Elucidating the controversial Drosera montana complex (Droseraceae): A taxonomic revision

Abstract

The species of the affinity of Drosera montana (Droseraceae) are reviewed taxonomically and the complex is redefined to include only D. montana, D. tentaculata, D. tomentosa var. tomentosa, D. tomentosa var. glabrata, and D. spirocalyx. The latter is a newly described narrow endemic species from the Serra do Cipó in central Minas Gerais state, Brazil. The mor- phological characters distinguishing these five taxa from each other and from other similar species are discussed together with habitat and ecological information. Detailed illustrations, photographs, distribution maps and an identification key are provided. A lectotype for D. tomentosa is here designated.

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Phytotaxa 172 (3): 141–175
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Copyright © 2014 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Duilio Iamonico: 7 May 2014; published: 18 Jun. 2014
http://dx.doi.org/10.11646/phytotaxa.172.3.1
141
Elucidating the controversial Drosera montana complex (Droseraceae): a taxonomic
revision
FERNANDO RIVADAVIA1, PAULO MINATEL GONELLA2*, PAULO TAKEO SANO3 & ANDREAS FLEISCHMANN4
1 Daniel Burnham Ct., San Francisco, 94109, USA; e-mail: fe_riva@uol.com.br
2 Programa de Pós-Graduação em Botânica, Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências,
Universidade de São Paulo, CEP 05508–900, São Paulo, Brasil; e-mail: paulogonella@uol.com.br
3 Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, CEP 05508–900,
São Paulo, Brasil; e-mail: ptsano@usp.br
4 Botanische Staatssammlung München, Menzinger Strasse 67, D-80638 Munich, Germany; e-mail: fleischmann@lrz.uni-muenchen.de
* author for correspondence
Abstract
The species of the affinity of Drosera montana (Droseraceae) are reviewed taxonomically and the complex is redefined to
include only D. montana, D. tentaculata, D. tomentosa var. tomentosa, D. tomentosa var. glabrata, and D. spirocalyx. The
latter is a newly described narrow endemic species from the Serra do Cipó in central Minas Gerais state, Brazil. The mor-
phological characters distinguishing these five taxa from each other and from other similar species are discussed together
with habitat and ecological information. Detailed illustrations, photographs, distribution maps and an identification key are
provided. A lectotype for D. tomentosa is here designated.
Keywords: Brazil, carnivorous plants, Drosera tentaculata, Drosera tomentosa, Drosera tomentosa var. glabrata, Drosera
spirocalyx, ecology, morphology, new species, lectotypification, taxonomy
Resumo
As espécies do complexo Drosera montana (Droseraceae) são revistas taxonomicamente e o complexo é redefinido, pas-
sando a incluir apenas D. montana, D. tentaculata, D. tomentosa var. tomentosa, D. tomentosa var. glabrata e D. spirocalyx.
A última é uma espécie aqui descrita pela primeira vez, sendo endêmica da Serra do Cipó, região central de Minas Gerais,
Brasil. Os caracteres morfológicos distinguindo estes cinco táxons entre si e de outras espécies similares são discutidos junto
a informações sobre habitat e ecologia. Ilustrações detalhadas, fotografias, mapas de distribuição e uma chave são apresen-
tados. Um lectótipo de D. tomentosa é aqui designado.
Palavras-chave: Brasil, Drosera tentaculata, Drosera tomentosa, Drosera tomentosa var. glabrata, Drosera spirocalyx,
ecologia, morfologia, novas especies, lectotipificação, plantas carnívoras, taxonomia
Introduction
The species complex of the Drosera montana Saint-Hilaire (1826: 260) affinity currently comprises a confusing list of
16 names proposed over nearly 200 years of taxonomic research on the genus Drosera in Brazil. This was initiated by
the French naturalist Auguste de Saint-Hilaire (1824, 1826) who described the first Brazilian Drosera taxa. Since then,
most botanists have applied a rather expansive taxonomic circumscription of D. montana, resulting in a classification
that synonymized under this taxon several previously segregated species (e.g. Diels 1906, Correa & Silva 2005). The
history of taxa associated with the D. montana complex is summarized below:
• Saint-Hilaire (1826) described D. montana, D. parvifolia Saint-Hilaire (1826: 263), D. tomentosa Saint-Hilaire
(1826: 261) var. tomentosa, D. tomentosa var. glabrata Saint-Hilaire (1826: 262), D. hirtella Saint-Hilaire (1826:
262) var. hirtella, and D. hirtella var. lutescens Saint-Hilaire (1826: 263), together with six other Drosera taxa of
different affinities (see Gonella et al. 2012, 2014).

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• Eichler (1872), when reviewing the genus Drosera for Martius’ Flora Brasiliensis, placed D. tomentosa var.
glabrata and D. hirtella var. lutescens under synonymy of D. tomentosa and D. hirtella, respectively. Furthermore,
he described D. communis var. pauciflora Eichler in Martius & Eichler (1872: 394).
• Chodat (1903) described D. montana forma parviflora Chodat (1903: 539) based on material from Paraguay
(Sierra de Maracayu).
• Diels (1906) considered Saint-Hilaire’s D. parvifolia a synonym of D. montana, while D. tomentosa and D.
hirtella were recombined at the rank of varieties of D. montana. In the same generic revision, D. montana var.
schwackei Diels (1906: 89) and D. montana var. roraimae Klotzsch ex Diels (1906: 90) were described.
• Diels (1914) published D. montana var. robusta Diels (1914: 136).
• Maguire & Laundon (1957) raised Diels’ D. montana var. roraimae to species rank, D. roraimae (Klotzsch ex
Diels) Maguire & Laundon (1957: 333), and placed D. montana var. robusta under its synonymy.
• Santos (1986) published D. pumilla E.Santos (1986: 305), alleged to be closely related to D. brevifolia Pursh
(1813: 211), differing by the presence of stipules and by scapes with long eglandular trichomes.
• Santos (1989), returning in part to the views of Saint-Hilaire (1826), accepted D. hirtella var. hirtella, D. hirtella
var. lutescens, and D. tomentosa, but not D. tomentosa var. glabrata. Strangely, no mention was made of Diels’ D.
montana var. schwackei in her treatment.
• Silva (1994) fully adopted the views of Diels (1906), accepting D. montana var. montana, D. montana var.
hirtella (A.St.-Hil.) Diels (1906: 89), D. montana var. roraimae, D. montana var. schwackei, and D. montana var.
tomentosa (A.St.-Hil.) Diels (1906: 89). Furthermore, D. pumilla was considered a dubious taxon with affinities
to D. montana var. schwackei.
• Silva & Giulietti (1997) considered D. roraimae at species status, following Maguire & Laundon (1957). Although
the authors strangely omitted almost all mention of synonyms under each taxon, it is here assumed that D. pumilla
was considered conspecific with D. montana var. schwackei, since collections of the former from Mato Grosso
were included under the latter taxon.
• Rivadavia (2003) published four new Drosera species, including one belonging to the D. montana complex: D.
tentaculata Rivadavia (2003: 79).
• Correa & Silva (2005) imprudently synonymized ten names under D. montana, considering them all conspecific
in their taxonomic treatment: D. communis var. pauciflora, D. hirtella, D. hirtella var. lutescens, D. montana var.
schwackei, D. parvifolia, D. tomentosa, D. tomentosa var. glabrata, D. pumilla, D. montana var. hirtella, and D.
montana var. tomentosa, but no reference was made to D. tentaculata.
• Silva (2007), in her floristic treatment of Droseraceae for the Flora of Bahia State, considered D. tentaculata as a
synonym of D. montana.
• Rivadavia (2008) elevated D. montana var. schwackei to species rank, D. schwackei (Diels) Rivadavia (2008: 39),
and also highlighted the distinctness of the four species D. hirtella, D. montana, D. tentaculata, and D. tomentosa,
adopting Saint-Hilaire’s original species concept.
• Ferrero & Mello-Silva (2011), in their treatment of Droseraceae for the Flora of Ibitipoca State Park, recognized
D. montana and D. tomentosa var. glabrata as distinct taxa.
• Rivadavia & Gonella (2011) and Gonella et al. (2012, 2014) followed the species concepts of Saint-Hilaire (1826)
and Rivadavia (2008), accepting D. hirtella, D. montana, D. schwackei, D. tentaculata, and D. tomentosa as
distinct species.
• Silva (2013), in the online “Lista da Flora do Brasil”, followed the controversial species concept of D. montana
proposed by Correa & Silva (2005), and also considered D. schwackei a synonym, but does not say under which
species (presumably to D. montana).
The goals of our study were to present a comprehensive revision of the D. montana complex, including:
I. Identify and circumscribe D. montana, as described by Saint-Hilaire (1826), in order to enable a critical and
comparative species delimitation of closely allied taxa;
II. Identify the taxa allied to D. montana based on morphological evidence, as well as cytological and phylogenetic data
(from literature), in order to redefine the complex in the view of a modern, comprehensive taxonomic revision;
III. Evaluate the distribution range of each taxon;
IV. Present a complete botanical description of all taxa recognized as belonging to this complex, including the
description of eventual new species.

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 143
Materials & Methods
Extensive field studies were undertaken by the first two authors between 1990 and 2014, studying hundreds of different
Drosera populations throughout South America, and especially eastern Brazil, including all type locations of taxa
involved with the D. montana complex—with the exception of the type location of D. montana, the Serra do Papagaio,
although live plants and pictures of D. montana from this location were examined (as well as the holotype material).
Herbarium specimens at ALCB, B, BHCB, CEN, CESJ, DIAM, ESA, F, GFJP, HB, HRB, HUEFS, M, MBM, MO,
NY, OUPR, P, PAMG, R, RB, RFA, SP, SPF, UB, UEC, and US were personally examined and annotated by at least
one of the authors (herbarium acronyms according to Thiers 2011). Type materials held at B, P, R, and SPF were
personally examined under a stereo dissecting microscope. High-resolution digital images of the type materials held at
K (http://apps.kew.org/herbcat/navigator.do), MPU and P (http://hvsh.cria.org.br/) were also examined. Descriptions
are based on fresh, dried and spirit material.
Distribution maps were prepared using georeferenced location data obtained from herbarium records (extrapolated
or approximated for a few old or vaguely specified localities) or our own field observations, and created with ArcMap
(ESRI 2011), based on the spatial cartographic data provided by the Brazilian Institute of Geography and Statistics
(IBGE: http://www.ibge.gov.br).
Under each species in the “Specimens Examined” section, states and departments are cited in geographical order,
from North to South.
The non-carnivorous indumentum present on leaves and scapes of Drosera often provides valuable taxonomic
characters for species delimitation. In the genus, this indumentum can be classified either as trichomes (non-vascularized;
present or absent on scapes, petioles and lamina abaxial surface) or as carnivorous emergences (vascularized; Seine
& Barthlott 1993), the latter often wrongly referred to as “glandular hairs” or “glands”. To avoid confusion, the
carnivorous, glandular vascularized emergences present on the margins and adaxial surface of the lamina of Drosera
leaves are here referred to as “tentacles”. Among the species treated below, three types of trichomes are found as
indumentum on the scapes, petioles and abaxial surface of the leaves, and the following terminology is employed
(adapted from Seine & Barthlott 1993):
− “eglandulartrichomes”:simple,non-glandulartrichomes;
− “glandulartrichomes”:glandulartrichomeswithconspicuousstalk,usuallylongerthan0.2mminlength.
− “sessileglands”:shortglandulartrichomeswithinconspicuousstalk,thegland-headusually0.03–0.05mmindiameter;
Results
Most of the taxonomic confusion surrounding the taxa historically allied to the D. montana complex originated from the
excessive weight placed on inflorescence indumentum by Diels (1906) in his delimitation of D. montana, which included
four varieties. Other important morphological characters, such as leaf shape, were mostly ignored by Diels (1906) and by
most of the subsequent works (e.g. Correa & Silva 2005, Silva 1994, 2007, 2013), in favor of scape indumentum. This left
few options for proper morphological identification of non-flowering specimens within this complex.
Herbarium specimens of small rosetted Drosera species are often badly preserved and may thus look very similar
superficially. The rosettes are usually pressed into dense columns, frequently requiring preparations of single leaves
(i.e. destructive examination of herbarium specimens) for careful study. Diels (1906) never had the opportunity to
examine Brazilian Drosera populations in situ, and thus based his taxonomic opinions on scarce evidence acquired
from the study of a few herborized specimens. In live plants of the D. montana complex, where foliar characters such
as leaf shape, orientation and coloration are clearly evident, we find it surprisingly easy to identify taxa even from
sterile rosette material alone. Sadly however, Diels’ (1906) narrow taxonomic circumscription for this group was
adopted and propagated over the past century, helped by the fact that ecological information has been mostly inexistent
or overlooked for the numerous Brazilian Drosera taxa.
The lumping of Brazilian Drosera taxa based exclusively on scape indumentum reached its apex with the treatment
of Droseraceae by Correa & Silva (2005) for the Flora Neotropica, where a single, generalized argument—that “the
indumentum of the scapes is very variable” (Correa & Silva 2005, page 48)—was used by the authors as the entire
foundation to justify the synonymization of ten names under D. montana (see above). Not only were the differences in
scape indumentum shape and distribution, as described by Saint-Hilaire (1826) and others, totally disregarded, but so
were all other vegetative characters distinguishing the multiple taxa therein.

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Furthermore, this synonymization contradicted the available phylogenetic data (Rivadavia et al. 2003), cytological
data (Rivadavia 2005), as well as morphological and ecological data (Santos 1989, Rivadavia 2003). All these published
data support the separation of the D. montana complex into several distinct taxa, and also that D. montana s.l. (sensu
Correa & Silva 2005) does not represent a monophyletic lineage. Moreover, in their floristic treatment, Correa & Silva
(2005) entirely overlooked four newly described species (Rivadavia 2003), one of them (D. tentaculata) belonging to
the taxonomic group of the D. montana complex.
A close examination of the type material of Saint-Hilaire’s D. parvifolia at P, and of live specimens at the locus
classicus near São João del Rey in the Serra de São José, Minas Gerais, showed that this taxon is closely allied to
D. communis Saint-Hilaire (1824: t. XV[1]), due to characters such as leaves that are distinctly petiolate-spatulate,
semi-erect and with geniculate-involute vernation; the papery, broadly triangular to rectangular stipules, divided into
several laciniae from the insertion point; a curved, ascending base of the scape; narrowly elliptic calyx lobes; and a
fusiform seed shape. The eglandular trichomes found near the base of the scape of D. parvifolia represent a previously
undescribed character for the D. communis complex. Nevertheless, it is here considered part of the wide natural
variation found within this group, and has been observed by the authors at several natural populations of D. communis
in central-southeastern Minas Gerais.
The type material of Chodat’s D. montana f. parviflora was also examined at P, and this taxon is considered
conspecific with D. communis, based on the spatulate leaves with distinctly narrow petioles, and a scape that is
arctuately curved from the base.
The holotype of D. pumilla was examined at R, and is here considered a member of the species affinity of
D. cayennensis Sagot ex Diels (1906: 86), due to characters such as short-petioled cuneate-spatulate leaves with
geniculate-involute vernation, short stiff scapes, large lanceolate calyx lobes, and foveolate testa ornamentation. Until
further studies can resolve the wide variability of overlapping characters observed, D. colombiana Fernández (1965:
226), D. panamensis Correa & Taylor (1977: 390), D. pumilla, and D. sanariapoana Steyermark (1952: 243) are
here considered conspecific with, and hence synonyms of D. cayennensis, based on the examination of the types and
other herbarium material, as well as of live plants in situ in Brazil (Mato Grosso, Goiás, Distrito Federal, Tocantins,
Roraima, Pará) and in Venezuela (Bolívar, Amazonas).
In accordance with Rivadavia (2008), D. schwackei is excluded from the D. montana complex, based on
morphological characteristics discerned from the study of herbarium specimens and during field studies of this species
in Minas Gerais. Drosera schwackei shares morphological characters with other relatively large Brazilian Drosera
of the tetraploid clade (Rivadavia et al. 2003), such as D. graminifolia Saint-Hilaire (1826: 269) and D. quartzicola
Rivadavia & Gonella (2011: 34) (Rivadavia 2008, Rivadavia & Gonella 2011, Gonella et al. 2012). These include the
semi-erect leaves which are yellow-green in color and oblong-lanceolate in shape, the presence of translucent-yellow
short-stalked globose trichomes, relatively thick and slightly succulent erect scapes with an overall woolly indumentum
of long patent hairs, and comparatively large oblong-ovate seeds with reticulate testa (see Rivadavia 2008).
A detailed examination of the holotypes of D. hirtella and D. hirtella var. lutescens at P revealed that these clearly
represent two distinct taxa belonging to a separate and well-defined group closely allied to the D. communis complex,
due to characters such as spatulate to cuneate-spatulate leaves of geniculate-involute vernation pattern, and the unique
eglandular hirsute indumentum of the scape. Field studies at numerous sites in the states of Bahia, Minas Gerais, São
Paulo, Mato Grosso do Sul, Mato Grosso, Goiás, Distrito Federal, and Tocantins, revealed that these two taxa are easily
distinguishable from each other and from all taxa of the D. montana complex (even in sterile specimens), with which they
sometimes occur sympatrically. Drosera hirtella var. lutescens has even been observed to form a newly reported natural
hybrid with D. communis at a few locations, supporting a closer relationship between these diploid taxa, but not with D.
montana, D. tentaculata, D. tomentosa, nor any other member of the Brazilian tetraploid clade (Rivadavia et al. 2003).
Leaf ontogeny and shape alone suggest a split among the numerous taxa historically allied to D. montana, where
two main types of leaf vernantion from bud are observed:
1. Geniculate-involute vernation: in budding leaves the lamina is initially folded over the petiole, and the lateral
margins of the lamina are rolled inwards. This character state is found in D. cayennensis, D. communis, D.
hirtella, and D. roraimae (including their synonyms mentioned above). The leaves of these taxa are also spatulate
to cuneate-spatulate in shape, with distinct petioles that are longer than or equalling the lamina in length.
2. Circinate vernation: in budding leaves the lamina and petiole are convolutely curled inwards in a circinate fashion
similar to fern fronds. Found in D. montana and the remaining taxa of this complex with the exception of D.
tentaculata (see discussion under the respective species description). The leaf shapes of these taxa vary between
oblong and obovate (rarely short-petioled spatulate), with petioles that are often indistinct or much shorter than
the lamina (lamina about twice as long as the petiole, or longer).

