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Phytotaxa 292 (3): 279–286
Copyright © 2017 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Lorenzo Peruzzi: 5 Jan. 2017; published: 27 Jan. 2017
Pinguicula pygmaea (Lentibulariaceae), a new annual gypsicolous species from
Oaxaca State, Mexico
FERNANDO RIVADAVIA1, EDWARD LLOYD READ2 & ANDREAS FLEISCHMANN3*
1185 SW 7th St, Miami, FL 33130, USA.
2California State University Fullerton, Department of Biological Science, 800 N. State College Blvd., Fullerton, CA 92834, USA.
3Botanische Staatssammlung München, Menzinger Strasse 67, D-80638 Munich, Germany; e-mail: firstname.lastname@example.org.
*author for correspondence
Pinguicula pygmaea (Lentibulariaceae), a new species from the Sierra Madre del Sur of western Oaxaca, Mexico is de-
scribed and illustrated. The morphological characteristics distinguishing this new species from other similar species are
discussed, together with its distribution and ecology.
Se describe e ilustra Pinguicula pygmaea (Lentibulariaceae), una nueva especie de la Sierra Madre del Sur de Oaxaca
Occidental, México. Se discuten las características morfológicas que distinguen esta nueva especie de otras especies
similares, así como su distribución y ecología.
Key words: butterworts, carnivorous plant, gypsophile, taxonomy
The genus Pinguicula Linnaeus (1753: 17) is one of three genera in the carnivorous plant family Lentibulariaceae
(Lamiales) and comprises c. 95 species, of which about 40% occur in Mexico (Legendre 2000, Cieslak et al. 2005,
Shimai & Kondo 2007, Lampard et al. 2016, Roccia et al. 2016). The state of Oaxaca in southern Mexico is home to
thirteen species of Pinguicula (including the taxon newly described here), eight of which are endemic, the majority
of them occurring on the Sierra Madre del Sur mountain range (Zamudio 2001, 2005, Zamudio & van Marm 2003,
Lampard et al. 2016).
The greater part of Mexican Pinguicula species are perennial plants, surviving a winter dry dormancy as succulent,
non-carnivorous “winter rosettes” or as compact, bulb-like subterraneous rosettes (so-called “heterophyllous growth
type”; Casper 1966, Roccia et al. 2016). However, a few homophyllous species evolved an annual life cycle in
response to the seasonally dry climate: Pinguicula crenatiloba Candolle (1844: 30), Pinguicula lilacina Schlechtendal
& Chamisso (1830: 94) [incl. Pinguicula sharpii Casper & Kondo (1977: 112); species concepts following Roccia
et al. 2016 and Lampard et al. 2016], and Pinguicula takakii Zamudio & Rzedowski (1986: 260). These short-lived
therophytes germinate towards the end of the rainy season and start to die back after anthesis and seed set, outlasting
the dry season as seeds dispersed on the ground.
These ephemeral annual Pinguicula species are generally much smaller and more delicate plants than their
perennial counterparts, with leaves translucent enough that one can see droplets of condensed water collected on the
undersides of the leaves. Among these annuals, P. crenatiloba is one of the smallest species in the genus, usually only
3–4 cm tall in flower, rarely exceeding 7 cm in height, and bearing minute flowers with a corolla of just about 4.5–6
mm in length (Ernst 1961, Casper 1966, Lampard et al. 2016).
During expeditions to western Oaxaca state in 2003 and 2016, a minute and hitherto unknown annual species of
Pinguicula was discovered and studied at three locations growing on gypsum soils. This taxon is described as a species
new to science, based on comparative herbarium studies and observations made in the wild.
RIVADAVIA ET AL.
280 • Phytotaxa 292 (3) © 2017 Magnolia Press
Material and Methods
Herbarium specimens (see also Appendix 1) were studied by at least one of the authors at IEB, M and MEXU (herbarium
acronyms following Thiers 2016), field observations were performed by FR & ELR in Mexico in November 2003 and
by FR in December 2003 and November 2016. The distribution map was created by DIVA GIS (Hijmans et al. 2005)
using free spatial geodata for Mexico.
