A revision of Drosera (Droseraceae) from the central and northern Andes, including a new species from the Cordillera del Cóndor (Peru and Ecuador)

Abstract

The Drosera species endemic to the central and northern Andes are revised here, including three species: the Venezuelan D. cendeensis as well as D. peruensis and D. condor sp. nov., from Peru and Ecuador. The latter is a new species endemic to the Cordillera del Cóndor that is here described and illustrated for the first time. The similarities and differences between these three taxa are discussed and the previously poorly known D. cendeensis and D. peruensis are provided with amended descriptions, illustrations and photographs. New records expand the known distribution range of D. peruensis through the sub-Andean cordilleras in Peru and Ecuador.

Full text

ORIGINAL ARTICLE
A revision of Drosera (Droseraceae) from the central and northern
Andes, including a new species from the Cordillera del Co
´ndor
(Peru and Ecuador)
Paulo M. Gonella
1
•Andreas Fleischmann
2
•Fernando Rivadavia
3
•
David A. Neill
4
•Paulo T. Sano
1
Received: 10 May 2016 / Accepted: 27 July 2016 / Published online: 30 August 2016
ÓSpringer-Verlag Wien 2016
Abstract The Drosera species endemic to the central and
northern Andes are revised here, including three species:
the Venezuelan D. cendeensis as well as D. peruensis and
D. condor sp. nov., from Peru and Ecuador. The latter is a
new species endemic to the Cordillera del Co
´ndor that is
here described and illustrated for the first time. The simi-
larities and differences between these three taxa are dis-
cussed and the previously poorly known D. cendeensis and
D. peruensis are provided with amended descriptions,
illustrations and photographs. New records expand the
known distribution range of D. peruensis through the sub-
Andean cordilleras in Peru and Ecuador.
Keywords Andes Drosera cendeensis Drosera condor 
Drosera peruensis New species Sub-Andean cordilleras
Introduction
Drosera L. (Droseraceae, non-core Caryophyllales) is a
carnivorous plant genus that comprises around 250 species
distributed worldwide, of which ca. 40 species are
currently recognized to occur in South America (Gonella
et al. 2015, Gonella et al. in prep.). In the Neotropics, the
genus is usually found in open vegetation types growing in
nutrient-poor sandy soils, with two main centers of diver-
sity: the Espinhac¸o Range in eastern Brazil and the Guiana
Highlands on the border between Venezuela, Guiana and
Brazil (Duno de Stefano 1995; Gonella et al. 2014; Fleis-
chmann et al. 2007).
Dominated by two regions with low Drosera diversity—
the Andes and the Amazon—Peru and Ecuador had only
two records of this genus: D. peruensis T.Silva &
M.D.Correa and D. chrysolepis Taub. (sensu Correa and
Silva 2005), both found in the sub-Andean cordilleras
(Duno de Stefano and Silva 2001; Silva and Correa 2002;
Correa and Silva 2005). These low-elevation mountain
chains scattered from northern Ecuador to southern Peru,
between the main Andes chain and the Amazon basin, have
attracted the attention of botanists over the past few dec-
ades because of their rich, unique, and poorly studied plant
diversity. Although uplifted concurrently with the Andean
chain several million years ago, these cordilleras differ
significantly from the Andes both geologically and floris-
tically (Schulenberg and Awbrey 1997). They consist
mostly of sandstones and limestones from Mesozoic and
early Cenozoic sediments and present a fascinating bio-
geographic and floristic connection at the generic level
with the more ancient tepui highlands of the Guiana Shield
(Huber 1995; Neill 2005; Ulloa and Neill 2006; Rodrı
´guez-
Rodrı
´guez et al. 2013).
Several new, locally endemic plant species and genera
have been recorded and described from these sub-Andean
highlands over the past two decades, especially from the
Cordillera del Co
´ndor, on the Peru-Ecuador border (Pruski
1998; Neill 2005; Clark et al. 2010; Neill and Asanza 2012;
Neill et al. 2012; Riina et al. 2014; Rodrı
´guez-Rodrı
´guez
Handling editor: Ricarda Riina.
&Paulo M. Gonella
paulogonella@uol.com.br
1
Laborato
´rio de Sistema
´tica Vegetal, Instituto de Biocie
ˆncias,
Universidade de Sa
˜o Paulo, Rua do Mata
˜o 277,
Sa
˜o Paulo 05508–900, Brazil
2
Botanische Staatssammlung Mu
¨nchen, Menzinger Strasse 67,
80638 Munich, Germany
3
1 Daniel Burnham Ct., San Francisco, CA 94109, USA
4
Herbario Amazo
´nico del Ecuador ECUAMZ, Universidad
Estatal Amazo
´nica, Puyo, Pastaza, Ecuador
123
Plant Syst Evol (2016) 302:1419–1432
DOI 10.1007/s00606-016-1341-3

