Pinguicula filifolia subsp. alba (Lentibulariaceae), a new subspecies with an extremely restricted distribution in Pinar del Río, Cuba

Abstract

The carnivorous plant Pinguicula filifolia occurs in western Cuba and shows a restricted distribution due to habitat specificity associated with poor wet soils. Its populations are distributed mainly in south Pinar del Río (Cuba) and in Isla de la Juventud. Plants from one isolated location in NW Pinar del Río shows morphological differences with respect to the rest of the known populations observed in nature. Subsequent detailed morphological analysis of specimens led to the description of Pinguicula filifolia subsp. alba as a new subspecies. It differs from the typical subspecies in several reproductive traits involving flower, fruit and seed characters and is so far known only from the type locality.

Full text

Accepted by Lorenzo Peruzzi: 24 Dec. 2013; published: 30 Jan. 2014 85
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 158 (1): 085–092
www.mapress.com/phytotaxa/Article
http://dx.doi.org/10.11646/phytotaxa.158.1.6
Pinguicula filifolia subsp. alba (Lentibulariaceae), a new subspecies with an
extremely restricted distribution in Pinar del Río, Cuba
YOANNIS DOMÍNGUEZ1,2*, CRISTINA M. PANFET VALDÉS3 & VITOR F.O. DE MIRANDA4
1Universidade de Mogi das Cruzes, Herbarium Mogiense – Núcleo de Ciências Ambientais, Av. Dr. Cândido Xavier de Almeida Souza
n. 200, CEP 08780-911 Mogi das Cruzes, SP, Brazil; e-mail: yoannisd@gmail.com
2Universidad Agraria de La Habana, Laboratorio de Botánica, Departamento de Biología y Sanidad Vegetal, Autopista Nacional Km.
23., San José de las Lajas CP 32700, Mayabeque,
Cuba
3Universidad de La Habana, Jardín Botánico Nacional de Cuba, Carretera del Rocío Km. 3., Calabazar, Boyeros CP 19230, La
Habana, Cuba
4UNESP – Univ Estadual Paulista, Faculdade de Ciências Agrárias e Veterinárias, Departamento de Biologia Aplicada à
Agropecuária, Labratório de Sistemática Vegetal e Palinologia, Via de Acesso Prof. Paulo Donato Castellane s/n, CEP 14884-900
Jaboticabal, SP, Brazil
*Author for correspondence.
Abstract
The carnivorous plant Pinguicula filifolia occurs in western Cuba and shows a restricted distribution due to habitat
specificity associated with poor wet soils. Its populations are distributed mainly in south Pinar del Río (Cuba) and in Isla
de la Juventud. Plants from one isolated location in NW Pinar del Río shows morphological differences with respect to
the rest of the known populations observed in nature. Subsequent detailed morphological analysis of specimens led to the
description of Pinguicula filifolia subsp. alba as a new subspecies. It differs from the typical subspecies in several
reproductive traits involving flower, fruit and seed characters and is so far known only from the type locality.
Keywords: morphology, taxonomy, western Cuba
Introduction
The carnivorous genus Pinguicula Linnaeus (1753: 17) comprises more than 100 recognized species (Rodondi et
al. 2010, Yildirim et al. 2012, Peruzzi & Gestri, 2013) which are mainly distributed throughout the northern
hemisphere and in tropical and subtropical America (Legendre 2000). At present 13 Pinguicula species are known
in Cuba (Domínguez 2012), all of which are endemic and show a very restricted distribution associated with poor
wet soils in the westernmost, central and easternmost parts of Cuban archipelago. Pinguicula filifolia C.Wright ex
Grisebach (1866: 162) is the species with the widest distribution in western Cuba; it occurs on Pinar del Río (island
of Cuba) and in Isla de la Juventud (Isle of Youth), the second largest island of the archipelago.