Description of the new species
Pinguicula pygmaea Rivadavia, E.L.Read & A.Fleischm., sp. nov. (Figs. 1–3)
Similar to Pinguicula crenatiloba DC. regarding habit, leaf shape, the bilabiate corolla with upper lip smaller than the lower lip, and the
overall diminutive size, but differs from that species by entire corolla lobes (crenate to emarginate in P. crenatiloba), by the upper
corolla lip only slightly separated from the lower lip, creating a short tubular throat (deeply bilabiate corolla, tubular corolla throat
almost absent), by the three subequal lobes of the lower corolla lip (median lobe much larger than the two lateral ones), and by a spur
which exceeds the corolla lower lip in length (spur shorter than to at maximum equal to the length of lower corolla lip).
Type:—MEXICO. Oaxaca: Município de Santo Domingo Tonalá, Highway 125, between Huajuapan and Tlacotepec, 1370 m, 20
November 2003, Rivadavia & Read 1814 (holotype IEB!, isotype M!).
Small delicate annual herb, forming a lax rosette spread flat on the ground. Roots few, poorly developed, unbranched.
Active green leaves 2–5, membranous, obovate to elliptic, 4–6(–8) mm long and 2.5–4(–5) mm wide; apex rounded,
base cuneate, short petiolate; margins strongly involute in the distal 2/3 of the leaf; lamina upper surface densely
covered with short-stalked adhesive and sessile digestive glands. Inflorescences 1-flowered, erect, densely covered
with stalked-glands, 1–5(–7) scapes per plant; scapes slightly arcuate and (5–)10–25 mm long at anthesis, terete, up to
0.5 mm in diameter, straight, erect and prolonged to 30–65 mm in fruit. Calyx pentamerous, bilabiate, densely covered
with glandular hairs on the outer surface; the three upper calyx lobes divided to 2/3 of their length, triangular, 1–1.5 mm
long and 0.5–1 mm wide; the two lower calyx lobes fused to about 3/4 of their length, c. 2 mm long and 1 mm wide.
Corolla pentamerous, bilabiate, tubular, white to very pale rose, white at the base of the corolla lobes near the throat,
with yellowish-green mark on the palate at the base of the lower lip and inside the throat and spur; corolla (3–)3.5–5
mm long including the spur; upper corolla lip bilobate, smaller than the lower lip, lobes spreading, 1–1.5 × 0.5–1 mm,
rectangular to ovate-elliptic in outline, apex obtuse to shallowly retuse; lower corolla lip distinctly trilobate, lobes
subequal, 1–1.5(–2) mm × 1–1.5 mm, the lateral lobes only slightly smaller than the median lobe, ovate to circular
in outline, apex obtuse, palate sparsely covered with short, subclavate to cylindrical, multicellular hairs c. 0.05–0.1
mm long; tubular throat widely open, very short, 0.5–0.7 mm long, up to 7 mm in diameter, conical to cylindrical;
spur yellowish-green, narrowly conical to cylindrical, narrowed towards an acute or obtuse apex, 1–2.5(–3) mm long,
0.3–0.5 mm in diameter; corolla outer surface glabrous, except single scattered glandular hairs on the spur. Anthers
2, filaments falcate, white, c. 0.7 mm long and up to 0.2 mm wide, papillate, thecae subequal. Ovary subglobose,
glandular; style short, subsessile, c. 0.5 mm long, stigma bilabiate, the lobes subequal, upper lobe scale like, c. 0.1 mm
long, the lower lip much larger, c. 0.4 mm long, suborbicular, papillate, reddish pink. Capsule globose, to 1.5 mm in
diameter, subequaling the persistent calyx lobes, wall membranous, capsule longitudinally bivalvate. Seeds numerous,
light brown, truncate-obovoid, testa reticulate.
