata for Mexico with WGS 1984 projection.
FIGURE 1. Pinguicula zamudioana. A, seed. B, corolla, side view. C, corolla variation. D, hairs from corolla. E, habit. F, leafs series. A,
C, D, F from D. Juárez, M.A. Muñiz & A. Nuño 210, B, E from photographs of plants in situ. Drawing by J.M. Ramírez and D. Juárez.

A NEW SPECIES OF PINGUICULA, P. ZAMUDIOANA Phytotaxa 372 (4) © 2018 Magnolia Press • 245
FIGURE 2. Distribution of Pinguicula zamudioana (star) and the related species P. oblongiloba (dots) and P. michoacana (diamond) in
Mexico (location data based on herbarium specimen records).
Description of the new species
Pinguicula zamudioana D.Juárez & Muñiz-Castro, sp. nov. (Figs. 1–3)
Similar to Pinguicula oblongiloba DC. but differs in having a homophyllous rosette with a single type of leaves (summer leaves), light
green leaves with glabrous petioles, short peduncles, calyx with triangular–lanceolate to lanceolate lobes, pink corolla with oblong
to suborbicular lobes and inhabiting calcium concretions.
Type:—MEXICO. Jalisco. Ixtlahuacán del Río municipality, Río Verde canyon, wet sandy wall in canyon background, 1234 m, 19
November 2014, D. Juárez, M.A. Muñiz & A. Nuño 210 (holotype IBUG!).
Perennial herbaceous plant, homophyllous. Leaves arranged in a lax basal rosette, with 4–8(10) membranaceous
leaves, petiolate; petiole slightly concave, 10–20(30) mm long, margin entire; lamina obovate–elliptic to suborbicular
(15)25–50(70) mm long and 15–50(60) mm wide, apex rounded, base cuneate to attenuated, margin involute, covered
by sessile and stipitate glands on the upper face. Peduncles 1–4, erect, (35)40–60(80) mm long, green, covered by
scattered stipitate glands, uniflorous. Flowers pentamerous, 40–55(65) mm long including spur. Calyx bilabiate, covered
by stipitate glands on the outer face and scant on the inner face, upper lip trilobed, lobes triangular–lanceolate, 2–4 mm
long and 1–3 mm wide, lower lip bilobed, lobes lanceolate, 2–4 mm long and 0.5–3 mm wide. Corolla bilabiate, pink,
with a small white macula at the base of the lower lip, upper lip bilobed, lobes oblong–elliptic to suborbicular, 10–15
mm long and 5–10 mm wide, lower lip trilobed, lobes obovate–oblong to suborbicular, 8–20(25) mm long and 5–10
mm wide, often superimposed and then undulated, apex rounded. Tube very short, infundibuliform, 2–5 mm long and
1.5–3 mm wide, slightly dyed pink, with capitated glandular hairs in the interior. Spur cylindric–subulate, 20–30(35)
mm long and 0.5–1 mm wide. Ovary subglobose, glandular–pubescent, with short stipitate glands, ± 1 mm in diameter,
stigma bilobed, pink, subsessile, lower lobe slightly larger than upper, oblate. Capsule subglobose, covered by short
stipitate glands, scattered, ±3 mm long and ±3 mm wide. Seeds numerous, ellipsoidal, ±0.5 mm long and ± 0.1 mm
wide.

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246 • Phytotaxa 372 (4) © 2018 Magnolia Press
FIGURE 3. A–C, Pinguicula zamudioana. A–B, habitat. C, habit, face view. D–E, P. oblongiloba. D, habitat. E, habit, lateral view. F–G,
P. michoacana. F, habitat. G, habit, face view. Photos A–C by A. Nuño, D by D. Juárez and E–G by S. Zamudio.

A NEW SPECIES OF PINGUICULA, P. ZAMUDIOANA Phytotaxa 372 (4) © 2018 Magnolia Press • 247
Distribution:—Pinguicula zamudioana is known only from three restricted locations close to each other that are
difficult to access in the Río Verde Canyon in Ixtlahuacán del Río and Zapotlanejo municipalities, which belong to
Jalisco’s central region, at 1100–1250 m elevation (Fig. 2).
Conservation status:—The distribution area of the new species is located within the protected natural area
“Formación Natural de Interés Estatal Barrancas de los Ríos Santiago y Verde” managed by Gobierno del Estado de
Jalisco (2016), however, according to the IUCN Red List Criteria (IUCN 2012) used to evaluate the conservation
status, P. zamudioana should be categorized as Critically Endangered, due to the fact that it has a known extent
of occurrence (EOO) <100 km2 (B1 criterion) and a known area of occupancy (AOO) <10 km2 (B2 criterion) with
severely fragmented locations [condition (a)] and an estimated continuing decline of area of occupancy [condition
(b)(ii)], quality of habitat (iii) and number of locations or subpopulations (iv). The 28-meter-high El Purgatorio dam is
under construction and will flood soon one of the three known subpopulations of P. zamudioana if it is not successfully
translocated to a safe place.