et al. 2006,2013; SERNANP 2012; Ulloa and Neill 2006;
Ulloa et al. 2012;Va
´zquez-Garcı
´a et al. 2012; Penneys
et al. 2015). The Cordillera del Co
´ndor extends for 150 km
along a north–south axis, rising up to 2900 m and hosting
one of the richest floras in the Neotropics (Schulenberg and
Awbrey 1997).
The genus Drosera is known to present some extreme
cases of geographic disjunction, which have been
explained by long-distance dispersal. Drosera meristo-
caulis Maguire & Wurdack, endemic to the Neblina Peak
at the border between Venezuela and Brazil, belongs to a
monophyletic group of sundews endemic to Australia and
New Zealand (Rivadavia et al. 2012). The sister pair D.
sessilifolia A.St.-Hil. (Neotropics) and D. burmannii Vahl
(Southeast Asia and Australasia) (Rivadavia et al. 2003).
Two boreal species, D. anglica Huds. and D. rotundifolia
L., show disjunct populations in subtropics and/or tropics
with the former occurring on Kauai Island (Hawai’i chain)
and the latter in the Philippines and New Guinea (Coritico
and Fleischmann 2016). Furthermore, some Neotropical
Drosera species, such as D. brevifolia Pursh, D. intermedia
Hayne, and D. capillaris Poir, occur in disjunct populations
across the American continent along the north–south axis.
Finally, D. montana A.St.-Hil., a species widely distributed
in the highlands of eastern and southern Brazil, presents a
disjunct population in the Bolivian Yungas (Rivadavia
et al. 2014).
While previous works have confirmed all the above
cases, one remained doubtful until now: the disjunct dis-
tribution of D. chrysolepis. This species has its core dis-
tribution in the campos rupestres (rupestrian fields—
montane savannas with quartzitic and sandstone rock out-
crops) along the Espinhac¸o Range, in the Brazilian states of
Minas Gerais and Bahia (Gonella 2012). However, Dro-
sera chrysolepis also occur in restingas (coastal vegetation,
including moist broadleaf forests and herbaceous/shrubby
fields over sandy soils) of coastal Bahia, representing an
interesting case of geographic and ecological separation
(Silva and Giulietti 1997; Giulietti and Pirani 1988). Oddly
enough, this species had also been reported from the Cor-
dillera del Co
´ndor, over 4000 km from its Brazilian range
(Duno de Stefano and Silva 2001; Silva and Correa 2002;
Correa and Silva 2005).
Rivadavia (2003), while describing D. camporupestris
Rivadavia, a species from central Minas Gerais state (SE
Brazil) and formerly treated under D. chrysolepis by Silva
and Giulietti (1997), mentioned similarities between the
newly described species and a single specimen examined
from Ecuador (Gentry 80306). However, the great geo-
graphical disjunction and the incomplete specimen exam-
ined prevented this author from making a decision regarding
the identity of the Ecuadorian specimen (Rivadavia 2003).
During a comprehensive study of the Drosera from
Brazil and South America, further herbarium specimens
from the Cordillera del Co
´ndor were uncovered, enabling
us to conduct a meticulous morphological analysis. Addi-
tionally, these studies revealed new records of D. peruen-
sis, extending its distribution within Peru and to Ecuador.
Materials and methods
Herbarium specimens held at ECUAMZ, F, HUEFS, M,
MO, NY, QCNE, SPF, UC, US, USM, and VEN (herbar-
ium acronyms according to Index Herbariorum: Thiers
2016) were personally studied and carefully examined
under a stereomicroscope, and the descriptions were based
on dried specimens and spirit material. In addition, speci-
men photographs from MER, MERC, MERF, and PORT
were also analyzed. Habitat information was extracted
from the specimens’ labels and literature, as well as from
personal observations in the field. Distribution maps were
prepared using georeferenced location data obtained from
herbarium records and generated with DIVA-GIS (Hijmans
et al. 2005).
Conservation status was assessed using IUCN Red List
Categories and Criteria (IUCN 2012). For the estimation of
the extent of occurrence (EOO) and area of occupancy
(AOO), the CAT tool (available at http://geocat.kew.org/)
was used as described in Bachman et al. (2011). For the
AOO estimation, the standard IUCN cell size of 4 km
2
was
employed.
Results
Based on the analysis of vegetative and generative mor-
phology of the several new specimens from Cordillera del
Co
´ndor, it became evident that the similarities with D.
chrysolepis and D. camporupestris were merely superficial
and that the plants from Cordillera del Co
´ndor actually
belonged to a distinct, unnamed species.
The new species belongs to Drosera section Drosera
(sensu Seine and Barthlott 1994), based on the three styles
forked at the base, and was found to be more similar to two
other little-known species: the Andean D. cendeensis
Tamayo & Croizat, from the Pa
´ramo de Cende
´and sur-
rounding areas in northwestern Venezuela and D. peruen-
sis, previously known only from Cordillera Yanachaga, a
sub-Andean cordillera in central Peru (Fig. 1). In light of
this morphologically similar new taxon from the Cordillera
del Co
´ndor and the availability of additional specimens of
D. cendeensis and D. peruensis, a close reinvestigation of
the two latter taxa became necessary.
1420 P. M. Gonella et al.
123