Distribution:—Pinguicula pygmaea is known from only two localities just south of Santo Domingo Tonalá (c.
1370–1400 m elevation) and one just north of Santiago Juxtlahuaca (c. 1680 m elevation), along highway 125 between
Huajuapan and Tlacotepec, in western Oaxaca state, Mexico. Although no further populations of P. pygmaea have
been discovered (or specifically searched for) so far, it is likely that the species occurs in at least some of the numerous
seemingly suitable habitats on gypsum hillsides observed in the vicinities of Tonalá and Juxtlahuaca (Fig. 2).
Conservation status:—Although very abundant at two of the three sites where it was observed, P. pygmaea
does not occur within any protected area, and it is not guaranteed to occur in other apparently suitable habitats
nearby. Moreover, easily accessible gypsum deposits are generally under potential threat from mining (IUCN Threats
Classification: 3.2). Therefore, this new species is considered Vulnerable (VU D2) according to the criteria of IUCN
PINGUICULA PYGMAEA (LENTIBULARIACEAE) Phytotaxa 292 (3) © 2017 Magnolia Press • 281
FIGURE 1. Pinguicula pygmaea. A, habit (left: specimen near the end of its growing cycle; right: specimen at the beginning of anthesis).
B, leaf. C, corolla, side view. D, corolla, face view. E, corolla, flower preparation (spur dissected from the corolla). F, multiseriate hairs
from corolla palate. G, calyx. H, seed. A, B, G, H from Rivadavia & Read 1814, E, F from Rivadavia et al. 2727, C, D from photographs
of plants in situ. Drawing by A. Fleischmann.
RIVADAVIA ET AL.
282 • Phytotaxa 292 (3) © 2017 Magnolia Press
FIGURE 2. Distribution of Pinguicula pygmaea (white stars), of the related P. crenatiloba (white boxes), and the similarly annual P.
lilacina (black dots) in southern Mexico (location data based on herbarium records and field observations). The fourth small Mexican
annual species, P. takakii, is endemic to San Luis Potosí state, which is outside the range of this map.
Etymology:—The specific epithet “pygmaea” (pygmy) denotes the minute size of this delicate annual, which
is among the smallest of all known Pinguicula species, only rivaled in its diminutive stature by the closely related P.
crenatiloba, as well as by the only distantly related, Arctic circumpolar perennial Pinguicula villosa Linnaeus (1753:
17) and dwarf specimens of the annual Pinguicula lusitanica Linnaeus (1753: 17) from Atlantic western Europe and
Habitat and ecology:—Annual therophyte and strictly gypsicolous. Pinguicula pygmaea grows on north-facing
gypsum walls and hillsides in xeric shrubland, accompanied by the poikilohydric perennial (“resurrection plant”)
Selaginella L., the perennial heterophyllous geophyte Pinguicula heterophylla Bentham (1840: 70), annuals such as
Centaurium Hill (Gentianaceae), and xerophytic and geophytic perennial plants such as species of Hechtia Klotzsch
(Bromeliaceae), Agave L. (Agavaceae), Oxalis L. (Oxalidaceae), Aristolochia L. (Aristolochiaceae), Opuntia Mill.,
Neobuxbaumia mezcalaensis (Bravo) Backeb. (Cactaceae), Fouquieria ochoterenae Miranda (Fouquieriaceae), Brahea
dulcis (Kunth) Mart. (Arecaceae), ferns, liverworts, and grasses. Pinguicula pygmaea is locally abundant at two of the
three sites studied in early and late November on north-facing hillsides, growing in open gypsum soil. At the third site,
only a small number of individuals was observed. At two of the three sites, P. pygmaea grows sympatrically with P.
heterophylla. The type location was revisited in December 2003, three weeks after the species’ initial discovery, and
almost all plants were already dead, possibly as a result of a frost event that occurred the night before, but not because
of drought. The remaining annual species in Mexico (P. lilacina, P. crenatiloba, and P. takakii) have been observed
alive and flowering well into the dry season at some locations (information from herbarium records, and from R.