Etymology:—The specific epithet honors Sergio Zamudio Ruiz, who has enthusiastically dedicated himself to
the study of the genus Pinguicula in Mexico and Central America.
Ecology and phenology:—The plant grows on limestone concretions over rocky and shaded walls with water
runoff, in deciduous tropical forest with few mesophilic elements such has Magnolia pugana (Iltis & Vázquez in
Vázquez (1994: 14)) Vázquez & Carvajal in Vázquez et al. (2002: 137), Oreopanax peltatus Linden (1859: 368) and
Trophis racemosa (Linnaeus 1753b: 1190) Urban (1905: 195). Flowering and fruiting occurs throughout the year. The
rosette develops continuously through the year, although in winter, when the temperature and the humidity decrease,
it reduces the rate of growth and the size of the leaves.
Taxonomic relationships:—According to the subgeneric classification by Casper (1966) Pinguicula zamudioana
is to be placed in Pinguicula subgenus Pinguicula, section Orcheosanthus Candolle (1844: 27) by its deeply bilabiate
corolla, corolla lobes greater than the tube, short tube, broadly infundibuliform, spur longer than the tube, and its
resemblance to P. oblongiloba Candolle (1844: 27) and P. michoacana Zamudio & Juárez in Zamudio et al. (2018:
16), sharing with them the frequently presence of oblong-obovate corolla lobes with rounded apex, corollas with pink
tones (pink-violet in P. oblongiloba and P. michoacana) and the presence of a white macula at the base of the lower
lip of the corolla (see Table 1 and Fig. 3). It is necessary to take into consideration that the heteromorphic rosette was
a distinctive trait of the group. However, Zamudio in Burelo et al. (2018) proposed a new amendment for P. section
Orcheosanthus, where the recently described P. olmeca Zamudio, Burelo & González in Burelo et al. (2018: 360) can
be placed due to their absence of a winter rosette, and now P. zamudioana confirms the absence of winter rosettes
for a part of the section. The isomorphic rosette in this species is likely to be a recent adaptation to stable weather
conditions and until now, only this character is known within the section for P. olmeca, P. zamudioana and partially
for P. moctezumae Zamudio & Ortega (1994: 58) and P. moranensis Kunth in Bonpland & Humboldt (1817: 226)
(Zamudio 2001, Lampard et al. 2016, Burelo et al. 2018).
TABLE 1. Comparison of P. zamudioana, P. oblongiloba and P. michoacana. Measurements were taken from examined
herbarium specimens and literature (Zamudio 2001, Zamudio et al. 2018). CF = Cypress forest. DTF = Deciduous tropical
forest. JF = Juniper forest. OF = Oak forest. PF = Pine forest. POF = Pine–Oak forest.
P. zamudioana P. oblongiloba P. michoacana
Winter leaves
number – (25)60–86 20–40
apex – acute–acuminate distinctly acuminate
margin – ciliate glabrous
Summer leaves
......continued on the next page

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TABLE 1 (Continued)
P. zamudioana P. oblongiloba P. michoacana
number (2)4–8(10) (2)3–4(8) 2–8
size (15)25–50(70) × 15–50(60) mm (16)20–50(75) × (6)10–55 mm (15)20–60(70) × 10–55 mm
color light green green to purple green
Petiole margin glabrous coarsely ciliate ciliate
Peduncles
size (35)40–60(80) mm (50)100–150(230) mm (40)50–120(150) mm
Flowers
form bilabiate bilabiate bilabiate
color pink pink to purple pink to purple
Calyx lobes
form triangular–lanceolate to
lanceolate lanceolate lanceolate
size 2–4 × 0.5–3 mm (1.5)4–6 × 1–3 mm 3–5 × 0.5–2 mm
accrescence absent present absent
Flowering continuous May–August March–September
Habitat DTF with mesophilic elements OF, PF, POF, JF, CF DTF
Geological substrate limestone igneous rocks limestone with interspersed
igneous rocks
Pinguicula zamudioana resembles P. oblongiloba and P. michoacana, but is easily distinguished from the other
two taxa, due to the lack of a winter rosette, i.e. isomorphic leaves during the whole year, constant growth and blooming
all year round, and flowers that are generally pink and petioles that lack cilia. The phenological stage of flowering
throughout the year is not exclusive to P. zamudioana in the section Orcheosanthus, there is another species within the
section, P. moctezumae, which also blooms throughout the year. The habitat is similar to that of P. michoacana, only
with scarce mesophilic elements and a constant water availability, as in this site a dry season is practically nonexistent
(Table 1, Fig.3).