Studies on Cuban Pinguicula have been done through traditional morphological analyses (Ernst 1961, Casper
1966, Casper 2003, 2004, 2007, Casper & Urquiola 2003, Domínguez et al. 2012, Domínguez et al. 2013), as well
as using molecular techniques (Shimai et al. 2007). Some of these studies have revealed a large morphological
variability in P. filifolia consistent with morphological variations observed during botanical explorations in western
Cuba. Such intraspecific variability can be found among plants occurring in south Pinar del Río and in Isla de la
Juventud; meanwhile, plants from a location toward the extreme of species distribution at the Ecological Reserve
“Los Pretiles” in Pinar del Río showed exclusive morphological features not shared with other populations. After
analysis and comparison of the plants collected in nature with a large number of specimens available in herbarium
collections as well as with type specimens of P. filifolia, plants from “Los Pretiles” (NW Pinar del Río) are here
described as a new subspecies of P. filifolia.

DOMÍNGUEZ ET AL.86 • Phytotaxa 158 (1) © 2014 Magnolia Press
Materials and Methods
Living material from natural populations of P. filifolia, herbarium specimens housed at HAJB, HAC (series LS,
PFC, Roig, SV; Regalado et al. 2008), HFA, HPPR and HUMC as well as digital images of types from GOET, MO,
NY and US (see also Appendix 1) were analyzed and compared with taxonomic literature on Cuban Pinguicula
(Grisebach 1866, Britton 1920, Alain 1957, Ernst 1961, Casper 1966, 2003, 2004, 2007, Casper & Urquiola 2003).
Morphological analyses of specimens were carried out using a stereo microscope, Motic SMZ-168 and
measurements were taken using the software Motic Image Plus 2.0ML (Motic China Group Co., Ltd.). Seed samples
were coated with gold, then analyzed and photographed at 15 kV using a scanning electron microscope (SEM)
JEOL JSM 5410. Qualitative characters were recorded on living plants from sampled populations.
Results
According to the morphological differences among examined specimens, P. filifolia was segregated into two
subspecies: P. filifolia subsp. filifolia C.Wright ex Griseb. and P. filifolia subsp. alba Y.Domínguez, Panfet &
V.Miranda subsp. nov (Fig. 1). Both subspecies share common morphological features, mainly those regarding
vegetative characters, such as linear-filiform erect leaves; although, they differ in several quantitative and
qualitative traits. P. filifolia subsp. alba differs from the typical subspecies in the number and length of leaves; it
also has up to ~13 inflorescences per plant which are larger than in subsp. filifolia. The new subspecies shows only
white flowers with oblong-spathulate lobes, a shorter and wider conical spur that forms an angle of 99°–110° with
the tube and smaller fruits (Table 1). Seeds are smaller and exotesta cells show no crests like those observed in the
typical subspecies (Fig. 2). P. filifolia subsp. alba is so far known only from the type locality in northwestern Pinar
del Río province in Cuba (Fig. 3).
Discussion
Morphological analyses allowed the identification of several characteristics for the circumscription of subspecific
taxa within P. filifolia (Table 1). Differences in vegetative features include the number and length of leaves which
are more numerous and larger in subsp. alba. Our results showed that floral characters are important features in the
identification of the two subspecies, this is in agreement with previous studies that have utilized such features for
species and subspecies discrimination in European Pinguicula (Conti & Peruzzi 2006, Peruzzi & Gestri 2013). In
particular, a larger flower with oblong-spathulate white lobes, a conical spur and the narrowest angle (99°–110°)
between spur and corolla tube marks a clear distinction of the new subspecies; whereas in subsp. filifolia flowers
are smaller, corolla lobes vary in shape and color, the spur is saccate and the angle is always wider (115°–140°).
Characters of calyx are somehow controversial in Cuban Pinguicula (Casper 2003, 2004, 2007); nevertheless,
Casper has used some of these features to segregate species from Pinguicula benedicta Barnh. in Britton (1920:
110). Complementing Casper’s observations in the P. benedicta-complex, both subspecies of P. filifolia show
exclusive different fusion patterns of calyx lower-lip lobes (Table 1).