Resendiz Torreblanca, pers. comm.), as late as February. This suggests that, unlike P. pygmaea, these other species
are seemingly unaffected by the light frosts of winter. It is suspected that cooler winter temperatures often allow these
annuals to persist into the mid or late dry season thanks to a combination of condensation at night and shade resulting
from the northern orientation of their habitats or from surrounding vegetation. Moreover, at least in the state of Oaxaca,
P. lilacina and P. crenatiloba appear to occur at lower elevations (250–1000 m, see other specimens examined in
Appendix 1 and Fig. 2) compared to P. pygmaea (1370–1680 m), hence will experience different climatic conditions.
However, P. lilacina, P. crenatiloba and P. takakii apparently die off as the weather heats up and soils dry out late in
the dry season, when the sun rises higher during spring (March to June), and before the rains return in late spring and
PINGUICULA PYGMAEA (LENTIBULARIACEAE) Phytotaxa 292 (3) © 2017 Magnolia Press • 283
early summer (June to July). The Sierra Madre del Sur highlands of southern Mexico (Oaxaca, Guerrero and southern
Michoacán) is rich in Pinguicula species, especially on its eastern portion in the state of Oaxaca. The high species
diversity of Pinguicula (and also other species-rich montane plant genera) observed on certain Mexican mountain
ranges has been explained by differing microclimates and especially a variety of edaphic conditions found in close
proximity, ranging from limestone and gypsum to igneous and volcanic rock, resulting in many geographically close
but fundamentally different habitat niches (Zamudio 2005, Mastretta-Yanes et al. 2015). Among the Mexican mountain
ranges, the Sierra Madre del Sur is geomorphologically the most complex, with a marked topography resulting in a
mosaic of soils and climatic gradients (Krasilnikov et al. 2011). Yet, no gypsum endemic Pinguicula species (following
the species concept of Lampard et al. 2016) was known thus far from the Sierra Madre del Sur before the discovery
of P. pygmaea—in contrast to the Sierra Madre Oriental of north-eastern Mexico (Coahuila, Nuevo Léon, Tamaulipas,
San Luis Potosí, Hidalgo, to northern Puebla and Querétaro states), likewise a Pinguicula diversity center providing
various soil types, including gypsum, that host six narrowly endemic gypsicolous species: P. debbertiana Speta &
Fuchs (1992: 375), Pinguicula gypsicola Brandegee (1911: 190), Pinguicula immaculata Zamudio & Lux (1992:
40), Pinguicula nivalis Luhrs & Lampard (2006: 4), Pinguicula rotundiflora Studnička (1985: 201) and P. takakii
(Zamudio & Lux 1992, Zamudio 2005, Lampard et al. 2016).
Taxonomic relationships:—Pinguicula pygmaea is closely related to P. crenatiloba, a delicate annual species,
with which it shares leaves with strongly involute margins (incurved up to 1 mm) in the apical 2/3 of the lamina, calyx
shape, and especially the bilabiate corolla with very short and only weakly pronounced tubular throat. However, the
former species is readily distinguished from the latter by its corolla, which has subequal lobes with an entire margin
and an obtuse to shallowly retuse apex. The corolla lobes of P. crenatiloba differ greatly between the upper and lower
lip: the lobes of the corolla upper lip are distinctly smaller, more or less rectangular in outline, with bifid apex and
acute (rarely obtuse) tips, while the lobes of the corolla lower lip are much larger, with undulate (rarely rotundate)
to emarginate apex. In contrast to the strict gypsicole P. pygmaea, the closely related P. crenatiloba, and the likewise
delicate annual P. lilacina (which is of different corolla shape, see Table 1 and Fig. 3) do not occur on gypsum soil, but
inhabit clayey soils overlying limestone or igneous rock (Casper 1966, Zamudio 2005).