On the other hand, the subdivision of Pinguicula in three subgenera [Pinguicula L. (Casper 1962: 60), Isoloba
Barnhart (1916: 47) and Temnoceras Barnhart (1916: 47)] proposed by Casper (1966) has been refuted in recent works
(Cieslak et al. 2005; Shimai & Kondo 2007, Shimai 2017; Fleischmann & Roccia 2018) because it is not natural
according to molecular phylogenetics, but the clades are rather clustered by geographic affinity within the genus.
De Candolle (1844), based on morphology and flower color, recognized three sections for the genus Pinguicula
[Orcheosanthus Candolle (1844: 27), Pionophyllum Candolle (1844: 28), and Brandonia Candolle (1844: 32)]. Section
Orcheosanthus was characterized by having a large purplish corolla, with five subequal lobes, with a significantly short
tube and a spur longer than the corolla tube. Since De Candolle diagnosis of Orcheosanthus, its circumscription has not

A NEW SPECIES OF PINGUICULA, P. ZAMUDIOANA Phytotaxa 372 (4) © 2018 Magnolia Press • 249
changed essentially, rather it is currently valid. A century later, Casper (1966) introduced some additional characters
to the diagnosis of P. section Orcheosanthus: heterophyllous leaves (two different shapes of rosette leaves during the
year), hibernacle absence, widely infundibuliform tube and deeply bilabiate corolla.
According to the taxonomic treatments based on morphology by Casper (1966), Luhrs (1995), Luhrs & Lampard
(2006), Speta & Fuchs (1982), Zamudio & Rzedowski (1991), Zamudio (2001), Zamudio et al. (2018) and Burelo et al.
(2018), up to 15 species have been recognized as valid taxa for P. section Orcheosanthus: twelve endemic to Mexico
[P. colimensis McVaugh & Mickel (1963: 138), P. cyclosecta Casper (1963a: 11), P. elizabethiae Zamudio (1999: 16),
P. gypsicola Brandegee (1911: 190), P. macrophylla Kunth in Bonpland & Humboldt (1817: 226), P. michoacana, P.
moctezumae, P. oblongiloba, P. olmeca, P. potosiensis Speta & Fuchs (1989: 100), P. rectifolia Speta & Fuchs (1989:
97) and P. zecheri Speta & Fuchs (1982: 111); and three endemic to Mexico and Central America [P. mesophytica
Zamudio (1997: 65), P. moranensis and P. orchidioides Candolle (1844: 27)]. Nevertheless, the last classification of
the genus Pinguicula proposed by Shimai (2017), mainly based on an extensive revision of morphological features
and a phylogenetic analysis, disagrees with the current classification of Casper (1966) based only on morphology.
According to the phylogenetic analysis of ITS sequences of Shimai (2017) [a phylogenetic method of choice suggested
by Degtjareva et al. (2006)], the genus Pinguicula segregates into nine infrageneric groups (clades) that mainly
correspond with geographical distribution. In that analysis the species normally assigned to P. section Orcheosanthus
were segregated in two clades: Clade VII, which consists of 18 taxa distributed from northwestern Mexico to Central
America (including P. colimensis P. moctezumae P. moranensis, P. oblongiloba, P. potosiensis, P. rectifolia P. zecheri
P. mesophytica, among others which did not previously belong to P. section Orcheosanthus); and Clade VIII, which
consists of 16 taxa (including P. cyclosecta P. gypsicola and P. macrophylla, among others which do not belong to P.
section Orcheosanthus) mostly distributed in northeastern Mexico. Within this context, P. zamudioana is proposed
to be allocated within the Clade VII. However, despite the wide morphological diversity, monophyly and genetic
homogeneity of these clades are well agreed, Shimai (2017) proposed eleven sections without any subgeneric ranks
for Pinguicula, in which the species of P. section Orcheosanthus are included in his section Mesoamericana Shimai
(2017: 350).
Additionally, Fleischmann & Roccia (2018) consider the three major clades evident from phylogenetic
reconstructions by Cieslak et al. (2005), Degtjareva et al. (2006) and Beck et al. (2008) to be subgenera, in which
by nomenclatural priority at subgenus and section rank, the species of P. section Orcheosanthus belong to P. section
Temnoceras Casper (1963b: 333) in P. subgenus Temnoceras.