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 145
Furthermore, the flower scapes are arcuately curved from the base (ascending) in D. hirtella var. hirtella, D. hirtella
var. lutescens, D. communis, and most specimens of D. roraimae, supporting a closer affinity between these taxa.
This character is absent from the core D. montana taxa, all of which have scapes that are erect from the base, and is
only found consistently in one taxon of the Brazilian tetraploid clade, D. ascendens and, to a lesser degree, in some
specimens of D. graomogolensis Silva (1997: 85), D. quartzicola and D. riparia Rivadavia & Gonella (in Gonella et
al. 2014: 31).
Our classification of the D. montana complex is additionally supported by cytological evidence, which shows
that D. cayennensis (including plants from the locus classicus of D. pumilla), D. communis, D. hirtella var. hirtella,
D. hirtella var. lutescens, D. roraimae, and D. viridis Rivadavia (2003: 87), all with geniculate-involute vernation, are
diploids with a chromosome number of 2n = 2x = 20 (Rivadavia 2005). In contrast, D. tentaculata and D. tomentosa
were found to be tetraploids with 2n = 4x = 40, together with several other taxa of the so-called “Brazilian tetraploid
clade” (Rivadavia et al. 2003): D. camporupestris Rivadavia (2003: 85), D. chrysolepis Taubert (1893: 505), D.
grantsaui Rivadavia (2003: 82), D. graomogolensis, D. latifolia (Eichler in Martius & Eichler 1872: 395) Gonella &
Rivadavia (in Gonella et al. 2014: 21), D. spiralis, and D. villosa Saint-Hilaire (1826: 267) (Rivadavia 2005) — all but
D. tentaculata with circinate vernation.
These two morphological and cytological groups are further supported phylogenetically by plastid DNA
sequence data (rbcL; Rivadavia et al. 2003), which shows them to belong to two consecutive sister clades. One clade
comprises tetraploid (and supposed tetraploid) Brazilian Drosera, including D. montana and D. tomentosa, as well
as D. chrysolepis, D. graminifolia, D. graomogolensis, D. latifolia, D. schwackei, and D. villosa (all with circinate
leaf vernation). Not yet examined in phylogenetic context, but assumedly also belonging to this clade based on
morphological and cytological evidence, are D. ascendens Saint-Hilaire (1826: 268), D. camporupestris, D. chimaera
Gonella & Rivadavia (in Gonella et al. 2014: 34), D. grantsaui, D. quartzicola, D. riparia, D. spirocalyx, and D.
tentaculata.
The above tetraploid phylogenetic clade is sister to another clade comprised by predominantly New World diploid
(and supposed diploid) taxa including D. hirtella, D. brevifolia, D. capillaris Poiret (1804: 299), D. felix Steyermark
& Smith (1974: 491), D. kaieteurensis Brummer-Dinger (1955: 137), and the type species of the genus, D. rotundifolia
Linnaeus (1753: 281) (Rivadavia et al. 2003), most of which bear distinctly petiolate, spatulate leaves with geniculate-
involute vernation. This diploid clade probably includes all North American Drosera taxa and most South American
taxa, but excludes all species in the above-mentioned tetraploid clade, as well as two species which are not directly
related (see Rivadavia et al. 2003, 2012) to either of the two clades above: D. sessilifolia Saint-Hilaire (1826: 259) and
D. meristocaulis Maguire & Wurdack (1957: 332).
In light of all the recent evidence cited above, the D. montana complex is here circumscribed as a monophyletic
group of tetraploid (2n = 40) Drosera species native to South America, geographically centered at the Cadeia do
Espinhaço highlands in the states of Minas Gerais and Bahia (eastern Brazil), possessing scapes that are erect from the
base, have circinate leaf vernation (except D. tentaculata, see discussion below), rosettes of decumbent leaves more
or less obovate in shape, and petioles much shorter than the lamina. Members of this complex are D. montana, D.
tomentosa, D. tentaculata, and D. spirocalyx.
The D. montana complex as defined here is assumed to be of very close affinity to the D. villosa complex (Gonella
et al. 2014) and possibly also to D. grantsaui, with which D. tomentosa is known to form natural hybrids (Rivadavia
2009). This larger group of taxa, in turn, appears to be most closely related to a natural affinity of robust Brazilian
species, including D. camporupestris, D. chrysolepis, D. graminifolia, D. quartzicola, D. schwackei, and D. spiralis.
Regarding the indumentum of carnivorous tentacles, the apical margin of the lamina in all species of the D.
montana complex bears bilaterally symmetric tentacles from a dilated base, each with an unifacial gland head. These
marginal tentacles are usually capable of relatively fast, notable movement upon mechanical stimulation, and are often
casually referred to as “snap tentacles” (Hartmeyer & Hartmeyer 2006). In D. tentaculata, these marginal tentacles
are conspicuously distinct, each with a prominent stalk and long head. In the remaining species however, the apical
tentacles are usually hardly distinguishable from the numerous radially symmetrical tentacles on the lamina surface,
especially in dried specimens. Within the clade of Brazilian tetraploid sundew species, the presence of snap tentacles is
limited to the species of the D. montana complex. Snap tentacles are more often observed in Drosera with decumbent
leaves due to a possible specialization for capturing crawling prey (Hartmeyer & Hartmeyer 2006), which may explain
their absence in the remaining large Brazilian tetraploid species, all of which have semi-erect leaves.
An unusual herbarium collection [Cordeiro et al. CFSC 7533 (SPF)] belonging to the D. montana complex
was discovered and examined in the early 1990’s at SPF. Lacking precise location data, it sparked extensive (but
unsuccessful) explorations of the Serra do Cipó by one of the authors (FR) over the following years. While looking

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through photographs of multiple Drosera taxa taken at the Serra do Cipó in the late 1990’s by Japanese botanist Ms.
Chiaki Shibata, one of the authors (FR) identified plants that resembled the herbarium specimen mentioned above.
Fortunately, precise location data was recorded for the photographs, making it possible to finally rediscover this
elusive plant in 1999. Since then, in situ studies at this and other newly discovered sites, have allowed a more complete
understanding of this taxon, which is described as a new species, D. spirocalyx Rivadavia & Gonella, in the following
treatment.
Following the rather broad taxonomic circumscription presented by Correa & Silva (2005), almost all central-
southern Brazilian Drosera species with decumbent leaves can be keyed out as D. montana. This fact led to a great
number of herbarium specimens of small rosetted South American Drosera species being casually identified as “D.
montana”. In order to facilitate the identification of specimens from this taxonomically difficult group, we provide the
following key to the Drosera species with decumbent leaves from Brazil south of the Equator.
Key to the Drosera species with decumbent leaves from Brazil south of the Equator (with extensions in South
America south of the Equator given in brackets)
1. Gynoecium 5-merous, styles 5, undivided; leaves with glandular trichomes only (in addition to tentacles and sessile glands) [also
in Bolivia] ......................................................................................................................................................................D. sessilifolia
- Gynoecium 3-merous, styles 3, bifurcated at the base (6 stigmata in total); leaves glabrous, or with eglandular trichomes only, or
with both glandular and eglandular trichomes (in addition to tentacles and sessile glands) ..............................................................2
2. Stipules absent or reduced to few fimbriae isolated on the margins of the leaf base [also in Argentina and Uruguay] D. brevifolia
- Stipules well developed, conspicuous, intrapetiolar ...........................................................................................................................3
3. Lamina with apex narrower or of similar width than the base (oblong, lanceolate, or ellipsoid in outline) .....................................4
- Lamina with apex distinctly broader than the base (obovate, oblanceolate, spatulate, cuneate, or sub-orbicular in outline) ..........5
4. Leavesoblong,petioleusually≤0.5mmwide(rarelyupto1mmwide);glandulartrichomesdenselycoveringthescapesfromthe
base; seeds ovoid to ellipsoid [also in Bolivia] ................................................................................................................D. montana
- Leavesoblong-lanceolatetolanceolate,usuallysemi-erect,petiole≥1mmwide;glandulartrichomesabsentonthebasalthirdof
the scape; seeds fusiform .................................................................................................................................................... D. latifolia
5. Scape erect at the base .......................................................................................................................................................................6
- Scape arcuatelly curved at the base ..................................................................................................................................................13
6. Petiole narrow, suddenly broadening into lamina; petiole longer or equal in length with the lamina; leaf vernation geniculate-
involute ...............................................................................................................................................................................................7
- Petiole gradually broadening into lamina; petiole length equal to or shorter than half lamina length; leaf vernation geniculate or
circinate .............................................................................................................................................................................................9
7. Sepals glabrous; seeds with papillose testa [also in Uruguay] ........................................................................................ D. capillaris
- Sepals glandular- and/or eglandular-pilose; seeds with foveolate or reticulate testa .........................................................................8
8. Scapes glabrous; sepals exclusively glandular-pilose; seeds rectangular with reticulate testa .............................................D. viridis
- Scapes pilose; sepals exclusively eglandular-pilose, or both glandular- and eglandular-pilose; seeds ovoid with foveolate testa [also
in Bolivia and Peru] ..................................................................................................................................................... D. cayennensis
9. Leaf obovate-cuneate to cuneate with geniculate vernation; apical tentacles conspicuously distinct, in number of 7–12, 4–9 mm
long, with head with unifacial gland tip ........................................................................................................................ D. tentaculata
- Leaf obovate, oblanceolate, or oblong-spatulate with circinate vernation; apical tentacles inconspicuously distinct, only up to 5
mm long ............................................................................................................................................................................................10
10. Adaxial petiole surface densely eglandular-pilose; sepals ovate to sub-orbicular with rounded (subobtuse) apex, in bud forming a
pyramidal projection at the fusion point between adjacent sepals .................................................................................D. spirocalyx
- Adaxial petiole surface glabrous or sparsely eglandular-pilose; sepals oblong-ovate to narrowly-ovate with acute apex, not forming
pyramidal projections as described above ........................................................................................................................................11
11. Leaves oblong to oblong-spatulate; petiole 0.2–0.5(–1) mm wide; glandular trichomes of similar length present throughout scape;
generally flowering from December to March (rainy season) [also in Bolivia] ...............................................................D. montana
- Leaves obovate to oblong-obovate; petiole 0.4–3.0 mm wide; glandular trichomes absent on the basal third of scape, but dense and
larger towards the apex; generally flowering from June to September (dry season) ..............................................D. tomentosa (12)
12. Scapes densely eglandular-pilose (villous) on the basal half, decreasing in density towards apex .......D. tomentosa var. tomentosa
- Scapes exclusively glandular-pilose from base to apex, without notable eglandular indumentum, occasionally also sparsely
eglandular-pilose mostly towards the basal third ......................................................................................D. tomentosa var. glabrata
13. Basal third of scape densely eglandular-pilose............................................................................................................ D. hirtella (14)
- Basal third of scape glabrous to glabrescent ....................................................................................................................................15
14. Lamina obovate; scape green to red; eglandular trichomes of scape red ........................................................ D. hirtella var. hirtella
- Lamina spatulate; scape yellow; eglandular trichomes of scape white [also in Bolivia] ............................. D. hirtella var. lutescens
15. Leaves decumbent, entirely green or with pinkish-red petiole and green lamina (even if growing in sunny habitats); seeds
rectangular .............................................................................................................................................................................D. viridis
- Leaves usually semi-erect, red (greener only in shaded habitats); seeds fusiform [also in Bolivia, Paraguay, and Argentina] ..........
.........................................................................................................................................................................................D. communis

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Taxonomic Treatment
Drosera montana Saint-Hilaire (1826: 260) (Figs. 1–4)
Type:—BRAZIL. Minas Gerais: Inverni in jugis altioribus montium dictorum Serra do Papagayo in parte australi provinciae Minas
Geraes, March, without date, Saint-Hilaire D-542 (holotype P!; isotypes P!, K(plant on the top right) image!, MPU image!).
= Drosera communis A.St.-Hil. var. pauciflora Eichler in Martius & Eichler (1872: 394)
Type:—BRAZIL. Minas Gerais: Santa Bárbara, Serra do Caraça, January 1825, Riedel 1463 (holotype B!; isotypes P!, K image!).
Description:—Perennial rosetted herbs, winter-dormant, acaulescent, rosettes lax, few-leaved, never forming
columns of accumulated dead leaves. Roots slightly succulent, sparsely branched, black, densely covered with root
hairs. Indumentum consists of white eglandular trichomes (pale brown when dried) on the abaxial leaf surface, more
rarely on the adaxial petiole surface and base of the scape; capitate glandular trichomes on scape, bracts, pedicels,
and sepals; and minute sessile glands ca. 0.03 mm in diameter on leaves and inflorescence parts. Leaves 4–17 mm
long, with circinate vernation, decumbent, oblong, oblong-spatulate or more rarely spatulate, apex obtuse, wine to
deep red, greenish to dull green in shadier habitats; petiole 1–7 mm long, 0.2–0.5(–1) mm wide, abaxial surface
sparse to densely eglandular-pilose, adaxial surface glabrous to sparsely eglandular-pilose; lamina 3–11 mm long,
1.5–4(–4.5) mm wide, abaxial surface glabrous to sparsely eglandular-pilose, the trichomes concentrated at the base
and along the lateral margins of the lamina, adaxial surface covered with numerous red, motile, capitate tentacles with
radial symmetry, the apical tentacles (and often the lateral marginal tentacles) inconspicuously distinct, with bilaterally
symmetrical, ellipsoid to oblong heads with unifacial gland tip; stipules 1–3 mm long, 0.4–1.3 mm wide at the base,
rectangular to triangular, laciniate, membranaceous, translucent white to reddish in color (drying bronze colored), the
apical 1/3–2/3 divided into several laciniae, or into 3 segments, the lateral ones longer, narrower, and poorly divided,
the central shorter and broader with fimbriate apex. Scapes 1–2(–3), erect from the base, 30–320(–420) mm long
(including floriferous part), terete, green to deep red in color, glandular trichomes present from the base, somewhat
uniform in size and increasing in density towards apex, occasionally sparsely eglandular-pilose on the basal third;
inflorescence a scorpioid cyme bearing 1–7(–10) flowers, never bifurcating; bracts caducous, usually absent on mature
scapes, 0.5–3 mm long, lanceolate to linear, densely glandular-pilose abaxially; pedicels 1–7(–9) mm long, inserted
(5–)6–20 mm apart, densely glandular-pilose; sepals 5, 2–6 mm long, 0.8–2 mm wide, ovate to oblong-ovate, united
at the basal 1/4–1/3 of length, apex acute, densely glandular-pilose abaxially; petals 5, 5–7 mm long, 2–3 mm wide,
oblong-obovate to broadly obovate, light to dark pink in color, or rarely bicolored with nearly white base, becoming
pink towards the apex; stamens 5, 1.2–3.5 mm long, filaments white to light pink, anthers 0.5–1 mm long, bithecate,
yellow; ovary 3-carpellate, fused, 0.9–1.8 mm in diameter, globose to ovoid, trilobed or hexalobed in outline; styles
3, forked at the base, 1.5–3 mm long (including stigmata), pale pink in color, style arms curving inwards at the end
of anthesis, stigmata flabellate to cordiform, or bifid; fruit a dry dehiscent capsule, 2–3 mm long, ellipsoid, 3-valvate;
seeds 0.3–0.6 mm long, 0.15–0.20 mm wide, ovoid to ellipsoid, black, testa reticulate.
Illustrations:—Silva (1994: 75, fig. 13A–C, E1, E2, H, I1, J–N, O1); Silva & Giulietti (1997: 93, fig. 9A–C, F,
G, L, O, Q–U); Correa & Silva (2005: 45, fig. 26D–F); Silva (2008: 395, fig. 1H–J); Ferrero & Mello-Silva (2011: 17,
fig. 2D–F).
Distribution and ecology:—Drosera montana is one of the most widespread Drosera taxa in South America
(together with D. cayennensis, D. communis, D. grantsaui, and D. sessilifolia) and it is found disjunctly in Brazil and
Bolivia.
In Brazil, it is known from the states of Bahia, Goiás, Distrito Federal, Minas Gerais, Rio de Janeiro, São Paulo,
Paraná, and Santa Catarina (Fig. 1). It is probably also present in Espírito Santo, since it is abundant on the Minas
Gerais (western) side of the Pico da Bandeira, which lies on the border between these two states. No records were seen
for the Brazilian states of Rondônia, Mato Grosso, and Mato Grosso do Sul, geographically intermediate between the
Brazilian and Bolivian distributions.
Drosera montana is also found along a somewhat continuous stretch of nearly 1000 km of the eastern slopes of
the Bolivian Andes, in south, central and western Bolivia, in the Departments of La Paz, Santa Cruz, Chuquisaca and
Tarija (Fig. 2). No records were found from the Department of Cochabamba, intermediate between the northern and
southern ranges of the species in Bolivia. Considering how close some collections were made to the borders with Peru
and Argentina, it is possible that D. montana is also native to these two countries.