The delicate habit and the corolla entire lobes and color of P. pygmaea are superficially similar to the also annual
P. takakii (see Table 1 and Fig. 3), which inhabits similar gypsum soils of the Sierra Madre Oriental in San Luís Potosí
state, north-eastern Mexico, often sympatric with P. gypsicola.
TABLE 1. Comparison of P. pygmaea, P. crenatiloba, P. lilacina, and P. takakii. Measurements are taken from examined
herbarium specimens and from literature (Ernst 1961, Casper 1966, Zamudio & Rzedowski 1986, Lampard et al. 2016).
P. pygmaea P. crenatiloba P. lilacina P. takakii
leaf obovate to elliptic, 4–8
× 2.5–5 mm; margins
ovate to obovate-oval, 5–14
× 3–7 mm; margins strongly
obovate to ovate, 15–55 ×
5–30 mm; margins slightly
spathulate to ovate, 5–
16 × 4–12 mm; margins
length of the corolla
(including the spur)
(3–)3.5–5 mm (3–)4.5–6(–7) mm 8–17 mm 6–12 mm
corolla shape bilabiate, with very
short tubular throat
that narrows into the
deeply bilabiate, generally
lacking a pronounced tubular
tubular throat well
expressed and separated
from the spur
5-merous, tubular throat
well expressed and
separated from the spur
corolla lobes subequal; lobes of
corolla upper and
lower lip entire, obtuse
to shallowly retuse
lobes of corolla upper lip
bifid, acute; lobes of lower
lip emarginate, undulate, or
subequal; lobes of both
corolla lips entire, obtuse or
subequal; lobes of
both corolla lips entire,
obtuse or retuse
habitat gypsum limestone, igneous rock, red
limestone or igneous rock gypsum
Other specimens examined (paratypes):—MEXICO. Oaxaca: Município de Tonala: Highway 125, between
Huajuapan and Tlacotepec, 1 km south of Tonalá, c.1380 m alt., 5 November 2016, F. Rivadavia, R. Resendiz
Torreblanca, J.I. Guerrero Argüelles, A. Pérez Ángeles & J. Gomez Landeros 2726 (XAL!); Highway 125, between
Huajuapan and Tlacotepec, 3 km south of Tonalá, c.1400 m alt., 5 November 2016, F. Rivadavia, R. Resendiz
Torreblanca, J.I. Guerrero Argüelles, A. Pérez Ángeles & J. Gomez Landeros 2727 (XAL!); Município de Tlacotepec:
on hills next to the Laguna Encantada, c.1680 m alt., 5 November 2016, F. Rivadavia, R. Resendiz Torreblanca, J.I.
Guerrero Argüelles, A. Pérez Ángeles & J. Gomez Landeros 2728 (XAL!).
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284 • Phytotaxa 292 (3) © 2017 Magnolia Press
FIGURE 3. Morphological comparison between Pinguicula pygmaea, P. crenatiloba, P. lilacina and P. takakii. All scale bars = 5 mm.
Photos by F. Rivadavia, except P. crenatiloba middle row and lower row right top image, P. lilacina middle and lower row (by R. Resendiz
Torreblanca) and P. crenatiloba lower row left image (by M. Welge).
We would like to thank Ruben Resendiz Torreblanca, Marlene Becerril Lopez and Adolfo Ibarra Vázquez for important
help researching Pinguicula in the field in Mexico between 2003 and 2004; Ruben Resendiz Torreblanca, José Israel
Guerrero Argüelles, Adrián Pérez Ángeles, Jonathan Gomez Landeros for important discussions about Pinguicula
species and their habitats in Mexico, as well as for their help with field research of this new species in 2016; Sergio
Zamudio Ruiz for providing valuable location information, as well as initial guidance on how and where to find
Pinguicula in Mexico; David Juárez for early discussions and additional information about this taxon; Ruben Resendiz
Torreblanca and Markus Welge for kindly providing images of cultivated plants of P. lilacina and P. crenatiloba;
curators of the herbaria IEB, MEXU and XAL are thanked for providing access to their collections; Jan Schlauer,
Yoannis Domínguez and Lorenzo Peruzzi are thanked for helpful comments on the manuscript.
