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Pinguicula pygmaea (Lentibulariaceae), a new annual gypsicolous species from Oaxaca State, Mexico

  • Botanische Staatssammlung München, Munich, Germany
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
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:
*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.
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.
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
north-western Africa.
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
strongly involute
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
strongly involute
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
5-merous, cylindrical,
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
rarely rotundate
subequal; lobes of both
corolla lips entire, obtuse or
shallowly retuse
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!).
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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286 Phytotaxa 292 (3) © 2017 Magnolia Press
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
(isotype IEB!).
... 60 species. However, Shimai et al. (2021) in their large-scale phylogeny retrieved four well-supported subclades within that lineage: one comprising all Caribbean species, one covering the Mexican annuals (represented by P. lilacina and P. sharpii in their phylogeny, two taxa which however are considered conspecific by Lampard et al. 2016, Roccia et al. 2016, and Rivadavia et al. 2017, and which are even given as synonyms in Appendix S1 of Shimai et al. 2021). Another lineage covers the species with very succulent carnivorous leaves with margins involute only near the very tip or not involute at all, i.e., the relatives of P. agnata and of P. esseriana, and additionally P. gypsicola was found in that group by Shimai et al. (2021). ...
... The autonymous type section of subgenus Temnoceras by definition is the lineage that includes the type species of that subgenus, i.e., P. crenatiloba. Although that species had not been included in any phylogenetic analyses yet, it is evidently closely related to the other delicate Mexican annuals, P. lilacina, P. takakii, and P. pygmaea (Rivadavia et al. 2017). Clade VI of Shimai et al. (2021) covers these annual species, represented in their sampling by P. lilacina and P. sharpii, or respectively, by two accessions of P. lilacina if you consider both species conspecific (Lampard et al. 2016;Rivadavia et al. 2017). ...
... Although that species had not been included in any phylogenetic analyses yet, it is evidently closely related to the other delicate Mexican annuals, P. lilacina, P. takakii, and P. pygmaea (Rivadavia et al. 2017). Clade VI of Shimai et al. (2021) covers these annual species, represented in their sampling by P. lilacina and P. sharpii, or respectively, by two accessions of P. lilacina if you consider both species conspecific (Lampard et al. 2016;Rivadavia et al. 2017). Based on morphological similarities (Rivadavia et al. 2017), P. crenatiloba is likely to fall into that clade as well. ...
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In this article, the morphology-based "traditional" infrageneric classification of the genus Pinguicula is opposed to "modern", phylogeny-based concepts. Both concepts are compared and illustrated, and a new infrageneric concept of four sections for the Central American evolutionary linage of Pinguicula is proposed.
... The genus Pinguicula Linnaeus (1753a: 17) is one of the three genera in the carnivorous plant family Lentibulariaceae (Lamiales), with ca. 98 species , Rivadavia et al. 2017, 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. ...
... America is the richest continent in Pinguicula species with ca. 78 species (Ulloa-Ulloa et al. 2017, Rivadavia et al. 2017, Burelo et al. 2018, having its main center of diversification in Mexico, with ca. 49 accepted species , Rivadavia et al. 2017, Fleischmann & Roccia 2018, Burelo et al. 2018. ...
... 78 species (Ulloa-Ulloa et al. 2017, Rivadavia et al. 2017, Burelo et al. 2018, having its main center of diversification in Mexico, with ca. 49 accepted species , Rivadavia et al. 2017, Fleischmann & Roccia 2018, Burelo et al. 2018. All taxa of the genus are herbaceous, rosetteforming carnivorous plants with typical lamialean, bilabiate, tubular and spurred flowers (Fleischmann & Roccia 2018). ...
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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.
... (Orchidaceae), Cephalocereus Pfeiff. (Cactaceae), Hechtia Klotzsch (Bromeliaceae), and Pinguicula L. (Lentibulariaceae) [35,[45][46][47][48][49]. ...