The rarely available fertile material of D. cendeensis,
especially in recent collections, hinders the study of the inner
flower organs, such as the stamens and gynoecium, rendering
the floral description of this species incomplete in the current
work. Seeds were also not observed in the studied material of
D. cendeensis or D. peruensis, but SEM images are available
in Correa and Silva (2005) for both species, although the seeds
appeared to be unripe. Among the South American repre-
sentatives of Drosera section Drosera, floral characters are
generally not taxonomically significant to distinguish species,
whereas leaf characters are useful for species delimitation (see
Gonella et al. 2012,2014,2015; Rivadavia et al. 2014).
Discussion
Drosera condor,D. cendeensis, and D. peruensis share the
circinate leaf vernation (Fig. 5b), a somewhat similar (very
narrowly spatulate) leaf shape, and the indumentum of
scape, pedicels, and sepals consisting exclusively of sessile
glands and eglandular trichomes (lacking long-stalked
glandular trichomes). Drosera condor and D. peruensis are
further united by the triangular stipules. This combination
of characters is not found in any other Neotropical Dro-
sera, making it very difficult to phylogenetically place
these three species based on morphology only.
The eglandular-pilose inflorescence parts lacking long-
stalked glandular trichomes are shared with only a few
species in the Neotropics, namely D. cayennensis Sagot ex
Diels, D. amazonica Rivadavia, A.Fleischm. & Vicent., D.
felix Steyerm. & L.B.Sm., D. kaieteurensis Brumm.-Ding.
and D. solaris A.Fleischm., and Wistuba & S.McPherson
(the last three endemic to the Guiana Shield), which are in
turn all easily distinguished from the three Andean species
by geniculate-involute leaf vernation and leaves with an
obovate to round lamina.
Circinate leaf vernation (Fig. 5c) is a widespread char-
acter among South American Drosera species, being found
in another 17 species of the so-called Brazilian tetraploid
clade (Gonella 2012; Gonella et al. 2012,2014,2015;
Rivadavia et al. 2014). Triangular stipules are also present
in six species of this clade. However, all species of the
Brazilian tetraploid clade possess some kind of glandular
trichomes on the scape. The leaf vernation pattern is a
valuable taxonomic character to distinguish between the
New World diploid and tetraploid Drosera species
(Gonella 2012; Rivadavia et al. 2014), yet further cyto-
logical and molecular data are still necessary to securely
assign the three Andean species to one of these two groups.
Drosera species are mostly missing from the main
Andean chain (Ferna
´ndez-Pe
´rez 1965; Fleischmann and
Duno de Stefano 2014; Rivadavia et al. 2014), with the
exception of D. cendeensis in NW Venezuela, D. uniflora
Willd. from the southernmost Andean foothills of Argen-
tinean and Chilean Patagonia, and D. montana from the
Bolivian Yungas (a disjunction from its core distribution in
central and eastern Brazil; Rivadavia et al. 2014). This low
diversity may be explained by the Andean geology, which
consists mainly of nutrient-rich volcanic soils. In fact, most