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FIGURE 1. Distribution map of D. montana in Brazil.
FIGURE 2. Distribution map of D. montana in Bolivia (departments: BNI = Beni, CHQ = Chuquisaca, CBB = Cochabamba, LPZ = La
Paz, ORU = Oruro, PND = Pando, PSI = Potosi, SCZ = Santa Cruz, TJA = Tarija).

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 149
FIGURE 3. Drosera montana. A, habit. B, C, detail of the indumentum of the base (B) and apex (C) of the scape. D, E, stipule. F–J, leaf,
abaxial surface (F, J) and adaxial surface (G, H). K, side view of the calyx. L, sepal. M, N, petal. O, stamen. P, gynoecium. Q, R, seed. A-C,
E, J, K based on Gonella et al. 361; D, H based on Rivadavia & Batista 2630; F, G based on Gonella et al. 94; L based on Rivadavia 432;
M–P based on photographs of living material; Q based on Rivadavia & Mullins 543; R based on Rivadavia 639. Drawn by P.M. Gonella.
A previous report of D. montana for Paraguay (Duno de Stefano & Silva 2001) could not be verified as no specific
references are cited. It is here assumed that this record was based on Chodat’s type of D. montana f. parviflora [Hassler
5271 (P)], which is here considered conspecific with D. communis (see Results). A previous record for Peru (Brako &
Zaruchi 1993) was verified to be based on a misidentified specimen of D. cayennensis [Gentry & Núñez 69485 (MO)],
a species that is here first recorded for that country.
Drosera montana is most often observed in sandy locations or on thin layers of soil over sandstone, but it has a
relatively broad ecological amplitude and can be found in a variety of different open habitats and nutrient-poor soil
types in montane to sub-alpine regions at elevations between 800–2500 m in Brazil, and between (1240–)1800–2310
m in Bolivia. In Brazil it is often seen growing as a ruderal species in disturbed areas such as trail sides, banks along
roads, and other open, freshly exposed, or eroded habitats. These are generally drier than those preferred by other
native Brazilian Drosera taxa. Drosera montana is also one of the rare Brazilian sundew species that regularly occurs
in reddish soils (Fig. 4B, C) or in white clayey substrates (Fig. 4A).

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A few Drosera species have been observed growing sympatric with D. montana, including D. brevifolia, D.
communis, D. hirtella var. hirtella, D. latifolia, D. villosa, as well as D. tomentosa and D. spirocalyx. Several other
taxa have been observed in neighboring habitats: D. ascendens, D. camporupestris, D. cayennensis, D. chrysolepis, D.
grantsaui, D. hirtella var. lutescens, D. spiralis, D. tentaculata, and D. viridis.
In Brazil, D. montana, D. hirtella, and especially D. cayennensis are very well adapted to seasonally wet habitats,
where they regularly enter a dormancy state in the drier months of the year, during which the plants completely
dissapear from the soil surface, growing back from the roots once the rains return months later.
Phenology:—Drosera montana is a seasonal geophyte, usually without vegetative parts present above the soil
during the dry season (which extends from approximately May to November in the species’ distribution in Brazil and
Bolivia), surviving as dormant tap roots—although in rare perennially humid habitats D. montana can produce leaves
throughout the year. The flowering period is concentrated in the wet season, from December to March in Brazil, and
from January to May in Bolivia, with a few individuals occasionally flowering until early dry season (June) in both
countries. Similar to almost all other Drosera taxa, the species of the D. montana complex produce flowers that open
for only one day, beginning early in the morning on sunny days and usually closing around midday.
IUCN Red List Category:—Drosera montana is widespread and common, occurring inside numerous protected
areas, sometimes even growing as a ruderal species in disturbed areas, and is thus considered here of Least Concern
(LC), according to the IUCN criteria (IUCN 2001).
Discussion:—The seasonal geophyte habit of D. montana immediately distinguishes it from other members of
the complex, an ecological adaptation that also results in an earlier flowering period (see Table 1). It can further be
identified by its relatively lax, delicate, few-leaved rosettes with oblong, oblong-spatulate or more rarely spatulate
leaves, the presence of uniformly-sized glandular trichomes from base to apex of the scapes, and by the long and broad
ovate to oblong-ovate sepals (2–6 × 0.8–2 mm). In contrast, the four perennially growing members of this complex
bloom during the drier months, when D. montana is usually absent above ground. This contrasting seasonal behavior
is particularly evident in mixed populations of D. montana and D. tomentosa at the Serra do Ibitipoca and along the
Espinhaço Range in Minas Gerais state: when the former is in bloom, the latter is sterile (wet season), and when the
latter is in bloom, the former is dormant (dry season).
TABLE 1. Comparative diagnostic characters between the species of the Drosera montana complex.
D. montana D. spirocalyx D. tentaculata D. tomentosa
Leaf shape Oblong to oblong-spatulate
(spatulate)
Oblanceolate to narrowly
obovate
Obovate, obovate-cuneate to
cuneate
Obovate to oblong-obovate
(oblong)
Leaf vernation Circinate Circinate Geniculate Circinate
Adaxial petiole
indumentum
Glabrous to sparsely eglandular-
pilose
Densely eglandular-pilose Glabrous to sparsely
eglandular-pilose
Glabrous to sparsely
eglandular-pilose
Leaf apical tentacles Inconspicuously distinct Inconspicuously distinct Conspicuously distinct Usually inconspicuously
distinct
Glandular trichomes
on scape
Very sparse to dense, uniform in
length along the scape
Very sparse, uniform in
length along the scape
Sparse, uniform in length
along the scape
Largest near apex, decreasing
in size to 2/3 of scape length,
absent on basal 1/3
Sepal shape Ovate to oblong-ovate Sub-orbicular to broadly ovate Oblong-ovate (ovate) Narrowly ovate to oblong-ovate
Sepal aestivation Valvate to slightly imbricate Spirally contorted, forming a
pyramidal projection where
adjacent sepals fuse
Valvate to slightly imbricate Valvate to slightly imbricate
Sepal apex Acute Rounded (subobtuse) Acute Acute
Sepal (length × width) 2–6 × 0.8–2 mm 2–4.2 × 1.7–2.8 mm 2–4.5 × 0.7–2 mm 1.5–3 × 0.6–1.3 mm
Main flowering period Wet season (December–March in
Brazil, January-May in Bolivia)
Dry season (July–November) Dry season (July–October) Dry season (June–September)
Dormancy period Dry season (May–November) None None None
Distribution Widespread in Brazil (Bahia, Goiás,
Distrito Federal, Minas Gerais, Rio de
Janeiro, São Paulo, Paraná, and Santa
Catarina), disjunct in Bolivia (Tarija,
Chuquisaca, La Paz, and Santa Cruz)
Endemic to the central-
southern region of the Serra
do Cipó, Minas Gerais, Brazil
Endemic to the Espinhaço
Range in Bahia and Minas
Gerais, Brazil
Widespread in Brazil
– Sergipe, Bahia, Goiás, and
Minas Gerais

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FIGURE 4. Drosera montana. A, habitat, in white clayey soil (blackened after a recent fire), at Chapada dos Veadeiros, Goiás (central-
western morphotype). B, habitat, in red clayey soil at São Gonçalo do Rio Preto, Minas Gerais (type morphotype). C, habit of the central-
western morphotype, at Brasília, Distrito Federal. D, habit of the typical morphotype, with young flower scape densely glandular-pilose,
at the Itatiaia National Park, Rio de Janeiro. E, detail of the base of the scape of the southern morphotype, presenting a dense indumentum
of white eglandular trichomes at the base, near Curitiba, Paraná. F, flower, at the Serra do Quiriri, Santa Catarina. G, flower, cultivated
specimen from the Chapada dos Veadeiros, Goiás. A, E by F. Rivadavia; B, D by P.M. Gonella; C by Henrique Moreira; F, G by Carlos
Rohrbacher.

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As a result of this ephemeral growth of the vegetative parts, D. montana does not form any stems with age, nor
do columns of leaves accumulate beneath the live rosettes. Hence, rosettes are fewer-leaved and appear more lax in
habit than those of the other species of the complex. The leaves of D. montana are also of a thinner texture and more
delicate (appearing more membranous when dried), similar to those of D. hirtella and D. cayennensis, which share the
same growth habit and similar habitats. Curiously, when a mature or spent flower scape of any of these three summer-
dormant taxa is pulled, it usually results in the rosette breaking away with the scape, leaving the roots firmly in place.
When the same act is performed with the more robust perennially growing D. tomentosa and D. tentaculata, the rosette
is usually left behind together with the roots.
Drosera montana is the most geographically widespread and ecologically flexible species of the complex. Hence,
it is no surprise that it also displays more morphological variation, especially in leaf shape, and petiole length plus
width. Much of the observed morphological variation is apparently not ecologically induced by differences in habitats
or environmental conditions (e.g., elevation or soil), since plants often remain stable when cultivated under identical
conditions. These variations are subtle and more or less continuous, showing a gradual overlap across the range of
D. montana, and therefore do not currently constitute sufficient evidence to support a taxonomic separation of D.
montana at infraspecific rank. Nevertheless, the notable variation is here recorded as regional morphotypes, similar to
the equally variable D. latifolia (Gonella et al. 2014):
1. Typical morphotype (Fig. 4B, D): leaves oblong to oblong-spatulate, petioles much shorter and distinctly narrower
than the lamina (equal to or less than 0.5 mm wide).
The geographical distribution of this morphotype comprises almost the entire range of the species in Brazil, being
found from the Serra da Mantiqueira, on the border between São Paulo, Minas Gerais and Rio de Janeiro states
(where the locus classicus is situated), south along the Serra do Mar in São Paulo and Paraná, west to the Serra
da Canastra region in Minas Gerais, northwest to the Chapada dos Veadeiros in Goiás, as well as north along the
Espinhaço Range in Minas Gerais and Bahia states. Plants from the campos rupestres of the Espinhaço Range,
Serra da Canastra and Chapada dos Veadeiros tend to have more glabrescent leaves.
2. Southern morphotype (Fig. 4E): leaves oblong, with short and broad petioles (up to 1 mm wide), often of width
similar to the lamina. The base of the scape more often covered with sparse to dense white eglandular trichomes
(usually absent in other morphotypes).
This morphotype is found in the southern range of D. montana, in the Brazilian states of Paraná and Santa Catarina.
Robust plants of this morphotype are reminiscent of diminutive D. latifolia, which often occurs sympatrically.
However, D. montana can be promptly distinguished from D. latifolia (differences given in parentheses) by the
decumbent leaves in a flat rosette (vs. leaves semi-erect), by the presence of a dense glandular indumentum on the
basal third of the scape (vs. absent or very sparse), and by the smaller ovoid to ellipsoid seeds, 0.3–0.6 mm long
(vs. large fusiform seeds, 0.8–1 mm long).
3. Central-western morphotype (Fig. 4A, C): leaves oblong-spatulate to spatulate, long and distinctly narrower
petioles compared to the lamina (equal to or less than 0.5 mm wide).
This morphotype is found in Goiás and Distrito Federal in Brazil, and disjunctly in Bolivia.
Herbarium specimens from Bolivia, including previous references of D. villosa for that country (Schulenberg &
Awbrey 1997), are here considered conspecific with D. montana, geographically separated from the nearest known
Brazilian collections of D. montana in Goiás state by a distance of over 1500 km.
The isotype of D. communis var. pauciflora was examined and, in accordance with Correa & Silva (2005), this
taxon is considered conspecific with D. montana due to the oblong-spatulate leaves with short petioles, the erect base
of the scape, the large and broad sepals, and the flowering period (January).
Specimens examined:—BOLIVIA. Tarija: Prov. José María Avilés, cerca Pinos, 19 January 1986, Bastian 455
(LPB, M); 04 February 1999, Beck et al. 26900 (LPB); road from San Andrés to San Pedro, 05 March 2000, Wood 15942
(LPB). Prov. Cercado, Valle de San Andrés, 07 February 1987, Ehrich 379 (LPB). Prov. Burdet O’Connor, Camino
Papachacra quebrada Tambo, 18 February 2006, Zenteno & Quevas 3859 (LPB, M). Chuquisaca: Prov. Sud Cinti, Cerro
Bufete, 16 May 1995, Holst, Arroyo & Serrano 4700 (USZ); Puca Pampa, en la parte NW del Río Alborniyoj, 12 February
2004, Gutiérrez et al. 571 (MO, NY). Prov. Belisario Boeto, aproximadamente 25 km de Villa Serrano en Nuevo Mundo,
02 March 2006, Wood et al. 22356 (LPB). Santa Cruz: Prov. Florida, ca. 165 km hacia Cochabamba, 24 March 1981,
Beck 6842 (LPB); Samaipata, Alrededor de El Fuerte, 03 May 1994, Ibisch 35Pl38 (LPB). Prov. Vallegrande, 5 km al este
de Santa Rosita, 28 February 1995, Vargas 3723 (NY). La Paz: Prov. Franz Tamayo, Apolo to Salinas trail, near Salinas,
05 April 1977, Boeke 1456 (LPB, NY); Parque Nacional-ANMI Madidi, 17 April 2003, Miranda et al. 721 (LPB, MO).