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Gypsum soils occur around the world, mainly in arid regions. These harsh environments promote unusual flora with high degrees of endemism. Mexico has extensive gypsum outcrops, but their flora has been poorly studied. However, the highest species richness and endemism are expected to be concentrated in Mexico's northern dry regions. To promote the study of this flora and its conservation, we estimate how well sampled it is, quantify species richness, identify centers of endemism, and detect which gypsum outcrops lie within federal protected natural areas (PNA). We conducted exhaustive literature and herbaria reviews to generate a database of botanical records on gypsum soils. The total species and gypsophyte richness were calculated using cell grids. Centers of endemism were identified using the corrected weighted endemism index (CWE). We mapped the gypsum outcrops within PNA polygons. The most collected sites are Cuatro Ciénegas (Coahuila) and Santo Domingo Tonalá (Oaxaca), which also had the highest total species richness. Nevertheless, gypsophyte richness was higher in Cuatro Ciénegas and Nuevo León. The CWE identified seven gypsophyte centers of endemism. Mexico hosts the most diverse gypsophile flora in the world, despite having been only partially studied and collected. The regions with the highest species richness and endemism are unprotected.
... Currently, there are around 110 recognized species in the genus Pinguicula, mainly distributed in the Americas. Mexico is considered the main center of diversification with 51 accepted species (Rivadavia et al. 2017, Burelo et al. 2018, Fleischmann & Roccia 2018, Crespo et al. 2020. Some studies in Pinguicula have discussed the relevance of hybridization in species diversification in the genus (Zamudio 2001, Cieslak et al. 2005, Degtjareva et al. 2006, Shimai & Kondo 2007, hence the study of micromorphological characters could shed light on additional characters indicating hybridogenesis. ...
Pinguicula (Lentibulariaceae) includes around 110 described species, 51 of which are distributed in Mexico. Recently, P. casperi was described and its authors observed that it has intermediate morphological characters between P. oblongiloba and P. parvifolia, its putative parents. The micromorphological study of seeds has previously defined the identity of some species, and this can provide evidence of hybridization. The objectives of the present study were 1) to describe the seed micromorphology of P. casperi, P. oblongiloba and P. parvifolia and 2) to provide an economical, easy, and practical method for observing seed micromorphology using confocal laser scanning microscopy (CLSM). Seed micromorphology was considerably different between P. casperi, P. oblongiloba, and P. parvifolia. Therefore, no evidence of hybridization was found. The seeds of the three species were autofluorescent. The process of obtaining the images of the seeds with CLSM was harmless, low cost, and practical. Additionally, the images obtained were high quality and had a high resolution, making it possible to observe and compare the microstructures on the seed surfaces. CLSM is a useful, practical alternative for future taxonomic studies of the genus Pinguicula.
... It is also the state where it is possible to find most Pinguicula species (Table 1). Fifteen species have been reported, including the recently described P. pygmaea (Rivadavia et al. 2017). Also, some of these species are controversial and under taxonomic discussion, i.e., P. rectifolia and P. medusina, whereas others have not been seen since they were first collected (P. ...
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After much planning and aligning our schedules, we finally defined the dates to visit populations of various Pinguicula species in northwestern Oaxaca, Mexico. It is essential to highlight that the state of Oaxaca has the greatest biodiversity in Mexico, as regard to vascular plants, it has 10,299 species, of which 4,071 are endemic to the state (Villaseñor 2016). It is also the state where it is possible to find most Pinguicula species (Table 1). Fifteen species have been reported, including the recently described P. pygmaea (Rivadavia et al. 2017). Also, some of these species are controversial and under taxonomic discussion, i.e., P. rectifolia and P. medusina, whereas others have not been seen since they were first collected (P. greenwoodii and P. utricularioides).
... Las especies de Pinguicula, al igual que muchas de las plantas consideradas carnívoras, se desarrollan en hábitats pobres en nutrientes (Givnisht 1989). México, con 48 especies registradas hasta ahora, es el país que reúne la mayor diversidad del género y es considerado el principal centro moderno de diversificación (Cheek 1994, Zamudio 1994, Zamudio 2005, esta idea se confirma por el hecho de que las exploraciones recientes siguen revelando la existencia de especies nuevas (Rivadavia et al. 2017, Zamudio et al. 2018. ...