The capsule in subsp. alba is globose and smaller than in subsp. filifolia. Seeds show the morphological pattern
previously described by Casper (1966) and Dwyer (1983), although some differences can be noted for each
subspecies. In subsp. alba, seeds are 339–509 µm length with a shorter micropylar appendage and the anticlinal
walls of exotesta cells are thickened, almost flat; whereas, in subsp. filifolia the seeds are larger and show
prominent thickened anticlinal walls like crests at transversal margins (Fig. 2) as described for other Pinguicula
species (Degtjareva et al. 2004). All of these characteristics together allow a clear morphological circumscription
of P. filifolia subsp. alba against the typical subspecies.
Apart from the morphological differences identified between the two subspecies, each one has different
distribution ranges in western Cuba. Subspecies filifolia can be found in the south of the Pinar del Río province
(Island of Cuba) and on Isla de la Juventud (Isle of Youth); on the other hand, subspecies alba occurs exclusively in
one locality in NW Pinar del Río (Fig. 3). Such distribution patterns could be related to the divergence of
populations at the extremes of the species’ distributional gradient. The current flora of western Pinar del Río and

Phytotaxa 158 (1) © 2014 Magnolia Press • 87
Pinguicula filifolia subsp. alba, a new subspecies
Isla de la Juventud comprises several plant families (e.g. Cyperaceae, Droseraceae, Eriocaulaceae,
Lentibulariaceae and Xyridaceae) associated with poor, wet soils that, at present, are restricted to small patches
(Urquiola et al. 2010) and show disjunct distribution patterns due to vicariance (Capote et al. 1988). Such
discontinuity may have led to intraspecific differentiation among geographically isolated populations as observed
in other species from this biogeographical region (Borhidi 1996, Urquiola et al. 2010).
TABLE 1. Morphological differences between the two subspecies of P. filifolia based on measurements from living
plants (subsp. filifolia: N = 52; subsp. alba: N = 120) and herbarium specimens (see also Appendix 1). Values are
presented as minimum, first quartile, third quartile and maximum, (min–)Q1–Q3(–max).
Taxonomic treatment
Pinguicula filifolia subsp. alba Y.Domínguez, Panfet & V.Miranda, subsp. nov. (Fig. 1)
Differs from P. filifolia subsp. filifolia in the larger number and length of leaves and inflorescences; white flowers with oblong-
spathulate lobes; conical spur, angle between spur and corolla tube of 99°–110°, calyx lower-lip lobes connate (0.5–0.8
mm); capsule globose; seeds smaller with anticlinal exotesta walls thickened and almost flat.
Typ e : —CUBA. Pinar del Río: Mantua, Playa “La Cana”, Reserva Ecológica “Los Pretiles”, pinares sobre arenas blancas, 4 m.
a.s.l., 22°26'21''N, 84°17'58''W, 4 January 2012, Y. Domínguez, C. Panfet & R. Novo 313 (holotype HAJB–PFC 87590!;
isotypes HUMC 5321!, HFA!, JABU!).
Characters subsp. filifolia subsp. alba
Leaf
Number 6–11(–13) 18–23(–30)
Length 7–8.5(–9.5) mm 9.5–10.5 mm
Width 0.1–0.15(–0.2) mm 0.2–0.3 mm
Inflorescence
Number 1–2(–3) (3–)6–8(–13)
Length (10–)12–13.5(–14) cm 15–18 cm
Corolla
Lobes shape Obovate, suborbiculate or oblong Oblong-spathulate
Lobes color Blue, violet, pale purple, rarely white White
Tube color Violet striated blue, pale purple striated violet
or greenish striated purple
Yellow striated brown
Spur
Shape, color and size Saccate, apex acute-rounded Conic, apex rounded-truncate
Color Blue, violet, purple or greenish Yellow or yellow-brownish
Size (5.5–)6–6.8 × 2.2–2.5(–2.8) mm 3.5–4(–4.5) × 3–3.5 mm
Calyx lower-lip Sepals 0.9–1 mm joined from the base Sepals 0.5–0.8 mm joined from the base
Angle spur/tube (115°–)120°–130°(–140°) 99°–106°(–110°)
Fruit Sub-globose, 2.4–2.8 × 2.5–3 mm Globose, 1.7–2 × 1.7–2 mm
Seed
Length (544–)579–595(–655) µm (339–)379–450(–509) µm
Appendage length (33–)58–65(–82) µm (30–)37–47(–56) µm
Anticlinal wall of exotesta cells Pronounced at transversal margins forming
crests
Not pronounced, without crests

DOMÍNGUEZ ET AL.88 • Phytotaxa 158 (1) © 2014 Magnolia Press
FIGURE 1. Pinguicula filifolia subsp. alba. A, habit; B–C, frontal and lateral view of the flower; D, corolla hairs from: i, lobes base;
ii, tube; iii, spur; E, fruit; F, seed. P. filifolia subsp. filifolia: G, lateral view of the flower; H, fruit; I, corolla hairs from: i, lobes base; ii
tube; iii, spur. (Drawings by Y. Domínguez; subsp. alba from the isotype at HUMC and subsp. filifolia from HUMC 5005).