Additional specimens examined (paratypes):—MEXICO. Jalisco. Ixtlahuacán del Río municipality, ravine
inside the Río Verde Canyon, 1140 m, 02 February 2013, M.A. Muñiz, A. Nuño & A. Nuño 1385 (IBUG!). Zapotlanejo
municipality, 1104 m, 01 April 2018, A. Nuño 54 (IBUG!).
Identification key to Pinguicula species occurring in western Mexico
The identification of plants of this genus from herbarium specimens can be complicated, since it is compulsory to use
flowers, as well as information on “winter” and “summer” leaves and rosettes.
1. Corolla subisolobate, lobes equal or almost equal, no clear distinction between superior and inferior lip, tube subcylindric and spur
shorter than the tube ..........................................................................................................................................................................2.
– Corolla bilabiate, lobes not equal, with a clear distinction between the superior and inferior lip, tube short, infundibuliform, spur
equal or longer than the tube .............................................................................................................................................................4.
2. Annual plants, with a single type of leaf (homophyllous); corolla lilac or white, palate sublentiform yellow, with some of the veins
tinted in purple ......................................................................................................................................................................P. lilacina
– Perennial plants, heterophyllous, winter rosette hypogeous, bulbous; corolla white, palate absent, veins of the tube not tinted in
purple .................................................................................................................................................................................................3.
3. Summer leaves ovate–elliptical, blades 15–60 mm long, by 11–40 mm wide, winter leaves ovate–lanceolate or lanceolate, 10–30
mm long, by 5–20 mm wide, apex acute to acuminated, margin ciliated; calyx lobes deltoid, accrescent during fructification; cap-
sule 4–5 mm long, covered by the calyx .............................................................................................................................. P. casperi
– Summer leaves oblong to elliptical, blade 15–50 mm long, by 7–15 mm wide, winter leaves oblong–lanceolate to lanceolate,
15–22 mm long, by 2–10 mm wide, apex acute–acuminated, margin lightly ciliated; calyx lobes oblong–lanceolate, not accrescent
during the fructification; capsule 3–3.5 mm long, not covered by the calyx .................................................................. P. parvifolia
4. Annual plant, small and fragile, with a single type of leaf (homophyllous); flower 3–7 mm long (including the spur); corolla white
with palate, inferior lip with a yellow spot at the base, lobes dentate, crenate or laciniate .......................................... P. crenatiloba
– Perennial plants, homophyllous or heterophyllous, flowers 28–120 mm long (including the spur); corolla pink, purple or lilac, with
entire lobes .........................................................................................................................................................................................5.

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5. Homophyllous plants that do not form a winter rosette and grow and bloom all year round, glabrous petioles ......... P. zamudioana
– Heterophyllous plants, winter and summer rosettes well differentiated, bloom only during spring or summer ..............................6.
6. Winter rosette extended over the surface or slightly covered by soil (epigeous or subepigeous) .....................................................7.
– Winter rosette, compact, bulbous, buried 1–3 cm below the surface (hypogeous) ...........................................................................8.
7. Corolla pink–purple; growing on gypsum hills, in deciduous rain forest, at 350–500 m elevation, only known from Colima ..........
......................................................................................................................................................................................... P. colimensis
– Corolla purple; growing on igneous rock walls or epiphyte in moist pine–oak forest or cloud forests, at 1600–2600 m elevation,
known from Guerrero and Michoacán .................................................................................................................................P. zecheri
8. Summer leaves suborbicular, 40–195 mm long, petioles 12–60 mm long, not ciliated; superior lip lobes oblong to ovate–cunei-
form ............................................................................................................................................................................. P. macrophylla
– Summer leaves obovate–elliptical, oblong–elliptical to suborbicular, 20–120 mm long, petiole 10–45 mm long, ciliated; superior
lip lobes oblong–elliptical, oblong–ovate, obovate–oblong, elliptical to suborbicular .....................................................................9.
9. Winter rosette covered by dry membranous (scarious) leaves, external winter leaves ciliated, grows in oak, pine, pine–oak or cy-
press forest .................................................................................................................................................................... P. oblongiloba
– Winter rosette not covered by dry membranous (scarious) leaves, winter leaves not ciliated, grows in deciduous rain forest ..........
...................................................................................................................................................................................... P. michoacana
Acknowledgments
We would like to thank Alberto Nuño Rubio, Alejandro Zabalgoitia Ibarra, Francisco Javier Ramos Gallegos, Francisco
Javier Rendón Sandoval and Jesús Padilla Lepe for their company and support during the field explorations; Rafael
Soltero Quintana and Pablo Israel Navarro Cuevas for their support to describe the new species; Arturo Castro Castro
and Jesús Guadalupe González Gallegos for their spot on notes and revisions on specimens from MICH herbarium;
José Manuel Ramírez Amezcua and Daniela Alejandra Cuevas Sanchez for the figure preparation; Mario Arnaldo
Méndez Brilanti for the revision of the manuscript. Two anonymous reviewers are thanked for helpful comments.