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 153
Prov. Nor Yungas, ca. 4 km al Sur de Coroico, 05 March 1989, Beck 14865 (LPB); Milluguayo, December 1917, Buchtien
s.n. (US). Prov. Sud Yungas, 1 km al W de Yanacachi, 30 May 1987, Seidel 985 (LPB); 1988, Richter s.n. (LPB, M).
Without attribution to administrative area, Yungas, 1890, Bang 337 (B, M, MO, NY, US). Without further location data,
without date, Britton & Rusby 337 (NY). BRAZIL. Bahia: Município de Abaíra, Campo de Ouro Fino, 19 January 1992,
Hind & Queiroz 50900 (SPF), 15 February 1992, Harley et al. 52073 (SPF), 25 February 1992, Laessoe & Sano 52326a
(SPF), 16 March 1992, Laessoe & Silva 52556 (SPF), 23 March 1992, Laessoe & Silva 53328 (SPF); cachoeira das
Anáguas, 24 February 1992, Stannard et al. 51557 (SPF); Tijuquinho, 04 March 1992, Sano & Laessoe 50870 (SPF);
Catolés, 29 July 1995, Rivadavia & Ganev 493 (SPF); Catolés, trilha para o Pico do Barbado e Campo da Mutuca, 28
April 2010, Gonella & Fleischmann 326 (SPF); Serra do Barbado, 14 July 2005, Rivadavia et al. 2015 (SPF). Município
de Barra da Estiva, Serra do Ouro, March 1980, Pinto 8920 (ALCB); S de Barra da Estiva, 22 November 1992, Arbo et
al. 5681 (SPF), 22 July 2005, Rivadavia et al. 2094 (SPF); ~8km ao norte de Barra da Estiva, 10 July 2005, Rivadavia et
al. 1973 (SPF); Estrada Ituaçu-Barra da Estiva, antes de Barra da Estiva, 18 July 1981, Giulietti et al. 1247 (SPF).
Município de Palmeiras, trilha para a cachoeira da Fumaça, 20 July 2005, Rivadavia et al. 2079 (SPF). Goiás: Município
de Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada Alto Paraíso-Teresina de Goiás, 19 March 1997, Rivadavia 658
(SPF), 24 June 2007, Rivadavia & Batista 2627 (SPF); estrada Alto Paraíso-São Jorge, 20 March 1997, Rivadavia &
Araújo 665 (SPF), 24 June 2007, Rivadavia & Batista 2630 (SPF); Parque Nacional da Chapada dos Veadeiros, 26
February 2014, Gonella & Andrino 665 (SPF). Município de Cavalcante, Parque Nacional da Chapada dos Veadeiros,
estrada Alto Paraíso-Teresina de Goiás, 26 February 2014, Gonella & Andrino 660 (SPF), Gonella & Andrino 662 (SPF).
Município de Corumbá de Goiás, salto do Corumbá, 21 March 1997, Rivadavia 667 (SPF); oeste da cachoeira do Corumbá,
01 July 2007, Rivadavia & Batista 2666 (SPF). Município de Cristalina, a 20 km da cidade, 29 April 1999, Rivadavia &
Sato 936 (SPF). Município de Mossâmedes, Reserva Biológica da Serra Dourada, 01 May 1999, Rivadavia & Sato 995
(SPF). Município de Pirenópolis, próximo aos Três Picos, 23 April 1999, Rivadavia & Sato 925 (SPF). Distrito Federal:
Município de Brasília, Fundação Zoobotânica, 20 February 1961, Heringer 7966 (UB) (mixed collection with D. hirtella
var. hirtella); Setor Industrial, 19 March 1966, Irwin et al. 14116 (NY); Chapada da Contagem, 31 January 1980, Kirkbride
1055 (UB); Reserva Ecológica do Guará, 28 December 1993, Silva 2172 (CEN); saída do Setor Habitacional Norte, 18
March 1997, Rivadavia 639 (SPF), 05 January 2001, Rivadavia 1225 (SPF); Fazenda Sucupira, 2 February 2007, Vale et
al. 294 (CEN, UB); BR-020 para Formosa, 29 June 2007, Rivadavia & Batista 2637 (SPF); Fazenda Água Limpa, 14
March 2009, Meneguzzo et al. 85 (UB), Meneguzzo et al. 86 (UB); Sobradinho, beira de estrada no lado do Parque
Nacional de Brasília, 10 April 2010, Meneguzzo 575 (UB) (mixed collection with D. hirtella); Jardim Botânico, cerro
Cristo Redentor, 14 May 2012, Carvalho-Silva et al. 1641 (UB, MA). Minas Gerais: Município de Alto Caparaó, Parque
Nacional do Caparaó, 03 March 1991, Leoni 1453 (R); Pico da Bandeira, 19 February 1996, Rivadavia et al. 524 (SPF).
Município de Araxá, km 584 da BR262, 04 August 2002, Rivadavia 1405 (SPF). Município de Belo Horizonte, Serra
Taquaril, 23 March 1933, Mello-Barreto 2438 (BHCB), 31 March 1933, Mello-Barreto 2440 (BHCB); Carlos Torres, 10
May 1935, Mello-Barreto 4553 (BHCB). Município de Brumadinho, Serra da Calçada, without date, without collector
204 (SPF); Retiro das Pedras, 19 March 2002, Viana 616 (BHCB). Município de Carangola, Serra da Grama, 20 April
1935, Kuhlmann 5068 (VIC). Município de Carrancas, estrada Itutinga-Carrancas, 09 June 2007, Rivadavia et al. 2592
(SPF). Município de Congonhas do Norte, Serra Talhada, 24 April 1982, Furlan et al. CFSC 8473 (SP). Município de
Datas, arredores da cidade, 23 February 1982, Cordeiro & Simonis 4065 (SPF). Município de Diamantina, road to
Mendanha, 28 January 1969, Irwin et al. 22761 (NY, UB); ao leste da cidade, 23 February 1992, Rivadavia 71 (SPF);
rodovia Diamantina – Araçuaí, 14 April 1995, Kirizawa & Watanabe 2986 (SP) (mixed collection with D. hirtella var.
hirtella); estrada para Tapetes Arraiolos, 02 March 1997, Rivadavia et al. 594 (SPF); ao sul da cidade, 24 July 2008,
Gonella et al. 166 (SPF), 24 July 2008, Albuquerque et al. 2 (UB), 20 April 2010, Gonella et al. 276 (SPF); estrada para
Milho Verde, 14 February 2011, Gonella et al. 404 (SPF); Parque Nacional das Sempre Vivas, 08 March 2013, Gonella
et al. 608 (SPF). Município de Itabirito, Gandarela, 17 July 1972, Emygdio 3450 (R); Itabirito, Pico do Itabirito, 50 km
SE of BH, 11 February 1968, Irwin et al. 19817 (UB), Irwin et al. 19983 (NY), 18 January 1994, Teixera s.n. (BHCB).
Município de Itamonte, Parque Nacional de Itatiaia, 19 November 2005, Rivadavia et al. 2130 (SPF), 19 November 2005,
Rivadavia et al. 2132 (SPF). Município de Jaboticatubas, km 128 da rodovia Lagoa Santa-Conceição do Mato Dentro, 20
August 1972, Joly & Semir CFSC 3011 (SP); km 138-9 da rodovia Lagoa Santa-Conceição do Mato Dentro, 8 September
1972, Joly & Müller CFSC 3434 (SP); km 142 da rodovia Lagoa Santa-Conceição do Mato Dentro, 22 August 1972, Joly
& Semir CFSC 3166 (SP); Serra do Cipó, Fazenda Serra do Cipó, 27 February 1992, Rivadavia 103 (SPF), 24 February
1996, Rivadavia & Mullins 543 (SPF); Serra do Cipó, trilha para o mirante, 06 July 1995, Rivadavia 446 (SPF). Município
de Joaquim Felício, Serra do Cabral, 06 March 1997, Rivadavia 616 (SPF), 21 April 2010, Gonella et al. 284 (SPF).
Município de Lagoa Santa, November 1915, Hoehne 6459 (R). Município de Lavras Novas, 25 January 1986, Wagner et
al. 41592 (SPF). Município de Lima Duarte, Parque Estadual Florestal da Serra de Ibitipoca, 25 February 1977, Coons

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154 • Phytotaxa 172 (3) © 2014 Magnolia Press
77-276 (VIC), 08 October 1987, Andrade & Renato 1049 (BHCB), 06 February 1989, Souza & Krieger 24296 (R), 29
October 1995, Rivadavia & Padovese 504 (SPF), 23 November 2001, Rivadavia 1321 (SPF), 24 November 2001,
Rivadavia 1330 (SPF), 26 February 2005, Rivadavia 1953 (SPF), 25 January 2010, Ferrero et al. 1 (SPF), 15 December
2010, Gonella et al. 361 (SPF). Município de Moeda, Serra da Moeda, 04 August 1987, Andrade et al. 217 (BHCB), 12
March 2002, Rivadavia 1338 (SPF), 26 April 2002, Rivadavia 1348 (SPF), 08 March 2007, Batista 1989 (BHCB).
Município de Nova Lima, Morro do Chapéu, 23 February 1982, Grandi 872 (BHCB), 18 March 1983, Andrade & TSMG
1350 (BHCB). Município de Ouro Branco, Serra de Ouro Branco, 15 April 1988, Grandi 17887 (BHCB). Município de
Ouro Preto, 1888, Bello 216 (mixed collection with D. communis) (R); cachoeira das Andorinhas, 06 March 1982, Grandi
903 (BHCB); caminho para Lavras Novas, 25 January 1986, Kirizawa 1579 (SP); Itacolomi, 12 May 1998, Stehmann &
Mota 2347 (BHCB); Serra de Lavras Novas, 05 May 1974, Badini 23250 (OUPR), 11 May 1974, Badini 21562 (OUPR)
(mixed collection with D. tomentosa). Município de Patrocínio, Morro das Pedras, 28 January 1970, Irwin et al. 25487
(NY, UB). Município de Presidente Kubitschek, estrada para Datas, 27 February 1997, Rivadavia & Pinheiro 569 (SPF).
Município de Santa Bárbara, Serra do Caraça, January 1825, Riedel 1463 (P; type of D. communis var. pauciflora), 01
May 1980, Tales & Telma 85 (BHCB); Parque Natural do Caraça, 06 March 1992, Rivadavia 127 (SPF), 16 January 1994,
Souza et al.13810 (SPF), 01 July1995, Rivadavia 432 (SPF), 10 April 2011, Gonella & Siniscalchi 418 (SPF). Município
de Santana do Riacho, Serra do Cipó, km 120, 14 February 1968, Irwin et al. 20033 (NY, UB); Serra do Cipó, 17 March
1969, Saddi et al. 1125 (RFA); distrito da Lapinha, 01 January 2005, Rivadavia 1946 (SPF); inicio da trilha para as
Vellozias Gigantes, 15 May 2008, Gonella et al. 74 (SPF); km 112-113 da MG-10, 15 May 2008, Gonella et al. 94 (SPF),
19 April 2010, Gonella et al. 263 (SPF). Município de São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, na base
do Morro Redondo, 06 February 2009, Gonella & Viana 212 (SPF); trilha para Morro Redondo, 06 February 2009,
Gonella & Viana 213 (SPF). Município de São Roque de Minas, Parque Nacional da Serra da Canastra, 01 April 1999,
Rivadavia & Peixoto 867 (SPF), 01 April 1999, Rivadavia & Peixoto 872 (SPF), 02 April 1999, Rivadavia & Peixoto 877
(SPF), 02 April 1999, Rivadavia & Peixoto 879 (SPF), 04 Apr 1999, Rivadavia & Peixoto 895 (SPF). Município de São
Tomé das Letras, ao norte da cidade, 20 April 1992, Rivadavia 137 (SPF). Município de Tiradentes, Serra de São José, 08
June 2007, Rivadavia et al. 2585 (SPF). Without attribution to administrative area: Campos alpinos da Serra do Cipó, 16
January 1951, Andrade-Lima 51-867 (IPA); Serra do Cipó, summit of Serra do Cipó, 07 August 1960, Maguire 49071
(NY); Serra Itabira do Campo, 12 September 1887, Schwacke 5898 (R); Serra do Papagayo, March, Saint-Hilaire D542
(P; type). Without location data: 23 June 1980, Menezes et al. 6386 (SPF). Rio de Janeiro: Without attribution of
administrative area: Parque Nacional de Itatiaia, 12 March 1947, Occhioni 949 (RFA); Serra de Itatiaia, February 1894,
Ule 48 (R), 20 April 1959, Peres 54 (R), 16 October 1971, Andrade 32 (R); S.Itatiaia, planalto, 07 January 1947, Aparicio
& Edmundo 885 (R); Serra de Itatiaia, base das Agulhas Negras, November 1950, Leritz s.n. (R); Serra de Itatiaia, encosta
da Macieira, 07 June 1956, Leritz 64 (R); Serra de Itatiaia, 20 April 1957, Emygdio 1435 (R). São Paulo: Município de
Campos de Cunha, Serra da Bocaina, 22 October 2002, Rivadavia & Morais 1421 (SPF). Município de Campos do
Jordão, 20 November 1937, Porto 3226 (R, UB); Parque Estadual, 26 May 1977, Trinta et al. 3328 (R); caminho do Pico
do Itapeva, 17 November 1991, Rivadavia 25 (SPF), 10 October 1999, Rivadavia et al. 1183 (SPF); Recanto São Cristóvão,
22 January 1992, Rivadavia 57 (SPF); perto do Recanto São Cristóvão, 10 October 1999, Rivadavia et al. 1180 (SPF);
Estrada São Paulo-Campos do Jordão, 20 February 1937, Porto 3226 (R). Município de Pedregulho, Parque Estadual
Furnas do Bom Jesus, 11 March 2006, Rivadavia 2183 (SPF). Município de São José do Barreiro, Serra da Bocaina,
March 1951, Segadas-Vianna 3310 (R) 22 January 1998, Freitas & Sazima 128 (SPF), 22 October 2002, Rivadavia &
Morais 1413 (SPF), 22 October 2002, Rivadavia & Morais 1418 (SPF), 22 October 2002, Rivadavia & Morais 1419
(SPF). Município de São Paulo, Vila Ema, December 1932, Brade 12218 (R). Serra da Bocaina, without attribution to
administrative area: March 1951, Segadas-Vianna 2693 (R); Fazenda Bonito, 05 February 1917, Lutz & Lutz 1181 (R);
Morro do Corisco, 30 April 1951, Brade 20810 (R); source or Paraíba River, March 1951, Segadas-Vianna 2693 (R);
trilha para cachoeira de Santo Izidro, 11 June 1999, Pinheiro 100 (SPF); trilha para Fazenda Barreiro, 12 June 1999,
Pinheiro 101 (SPF); Fazenda Barreiro, 13 June 1999, Pinheiro 102 (SPF). Paraná: Município de Balsa Nova, divisa entre
os municípios de Balsa Nova e Palmeira, Rio dos Papagaios, 12 October 2010, Alves 3 (SPF); estrada que dá acesso ao
Morro do Cristo, 07 October 2012, Alves 27 (SPF); Serra do Purunã, 07 October 2012, Alves 28 (SPF), 07 October 2012,
Alves 29 (SPF). Município de Campo Largo, Fazenda São José, 02 October 2011, Alves 13 (SPF), 02 October 2011, Alves
16 (SPF) 02 October 2011, Alves 17 (SPF). Município de Palmeira, fazenda Padre Inácio, 31 October 1968, Hatschbach
20152 (MBM); Rio dos Papagaios, 21 April 1995, Rivadavia et al. 398 (SPF). Município de São José dos Pinhais, Rio
Pequeno, 20 October 1994, Ribas & Deodato 683 (SPF), 18 October 1998, Ribas 2742 (SPF); km 54 da rodovia BR-277,
12 September 2004, Rivadavia 1867 (SPF). Santa Catarina: Município de Campo Alegre, Morro Iquererim, 08 November
1956, Smith & Klein 7401 (R); Serra do Iquererim, 10 January 1992, Ribas & Guimarães 441 (MBM); Rancho Queimado,
Chapadão Serra Boa Vista, 27 April 2007, Rivadavia 2508 (SPF).

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 155
Drosera spirocalyx Rivadavia & Gonella, sp. nov. (Figs. 5, 6, 7)
Type:—BRAZIL. Minas Gerais: Santana do Riacho, trilha para velózias gigantes, 12 September 1999, Rivadavia 1165 (holotype SPF!;
isotypes BHCB!, P!).
Diagnosis:—Species similar to Drosera tentaculata Rivadavia, but differs in the oblanceolate to narrowly obovate
leaves densely covered by white eglandular trichomes on the adaxial petiole surface and whole abaxial leaf surface,
apical tentacles much less developed, shorter scapes, and sepals sub-orbicular to broadly ovate in shape with rounded
apex, overlapping (torsive or spirally contorted aestivation) when in bud and the lateral margins of adjacent sepals
forming a pyramidal projection where fused at their bases.
Description:—Perennial rosetted herbs, often forming short upright or prostrate stems up to 3 cm in length,
densely covered by persistent dead leaves. Roots slightly succulent, sparsely branched, black, densely covered with
root hairs. Indumentum consists of a dense cover of white eglandular trichomes (pale brown when dried) on the abaxial
leaf surface, adaxial petiole surface and base of the scape; capitate glandular trichomes on scape, bracts, pedicels and
sepals; sparse minute sessile glands ca. 0.03 mm in diameter on leaves and inflorescence parts. Leaves 3.5–15 mm
long, with circinate vernation, decumbent, oblanceolate to narrowly obovate, apex obtuse to sub-truncate, green to
wine red in color, usually bicolored with the petiole greener than the lamina; petiole 1.3–6.1 mm long, 0.4–1.6 mm
wide, densely eglandular-pilose (villous) on both surfaces; lamina 2.2–8.9 mm long, 1.5–3.6 mm wide, abaxial surface
densely eglandular-pilose, decreasing in density towards apex, adaxial surface covered with numerous red, motile,
capitate tentacles with radial symmetry, the apical tentacles inconspicuously distinct, 6–8 in number, 3–4 mm long,
stalk up to 0.2 mm wide at base, the head oblong, up to 0.3 mm long with unifacial gland tip; stipules 2–4 mm long,
0.3–1.1 mm wide at the base, rectangular, membranaceous, translucent white to reddish (drying bronze colored), the
apical 2/3–1/2 divided into 3 segments, the lateral ones longer, narrower, and poorly divided or entire, the central one
shorter, broader, and with fimbriate apex. Scapes 1–3, erect from the base, 14–82(–105) mm long (including floriferous
part), terete, green to deep red in color, covered with glandular trichomes uniform in size, sparse on the basal half,
increasing in density towards apex, often sparsely eglandular-pilose in the basal third; inflorescence a scorpioid cyme
bearing 1–4 flowers, rarely bifurcating; bracts caducous, usually absent on mature scapes, 0.2–1.5 mm long, linear,
glandular-pilose abaxially; pedicels 0.6–3 mm long, inserted 3.3–7.5(–9.5) mm apart, densely glandular-pilose; sepals
5, 2–4.2 mm long, 1.7–2.8 mm wide, sub-orbicular to broadly ovate, united at basal 2/5 of length, overlapping (torsive
or spirally contorted) when in bud with the lateral margins forming a pyramidal projection at the fusion point between
adjacent sepals, apex rounded (rarely subobtuse), sparse to densely glandular-pilose abaxially; petals 5, (3.5–)5–7.5
mm long, 4.8–6.1 mm wide, obovate, light to dark pink in color; stamens 5, 2–3.8 mm long, filaments white to light
pink, anthers 0.5–1 mm long, bithecate, yellow; ovary 3-carpellate, fused, 1–2 mm in diameter, ovoid, subglobose
to trilobed in outline; styles 3, forked at the base, 1.4–3 mm long (including stigmata), pale pink in color, style arms
curving inwards at the end of anthesis, stigmata 2–3 lobed or cordiform; fruit a dry dehiscent capsule, 2–3 mm long,
ellipsoid, 3-valvate; seeds ca. 0.7 mm long, ca. 0.25 mm wide, ellipsoid to oblong, black, testa reticulate.
Distribution and ecology:—Drosera spirocalyx is a narrow endemic species, known only from the central-
southern region of the Serra do Cipó, in Minas Gerais state (Fig. 5). It has been observed growing in campo rupestre
vegetation near or sympatric with D. chrysolepis, D. quartzicola, D. hirtella var. hirtella, D. montana, D. tentaculata and
D. tomentosa, occupying narrowly defined habitats at elevations between 1000–1350 m. Only around 15 populations
of D. spirocalyx are currently known, most of which had ca. 50–100 individuals, although a few large populations
contained hundreds of plants.
The majority of D. spirocalyx populations were found in sandy quartzitic soils or borderline habitats where sandy
soils meet mounds of white quartz gravel mixed with fine sand (a habitat also typical of D. quartzicola, D. schwackei,
and D. tentaculata). A few populations were observed in disturbed areas along trail sides, growing in fine white sandy
soil, or on the margins of natural water drainages.
Compared to other Drosera species, D. spirocalyx occupies relatively dry habitats far from springs and other more
reliable water sources. Although their habitats may receive plenty of rain in the summer wet season, the main source
of water in the dry season appears to be condensation at night. As a result, the rosettes of D. spirocalyx become less
vigorous in the dry season, producing shorter leaves that often lack mucilage on the carnivorous tentacles. However,
this species surprisingly never retreats into complete dormancy underground, as does D. montana.
The flat rosettes of D. spirocalyx (as well as those of D. montana and D. tentaculata) are often observed somewhat
covered with sand adhering to the stipules and hairs on the petioles, which may possibly act to protect the plants from
excessive heat exposure and evaporative loss.