Phytotaxa 158 (1) © 2014 Magnolia Press • 89
Pinguicula filifolia subsp. alba, a new subspecies
FIGURE 2. SEM microphotographs of seed and details of seed surface; A–B, P. filifolia subsp. alba; C–D, P. filifolia subsp. filifolia.
Annual rosulate herb. Rhizome short, with numerous adventitious roots. Leaves 18–23(–30), 9.5–10.5 × 0.2–0.3
mm, linear-filiform, erect, green-yellowish, margin entire, plane, apex acute, circinate, base cuneate, adaxial
surface moderately covered with stalked and sessile glands, abaxial surface glabrous. Plantlets leaves 4–8, 4–5.5 ×
1.5 mm, flat to suberect, spathulate, base cuneate, apex rounded to obtuse, margin entire, planar. Inflorescence (3–
)6–8(–13), erect, 15–18 cm tall, 1 mm in diameter, 1-flowered, sparsely covered with stalked and sessile glands.
Flower (12.5–)13.5–15(–18) mm long (spur included). Calyx 2-lipped, sepals oblong, apex rounded, green-
yellowish, sparsely covered with stalked glands; upper-lip sepals 3, each 1.5–2 × 1 mm, entirely divided; lower-lip
sepals 2, each 1–1.5 × 0.5 mm, connate 0.5–0.8 mm from the base. Corolla subisolobate, lobes 3–4.3(–5) × (2.5–
)3–3.7(–4) mm, white, oblong-spathulate, apex rounded to truncate; tube (3.5–)4–4.5(–5) × 1.8–2.5(–3.2) mm,
cylindric, outside yellow with brown stripes, inside yellow, covered with long-stalked multicellular-headed hairs;
spur 3.5–4(–4.5) × 3–3.5 mm, conic, apex rounded-truncate, yellow-brownish, forming an angle of 99°–106°(–
110°) with the tube. Capsule 1.7–2 × 1.7–2 mm, globose, ± half covered by the persistent calyx. Seeds (339–)379–
450(–509) µm × (117–)169–190(–229) µm, elliptic-oblong, dark brown; exotesta cells ± rectangular-polygonal,
arranged in longitudinal-spiral rows, anticlinal walls thickened, not pronounced; micropylar appendage (30–)37–
47(–56) µm (Fig. 2A–B).
Etymology:—The subspecific epithet refers to the invariable white color of the corolla.
Distribution and habitat:—Endemic to western Cuba (Fig. 3). It occurs in quartz sand planes (savannas) and
open areas of Pinus forests at 0–10 m a.s.l., in the Ecological Reserve “Los Pretiles”, province Pinar del Río.
Conservation status:—The subspecies is proposed as Critically Endangered (CR) according to IUCN (2013)
criteria B1ab(i,ii,iii,iv)+2ab(i,ii,iii,iv) due to its restricted geographic and spatial distribution within the area of
occupancy (~10 km2).