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APPENDIX 1: Examined specimens of related species from Nueva Galicia region (P. michoacana and
P. oblongiloba)
P. michoacana:—MEXICO. Michoacán: Jiquilpan: 06 March 2014, S. Zamudio & I. García 16686 (paratypes
MEXU!, UAMIZ!); 18 September 2014, S. Zamudio & I. García 16860 (holotype UAMIZ!, isotypes CIIDIR!, ENCB!,
EBUM!, IBUG!, MEXU!, UAMIZ!).
P. oblongiloba:—MEXICO. Aguascalientes: Calvillo: 26–28 August 1960, R. McVaugh, C. Feddema & R.W. Pippen
18387 (MICH!); 13 August 1976, S. Correa 170 (MICH!); 18 July1996, C. Medina 3325 (IEB!); 07 December 2010,
G. González 300 (HUAA!). San José de Gracia: 15 July 1988, G. Garcia 4195 (HUAA!); 19 June 2000, G. González
391 (HUAA!); 19 June 2000, G. González 393 (HUAA!); 19 June 2000, G. González 396 (HUAA!); 24 July 2000, G.
González 404 (HUAA!); 24 July 2000, G. González 416 (HUAA!). Chihuahua: Camargo: 11 July 1946, W.P. Hewitt
140 (GH!). Guadalupe y Calvo: 14 July 1946, W.P. Hewitt 139 (GH!). Balleza: 21 August 1970, C.W. Pennington 91
(TEX!). Ocampo: 31. July 1988, R. Spellenberg, R. Corral, J. Brunth & L. Huenneke 9558 (IEB!, MEXU!, NMC!).
Temósachi: 28 July 1988, J.E. Laferriere 1574 (MEXU!). Urique: 14 July 1958, I.W. Knoblock 856 (MIC!). Uruachi:
24 July 1974, R.A. Bye 6443 (MEXU!) Colima: Comala: 06 August 1992, R. Ramírez 2901 (IEB!); 31 July1993, S.
Zamudio 9139 (IEB!, MEXU!). Minatitlán: 04 July 2004, Y. Ramírez 337 (IEB!). Durango: Canelas: 21 August 1990,
A. Benítez 1748 (MEXU!). Durango: 24 July 1958, D.S. Correll & I.M. Johnston 20102 (TEX!); 01 August 1970, D.
Flyr 1518 (LL!); 22 July 1975, D. LeDoux 1955 (ENCB!, UMO!); 13 July 1990, A. García, M. González & S. Acebedo
452 (CIIDIR!, IEB!); 14 July 1990, A. García 498 (CIIDIR!, IEB!); 17 August 1991, A. García & S. Acebedo 1019
(ENCB!, IEB!, MEXU!); 20 July 2001, R. Carrillo 100 (CIIDIR!, IEB!); 05 July 2002, I. Ortega & F. Pacheco 48
(CIIDIR!, IEB!). Mezquital: 15 July 1985, A. Torres 18 (CIIDIR!, IEB!); 21 July 1985, M. Gonzalez 1789 (IEB!); 22
July1985, M. González 1814 (MEXU!); 08 July1988, I. Solís 929 (MEXU!); 18 October 1985, S. Gonzalez 3774
(ENCB!, MEXU!). Pueblo Nuevo: 28 August 1934, F.W. Pennell 18326 (PH!); 28 June 1950, J.H. Maysilles 7153
(MEXU!, MICH!, TEX!); 28 June 1950, J.H. Maysilles 7188 (NY!); 26 July 1950, J.H. Maysilles 7489 (MICH!); 16
July 1955, J.H. Maysilles 7858 (MICH!); 08 August 1957, R.L. Dressler 2153 (MO!); 09 July 1961, L.E. Detling 8446
(WIS!); 23 June 1964, G. Mick & K. Roe 95 (MICH!, WIS!); 21 July 1969, B. Marcks & C. Marcks 1226 (ENCB!,
MICH!, WIS!); 27 August 1974, M.L. Roberts & D. Keil 10290 (NY!); 25 July 1981, S. Gonzalez & M. Gonzalez 1772
(CHAPA!, ENCB!); 26 June 1982, P. Tenorio, C. Romero & R. Hernández 644 (ENCB!, MEXU!); 01 July 1982, C.
González 58 (CHAPA!, ENCB!); 06 July 1982, R. Hernández & P. Tenorio 7688 (MEXU!, RSA!); 30 June 1984, P.