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FIGURE 5. Distribution map of D. spirocalyx (squares) and D. tentaculata (circles) in eastern Brazil.
An interesting feature observed in some D. spirocalyx populations is the presence of globose clumps of plants
containing dozens of rosettes (Fig. 7A), suggesting a possible clonal origin by lateral budding from the roots.
Phenology:—Drosera spirocalyx produces flowers during the height of the dry season, extending into the early
wet season, between July to November, which corresponds to late winter and early spring.
Etymology:—The specific epithet “spirocalyx” was chosen due to the characteristic broad sepals of this new
species, which spirally overlap each other in the flower bud and often after anthesis (Figs. 6G–H; 7C, E).
IUCN Red List Category:—Although locally somewhat common in the central-southern region of the Serra
do Cipó, D. spirocalyx presents a very restricted area of occurrence. Most of the known populations lie outside the
protected area of the national park and are vulnerable to habitat destruction by cattle farming and mining activities,
possibly facing a threat in the near future. Thus, D. spirocalyx is here considered as Vulnerable (VU) according to the
IUCN criteria B2a, biii (IUCN 2001).
Discussion:—Drosera spirocalyx can be easily recognized by the often bicolored leaves with green petioles and
wine red lamina (a characteristic shared with D. tentaculata), the villous adaxial petiole surface and whole abaxial
leaf surface, the short scapes with few flowers (up to 4), and by the sub-orbicular to broadly ovate sepals with rounded
(rarely subobtuse) apex. The sepals of this species are unique among New World Drosera as their margins partially
overlap in bud (contorted aestivation), forming a pyramidal projection at the fusion point between adjacent sepals, a
character only found in one other, unrelated, species of Drosera, D. stenopetala Hooker (1853: 19) from New Zealand
(A.F. pers. obs.).
The relatively broad sepals of D. spirocalyx are more rigid than those of other species in the D. montana complex,
and are likely to aid in seed dispersal. When the fruit matures, the capsule and calyx open in a way similar to (but not as
pronounced as) the splash-cups found in D. felix, D. kaieteurensis, and D. solaris Fleischmann, Wistuba & McPherson
(2007: 1) (Rivadavia 1999; Fleischmann et al. 2007).
Drosera spirocalyx is apparently most closely related to D. tentaculata, sharing characteristics such as bicolored
leaves with apex often sub-truncate, occasionally forming columns of dead leaves, as well as similar habitats and
flowering season (for distinguishing characters, see Table 1).

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 157
FIGURE 6. Drosera spirocalyx. A, habit. B, C, detail of the indumentum at the base (B) and apex (C) of the scape. D, leaf, abaxial surface.
E, leaf, adaxial surface. F, stipule. G, calyx showing actual shape of sepals. H, calyx showing the spirally contorted overlap of the sepals.
J, petal. K, gynoecium and one stamen seen from above. L, seed. Based on the holotype. Drawn by P.M. Gonella.

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FIGURE 7. Drosera spirocalyx. A, habitat, forming a circular clump of rosettes, at the locus classicus. B, a flowering specimen. C, an
open flower seen from below, highlighting the sepals. D, detail of a sterile rosette. E, side view of a flower bud, showing the contorted,
overlapping sepals with obtuse apex, forming a pyramidal projection where adjacent sepals fuse. All by P.M. Gonella.
Specimens of D. spirocalyx were considered as either D. montana var. montana or D. montana var. tomentosa by
Silva (1994).
Additional specimens examined (paratypes)—BRAZIL. Minas Gerais: Município de Conceição do Mato
Dentro, Parque Natural Municipal do Ribeirão do Campo, 13 September 2002, Mota & Viana 1913 (BHCB). Município
de Santana do Riacho, Fazenda Palácio, 22 December 1948, Palacios et al. 3601 (R); km 129 da rodovia Belo Horizonte-
Conceição do Mato Dentro, 6 October 1981, Cordeiro et al. CFSC 7533 (SP, SPF); trilha para as Velózias gigantes, 09
July 1999, Rivadavia et al. 1095 (SPF), 07 September 2002, Rivadavia 1410 (SPF), 12 November 2007, Gonella et al.
44 (SPF), 15 May 2008, Gonella et al. 73 (SPF), 15 May 2008, Gonella et al. 80 (SPF); acampamento Serra Morena,
28 July 2002, Rivadavia & Gibson 1361 (SPF); 2.5 km após acampamento Serra Morena, 28 July 2002, Rivadavia
& Gibson 1363 (SPF); após acampamento Serra Morena, 07 September 2002, Rivadavia 1411 (SPF), 04 April 2003,
Rivadavia 1541 (SPF), 29 January 2005, Rivadavia 1937 (SPF), 23 May 2009, Gonella et al. 234 (SPF), Gonella et al.

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 159
235 (SPF), Gonella et al. 236 (SPF), Gonella et al. 245 (SPF), Gonella et al. 246 (SPF); km 112 da MG-10, trilha para
o canyon do Travessão, 04 April 2003, Rivadavia 1545 (SPF), 29 January 2005, Rivadavia 1939 (SPF); 16 May 2008,
Gonella et al. 91 (SPF), 20 July 2008, Gonella et al. 134 (SPF), 20 July 2008, Gonella et al. 152 (SPF), 22 July 2008,
Gonella et al. 162 (SPF), 06 April 2009, Gonella et al. 219 (SPF), Gonella et al. 223 (SPF), 19 April 2010, Gonella et
al. 260 (SPF), 02 September 2011, Gonella et al. 422 (SPF); entre sede do IBAMA e Vellozias gigantes, 21 July 2008,
Gonella et al. 139 (SPF), Gonella et al. 140 (SPF).
Drosera tentaculata Rivadavia (2003: 79) (Figs. 5, 8, 9)
Type:—BRAZIL. Minas Gerais: Diamantina, no alto da cachoeira dos Cristais, estrada para Biribiri, c. 1000 m alt., 31 July 2002, Rivadavia
& Gibson 1376 (holotype SPF!).
FIGURE 8. Drosera tentaculata. A, habit. B, C, detail of the indumentum at the base (B) and apex (C) of the scape. D, leaf, adaxial
surface. E, leaf, abaxial surface. F, stipule. G, calyx. H, petal. J, gynoecium and one stamen seen from above. K, seed. A–D and G–J based
on the holotype; E based on Gonella et al. 127; K based on Gonella et al. 135. Drawn by P.M. Gonella.

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FIGURE 9. Drosera tentaculata. A, habitat, in sandy soil with white quartz gravel at the Serra do Cipó, Minas Gerais. B, habitat, in
cracks of sandstone, at Diamantina, Minas Gerais. C, rosette seen from above. D, detail of juvenile leaf, with seven differentiated apical
tentacles. E, detail of the center of the rosette, showing the leaf buds. F, detail of flowers in post-anthesis, showing the calyx. G, flower.
All by P.M. Gonella.

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 161
Description:—Perennial rosetted herbs, often forming short upright or prostrate stems up to 2.5 cm in length, densely
covered by persistent dead leaves. Roots slightly succulent, sparsely branched, black, densely covered with root hairs.
Indumentum consists of white eglandular trichomes (pale brown when dried), on the abaxial leaf surface, more rarely
on the adaxial petiole surface and base of the scape; capitate glandular trichomes on scape, bracts, pedicels and sepals;
and minute sessile glands ca. 0.03 mm in diameter on leaves and inflorescence parts. Leaves 3–13(–17) mm long,
with geniculate vernation, decumbent, obovate to obovate-cuneate to cuneate, apex obtuse to sub-truncate, orange-
red to deep red in exposed habitats, green in shadier habitats, usually bicolored with the petiole greener than the
lamina; petiole 0.5–4(–8) mm long, 0.8–1.5 mm wide, abaxial surface sparsely (rarely densely) eglandular-pilose
adaxial surface glabrous to sparsely eglandular-pilose; lamina 2–9 mm long, 1.5–4(–6.5) mm wide, abaxial surface
glabrescent, eglandular trichomes concentrated on basal lateral margins, adaxial surface covered with numerous red,
motile, capitate tentacles with radial symmetry, the apical tentacles conspicuously distinct, 7–9(–12) in number, 4–
9 mm long, stalk horizontally flattened and narrowly-obcuneiform in shape, 0.4–0.5(–0.6) mm wide at base, the
head linearly-rectangular, 0.7–1.2(–1.5) mm long with unifacial gland tip; stipules 2–4 mm long, 0.8–1.5 mm wide,
rectangular, membranaceous, translucent white in color (drying bronze colored), apical 1/2–2/3 divided into 3 segments,
the lateral ones longer, narrower, poorly divided or entire, the central one shorter, broader, and with fimbriate apex.
Scapes 1–3, erect from the base, 30–160(–185) mm long (including floriferous part), terete, green to deep red in color,
erect from the base, glandular trichomes present from the base, somewhat uniform in size and increasing in density
towards apex, usually sparsely eglandular-pilose on lower third of the scape; inflorescence a scorpioid cyme bearing
(1–)2–8(–10) flowers, never bifurcating; bracts caducous, usually absent on mature scapes, 1–2.5 mm long, linear,
densely glandular-pilose abaxially; pedicels 0.7–5(–10) mm long, inserted 2–20 mm apart, densely glandular-pilose;
sepals 5, 2–4.5 mm long, 0.7–2 mm wide, oblong-ovate, rarely ovate, united at basal 1/3 of length, apex acute, densely
glandular-pilose abaxially; petals 5, 5.5–8.5 mm long, 3–4 mm wide, obovate, light to dark pink in color; stamens 5,
2.5–4 mm long, filaments white to light pink, anthers 1–1.6 mm long, bithecate, yellow; ovary 3-carpellate, fused,
1–1.7 mm in diameter, globose, subglobose to slightly trilobed in outline; styles 3, forked at the base, 2–4 mm long
(including stigmata), pale pink in color, style arms curving inwards at the end of anthesis, stigmata indistinct, bilobed,
basally bifurcate, or simply thicker than the style; fruit a dry dehiscent capsule, 2–2.5 mm long, ellipsoid, 3-valvate;
seeds 0.3–0.45 mm long, 0.15–0.2 mm wide, ovoid-ellipsoid, black, testa reticulate.
Illustrations:—Rivadavia (2003: 80, fig. 1).
Distribution and ecology:—Drosera tentaculata is endemic to the Espinhaço Range (Fig. 5), occurring in Minas
Gerais from the Serra do Cipó in the south to Itacambira in the north, and again disjunctly over 400 km to the north at
the Chapada Diamantina in central Bahia (Rio de Contas, Piatã, and Catolés). It is especially abundant at the Serra do
Cipó and on the Diamantina Plateau, which correspond to the core distribution area of this species, where large and
dense populations are commonly observed. In other regions (Itacambira, Serra do Cabral, Chapada Diamantina) D.
tentaculata is only know from small and scattered populations, and it is possibly also present on any of the numerous
and still poorly explored mountains between Itacambira and the Chapada Diamantina.
Drosera tentaculata occurs in campo rupestre vegetation in montane to high-montane habitats, at elevations
between 800–1600 m, where it is usually found in fine white sand with quartz gravel (Fig. 8A), but also on peaty-
sandy soil, in cracks on sandstone (Fig. 8B), or over a thin layer of soil over sandstone. During the summer wet season,
frequent rainfalls keep D. tentaculata habitats humid, but these become desiccated during the winter dry season,
which coincides with the flowering period of this species. At a few sites on the Diamantina Plateau D. tentaculata was
observed growing in year-round flooded areas, where it can present more robust rosettes with longer and wider leaves
bearing a few extra apical tentacles (these exceptional sizes and numbers are given in parentheses in the description).
Drosera tentaculata was found growing sympatric with D. ascendens, D. camporupestris, D. chrysolepis, D. montana,
D. quartzicola, D. schwackei, D. spiralis, D. spirocalyx, and D. tomentosa (both varieties).
Phenology:—Drosera tentaculata flowers primarily in the dry season, extending to the beginning of the wet
season, between July and October.
IUCN Red List Category:—Drosera tentaculata is widespread along the Espinhaço Range and especially
common in the southern part of its range. Because it is protected by national and state parks, D. tentaculata is here
considered of Least Concern (LC), according to the criteria of the IUCN Red List (IUCN 2001).
Discussion:—Drosera tentaculata can be distinguished by its obovate, obovate-cuneate or cuneate leaves with
7–9(–12) conspicuous apical retentive tentacles 4–9 mm long, 0.4–0.5(0.6) mm wide at the base, each with a narrowly-
obcuneiform stalk and a linearly-rectangular head 0.7–1.2(–1.5) mm long. Furthermore, it differs from other taxa in
this complex by the usually longer styles (up to 4 mm vs. up to 3 mm in the others), and less differentiated stigmata.