DOMÍNGUEZ ET AL.90 • Phytotaxa 158 (1) © 2014 Magnolia Press
Other specimens examined (paratypes):—CUBA. Pinar del Río: Guane, Dimas, Playa La Cana cerca de
Camarones, arenas blancas con pinos, September 1975, A. Areces et al. (HAC–PFC 40254!); Mantua, Dimas,
Playa La Cana cerca de Camarones, arena blanca con Pinus, October 1975, A. Areces et al. (HAJB–PFC 28567!);
Mantua, Playa la Cana SO de Dimas sobre arena blanca, 12 July 1978, J. Bisse et al. (HAJB–PFC 38731!);
Mantua, Playa La Cana, al oeste del Arroyo Camarones arena silícea, veg. Pinar, 1–2 m., 5 December 1979, J.Bisse
et al. (HAJB–PFC 41285!); Mantua, Playa La Cana, al este del Arroyo Camarones, arena silícea, pinar, 1–2 m., 5
December 1979, J. Bisse et al. (HAJB–PFC 41355!); Mantua, Playa La Cana, parte occidental del Pinar, 2 m., 29
March 1982, J. Bisse et al. (HAJB–PFC 46602!); Mantua, Isla de “Los Pretiles”, Reserva Ecológica “Los Pretiles”,
Pinares sobre arenas blancas, 5 m. a.s.l., 22°25'40''N, 84°18'32''W, 4 January 2012, Y. Domínguez et al. 313
(HAJB–PFC 87592!, HFA!, HUMC 5322!).
FIGURE 3. Distribution of P. filifolia subsp. alba (star) and P. filifolia subsp. filifolia (circle) in western Cuba based on herbarium
records.
Identification key to the subspecies of P. filifolia
1. Corolla white, angle between spur and corolla tube 99°–110°, spur conic 3.5–4.5 × 3–3.5 mm, seeds 339–509 µm long,
without crests.......................................................................................................................................................... subsp. alba
- Corolla blue, violet or purple (rarely white), angle between spur and corolla tube 115°–140°, spur saccate 5.5–6.8 × 2.2–
2.8 mm, seeds 544–655 µm long, with crests ...................................................................................................subsp. filifolia
Acknowledgements
We would like to sincerely thank Roberto Novo, the staff of the Ecological Reserve “Los Pretiles” and Pinar del
Río Botanic Garden for granting permissions in Pinar del Río, Yosbel Lazo and Daniel Ryed for granting
permissions and for fieldwork assistance in Isla de la Juventud, the staffs of herbaria HAJB, HAC and HPPR.
Thanks are also due to the staff of Laboratory of Plant Systematics and Palynology (LSVP)—UNESP for helpful

Phytotaxa 158 (1) © 2014 Magnolia Press • 91
Pinguicula filifolia subsp. alba, a new subspecies
discussions and suggestions and to the anonymous referees for helpful comments. This paper is part of Y.
Domínguez PhD project supported by CAPES (grant. No. 23038.001163/2010-68), and UNESP/PROPe, Univ
Estadual Paulista, Brazil.
References
Alain, Hno. (1957) Flora de Cuba IV. Dicotiledóneas: Melastomataceae a Plantaginaceae. Contribuciones Ocasionales del
Museo de Historia Natural del Colegio “De La Salle” 16, La Habana, 556 pp.
Borhidi, A. (1996) Phytogeography and vegetation ecology of Cuba. 2nd ed. Akadémiai Kiadó, Budapest, 752 pp.
Britton, N.L. (1920) Description of Cuban plants new to science. [Pinguicula benedicta Barnhart, J.H. p. 110]. Memoirs of the
Torrey Botanical Club 16(2): 7–118.
Capote, R.P., Berazaín, R. & Leiva, A. (1988) Cuba. In: Campbell, D.G. & Hammond, H.D. (eds.) Floristic inventories of
tropical countries: the status of plant systematic, collections, and vegetation, plus recommendations for the future. The
New York Botanical Garden, New York, pp. 315–335.
Casper, S.J. & Urquiola, A.J. (2003) Pinguicula cubensis (Lentibulariaceae), a new insectivorous species from western Cuba
(Cuba occidental). Willdenowia 33: 167–172.
Casper, S.J. (1966) Monographie der gattung Pinguicula L. Bibliotheca Botanica 127: 1–209.
Casper, S.J. (2003) Two new Pinguicula species (Lentibulariaceae) from east Cuba (Cuba oriental). Haussknechtia 9: 141–155.