Tenorio, C. Romero & T.P. Ramamoorthy 6078 (MEXU!); 28 July 1991, M. Flores, A. Espejo & A.R. López 526 (IEB!,
UAMIZ!); 26 July 2003, A. Espejo 6562 (IEB!, UAMIZ!). Rodeo: 24 July 1976, D.E. Breedlove 39517 (DS!). San
Dimas: 10 August 1955, J.H. Maysilles 8321 (MICH!); 25 July 2000, R. Alvarado 19 (IEB!). Santiago Papasquiaro:
15 July 1982, R. Hernández 7924 (MEXU!, UAT!); 05–08 July 2008, D. Tejero (MEXU!). Súchil: 03 July 1957, H.L.
Short s.n. (K!); 26 June 1982, M.A. Morales 15 (CHAPA!, ENCB!, MEXU!); 14 August 1984, F. Chávez 51 (CIIDIR!,
IEB!, MEXU!); 23 July 1990, R. Spellenberg & S. González 10280 (MEXU!); 10 August, A. García, S. Gonzalez & A.
Acevedo 128 (CIIDIR!, IEB!, MEXU!). Topia: 09 July 1983, R. Corral & R.D. Worthinton 227 (MO!, TEX!).
Guanajuato: Pénjamo: 12 July 2009, I. Guardián 288 (IEB!). San Felipe: 08 August 1987, R. Galván & J. Galván
2817 (IEB!, MEXU!); 28 June 1988, R. Galván & S. Galván 4891 (ENCB!). Jalisco: Ahualulco de Mercado: 24 June
1984, L. González 2007 (IBUG!); 10 July1993, M. Cházaro, M. Negrete, L. Vázquez & P. Corona 7172 (IEB!, MEXU!,
XAL!); 05 July 2012, D. Juárez & M. García 38 (IBUG!); 05 July 2012, D. Juárez & M. García 42 (IBUG!). Ameca:
06 September 2014, J. Hernández & A. Rodríguez 2 (IBUG!). Autlán de Navarro: 28 June 1949, R. Wilbur & C. Wilbur
1403 (MICH!); 15 July 1949, R. Wilbur & C. Wilbur 1744 (MICH!); 01 August 1985, A. Vázquez 3407 (MEXU!); 03
July1986, R. Cuevas 1454 (WIS!); 13 July1988, R. Cuevas 3020 (WIS!); 13 July 1992, A. Leinberger 74 (WIS!); 16
June 1994, F. Santana & L. Guzmán 6780 (IBUG!). Ayotlán: 09 July 1892, M. Jones 87 (POM!). Bolaños: 16 June1990
A. Flores, G. Martinez & N. Ramos 1895 (IEB!, MEXU!); 01 June 1991, A. Flores & J. Flores 2726 (IEB!); 24
July1991, M. Flores 475 (IEB!, UAMIZ!); 30 June 1996 G. Flores & J. Calonico 4595 (MEXU!); 02 July 1996, J.
Calonico & G. Flores 2411 (MEXU!); 06 July 1996, J. Calonico & G. Flores 2501 (IBUG!, MEXU!); 16 July 1997,
G. Flores & J. Calonico 4841 (MEXU!). Casimiro Castillo: 29 July 1987, F. Santana, R. Soltero & R. Ramírez 2787
(IBUG!). Cihuatlán: 22 July 1988, R. Cuevas 3184 (WIS!). Ciudad Guzmán: 14 July1985, O. Alfaro 48 (IBUG!); 15
September 1990, R. Ramírez, R. Soltero & C. Ramírez 2283 (IBUG!). Ejutla: 20 July 2001, P. Carrillo 2286 (IEB!).
Gómez Farías: 25 June 1980, F. Trujillo s.n. (IBUG!). Guadalajara: 22 May 1889, C. Pringle 2554 (MEXU!, MO!,
NY!, PH!, RSA!, WU!); 10 July 1902, C. Pringle s.n. (MEXU!). Huejuquilla el Alto: 19 July 1992, J. Reynoso 939
(IEB!). Jesús María: 20 June1983, J. Pérez s.n. (IBUG!). Jocotepec: 28 June 1964, L. Villarreal s.n. (MICH!); 12 June

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1966, L. Villarreal 328 (ENCB!, IBUG!); 12 June1966, L. Villarreal 329 (IBUG!); 28 July 1991, J. Machuca & M.
Cházaro 6669 (IEB!); 05 October 2011, D. Juárez, A. Rodríguez & M. Carrasco 4 (IBUG!). Mascota: 13 June1974, C.
Díaz 5136 (MICH!); 14 June1974, C. Díaz s.n. (HUMO!); 21 July 2011, J. González, A. Castro, R. Guerrero, I.