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A closer inspection of leaf vernation has revealed what is one of the most distinguishing features of D. tentaculata
from other members of the species complex. In D. montana, D. spirocalyx, D. tomentosa, and other Brazilian tetraploid
Drosera taxa, the developing leaf is rolled coil-like in bud and unfurls like a fern frond as it matures, representing
circinate vernation. More common among taxa of Drosera sect. Drosera is geniculate-involute vernation, where the
lamina is folded once over the petiole, and the leaf margins are folded inwards at an acute angle towards the apex of
the leaf. In D. tentaculata the vernation has been found to be geniculate, with the apical tentacles initially bent flat
against the lamina, and the lamina folded once over the petiole (see Fig. 8E), yet the margins of the lamina are not
folded inwards. This gives the young lamina of D. tentaculata a semi-circular shape, instead of the typical triangular
shape seen in geniculate-involute vernation. This mode of leaf vernantion is unique among this species affinity, and it
is likely to be correlated with the exceptionally large unifacial tentacles (“snap tentacles”) present in D. tentaculata.
In plants from Serra do Cipó, Minas Gerais, Rivadavia (2003) reported and illustrated styles with constrictions.
Several specimens were re-analyzed for the current study and this characteristic was not observed in any plant from
that region or any other area. Thus, the depicted styles with constrictions were possibly based on malformed or unripe
structures.
In previous floristic studies, D. tentaculata was referred to as D. montana var. montana (Silva, 1994; Silva &
Giulietti, 1997), probably due to the glabrous to sparsely eglandular-pilose base of the peduncle.
Specimens examined:—BRAZIL. Bahia: Município de Piatã, estrada para Inúbia, 16 July 2005, Rivadavia et al.
2033 (SPF), 01 August 2012, Gonella et al. 534, 538 (SPF). Município de Rio de Contas, lower NE slopes of the Pico
das Almas, 17 February 1977, Harley et al. 19568 (SPF); Pico das Almas, Campo do Queiroz, 11 November 1988,
Harley et al. 26353 (SPF); trilha subindo o Pico das Almas, 17 July 2005, Rivadavia et al. 2052 (SPF); trilha para o
alto do Pico do Itobira, 18 July 2005, Rivadavia et al. 2070 (SPF). Minas Gerais: Município de Buenópolis, alto da
Serra do Cabral em direção ao Brejo Grande, 03 July 2003, Rivadavia & Menezes 1651 (SPF); Parque Nacional das
Sempre Vivas, próximo ao alojamento de guarda-parques, 04 September 2011, Gonella et al. 449 (SPF), Gonella et al.
456 (SPF). Município de Diamantina, 03 June 1986, Brandão 11688 (PAMG); Planalto, 15 July 1970, Duarte 12983
(BHCB); campus II da UFVJM, 29 August 2005, Nunes & Costa 8 (DIAM); 11 December 2007, Vasconcelos et al. 2
(DIAM); 16 km de Diamantina para Mendanha, 06 July 1996, Souza et al. 11952 (SPF); estrada para Mendanha, 10
August 2010, Gonella et al. 353 (SPF); 12 October 1984, Mello-Silva et al. CFCR 5379 (SPF); estrada para Conselheiro
Mata, 23 September 1994, Splett 638 (UB), 29 November 2000, Prata et al. 1035 (SP); estrada Guinda-Conselheiro
Mata, 25 July 1998, Hatschbach et al. 68266 (SPF); estrada que sai do Bairro Arraiola, 08 February 2008, Gonella &
Viana 216 (SPF); ao sul da cidade, 24 July 2008, Gonella et al. 165 (SPF), 24 July 2008, Albuquerque et al. 4 (UB),
25 July 2008, Gonella et al. 173 (SPF), 25 July 2008, Albuquerque et al. 6 (UB); estrada para Curvelo, 27 February
1997, Rivadavia & Pinheiro 571 (SPF); estrada para Tapetes Arraiolos, 27 February 1997, Rivadavia & Pinheiro 580
(SPF), 10 July 1999, Rivadavia, et al. 1106 (SPF); estrada Diamantina-Conselheiro Mata, 29 June 2003, Rivadavia
1630 (SPF); estrada para Barão de Guacuí, 11 May 2007, Rivadavia 2518 (SPF); estrada para a Gruta do Salitre, 13
May 2007, Rivadavia 2542 (SPF); estrada para Milho Verde, 13 May 2007, Rivadavia 2555 (SPF); trilha dos Escravos,
10 August 2010, Gonella et al. 342 (SPF); Biri-Biri, cachoeira dos Cristais, 07 July 1995, Rivadavia & Pinheiro 451
(SPF), 31 July 2002, Rivadavia & Gibson 1376 (SPF; type); Biri-Biri, ao lado da estrada, 10 August 2010, Gonella
et al. 343 (SPF); Biri-Biri, Cachoeira da Sentinela, 10 August 2010, Gonella et al. 346 (SPF); São João da Chapada,
estrada para Macacos, 04 September 2011, Gonella et al. 425 (SPF), 04 September 2011, Gonella et al. 435 (SPF).
Município de Gouveia, estrada Diamantina-Curvelo, 02 August 2002, Rivadavia & Gibson 1402 (SPF), 06 April
2003, Rivadavia 1591 (SPF), 03 September 2011, Gonella et al. 424 (SPF). Município de Itacambira, estrada para
Montes Claros, 05 March 1997, Rivadavia 614 (SPF), 14 July 1999, Rivadavia & Pinheiro 1143 (SPF), 29 July 2002,
Rivadavia & Gibson 1364 (SPF), 22 April 2010, Gonella et al. 287 (SPF), 13 February 2011, Gonella et al. 401 (SPF).
Município de Jaboticatubas, Serra do Cipó, km 142 da rodovia Lagoa Santa-Conceição do Mato Dentro, 27 May
1972, Joly et al. CFSC 2111 (SP); estrada da usina, 21 August 1972, Joly & Semir CFSC 3067 (SP); km 138-9 rodovia
Lagoa Santa-Conceição do Mato Dentro, 08 September 1972, Joly & Müller CFSC 3433 (SP); 10 April 1980, Barros
200 (SP); trilha para mirante do canyon Bocaina, 05 June 1994, Rivadavia 289 (SPF); Fazenda da Serra do Cipó, 11
September 1994, Rivadavia 305 (SPF); estrada para o mirante do canyon Bocaina, 23 February 1996, Rivadavia &
Mullins 541 (SPF, MO); km 112-113 da estrada para Conceição do Mato Dentro, 09 July 1999, Rivadavia, et al. 1096
(SPF); km 112 da MG-10, 12 November 2007, Gonella et al. 54 (SPF), 12 November 2007, Gonella et al. 55 (SPF);
trilha do Travessão, 22 July 2008, Albuquerque 22 (UB), Albuquerque 25 (UB). Município de Joaquim Felício, alto
da Serra do Cabral, 07 March 1997, Rivadavia 629 (SPF); entre Joaquim Felício e Francisco Dumont, 03 July 2003,
Rivadavia 1662 (SPF), 04 July 2003, Rivadavia 1676 (SPF); entre Joaquim Felício e Augusto de Lima, 04 July 2003,
Rivadavia 1672 (SPF). Município de Santana do Riacho, ao lado da represa, 20 June 1990, Brandão 17283 (PAMG);

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 163
caminho da Lapinha, 11 February 1991, Arbo et al. 4851 (SPF); rodovia Belo Horizonte-Conceição do Mato Dentro,
06 September 1980, Forero et al. 7762 (SPF) (mixed collection with D. tomentosa var. tomentosa); 2.5 km após
acampamento Serra Morena, 28 July 2002, Rivadavia & Gibson 1362 (SPF); início da trilha para o Pico do Breu, 30
January 2005, Rivadavia 1944 (SPF); trilha para o Velozieto, 15 May 2008, Gonella et al. 84 (SPF), 08 August 2010,
Gonella et al. 336 (SPF); km 112 da MG-10, trilha para o canyon Travessão, 19 April 2010, 16 May 2008, Gonella
et al. 95 (SPF), Gonella et al. 98 (SPF), 20 July 2008, Gonella et al. 127 (SPF), Gonella et al. 135 (SPF), 22 July
2008, Gonella et al. 145 (SPF), Gonella et al. 154 (SPF), Gonella et al. 156 (SPF), Gonella et al. 161 (SPF), 06 April
2009, Gonella et al. 220 (SPF), 19 April 2010, Gonella et al. 261 (SPF), Gonella et al. 268 (SPF), 02 September 2011,
Gonella et al. 421 (SPF). Município de São Gonçalo do Rio Preto, PERP, trilha para o Pico Dois Irmãos, 05 February
2009, Gonella & Viana 198 (SPF). Município de Serro, estrada para Gouvea, 11 October 1996, Marcondes-Ferreira et
al. 1355 (UEC); Milho Verde, 24 July 2002, Mota 1643 (BHCB) (mixed collection with D. tomentosa var. glabrata);
trilha para a cachoeira do Arco-Íris, 05 April 2003, Rivadavia & Neves 1561 (SPF); planície ao lado da cidade, 05 April
2003, Rivadavia & Neves 1570 (SPF); estrada de Milho Verde para Capivari, 13 May 2007, Rivadavia 2557 (SPF).
Drosera tomentosa Saint-Hilaire (1826: 261) (Figs. 10–14)
Type (lectotype here designated):—BRAZIL. Minas Gerais: Crescit in paludosis montium propè Itambé, without date, Saint-Hilaire M-
378 (P-749166!, fifth specimen from the left (Fig. 14); isolectotypes P!, K(the two plants on the top right) image!, MPU image!).
≡Drosera montana A.St.-Hil. var. tomentosa (Saint-Hilaire 1826: 261) Diels (1906: 89).
Description:—Perennial rosetted herbs, often forming short upright or prostrate stems up to 2 cm in length, densely
covered by persistent dead leaves. Roots slightly succulent, sparsely branched, black, densely covered with root hairs.
Indumentum consists of white eglandular trichomes (pale brown when dried), on the abaxial leaf surface, often on the
adaxial petiole surface and from base to apex of scape, rarely on pedicels and sepals; capitate glandular trichomes on
scape, bracts, pedicels and sepals; and minute sessile glands ca. 0.03 mm in diameter on leaves and inflorescence parts.
Leaves (2.5–)4–24 mm long, with circinate vernation, decumbent (the youngest leaves sometimes semi-erect), obovate
to oblong-obovate, rarely oblong, apex obtuse, green to wine red in color; petiole (0.5–)1–8 mm long, 0.4–3 mm wide,
abaxial surface sparse to densely eglandular-pilose, adaxial surface glabrous to sparsely eglandular-pilose (very rarely
densely eglandular-pilose); lamina (2–)3–17 mm long, 1.3–6 mm wide, abaxial surface sparsely eglandular-pilose,
trichomes concentrated towards the base and along the margins, adaxial surface covered with numerous red, motile,
capitate tentacles with radial symmetry, the apical tentacles usually inconspicuously distinct, 5–9 in number, up to 5
mm long, stalk up to 0.3 mm wide at base, the head ellipsoid to rectangular-linear, up to 0.6 mm long with unifacial
gland tip; stipules 1.5–4 mm long, 0.5–2 mm wide at the base, rectangular, membranaceous, translucent white in color
(drying bronze colored), the apical 1/2–1/3 divided into several laciniate segments, the middle segments shorter than
the lateral ones. Scapes 1–3, erect from the base, (10–)28–330 mm long (including floriferous part), terete, green to
deep red in color, erect at the base, glandular trichomes absent on the basal third (only sessile glands present), but
increasingly denser and longer towards the apex, eglandular trichomes absent to dense from base to apex, usually
decreasing in density towards the apex; inflorescence a scorpioid cyme bearing 1–23 flowers, rarely bifurcating; bracts
caducous, usually absent on mature scapes, (0.4–)0.7–2 mm long, lanceolate to narrowly ellipsoid, densely glandular-
pilose and rarely sparsely eglandular-pilose abaxially; pedicels 0.7–6 mm long, inserted 2–20 mm apart, densely
glandular-pilose rarely bearing sparse eglandular trichomes; sepals 5, 1.5–3 mm long, 0.6–1.3 mm wide, narrowly
ovate to oblong-ovate, united at basal 1/3–1/5 of length, apex acute, densely glandular-pilose abaxially, rarely with
sparse eglandular trichomes; petals 5, 2–5 mm long, 2.5–4 mm wide, narrowly to broadly obovate or cuneate, light
to dark pink in color; stamens 5, 1.5–4 mm long, filaments white to light pink, anthers 0.4–0.8mm long, bithecate,
yellow; ovary 3-carpellate, fused, 0.5–1.3 mm in diameter, ellipsoid to globose, trilobed or hexalobed in outline; styles
3, forked at the base, 1.2–3 mm long (including stigmata), pale pink in color, style arms curving inwards at the end
of anthesis, stigmata flabellate, cupullate, bilobed, or more rarely bifurcate; fruit a dry dehiscent capsule, 1.1–2.3mm
long, ellipsoid, 3-valvate; seeds 0.45–0.7 mm long, 1.5–0.25 mm wide, ovoid to ellipsoid, dark brown to black, testa
reticulate.
Illustrations:—Ferrero & Mello-Silva (2011: 15, fig. 1D–F—D. tomentosa var. glabrata).
Distribution and ecology:—Drosera tomentosa is endemic to Brazil, occurring in the states of Sergipe, Bahia,
Goiás, and Minas Gerais (Fig. 10), where it mostly grows in montane to high-montane habitats at elevations between
700-2050 m. It is widespread and abundant in campo rupestre vegetation in central-eastern Brazil and is especially
common along the Espinhaço Range, from southern Minas Gerais to the Chapada Diamantina, in Bahia. In Minas Gerais
it furthermore spreads east to the Pico da Aliança, southwest to the Serra da Canastra and southeast to the Serra da

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Mantiqueira (Aiuruoca and Serra de Ibitipoca), possibly extending just over the border in São Paulo and Rio de Janeiro
states. In Goiás, D. tomentosa occurs disjunctly at the Chapada dos Veadeiros, in the northeastern part of the state.
FIGURE 10. Distribution map of D. tomentosa and its varieties in eastern Brazil.
An isolated collection of D. tomentosa is also known from the Serra de Itabaiana highlands in the state of Sergipe,
northeastern Brazil. This is a sandstone escarpment reaching nearly 700 m altitude, located in a relatively hot region
of Brazil, and it represents the northeastern distribution limit of D. tomentosa.
Drosera tomentosa, D. chrysolepis, and D. communis share a very interesting pattern of disjunct distribution
(geographically and ecologically). Although most populations in Brazil occur on relatively temperate highlands, above
700 m elevation, these three taxa also extend into hot coastal restinga habitats just above sea level, at several locations
along the northern coast of Bahia (and D. communis furthermore into Pernambuco and Paraíba).
Drosera tomentosa is one of the most common species of sundews in campos rupestres of Brazil, where it is
found in a variety of habitats, often forming large populations. It has been observed growing sympatric with nearly
all other Drosera species growing within its geographical range: D. ascendens, D. camporupestris, D. chrysolepis, D.
communis, D. graminifolia, D. grantsaui, D. graomogolensis, D. hirtella var. hirtella, D. intermedia, D. latifolia, D.
montana, D. riparia, D. spiralis, D. spirocalyx, D. tentaculata, and D. villosa.
Drosera tomentosa prefers growing in sandy-peaty soils around seepages or along streams on islands of vegetation
and in cracks on quartzitic rocks. Although it does not usually go dormant in the dry season, growth may slow down
and leaves sometimes reduce in size when growing in drier habitats. The largest D. tomentosa specimens are usually
found in the wettest soils, sometimes even growing semi-aquatically, while the smaller specimens are often observed
in the drier sandy habitats or on bare sandstone surfaces.
Drosera tomentosa is one of the few species of the genus in Brazil that can be found forming healthy populations
(albeit less vividly colored) when growing in shaded areas under riparian vegetation, the others being D. communis, D.
grantsaui, D. latifolia, D. riparia, and D. viridis.
Previous records of D. tomentosa from São Paulo state (Silva & Giulietti 1997; Silva 2002) result from a
misidentified specimen of D. hirtella var. lutescens [eg. Marcondes-Ferreira et al. 1167 (SPF)].
Drosera tomentosa has been reported to hybridize with D. grantsaui when both species are found sympatrically,
often forming large (and possibly fertile) hybrid populations, the hybrid being known as D. × fontinalis Rivadavia

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 165
(2009: 121). More rarely, D. tomentosa has also been observed to hybridize with D. latifolia and D. villosa (Gonella et
al. 2014).
Phenology:—Although D. tomentosa may occasionally be found in flower year round, most populations are
observed to begin flowering synchronously during the dry season, between June and September, usually lasting for
about one or two months (or longer if plants produce two or more scapes).
IUCN Red List Category:—Drosera tomentosa is widespread and abundant across its range, occurring inside
several protected areas. It is thus here considered of Least Concern (LC), according to the IUCN criteria (IUCN 2001).
Discussion:—Drosera tomentosa is distinguished from other members of the complex by its obovate to oblong-
obovate leaves (rarely oblong), with wide petioles (0.4–3 mm in width), apical tentacles usually inconspicuously
distinct in number of 5–9, up to 5 mm long, with a tentacle stalk up to 0.3 mm wide at the base and with a ellipsoid
to rectangular-linear gland head up to 0.6 mm long, glandular trichomes absent in the basal third of scape, usually
multiple flowered scapes (up to 23 flowers), anthesis in the dry season (June to September), and by the narrowly ovate
to oblong-ovate sepals (1.5–3 × 0.6–1.3 mm).
There is great variation in the degree of differentiation of the apical tentacles in D. tomentosa, from almost
indistinct (with ellipsoid head) to clearly longer, distinct, and with a rectangular head. However, even when distinct, the
apical tentacles of D. tomentosa are always much narrower at the base and have a much shorter head when compared
to the large marginal tentacles of D. tentaculata. The D. tomentosa variant with longer and distinct rectangular tentacle
heads has been observed growing sympatric with D. tentaculata at the Serra do Cipó and Diamantina regions, raising
the hypothesis of hybridization between these species. However, this D. tomentosa variant also occurs vicariantly from
D. tentaculata (e.g. in the Serra do Ibitipoca, Serra de São José, and the Chapada dos Veadeiros), suggesting that the
larger marginal tentacles of the former may not result from introgression with D. tentaculata, but simply represent
natural variation within D. tomentosa.
Although more similar to D. montana or D. tentaculata in most morphological characters, the overall habit of D.
tomentosa often significantly resembles diminutive sterile specimens of the taxa in the D. villosa complex (especially
D. latifolia), some of which are known to occasionally hybridize with D. tomentosa (Gonella et al. 2014). The floral
parts (including seeds) of D. tomentosa var. tomentosa are virtually indistinguishable from those of D. chimaera,
however the vegetative parts are very distinct, as the leaves of D. chimaera are narrowly to broadly oblong-lanceolate,
semi-erect and completely lack snap tentacles.
Drosera tomentosa is extremely variable in regards to the distribution of eglandular indumentum on the scape and
inflorescence, although this character is usually uniform and stable within populations. These white trichomes (pale
brown and often deciduous in old or dried specimens) may vary from completely absent to very densely present on
entire scapes, from base to apex, including pedicels and sepals.
Plants with scapes displaying a denser eglandular indumentum are more abundant in the northern part of the
D. tomentosa range, but variations in scape indumentum are not very continuous geographically. In central Minas
Gerais (and more rarely in central Bahia), the morphotype with dense eglandular-pilose indumentum is sometimes
found growing sympatrically with the morphotype bearing few or no eglandular trichomes on its scapes – yet both
morphotypes remain distinct and easily distinguishable in these populations. This is even evident from the type sheet
of D. tomentosa (Saint-Hilaire M-378), which represents a mixed collection of both morphotypes (and even a putative
scape of D. latifolia – see Gonella et al. 2014). Artificial crosses between the two morphotypes have been successfully
made in cultivation by the authors, and have shown to be fertile. Rarely, intermediate specimens can be observed in
mixed wild populations, suggesting that the two morphotypes may also hybridize naturally.
The above observations seem to support the original description by Saint-Hilaire (1826), splitting D. tomentosa
into two varieties based on scape eglandular indumentum: D. tomentosa var. tomentosa with hairy scapes and D.
tomentosa var. glabrata with glabrous to subglabrous scapes (Saint-Hilaire 1826). Therefore, in agreement with Saint-
Hilaire (1826), two infraspecific taxa are here accepted at the varietal rank, due to the huge overlap in their geographical
ranges, possible hybridization, and the fact that the sole distinguishing character (scape eglandular indumentum) is
not easily quantifiable. Furthermore, the presence or absence, as well as the density, of eglandular trichomes on scapes
of native Brazilian Drosera taxa has caused too much taxonomic confusion in the past, and is also currently known
to be a variable character (although to lesser degree) when comparing geographically separated populations of D.
cayennensis, D. communis, D. grantsaui, D. latifolia, D. montana, and D. tentaculata. Therefore, any status above rank
of variety for the two morphotypes observed in D. tomentosa would require further and more quantifiable data.
Taking into consideration the numerous individuals studied as herbarium specimens and seen in the wild across
the range of D. tomentosa, we provide a more detailed treatment of the two varieties below. As the type of D. tomentosa
was found to represent a mixed collection of both varieties and possibly D. latifolia, the designation of a lectotype