Casper, S.J. (2004) Two new Pinguicula species (Lentibulariaceae; P. benedicta-group) from the eastern mountain range of
Cuba (Greater Antilles) with reddish flowers. Wulfenia 11: 1–13.
Casper, S.J. (2007) Pinguicula lippoldii nova spec. and Pinguicula toldensis nova spec. – two endemic Pinguicula species
(Lentibulariaceae) from East Cuba new to science. Wulfenia 14: 75–96.
Conti, F. & Peruzzi, L. (2006) Pinguicula (Lentibulariaceae) in central Italy: taxonomic study. Annales Botanici Fennici 43:
321–337.
Degtjareva, G., Casper, J.S., Hellwig, F. & Sokoloff, D. (2004) Seed morphology in the genus Pinguicula (Lentibulariaceae)
and its relation to taxonomy and phylogeny. Botanische Jahrbücher für Systematik 125: 431–452.
http://dx.doi.org/10.1127/0006-8152/2004/0125-0431
Domínguez, Y. (2012) Alla ricerca delle pinguicole di Cuba occidentale / Searching for West Cuban Pinguicula. AIPC
Magazine 27: 10–35.
Domínguez, Y., Panfet, C.M. & Miranda, V.F.O. (2012) New features on Cuban endemic Pinguicula filifolia (Lentibulariaceae)
and considerations on its habitat and ecology. Flora 207: 838–841.
http://dx.doi.org/10.1016/j.flora.2012.09.009
Domínguez, Y., Silva, S.R., Panfet, C.M. & Miranda, V.F.O. (2013) Inter- and intra-specific diversity of Cuban Pinguicula
(Lentibulariaceae) based on morphometric analyses and its relation with geographic distribution. Plant Ecology &
Diversity
http://dx.doi.org/10.1080/17550874.2013.842616
Dwyer, T.P. (1983) Seed structure of carnivorous plants. Carnivorous Plant Newsletter 12: 8–23.
Ernst, A. (1961) Revision der Gattung Pinguicula. Botanische Jahrbücher für Systematik 80: 145–194.
Grisebach, A. (1866) Catalogus plantarum cubensium exhibens collectionem Wrightianam aliasque minores ex insula Cuba
missas. Apud Guilielmum Engelmann, Lipsiae, 301 pp.
http://dx.doi.org/10.5962/bhl.title.177
IUCN. (2013) Guidelines for Using the IUCN Red List Categories and Criteria, version 10. Prepared by the Standards and
Petitions Subcommittee, 87 pp.
Legendre, L. (2000) The genus Pinguicula L. (Lentibulariaceae): an overview. Acta Botanica Gallica 147: 77–95.
Linnaeus, C. (1753) Species Plantarum. Impensis Laurentii Salvii, Stockholm, 1200 pp.
Peruzzi, L. & Gestri, G. (2013) A new butterwort species (Pinguicula, Lentibulariaceae) from Northern Apennine (Italy). Plant
Biosystems 147: 692–703.
http://dx.doi.org/10.1080/11263504.2012.756073
Regalado, L., Ventosa, I. & Morejón, R. (2008) Revisión histórica de los herbarios de Cuba con énfasis en las series de
especímenes. Revista del Jardín Botánico Nacional 29: 101–138.
Rodondi, G., Beretta, M. & Andreis, C. (2010) Pollen morphology of alpine butterworts (Pinguicula L. Lentibulariaceae).
Review of Paleobotany and Palynology 162: 1–10.
Shimai, H., Masuda, Y., Panfet, C.M. & Kondo, K. (2007) Phylogenetic analysis of Cuban Pinguicula (Lentibulariaceae) based
on internal transcribed spacer (ITS) region. Chromosome Botany 2: 151–158.
http://dx.doi.org/10.3199/iscb.2.151
Urquiola, A.J., González-Oliva, L., Novo, R. & Acosta, Z. (2010) Libro rojo de la flora vascular de la provincia Pinar del Río.
Publicaciones Universidad de Alicante, Alicante, 457 pp.