Guerrero & C. Beltrán 1041 (IBUG!). Mazamitla: 20 September 1952, R. McVaugh & J. Sooby 13094 (MICH!); 17
July 1956, D. Gregory & G. Eiten 73–3 (MICH!, MEXU!, MO!); 30 July 1960, H. Iltis, R. Koeppen & F. Iltis 573
(MICH!, WIS!); 20 June 1990, J. Villa & I. Tejeda 764 (IEB!); 13 July 2014, J. Hernández & I. Terrones 1 (IBUG!).
Mezquitic: 09 July 1989, R. Ramírez & R. González 1299 (IBUG!). San Gabriel: 16 October 1952, R. McVaugh & J.
Sooby 13568 (MICH!); 20 June 1957, R. McVaugh, J. Mickel & M. McVaugh 14935 (MICH!); 29 June 1981, E. Lott,
A. Solís & A. Delgado 414a (MEXU!); 23 July 1992, A. Rodríguez & O. Vargas 2472 (IBUG!, IEB!, MEXU!); 07
August1994, L. Villarreal & Suárez 16611 (IBUG!). San Sebastián del Oeste: 06 July 2014, J. Hernández & L.
Hernández 22 (IBUG!). Tala: 20 June1986, A. Rodríguez 283b (IBUG!); 02 July 1988 A. Rodríguez & J. Reynoso 1260
(IBUG!). Talpa de Allende: 23 July 1973, R. González 914 (CHAPA!, MEXU!, MICH!); 12 August 1994, J. Lomelí,
M. Cházaro & A. García 2314 (MEXU!); 15 July 2012, J. González, A. Castro & G. Munguía 1221 (IBUG!, IEB!).
Tapalpa: 10 June 1892, M. Jones s.n. (POM!); 05–07 August 1960, H. Iltis, R. Koeppen & F. Iltis 781 (ENCB!, MICH!,
WIS!); 01 November 1960, R. McVaugh, C. Feddema & R. Pippen 20587 (MICH!); 19 July 1975, L. Villarreal 7670
(IBUG!); 29 June 1981, E. Lott, A. Solís & A. Delgado 409 (MEXU!, MO!); 18 June 1984, H. Iltis 29167 (IBUG!,
IEB!, MICH!, WIS!); 15 July 2001, P. Carrillo 2162 (IBUG!, IEB!). Tecalitlán: 23 June 1957, R. McVaugh, J. Mickel
& M. McVaugh 15011 (MICH!); 14 August 1957, R. McVaugh, J. Mickel & M. McVaugh 16210 (MICH!); 31 July
1960, H. Iltis, R. Koeppen & F. Iltis 595 (WIS!); 17 July 1986, A. Rodríguez 404 (FCME!, IBUG!); 07 July 1988, J.
Gaona 398 (MICH!). Tecolotlán: 23 June 2000, M. Cházaro & J. Luna 8079 (IBUG!, IEB!). Tequila: 17 July 1976, L.
Villarreal 15628 (IBUG!); 25 June 1987, A. Rodríguez & J. Suarez 866 (IBUG!, MEXU!); 31 July 1989, A. López &
A. Espejo 953 (CHAPA!, IEB!, MEXU!, UAMIZ!). Tlajomulco de Zúñiga: 19 July 1986, J. Machuca 2683 (IEB!).
Yahualica de González Gallo: 22 June 2008, L. González & A. Frías 5024 (IBUG!). Zapopan: 23 June 1967, L.
Villarreal 4087 (IBUG!); 25 June 1967, L. Villarreal 2113 (IBUG!); 05 July 1981, J. Vázquez 255 (IBUG!); 5 July
1986, A. Rodríguez & L. Guzmán 323 (IBUG!, WIS!); 06 July 1988, R. Ramírez & R. Soltero 929 (IBUG!); 05 July
1997, M. Harker 864 (IBUG!); 31. August 2011, A. Rodríguez, D. Juárez & L. Maya 6329 (IBUG!); 14 September
2011, D. Juárez & L. Maya 3 (IBUG!); 03 July 2012, D. Juárez & A. Castro 34 (IBUG!). México: Nicolás Romero:
27 June 1963, J. Rzedowski 16849 (ENCB!); 03 June 1979, R. Bracho 225 (ENCB!, IEB!). Temascaltepec: 17 July
1940, C.L. Hitchcock & L.R. Standford 7224 (GH!). Michoacán: Aguililla: 16 June 1945, A.J. Sharp 45504 (MEXU!);
12 July 1985, J. Soto, A. Román & F. Soto 9316 (MEXU!). Charapán: 28 May 1984, J. Labat 857 (MEXU!). Cherán:
20 July 1966, F.A. Barkley 36062 (GH!); 10 June 1987, M. Pérez 54 (IEB!); 27 June 1987, M. Pérez 64 (IEB!).
Coalcomán de Vázquez Pallares: 24 June 1939 G. Hinton 13841 (GH!, NY!, RSA!); 18 June 1999, E. Carranza & A.