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became necessary. Based on the original description by Saint-Hilaire (1826) and a careful examination of the type
material, we here select the fifth specimen from the left (Fig. 14) as the lectotype.
Of the three accessions used for cytological studies to determine the chromosome number of D. tomentosa
(Rivadavia 2005), the vouchers Rivadavia 298 and Rivadavia 449 represent D. tomentosa var. tomentosa, while
Rivadavia 439 represents D. tomentosa var. glabrata. Both varieties are tetraploids with 2n = 40 chromosomes.
Because a clear distinction between D. tomentosa var. tomentosa and D. tomentosa var. glabrata was not always
possible for all material studied (either because these contained only sterile specimens, or due to the deciduous nature
of the scape indumentum in old herbarium sheets), some of the specimens examined are tentatively listed under D.
tomentosa s.l. at the end of the specimen list, without recognition of either variety.
Drosera tomentosa A.St.-Hil. var. tomentosa. (Figs 10, 11C–E, G, J, O, P; 12C, D; 13A–C, G)
Description:—Scapes densely eglandular-pilose (villous) on the basal half, decreasing in density towards apex;
pedicels and sepals only glandular-pilose or also sparsely to densely eglandular-pilose.
Distribution:—Drosera tomentosa var. tomentosa has its core distribution on the Espinhaço Range in Minas
Gerais and Bahia states, extending east to the Pico da Aliaça in eastern Minas Gerais and northeast to the Serra de
Itabaiana highlands, in central Sergipe. It is the predominant variety in the northern part of the species’ range, especially
on the Chapada Diamantina in Bahia where specimens with the densest eglandular-pilose (villous) scapes are found.
From there, this variety extends south along the Cadeia do Espinhaço into Minas Gerais state, through the Northern
Mountain Complex (spanning the mountains from Monte Azul to Itacambira), the Diamantina Plateau (including the
Serra do Cabral), the Serra do Cipó, and the Serra do Ouro Branco, which represents the southernmost range limit of
D. tomentosa var. tomentosa (Fig 10).
Discussion:—The autonymous type variety, Drosera tomentosa var. tomentosa, represents plants with scapes bearing
eglandular trichomes that are longer and more densely distributed on the basal half, especially towards the base of the
scape, but are also present (although usually more sparsely) all the way to the apex, often even on pedicels and sepals.
Specimens Examined:—BRAZIL. Sergipe: Município de Itabaiana, Serra de Itabaiana, 13 September 1981,
Andrade-Lima et al. 81-23 (IPA). Bahia: Município de Abaíra, caminho Jambeiro–Belo Horizonte, 14 July 1994,
Ganev 3531 (HUEFS); Catolés, 26 July 1995, Rivadavia 474 (SPF); Abaíra, Catolés, 29 July 1995, Rivadavia & Ganev
490 (SPF); Serra do Barbado, 14 July 2005, Rivadavia et al. 2007 (SPF). Município de Ibicoara, trilha para a cachoeira
do Buracão, 12 July 2005, Rivadavia et al. 1987 (SPF), 12 July 2005, Rivadavia et al. 1993 (SPF). Município de
Lençóis, estrada Lençóis–Seabra, 14 February 1994, Harley et al. 14070 (SPF), Harley et al. 14072 (SPF). Município
de Mucugê, campo defronte ao cemitério, 20 July 1981, Giulietti et al. 1396 (SPF); a 5 km de Mucugê, 07 September
1981, Furlan et al. 1990 (SPF); 3-5 km N da cidade, 20 February 1994, Harley et al. 14296 (SPF); alguns km fora
de Mucugê, 23 July 1995, Rivadavia 467 (SPF); estrada para Barra da Estiva, 11 July 2005, Rivadavia et al. 1978
(SPF). Município de Palmeiras, trilha para a cachoeira da Fumaça, 20 July 2005, Rivadavia et al. 2082 (SPF); Morro
do Pai Inácio, 28 April 1997, Conceição 544 (SPF). Município de Piatã, Serra da Tromba, 25 August 1992, Ganev 950
(HUEFS); Serra do Santana, 23 July 1995, Rivadavia & Ganev 482 (SPF), 13 July 2005, Rivadavia et al. 1999 (SPF);
ao lado da capelinha no alto da serra, 29 April 2010, Gonella & Fleischmann 332 (SPF). Município de Rio de Contas,
Pico das Almas, 10 April 1999, Forzza et al. 1139 (SPF), 17 July 2005, Rivadavia et al. 2041 (SPF), Rivadavia et al.
2050 (SPF); estrada do Pico das Almas para Rio de Contas, 17 Jul 2005, Rivadavia et al. 2055 (SPF); estrada para
Caiambola, 18 July 2005, Rivadavia et al. 2059 (SPF); Pico do Itobira, 18 July 2005, Rivadavia et al. 2064 (SPF),
Rivadavia et al. 2071 (SPF). Minas Gerais: Município de Botumirim, Rio do Peixe, 06 September 2011, Gonella et
al. 475 (SPF). Município de Buenópolis, alto da Serra do Cabral, 03 July 2003, Rivadavia & Menezes 1649 (SPF);
Parque Nacional das Sempre Vivas, atrás do alojamento de guarda-parques, 04 September 2011, Gonella et al. 457
(SPF). Município de Couto de Magalhães de Minas, c. 30 km ao leste de Diamantina, 12 May 2007, Rivadavia 2525
(SPF). Município de Diamantina, estrada de Tapetes Arraiolos, 31 July 2002, Rivadavia & Gibson 1374 (SPF); Biri-
Biri, cachoeira da Sentinela, 12 July 1999, Rivadavia et al. 1121 (SPF); ao sul da cidade, 25 July 2008, Gonella et
al. 174 (SPF), 09 August 2010, Gonella et al. 339 (SPF); Trilha dos Escravos, 10 August 2010, Gonella et al. 341B
(SPF); estrada para Mendanha, 10 August 2010, Gonella et al. 352 (SPF); São João da Chapada, estrada para Macacos,
04 September 2011, Gonella et al. 427 (SPF); Inhaí, estrada que sai por trás de Inhaí, 05 September 2011, Gonella
et al. 463 (SPF); Campus II da UFVJM, 06 March 2013, Gonella & Baleeiro 587 (SPF). Município de Gouveia,
estr. Curvelo–Diamantina, 09 July 2001, Costa et al. 304 (SPF), 02 August 2002, Rivadavia & Gibson 1399 (SPF).
Município de Grão Mogol, Morro do Jambeiro, 08 September 1994, Rivadavia 298 (SPF), 14 October 1995, Rivadavia
et al. 554 (SPF), 12 October 2001, Rivadavia 1265 (SPF); Grão Mogol, estrada Grão Mogol-Montes Claros, 12 August

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2010, Gonella 355 (SPF), 07 September 2011, Gonella et al. 484 (SPF). Município de Itacambira, km 91 da estrada
para Montes Claros (MG-308), 07 March 2013, Gonella et al. 593, 596 (SPF). Município de Jaboticatubas, Serra do
Cipó, Fazenda Serra do Cipó, 11 September 1994, Rivadavia 306 (SPF), 05 July 2003, Rivadavia 1679 (SPF); trilha
para o mirante, 06 July 1995, Rivadavia 449 (SPF); estrada para Conceição do Mato Dentro, 09 July 1999, Rivadavia
et al. 1098 (SPF), 04 April 2003, Rivadavia 1548 (SPF); km 112 da BR-010, 12 November 2007, Gonella et al. 47
(SPF). Município de Joaquim Felício, Serra do Cabral, 21 April 2010, Gonella et al. 279 (SPF). Município de Monte
Azul, estrada para Sucuruiu/ Riacho Grande/ Brejo Grande, 09 September 2011, Gonella et al. 506 (SPF). Município
de Ouro Preto, km 16 estrada para Ouro Branco, 16 September 1999, Rivadavia 1170 (SPF). Município de Porteirinha,
alto da cachoeira Véu da Noiva, 10 September 2011, Gonella et al. 510 (SPF). Município de Santana do Riacho, Serra
do Cipó, km 112-113 da MG-10, 16 May 2008, Gonella et al. 97 (SPF), 20 July 2008, Gonella et al. 128 (SPF), 22
July 2008, Gonella et al. 160 (SPF); trilha para o Travessão e Cachoeira Congonhas, 22 July 2008, Gonella et al. 149
(SPF); acima da entrada para a Usina, 06 September 1987, Simão 10425 (SPF). Município de Serranópolis de Minas,
trilha que atravessa a Cadeia do Espinhaço, 17 July 1999, Rivadavia & Pinheiro 1150 (SPF); Canyon do Talhado,
Cachoeira Sete Quedas, 08 September 2011, Gonella et al. 492 (SPF), 08 September 2011, Gonella et al. 498 (SPF).
Without attribution to administrative area: Crescit in paludosis montium propè Itambé, Saint-Hilaire M378 (P; type).
Drosera tomentosa A.St.-Hil. var. glabrata Saint-Hilaire (1826: 262) (Figs. 10, 11A, B, F, H, L, M; 12A, B; 13D–F)
Type:—BRAZIL. Minas Gerais: Milho Verde, without date, Saint-Hilaire M-476 (holotype P!; isotypes P! MPU image!).
Description:—Scapes exclusively glandular-pilose from base to apex, without notable eglandular indumentum,
occasionally also sparsely eglandular-pilose on the basal one-third, with scattered eglandular trichomes rarely occurring
further up the scapes and even on pedicels.
Distribution:—Drosera tomentosa var. glabrata is the more widespread of the two varieties, occurring on the
Serra da Mantiqueira in the south, extending northwest to the Serra da Canastra (southwestern Minas Gerais) and north
along the Cadeia do Espinhaço from southern Minas Gerais to central Bahia. It also occurs disjunctly at the Chapada
dos Veadeiros (northeastern Goiás) and along the northern coast of Bahia. On the Espinhaço Range, this variety is more
common and abundant in the southern region (especially on the Serra do Cipó and Diamantina Plateau), but relatively
rare in the north (Fig. 10).
Discussion:—Eglandular trichomes are often completely absent on the scapes of D. tomentosa var. glabrata.
When present, the eglandular indumentum is overall sparse and scattered in comparison to D. tomentosa var. tomentosa,
and the trichomes are also shorter and thinner. Although usually only present on the basal 1/3 of the scapes, sparse
eglandular trichomes are occasionally found on the upper 2/3 in some D. tomentosa var. glabrata populations, but are
still much less dense than in the type variety. Distinguishing the two varieties becomes especially challenging when
examining old scapes of dried specimens, because the eglandular trichomes are often deciduous.
Specimens Examined:—BRAZIL. Bahia: Município de Camaçari, estrada para Arembepe, 18 June 1997, Grantsau &
Lima 2 (SPF); estrada para o litoral (BA-512), 30 November 2002, Rivadavia 1435 (SPF), Rivadavia 1446 (SPF),
Rivadavia 1447 (SPF), 18 January 2003, Rivadavia & Lima 1519 (SPF), 14 August 2003, Rivadavia & Lima 1681 (SPF).
Município de Palmeiras, trilha para a cachoeira da Fumaça, 20 July 2005, Rivadavia et al. 2088 (SPF). Goiás: Município
de Alto Paraíso de Goiás, estrada Alto Paraíso–Terezina de Goiás, 10 October 1979, Heringer et al 2313 (IBGE), 20
March 1997, Rivadavia & Araújo 661 (SPF), 23 June 2007, Rivadavia & Batista 2625 (SPF); km 196 da BR118, 23 June
2007, Rivadavia & Batista 2604 (SPF), 7 September 2008, Albuquerque 31 (UB). Minas Gerais: Município de Botumirim,
Serra da Canastra, 20 July 1994, Rivadavia 332 (SPF), 19 November 1992, Mello-Silva et al. 673 (SPF), 13 October 2001,
Rivadavia 1266 (SPF), Rivadavia 1273 (SPF). Município de Buenópolis, Parque Nacional das Sempre Vivas, atrás do
alojamento de guarda-parques, 04 September 2011, Gonella et al. 450 (SPF). Município de Carrancas, Poço da Rasquinha,
09 June 2007, Rivadavia et al. 2593 (SPF); cachoeira do Tico-Tico, 09 June 2007, Rivadavia et al. 2595 (SPF); ao oeste
da cidade, 09 June 2007, Rivadavia et al. 2599 (SPF). Município de Conceição do Mato Dentro, Parque Natural Municipal
de Ribeirão do Campo, 01 August 2002, Silva et al 1642 (BHCB); Serra do Cipó, Campo Redondo, caminho para
nascentes do ribeirão do campo, 17 May 2008, Gonella et al. 112 (SPF), Gonella et al. 113 (SPF). Município de Congonhas
do Norte, Serra Talhada, 24 April 1982, Furlan et al. CFSC 8465 (SPF), 23 June 2013, Gonella et al. 621 (SPF); Serra da
Mangabeira, 26 February 1997, Rivadavia & Pinheiro 568 (SPF). Município de Couto de Magalhães de Minas, c. 40 km
ao leste de Diamantina, 12 May 2007, Rivadavia 2529 (SPF). Município de Diamantina, estrada para Conselheiro Mata,
18 July 1987, Pirani & Mello-Silva CFCR 11047 (SPF); ao leste da cidade, 23 February 1992, Rivadavia 74 (SPF); Biri-
Biri, cachoeira da Sentinela, 10 August 2010, Gonella et al. 347 (SPF); cachoeira da Toca, 25 February 1992, Rivadavia