Yildirim, H., Şenol, S.G. & Pirhan, A.F. (2012) Pinguicula habilii (Lentibulariaceae), a new carnivorous species from South-
West Anatolia, Turkey. Phytotaxa 64: 46–58.

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Appendix 1
Examined specimens of Pinguicula filifolia subsp. filifolia:—CUBA. Cuba occ., 1860–1865, C. Wright 560=2886
(type GOET 004568!; isotypes HAC–LS!, MO 1764199!, NY 00111445!, US 00048178!); Pinar del Río: Jovero, San
Julian, Guanes, January 1938, Acuña 10720 (HAC–SV!); Sabanas del Sur de Pinar del Río, Km. 13 de la carretera de la
Coloma, 29 September 1955, Alain 2588 (HAC–LS!); Pine barren., Hacienda San Julian al Sur de Guane, 26–27
December 1916, León & Roca 6915 (HAC–LS!); Pinar en el Km. 11 de la carretera de la Coloma, 5 January 1940, León
17450 (HAC–LS!); Laguna de la Máquina, Km. 13 de la carretera de la Coloma, 28 November 1940, León & Alain
19493 (HAC–LS!); Pinar del Río, Km. 13 carretera de “La Coloma”, zanjas, 25 November 1926, Hitch et al. 4076
(HAC–Roig!); Pinar del Río, Puerta de Golpe, terrenos bajos, 31 January 1915, Van Herman & Muñoz 1234 (HAC–
Roig!); Pinar del Río, Sabanas, Km. 13 carretera de la Coloma, 29 September 1952, Alain 2653 (HAC–LS!); Pinar del
Río, Jovero San Julián, January 1938, J. Acuña 10720 (HAC–SV!); Pinar del Río, Km. 10 carretera La Coloma, zanjas,
25 November 1926, Hitch et al. 12917 (HAC–SV!); Pinar del Río, Pinares próximos Lag. Sta. María, San Luis,
Moldenke et al. 15303 (HAC–SV!); Guanes, laguna Sta María, sabana arenosa, 21 January 1967, J. Bisse & L. Rojas
(HAJB–PFC 1349!); Guane, Laguna Alcatraz Chico, sabana arenosa, 23 January 1967, J. Bisse & L. Rojas (HAJB–PFC
1401!); Las Ovas, sabana de arena blanca y lagunas, 26 February 1972, J. Bisse (HAJB–PFC 21624!); Guanes: El sabalo,
pinar de Sabana La Mar, arena blanca, October 1975, A. Areces et al. (HAJB–PFC 28374!); Las Ovas, terreno de arena
blanca al sur-oeste del pueblo, December 1975, A. Areces et al. (HAJB–PFC 29098!); Guane, cercanias de la Laguna
Cordero arena silicea, 4 December 1919, J. Bisse et al. (HAJB–PFC 41226!); Ovas, zona de desague de la presa El
Punto, vegetación secundaria, 17 December 1982, J. Bisse et al. (HAJB–PFC 48521!); San Luis, Sta María, potreros
cerca del terraplén a San Luis, 19 December 1982, J. Bisse et al. (HAJB–PFC 48613!); Guane, Laguna al norte de la
presa El Punto, 19 October 1983, A. Álvarez et al. (HAJB–PFC 50911!); El Pitirre, en el terraplen, desde hermanos
barcón hasta la carretera La Coloma, 19 October 1983, A. Álvarez et al. (HAJB–PFC 50974!); Ovas, alrededores de la
presa El Punto, 22 October 1988, L. González (HAJB–PFC 66381!); Guane, Santa Teresa, 13 March 2005, A. Urquiola
& H. Zeiger (HAJB–PFC 83214!); En suelo húmedo de laguna de la presa El Punto, 22 June 1988, E. Köhler et al.
(HPPR 4668!); Desague Presa “El Punto”, Las Ovas, 20 May 1982, J. A.Moreno (HPPR 143!); Laguna y cañada de la
presa El Punto, Ovas, 29 April 1989, A. Urquiola et al. (HPPR 5317!); Herradura, 05 August 1926, Van Herman 716
(HAC–SV!); Isla de la Juventud: Playa Roja, Bahía de Siguanea, Sabanas along road to Playa, 6 March 1953, E. P.