Blanco 5746 (IEB!); 28 June 2003, V. Steinmann & M. Fishbein 3251 (IEB!). Coeneo: 04 July 1987, J. Rzedowski
43538 (ENCB!, IEB!, MICH!); 09 July 1988, P.X. Ramos 91 (ENCB!, IEB!). Contepec: 27 July 2005, M.G. Cornejo
1229 (IEB!, MEXU!); 15 September 2005, M.G. Cornejo 1524 (IEB!). Erongarícuaro: 07 August 1992, J.M. Escobedo
2411 (CAS!, IEB!, MEXU!, XAL!). Jiménez: 27 June 1991, E. Pérez 2150 (IEB!, MEXU!). La Huacana: 09 July 2005,
V.W. Steinmann 5160 (IEB!). Marcos Castellanos: 19 July 1989, I. García & M. Carrión 2815 (IEB!). Morelia: 01 July
1909, B.G. Arsene 2618 (MO!, P!); 01 June 1986, J. Rzedowski 39757 (IEB!); 16 June 1986, V.M. Huerta 549 (IEB!);
05 July 1986, V.M. Huerta 567 (EBUM!, IEB!); 26 June 1988, C. Medina 1107 (EBUM!); 07 July 1988, C. Medina
1203 (IEB!); 19 August 1988, S. Zamudio 6698 (IEB!); 22 June 1991, H. Díaz 6708 (IEB!); 22 June 1991, M. Flores
432 (IEB!, UAMIZ!); 17 June 1998, J.L. Linares 4354 (MEXU!); 15 August 2008, M.G. Cornejo 2820 (IEB!, MEXU!).
Pátzcuaro: 17 June 1986, H. Díaz 2319 (ENCB!, IEB!, MEXU!, XAL!); 26 June 1986, S. Zamudio & H. Díaz 3990
(ENCB!, IEB!, MICH!, XAL!); 12 July 1986, J.M. Escobedo 1050 (ENCB!, IEB!, MEXU!); 24 June 1994, S. Zamudio
& E. Pérez 9274 (IEB!, MEXU!); 26 June 1995, S. Zamudio 9520 (IEB!, MEXU!); 23 July 1995, S. Zamudio 9522
(IEB!); 17 September 1995, S. Zamudio 9540 (IEB!); 10 October 1995, S. Zamudio 9563 (IEB!); 04 June 1996, S.
Zamudio 9817 (IEB); 28 June 2003, M.E. Molina & S. Zamudio 104 (IEB!). Quiroga: 23 July 1960, H. Iltis, F. Iltis &
R. Koeppen 359 (MICH!, WIS!). Tancítaro: 30 July 1940, W.M.C. Leavenworth 396 (F!). Tangancícuaro: 02 July 1989,
L. Torres 250 (EBUM!). Tlalpujahua: 23 July 1998, E. Pérez 3834 (IEB!). Tuxpan: 31 May 1945, E.L. Little s.n.
(MICH!). Tzintzuntzan: 25 June 1978, J. Caballero & C. Mapes 225 (IEB!, MEXU!); 14 June 1986, J. Espinosa 2121
(EBUM!, ENCB!, IEB!, MEXU!, XAL!); 04 July 1987, J. Rzedowski 43518 (ENCB!, IEB!). Uruapan: 21 June 1991,
H. Díaz 6733 (IEB!); 21 July 1991, S. Zamudio 8200 (IEB!); 07 July 2003, V.W. Steinmann & M. Fishbein 3359 (IEB!,
MEXU!). Zinapécuaro: 16 June 1998, J.L. Linares 4350 (MEXU!). Zitácuaro: 07 July 1985, J.C. Soto, A. Román & F.
Soto 9017 (MEXU!). Without defined locality: 16 July 1941, R.W. Schery 154 (MO!). Nayarit: El Nayar: 03 August
1990, G. Flores, R. Ramírez & O. Téllez 2234 (MEXU!). Sinaloa: Concordia: 18 July 1976, S. Walker 76H08 (MO!).

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Sonora: Álamos: 07–08 August 1935, F.W. Pennell 19585 (PH!). Yécora: 15 August 1998, R. Spellenberg 12618
(IEB!). Zacatecas: Tlaltenango de Sánchez Román: 02 August 1971, C. Díaz 2369 (ENCB!, MEXU!); 05 July 2006,
M. Fishbein 5848 (IEB!).