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81 (SPF); cachoeira dos Cristais, 07 July 1995, Rivadavia & Pinheiro 450 (SPF); estrada para Biri-Biri, 14 July 1996,
Parra et al. 127 (SPF), 10 August 2010, Gonella et al. 344 (SPF); estrada para São João da Chapada, 11 December 1997,
Lima et al. 31 (SPF); estrada para Tapetes Arraiolos, 12 July 1999, Rivadavia & Pinheiro 1116 (SPF), Rivadavia &
Pinheiro 1117 (SPF), Rivadavia & Pinheiro 1118 (SPF), 31 July 2002, Rivadavia & Gibson 1373 (SPF); estrada para a
Gruta do Salitre, 13 May 2007, Rivadavia 2541 (SPF); ao sul da cidade, 25 July 2008, Gonella et al. 170 (SPF), 20 April
2010, Gonella et al. 273 (SPF); trilha dos Escravos, 10 August 2010, Gonella et al. 341A (SPF); São João da Chapada,
estrada para Macacos, 04 April 2011, Gonella et al. 434 (SPF); Parque Nacional das Sempre-Vivas, 08 March 2013,
Gonella et al. 605 (SPF). Município de Furnas, estrada Passos–Piumhi, 08 March 1997, Rivadavia 637 (SPF). Município
de Grão Mogol, ao sul de Grão Mogol, without date, Magalhães 18091 (RFA); córrego das Mortes, 03 September 1985,
Zappi et al. 8310 (SPF); Vale do ribeirão das Mortes, 04 September 1986, Cordeiro & Mello-Silva CFCR 10096 (SPF);
morro do Papo da Ema, 06 September 1990, Silva et al. CFCR 13428 (SPF), 06 September 1990, Silva et al. CFCR 13429
(SPF); Trilha da Tropa, 02 June 1994, Rivadavia 276 (SPF). Município de Itacambira, alguns km antes da cidade, 16
December 1994, Rivadavia 310 (SPF); ao sul da cidade, 18 December 1994, Rivadavia 326 (SPF); estrada para Montes
Claros, 05 March 1997, Rivadavia 602 (SPF), 13 October 2001, Rivadavia & Pinheiro 1292 (SPF), 13 February 2010,
Gonella et al. 402 (SPF), 22 April 2010, Gonella et al. 290 (SPF). Município de Jaboticatubas, Serra do Cipó, Fazenda
Serra do Cipó, 26 February 1992, Rivadavia 100 (SPF), 06 July 1995, Rivadavia 448 (SPF); entre Juquinha e sede do
IBAMA, 04 July 1995, Rivadavia 439 (SPF); estrada Cardeal Mota–Conceição do Mato Dentro, 21 July 2008, Albuquerque
et al. 14 (UB); trilha do Travessão, 22 July 2008, Albuquerque et al. 17 (UB), Albuquerque et al. 23 (UB), Albuquerque et
al. 26 (UB). Município de Joaquim Felício, Serra do Cabral, 07 March 1997, Rivadavia 631 (SPF), 03 July 2003, Rivadavia
1663 (SPF). Município de Lima Duarte, Conceição de Ibitipoca, Parque Estadual Florestal de Ibitipoca, 12 September
1991, Verardo 25371 (UFJF), 18 July 1992, Rivadavia 141 (SPF), 28 October 1995, Rivadavia & Padovese 497 (SPF), 29
October 1995, Rivadavia & Padovese 505 (SPF), 24 November 2001, Rivadavia 1324 (SPF), Rivadavia 1329 (SPF), 26
February 2005, Rivadavia 1955 (SPF); Pico da Lombada, 16 December 2011, Gonella et al. 365 (SPF). Município de Rio
Vermelho, Serra da Torre, 11 July 1999, Rivadavia et al. 1108 (SPF). Município de Santa Bárbara, Parque Natural do
Caraça, Pico do Sol, 04 March 1992, Rivadavia 117 (SPF), 05 April 1993, Rivadavia 209 (SPF), 02 July 1995, Rivadavia
434 (SPF); Pico da Carapuça, 08 September 1992, Rivadavia 149 (SPF), 03 July 1995, Rivadavia 436 (SPF); entre
Carapuça e Pico do Sol, 08 September 1992, Rivadavia 150 (SPF); Pico do Canjerana, 01 July 1995, Rivadavia 431 (SPF);
trilha para Campo de Fora, 01 July 1995, Rivadavia 433 (SPF); Cascatinha, 02 July 1995, Rivadavia 435 (SPF); trilha da
bica, 03 July 1995, Rivadavia 437 (SPF); Tanque Grande, 03 July 1995, Rivadavia 438 (SPF); Cachoeira Bocaina, 10
April 2011, Gonella & Siniscalchi 417 (SPF). Município de Santana do Riacho, rodovia BH–Conceição do Mato Dentro,
11 July 1987, Dokkedal et al. 10385 (SPF), 02 August 1990, Sakuragui et al. 21 (ESA), 26 April 1991, Pirani et al. 12247
(SPF), 08 June 1993, Roque & Nunes 13151 (SPF); Fazenda Cachoeira da Capivara, 11 July 1987, Horta & Andrade 141
(BHCB); rodovia Lagoa Santa–Conceição do Mato Dentro, 07 June 1989, Barbosa 409 (HUFU); Alto do Palácio, 16 May
1990, Arbo et al. 4275 (SPF); antes da bifurcação para Morro do Pilar, 23 February 1996, Rivadavia & Mullins 538 (SPF),
20 July 1993, Pirani et al. 13195 (SPF), 12 September 1999, Rivadavia 1167 (SPF), 12 November 2007, Gonella et al. 42
(SPF), 17 May 2008, Gonella et al. 117 (SPF), 21 July 2008, Gonella et al. 141 (SPF), 21 July 2008, Albuquerque et al.
12 (UB), 19 April 2010, Gonella et al. 270 (SPF); distrito da Lapinha, 09 July 1999, Rivadavia et al. 1104 (SPF); Vellozias
Gigantes, 15 May 2008, Gonella et al. 76 (SPF), Gonella et al. 79 (SPF); trilha para o Velozieto, 15 May 2008, Gonella
et al. 81 (SPF), Gonella et al. 85 (SPF), 08 August 2010, Gonella et al. 334 (SPF); estátua do Juquinha, 15 May 2008,
Gonella et al. 87 (SPF); km 108 da MG-10, 16 May 2008, Gonella et al. 108 (SPF); km 112-113 da MG-10, 20 July 2008,
Gonella et al. 125 (SPF); atrás da sede do IBAMA no alto da serra, 21 July 2008, Gonella et al. 136 (SPF); trilha para o
Travessão e Cachoeira Congonhas, 22 July 2008, Gonella et al. 150 (SPF), Gonella et al. 157 (SPF), Gonella et al. 159
(SPF). Município de São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 28 June 2003, Rivadavia & Deco 1615
(SPF), Rivadavia & Deco 1618 (SPF), Rivadavia & Deco 1625 (SPF), Rivadavia & Deco 1629 (SPF); trilha para o alto
da chapada, 05 February 2009, Gonella & Viana 189 (SPF); trilha para o Pico Dois Irmãos, 05 February 2009, Gonella &
Viana 197 (SPF); trilha para o Morro Redondo, 06 February 2009, Gonella & Viana 207 (SPF). Município de São Roque
de Minas, Parque Nacional da Serra da Canastra, 14 July 1997, Lombardi 1898 (BHCB), 01 April 1999, Rivadavia &
Peixoto 865 (SPF), 02 April 1999, Rivadavia & Peixoto 876 (SPF), Rivadavia & Peixoto 882 (SPF), 03 April 1999,
Rivadavia & Peixoto 892 (SPF), 20 December 1991, Rivadavia 39 (SPF). Município de Serro, Milho Verde, Saint-Hilaire
M-476 (P; type); Pico do Itambé, 09 July 1995, Rivadavia & Pinheiro 457 (SPF), 01 August 2002, Rivadavia & Gibson
1382 (SPF), Rivadavia & Gibson 1386 (SPF), Rivadavia & Gibson 1388 (SPF), Rivadavia & Gibson 1393 (SPF),
Rivadavia & Gibson 1395 (SPF); estrada Milho Verde-Diamantina, 06 April 2003, Rivadavia 1584 (SPF); estrada Serro-
Diamantina, nascente do Rio Jequitinhonha, 09 August 2011, Gonella et al. 337 (SPF). Município de Tiradentes, Serra de
São José, 08 June 2007, Rivadavia et al. 2586 (SPF). Without attribution to administrative area: Crescit in paludosis
montium propè Itambé, Saint-Hilaire M378 (P, specimens labeled as ‘g’ in Fig. 14A).

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FIGURE 11. Drosera tomentosa. A, habit. B, C, stipule. D–G, leaf, abaxial surface (E, F), adaxial surface (D, G). H, J, side view of the
calyx. K, L, petal. M, gynoecium and one stamen seen from above. N–P, seed. A, B, F, H, L, M correspond to D. tomentosa var. glabrata.
C–E, G, J, O, P correspond to D. tomentosa var. tomentosa. A, F based on Gonella et al. 365; B on Gonella et al. 434; C, G on Gonella et
al. 47; H on Gonella et al. 141; J on Ganev 3531; D, E on Rivadavia & Ganev 482; K–M on photographs of living plants; N on Rivadavia
435; O on Gonella et al. 475; P on Gonella et al. 492. Drawn by P.M. Gonella.

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FIGURE 12. Drosera tomentosa. Indumentum of the scape at the base (1), middle (2), and apex (3). A, B, D. tomentosa var. glabrata.
C, D, D. tomentosa var. tomentosa. A based on Gonella et al. 141; B based on Gonella et al. 125; C based on Gonella et al. 427; D based
on Ganev 3531. Drawn by P.M. Gonella.
Other Specimens examined of D. tomentosa s.l. (not assigned to either variety)—BRAZIL. Bahia: Município
de Lençóis, 28 June 1983, Santos 15993 (ALCB); Rio Capivara, 12 October 1999, Stradmann et al. 679 (ALCB);
trilha para a cachoeira do Sossego, 21 July 2005, Rivadavia et al. 2089 (SPF). Município de Palmeiras, Pai Inácio, 27
June 1995, Guedes et al. 1926 (ALCB), 26 September 1994, Giulietti et al. 782 (ALCB). Município de Rio de Contas,
Serra do Marcelino, 02 February 1997, Passos et al. 4857 (ALCB). Goiás: Município de Alto Paraíso de Goiás, km
10 da estrada Alto Paraíso-São Jorge, 25 June 2007, Rivadavia & Batista 2635 (SPF). Minas Gerais: Município de
Carmo do Rio Claro, Fazenda Córrego Bonito, 05 September 1961, Andrade & Emmerich 1004 (R). Município de
Conceição do Mato Dentro, Serra do Cipó, 07 August 1933, Mello-Barreto 2436 (BHCB). Município de Congonhas do
Norte, Serra Talhada, 24 April 1982, Furlan CFSC et al. 8473 (SPF). Município de Diamantina, Guinda, 05 November
1937, Mello-Barreto 9493 (BHCB); subida para o Cruzeiro, 12 January 1963, Duarte & Graziela 7922 (R); ao leste
da cidade, 22 February 1992, Rivadavia 59 (SPF). Município de Grão Mogol, morro do Papo da Ema, 10 July 1985,
Wanderley et al. 855 (SP); Vargem do Quartel, 10 July 1985, Cerati et al. 263 (SP), Graças et al. 855 (SP); Cachoeira
Véu das Noivas, 11 February 2011, Gonella et al. 381 (SPF). Município de Itabirito, Serra de Itabirito, 14 June 1976,
Badini & Furlo 24776 (OUPR), 27 July 1978, Badini 24682 (OUPR). Município de Itacambira, estrada para Montes
Claros, 14 July 1999, Rivadavia & Pinheiro 1131 (SPF). Município de Jaboticatubas, Serra do Cipó, rodovia Lagoa
Santa–Conceição do Mato Dentro, 07 June 1970, Joly et al. CFSC 1220 (SP), 14 December 1971, Semir & Saziman
537 (SP), 17 April 1972, Joly et al. CFSC 1883 (SP), 06 September 1980, Forero et al. 7762 (SP); Fazenda da Serra do
Cipó, 26 February 1992, Rivadavia 88 (SPF). Município de Joaquim Felício, Serra do Cabral, 29 June 2003, Rivadavia
1636 (SPF), 04 July 2003, Rivadavia 1675 (SPF). Município de Lima Duarte, Serra de Ibitipoca, ribeirão do Salto,
13 September 1940, Magalhães 386 (BHCB); Serra de Ibitipoca, 15 October 1986, Sousa 14858 (BHCB). Município
de Ouro Preto, Itacolomi, 25 June 1884, Glaziou 14482 (mixed collection with D. schwackei) (R), 05 May 1893,
without collector (OUPR), 1894, Gonçalves 5953 (OUPR), without date, Barbosa 1018 (R), without date, Damazio
979 (OUPR); Parque do Itacolomi, 1894, Miranda 4206 (OUPR), 02 June 1999, Paula 23532 (VIC); Serra de Lavras
Novas, 11 May 1974, Badini 21562 (OUPR) (mixed collection with D. montana), 27 July 1977, Badini et al. 24041
(OUPR); Serra da Chapada, 21 September 1990, Alves & Becker 1571 (R); Morro de São Sebastião, without date,
Clemen 3471 (OUPR); Serra de Ouro Preto, without date, Damazio 978 (OUPR); without location data, 1904, L.D. 508
(BHCB). Município de Prados, Serra de São José, 07 July 1991, Stehmann s.n. (BHCB). Município de Santa Bárbara,
Serra do Caraça, 1912, Damazio 2098 (mixed collection with D. latifolia) (R). Município de Santa Luzia, Serra do
Cipó, km 122, 14 April 1935, Mello-Barreto & Brade 1261 (BHCB), 06 July 1933, Mello-Barreto 2435 (BHCB).

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FIGURE 13. Drosera tomentosa. A, habitat, dense population growing in sandy soil of seepage at the Parque Nacional das Sempre-vivas,
Minas Gerais. B, detail of the apex of the inflorescence. C–E, rosettes with emerging inflorescences, showing the variation in eglandular
indumentum of the scape. F–G, flowers. A–C and G represent D. tomentosa var. tomentosa; D–F represent D. tomentosa var. glabrata.
All by P.M. Gonella.

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172 • Phytotaxa 172 (3) © 2014 Magnolia Press
FIGURE 14. A, type sheet of D. tomentosa, with the selected lectotype highlighted inside a red box; t, D. tomentosa var. tomentosa;
g, D. tomentosa var. glabrata; l, D. latifolia. B, detail of the lectotype of D. tomentosa. Image extracted from the Saint-Hilaire Virtual
Herbarium (Pignal et al. 2013).
Município de Santana do Riacho, Serra do Cipó, May 1933, Costa 156 (R), 24 July 1966, Emygdio et al 2248 (R),
Emygdio et al 2249 (R), 27 May 1970, Occhioni et al. 11407 (RFA), 28 May 1970, Occhioni et al. 11409 (RFA), 26
June 1972, Occhioni 51217 (RFA); rodovia BH-Conceição do Mato Dentro, 06 September 1980, Forero et al. 7762
(SPF) (mixed collection with D. tentaculata); Serra do Cipó, 01 April 1989, Delgado & Costa 3426 (BHCB); distrito
da Lapinha, 01 January 2005, Rivadavia 1945 (SPF); km 108 da MG-10, 16 May 2008, Gonella et al. 106 (SPF); km
112-113 da MG-10, 18 April 2010, Gonella et al. 258 (SPF); trilha para o Canyon das Bandeirinhas, 13 November
2007, Gonella & Lovo 61 (SPF). Município de São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 04 February

DROSERA MONTANA COMPLEX (DROSERACEAE) Phytotaxa 172 (3) © 2014 Magnolia Press • 173
2009, Gonella & Viana 180 (SPF). Município de São João del Rey, Serra do Lenheiro, 08 July 1936, Mello-Barreto
4684 (BHCB). Município de Serro, Milho Verde, 05 April 2003, Rivadavia & Neves 1555 (SPF), Rivadavia & Neves
1565 (SPF). Without attribution to adiminstrative area, Serra do Treituba, entre Ubá e Serraria, without date, Magalhães
296 (R); Serra Itabira do Campo, without date, Damazio 980 (OUPR).
Conclusions
The presence or absence, as well as the distribution, of eglandular trichomes on scapes are alone insufficient in
delimiting clear taxonomic boundaries between taxa historically included in the D. montana complex. A combination
of morphological, ecological, cytological, and phylogenetic characters have been found necessary for a more complete
understanding of the taxa most closely allied to D. montana.
The D. montana complex, as classified here, represents a group of tetraploid (2n = 4x = 40) species native to South
America (centered on the Espinhaço Range) with circinate or geniculate leaf vernation, and rosettes of flat decumbent
leaves with short petioles gradually widening to a more or less oblong or obovate lamina. Taxa included in this affinity
are D. montana, D. spirocalyx, D. tentaculata, D. tomentosa var. tomentosa, and D. tomentosa var. glabrata.
The type of D. tomentosa was found to be a mixed collection of D. tomentosa var. tomentosa, D. tomentosa var.
glabrata, and possibly D. latifolia, making it necessary to designate a single individual of Saint-Hilaire’s syntypes as
the lectotype.
Most of the taxa previously associated with D. montana are excluded from the complex, based on differences in
leaf shape, scape morphology and indumentum, seed shape and ornamentation, geniculate-involute leaf vernation,
molecular phylogeny data, and a diploid karyotype of 2n = 20. Drosera colombiana, D. panamensis, D. pumilla, and
D. sanariapoana are considered conspecific with D. cayennensis, and should be treated as synonyms. Drosera hirtella
var. hirtella and D. hirtella var. lutescens are considered well-delimited and distinct taxa, as is also D. roraimae.
Drosera parvifolia and D. montana f. parviflora are considered conspecific with D. communis. Drosera schwackei
is a very distinctive species, not belonging to this alliance of species, but to a closely related group including D.
graminifolia and other large Brazilian tetraploid species with a unique type of indumentum present on leaves and scapes.
Finally, D. communis var. pauciflora is considered a synonym of D. montana, based on leaf and sepal morphology, and
phenology.
Acknowledgements
We would like to thank Nílber Gonçalves Silva for helping with field work, important observations, aid in choosing the
name of D. spirocalyx, measurements, and early reviews of the present work; Chiaki Shibata for helping to rediscover
D. spirocalyx in 1999; François Mey and Laurent Legendre for helping with photographs of herbarium specimens
at P; Adilson Peres for helping with field work, as well as providing photographs and information of D. montana at
the type location; Stephan Beck for providing essential information, photographs and specimens from Bolivia; Rolf
Grantsau for preliminary botanical drawings; Rita Pereira for sending photographs of specimens from IPA herbarium;
Carlos Rohrbacher for helping with pictures and field work; the directors, curators, and staff of the institutions visited
for the study of herbarium specimens; Anderson Alves for sending valuable herbarium specimens from Paraná state;
Reginaldo Vasconcelos for finding D. tomentosa var. tomentosa on the Pico da Aliança and providing a picture;
Henrique Moreira for providing helpfull photographs; an anonymous reviewer is thanked for helpful comments on the
manuscritp; ICMBio and Santuário do Caraça for providing licenses for field collections; and numerous other friends
who have helped with discussions and/or field studies of the D. montana complex over the past 24 years. This work is
part of the M.Sc. thesis of PMG, financially supported by the Conselho Nacional de Pesquisa e Tecnologia (CNPq).

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174 • Phytotaxa 172 (3) © 2014 Magnolia Press
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