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APPENDIX 1. Examined specimens of related species (P. crenatiloba and P. lilacina only considered for southern
P. crenatiloba:—MEXICO. Oaxaca: Santiago Lachiguiri, Media Loma [c. 800 m], 6 Dezember 1955, T. MacDougall
s.n. (MEXU 252915!); Santiago Lachiguiri, Tehuantepec, N de Crucero Buenavista [c. 1000 m], 25 January 1992,
V. Alvaro Campos 4293 (MEXU!); Santa María Chimalapa, S de Santa María por la vereda a la Gloria, 350 m, 14
December 1984, H. Hernández G. 691 (IEB!, MEXU!); San Felipe Usila, senda para la pista de Santiago Tlatepusco,
24 September 1990, J. Ismael Calzada 16440 (MEXU!); Guerrero: Chilpancingo, vicinity of Acahuizotla between
Chilpancingo and Acapulco, 17 October 1959, H. E. Moore Jr. 8122 (MEXU!); W de El Ocotito, camino a Jaleaca,
700 m, 24 November 1983, E. Martínez S. & F. Barrie 5736 (MEXU!); Michoacán: 25 al SW de Arteaga, 900 m, 30
November 1968, Rzedowski 26636 (MEXU!); México: Tejupilco de Hidalgo, puente sobre el Río Chilero, 1700 m, 20
October 1988, A.R. López Ferraria et al. 792 (IEB!).
P. lilacina:—MEXICO. Oaxaca: Santa María Chimalapa, E de Santa María por la vereda a Paso Venado, cerca del
entronque don la vereda al Río Piñal, 250 m, 27 October 1984, H. Hernández G. 536 (IEB!, MEXU!); por la vereda a
la cabecera del Río Escolapa, 400 m, 8 February 1986, H. Hernández G. 2067 (IEB!, MEXU!); cerca de la vereda al
Paso de la Cueva del Río del Corte, 250 m, 25 January 1986, H. Hernández G. 2026 (IEB!); Juchitán, Lázaro Cárdenas,
sobre camino a Santa María Chimalapa, 450 m, 1 March 1981, T. Wendt & A. Villalobos C. 2961 (MEXU!); Santiago
Lachiguiri, Tehuantepec, N de Crucero Buenavista, 25 January 1992, Alvaro Campos V. 4291 (MEXU!); Santiago
Lachiguiri, Media Loma, 6 December 1955, T. MacDougall s.n. (MEXU 253998!); Veracruz: Totutla, Encinal, 750
m, 14 December 1972, F. Ventura A. 7628 (MEXU!); Xalapa, Tronconal, 1100 m, 28 January 1980, P. Ventura A.
16783 (MEXU!); Xalapa, hills nearby the Languna del Castillo, 1150 m, 22 December 1989, M. Chazaro B. et al.
6062 (MEXU!); Xalapa, c. 15 km al SE de Xalapa a camino de Chavarillo, 900 m, 14 February 2016, F.Rivadavia et
al. 2707 (XAL!).
P. takakii:—MEXICO. San Luis Potosí: Minas de San Rafael y Guaxcama, 1350 m, 10 November 1965, F. Takaki
2057 (IEB!); Villa Juárez, Buenavista, 1350 m, 20 Dezember 1980, S. Zamudio Ruiz 3824 (IEB!); Villa Juaréz, 4
km from Buenavista, c.1420 m, 8 November 2003, F. Rivadavia, R. Resendiz & A. Ibarra 1806 (MEXU!); Minas de
Guascamá, 2 km al SE de Buenavista, municipio de Villa Juárez, 1400 m, 18 November 1989, S. Zamudio Ruiz 3789