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Antecedentes: Como parte del proyecto Flora de Tabasco, en los años recientes se han explorado las regiones montañosas del sur del estado. Recientemente, en un cerro poco conocido en el extremo suroeste se encontró por primera vez una población de plantas del género Pinguicula. Pregunta: ¿Puede un análisis taxonómico crítico de caracteres morfológicos demostrar que la especie en cuestión no corresponde a ninguna de las entidades de Pinguicula antes descritas para México y Centroamérica? Taxón: Lentibulariaceae, Pinguicula sp. nov. Sitio de estudio: Ejido Villa de Guadalupe, municipio de Huimanguillo, Tabasco Métodos: Se realizaron varias colectas de las plantas para elaborar muestras botánicas; posteriormente se realizó un análisis crítico de los caracteres morfológicos de las plantas colectadas y se compararon con las especies conocidas del sureste de México y Centroamérica para contrastarlas. Resultados: A partir del material colectado se describe Pinguicula olmeca como una especie nueva para la ciencia. Conclusiones: La nueva especie se ubica en la sección Orcheosanthus, dentro de ésta es parecida a P. moranensis y P. zecheri, pero se diferencia de éstas por producir un solo tipo de hojas durante todo el año y por tanto no forma rosetas de invierno, por el pedúnculo y cáliz glabros y por el espolón muy largo, tan largo como el pedúnculo o ligeramente más corto. Esta especie representa el primer registro del género Pinguicula para Tabasco.
... El estudio del género Pinguicula (Lentibulariaceae) en el siglo XXI ha tenido como resultado el descubrimiento de 21 especies nuevas en todo el mundo; de éstas, ocho son de México (Zamudio 2001, Zamudio y Studnicka 2001, Zamudio y van Marm 2003, Luhrs et al. 2004, Zamudio 2005, Luhrs & Lampard 2006, Rivadavia, et al. 2017, siete de Cuba, cuatro de Italia, una de Bolivia y una de Turquía. No debe sorprendernos el hecho de que la mayoría de los taxa nuevos provengan de México, pues diversos autores ya han comentado que este país posee la mayor diversidad de especies del género en el mundo y también ha sido señalado como el principal centro moderno de diversificación del género. ...
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Four new, narrowly endemic species of Pinguicula are described and illustrated. Two are from a small area of the Sierra Madre Oriental: Pinguicula robertiana Zamudio & Hernández Rendón sp. nov. (subg. Isoloba, sect. Heterophyllum), from San Luis Potosí, is characterized by a white corolla, membranous spathulate winter leaves with rounded apex, and linear summer leaves; P. rzedowskiana Zamudio & D. Juárez sp. nov. (subg. Pinguicula, sect. Longitubus), from Querétaro, is distinctive in its red-purple corolla, spathulate membranous winter leaves, and linear summer leaves. The other two are from the western region of the country: P. casperi D. Juárez & Zamudio sp. nov. (subg. Isoloba, sect. Heterophyllum), from Jalisco and Durango, has white to slightly lilac flowers, succulent (thick) winter leaves with acuminate apex, and petiolate, ovate-elliptic summer leaves; P. michoacana Zamudio & D. Juárez, sp. nov. (subg. Pinguicula, sect. Orcheosanthus), from Michoacán, has pink to purple corollas, thick winter leaves with broadly acuminate apex, and petiolate, obovate-elliptic to suborbicular summer leaves.
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The carnivorous genus Pinguicula, which currently consists of about 74 species, is part of the Lentibulariaceae family. All members are active animal and plant organ trappers. The leaves are sticky and constitute flypaper—type traps capable of slow motions. The present review focusses on several aspects of the research activities conducted on this genus. These include the study of its evolution, carnivory, trapping and digestion mechanisms, with a special emphasis on the key role played by the mucilage, its ecology and its pharmacological and culinary utilization. This review also stresses the need for a global taxonomic revision of the genus. Finally, it aims to generate greater interest on these very fascinating plants which are in urgent need of protection.
(1) To synthesize data on the physical and phylogeographical history of the Mexican highlands, with a focus on the Trans-Mexican Volcanic Belt (TMVB), and (2) to propose approaches and analyses needed for examining the interaction of climate and volcanism.