Killip 43002 (HAC–LS!); Siguanea region, White-sand sabanas, 18 March 1954, E. P. Killip 43705 (HAC–LS!);
Between Mina de Oro and Playa del Soldado; white-sand sabanas, 12 January 1956, E. P. Killip 45398 (HAC–LS!); Near
Nueva Gerona, W.L., 19 December 1903, A.H. Curtis 243 (HAC–SV!); Santa Bárbara, white-sand sabanas, 27
December 1951, Alain & E. P. Killip 2150 (HAC–LS!); Santa Fé, 19–22 June 1952, J. Acuña 23934 (HAC–SV!);
Between San Francisco de las Piedras and Cerro La Cañada, Moist sandy soil, 27 January 1955, E. P. Killip 44608
(HAC–LS!); Río Júcaro region, White sand sabanas, 27 February 1953, E. P. Killip 42914 (HAC–LS!); ibidem E. P.
Killip 42915 (HAC–LS!); Along road to San Francisco de las Piedras, gray-sand sabanas, 21 January 1955, E. P. Killip
44577 (HAC–LS!); San Pedro region, White-sand sabanas, 18 March 1954, E. P. Killip 43685 (HAC–LS!); Santa
Bárbara, sabanas 3–4 km. South of town, Marshy ground, 13 February 1953, E. P. Killip 42718 (HAC–LS!); ibidem, E.
P. Killip 42719 (HAC–LS!); Along road to San Francisco de las Piedras, Wet bank of stream above pond, 17 January
1966, E. P. Killip 45438 (HAC–LS!); Sabana de Los Indios, 7 February 1940, León & W.Seifriz 17487 (HAC–LS!);
Sabana arenosa 3 Km al este de Siguanea, 25 December 1966, J. Bisse (HAJB–PFC 829!); Sabana arenosa de Los Indios,
February 1969, J. Bisse & H. Lippold (HAJB–PFC 12716!); Sabana arenosa cerca del hotel El Colony, February 1969, J.
Bisse & H. Lippold (HAJB–PFC 12939!); Arenales silíceos a 2 Km Noroeste del Hotel Colony, 28 October 1976, A.
Areces et al. (HAJB–PFC 32989!); La Esperanza, cerca de la masa semillera de Pinus, en arenales silíceos, 23 October
1976, A. Areces et al. (HAJB–PFC 33065!); Las Delicias, Zona de arenas blancas, Mina de Oro, veg. Pinar abierto arena
silícea, 6 April 1980, A. Álvarez et al. (HAJB–PFC 42024!); 1 Km al NO Hotel Colony, pinares destruidos, 6 November
1981, A. Álvarez et al. (HAJB–PFC 45825!); Las Delicias, entre la Planta eléctrica y la costa, pinares sobre arenas
blancas, 7 November 1981, A. Álvarez et al. (HAJB–PFC 45942!); Camino de Cocodrilo hacia sabanas de Capé, suelo
turba, 16 November 1982, R. Berazaín et al. (HAJB–PFC 48397!); Cercanías del Hotel Colony a unos 2 Km, pinar
degradado, suelo arenoso, 15 November 1982, R. Berazaín et al. (HAJB–PFC 48408!); Pinares de Los Indios, 10 m. 19
October 1991, A. Urquiola et al. (HPPR 7517!); Reserva Biológica Los Indios (Indio Centro), Pinares sobre arenas
blancas 21°41'35''N, 82°59'40''W, 29 July 2009, Y. Domínguez et al. 97 (HAJB–PFC!, HFA!); same locality, 16
December 2009, Y. Domínguez et al. 118 (HAJB–PFC!, HFA!); Sabana noroeste del Hotel Colony, Pinares sobre arenas
blancas, 21°38'14''N, 82°58'32''W, 30 July 2009, Y. Domínguez et al. 100 (HAJB–PFC!, HFA!, HUMC 5005!); ibidem,
15 December 2009, Y. Domínguez et al. 103 (HAJB–PFC!, HFA!).