Aim (1) To synthesize data on the physical and phylogeographical history of the Mexican highlands, with a focus on the Trans-Mexican Volcanic Belt (TMVB), and (2) to propose approaches and analyses needed for examining the interaction of climate and volcanism. Location Mexico. Methods We performed a literature and data survey of the climatic, geological and phylogeographical history of the Mexican highlands. We then assessed how the expected effects of topographic isolation, co-occurring palaeoclimatic fluctuations and volcanism can be tested against the distribution of genetic diversity of high-elevation taxa. Results The Mexican highlands present a complex biogeographical, climatic and geological history. Montane taxa have been exposed to a sky-island dynamic through climate fluctuations, allowing for long-term in situ population persistence, while also promoting recent divergence and speciation events. Volcanic activity transformed part of the Mexican highlands during the Pleistocene, mainly in the TMVB, leading to co-occurring climate and topographical changes. The TMVB highlands provide a suitable template to examine how low-latitude mountains can facilitate both the long-term persistence of biodiversity as well as allopatric and parapatric speciation driven by climatic and geological events. Main conclusions Climate fluctuations, together with recent volcanism, have driven the diversification and local persistence of biodiversity within the Mexican highlands. The climate–volcanism interaction is challenging to study; however, this can be overcome by coupling genomic data with landscape analyses that integrate the geological and climatic history of the region.
Se describen tres especies nuevas de Pinguicula (Lentibulariaceae) de México: P. barbata (Subg. Temnoceras) de Chiapas, P. emarginata (Subg. Temnoceras) de Veracruz y Puebla y P- takakii (Subg. Isoloba) de San Luis Potosí.
The internal transcribed spacer (ITS) region of 18S-26S nuclear ribosomal DNA (nrDNA) in 36 species of Pinguicula(Lentibulariaceae) in Mexico and Central America, most of which are endemic to Mexico, was sequenced. Based on the ITS results, those species of Pinguicula studied were divided into three clades. In general, these cledes do not agree with the current conventional classification proposed by various authors except for Clade I. Clade I includes only two species belonging to section Isoloba, which are widely but sparsely distributed in Mexico and Central America. Clade II included 18 species, which have been conventionally classified in the sections Heterophyllum (six species), Longitubus (three species), Orcheosanthus (eight species), and Temnoceras (one species). Clade III included 16 species, which have been conventionally placed in section Agnata (four species), Crassifolia (four species), Heterophyllum (two species), Microphyllum (three species) and Orcheosanthus (three species). Clade II is widely distributed in the Sierra Madre Occidental, the Sierra Madre del Sur to the southern part of the Sierra Madre Oriental regions and extends down south to Honduras, while the species in clade III were mostly confined to the northern part of the Sierra Madre Oriental regions. In general, the ITS results were largely consistent with biogeographic patterns, but showed disagreements with the current morphological classification.
The paper gives an analysis of the pattern of soil cover of the Sierra Madre del Sur, one of the most complex physiographic regions of Mexico. It presents the results of the study of four latitudinal traverses across the region. We show that the distribution of soils in the Sierra Madre del Sur is associated with major climatic gradients, namely by vertical bioclimatic zonality in the mountains and by the effect of mountain shadow. Altitudinal distribution of soil-bioclimatic belts is complex due to non-uniform gradients of temperature and rainfall, and varies with the configuration of the mountain range. The distribution of soils is associated with the erosion and accumulation rates both on mountain slopes and in river valleys. The abundance of poorly developed soils in (semi)arid areas was ascribed to high erosion rate rather than to low pedogenetic potential. The formation of soil mosaic at a larger scale might be ascribed to the complex net of gully erosion and to the system of seismically triggered landslides of various ages. In the valleys, the distribution of soils depends upon the dynamics of sedimentation and erosion, which eventually exposes paleosols. Red-colored clayey sediments are remains of ancient weathering and pedogenesis. Their distribution is associated mainly with the intensity of recent slope processes. The soil cover pattern of the Sierra Madre del Sur cannot be explained by simplified schemes of bioclimatic zonality. The soil ranges can be explained by the distribution of climates, lithology, complex geological history of the region, and recent geomorphological processes.
Pinguicula rotundiflora (Lentibulariaceae), a new species from southern Mexico related toP. parvifolia Robinson is described and illustrated.
A new species ofPinguicula from Mexico,P. sharpii, is described. It belongs in SubgenusIsoloba, SectionIsoloba, as defined in Casper’s monograph of the genusPinguicula (1966a). The somatic chromosome number ofPinguicula sharpii is2n = 16, which is diploid. The basic chromosome numberx = 8 is found in many members of SubgenusIsoloba ofPinguicula, while that ofx = 11 is also found in some members of the subgenus.