Neotropical Drosera prefers nutrient-poor, quartzitic soils.
If the three Andean Drosera species described in this
work are revealed to be closely related to the tepui species
from the Guiana Shield, the occurrence of Drosera in the
sub-Andean cordilleras could corroborate the hypothesis of
a biogeographical connection between the two highland
areas, already supported by several disjunct genera (Neill
2005). Nevertheless, further phylogenetic studies are nec-
essary to test this hypothesis in the case of Drosera.
Taxonomic treatment
Drosera condor Gonella, A.Fleischm. & Rivadavia, sp.
nov.—TYPE: Ecuador. Morona-Santiago: Gualaquiza
Canto
´n, Campamento Achupalla, Cordillera del Co
´ndor,
15 km east of Gualaquiza, 2090 m, 21 Jul 1993, Gentry
80306 (holotype: QCNE–92866!; isotypes: MO–
04984889!, MO–04604340!, PMA [n.v.], SPF–109890!)
(Figs. 1,2,3).
Etymology: The scientific epithet is a noun in apposition
referring to the condor, the great vulture that lends its name
to the cordillera where this species grows.
Diagnosis: The new species is most similar to Drosera
cendeensis, but differs from the latter in its isomorphic
leaves that are narrower, more pilose, and fewer per
rosette; its larger triangular stipules (6.5–10 mm long) with
two abaxial longitudinal, sub-parallel ridges along their
whole length, each terminally laciniate and dividing the
apical third of the stipule into three segments, the lateral
segments being laciniate and longer, the central shorter and
divided into a few fimbriae.
Description: Perennial rosetted herb, rosettes lax, forming
short upright stems up to 15 mm covered by the persistent
dead leaves, stems not branching. Roots terete, dark, spar-
sely branched. Indumentum of white eglandular trichomes
(bronze colored when dried) up to 2.5 mm long, on abaxial
surfaces of petioles, laminae, and sepals, as well as on
scapes and pedicels; and minute sessile glands c. 0.03 mm in
diameter on both leaf surfaces, scapes, pedicels, and sepals.
Leaves 40–70 mm long, active leaves only 2–4 per plant,
with circinate vernation, erect to semierect, very narrowly
spatulate, red, apex rounded; petioles 30–55 90.4–0.8 mm,
linear, margins slightly revolute, adaxial surface glabrous,
abaxial surface sparsely eglandular-pilose; lamina
10–20 90.7–1.5 mm, narrowly oblanceolate, adaxial
A revision of Andean Drosera (Droseraceae) 1421
123

surface covered with numerous red, carnivorous, capitate
tentacles of radial symmetry, abaxial surface sparsely
eglandular-pilose; stipules 6.5–10 92–4 mm, triangular,
membranaceous, reddish in color (drying golden), with two
abaxial longitudinal ridges along their whole length, sub-
parallel, each terminating in a lacinia and dividing the apical
third into three segments, the lateral segments laciniate and
longer, the central segment shorter and divided into a few
fimbriae. Scape 1 per plant, 90–220 mm long (including
floriferous part), terete, 0.4–1 mm in diameter at the base,
scape erect from the very base, sparsely eglandular-pilose
from base to apex, red in color; inflorescence a simple
scorpioid cyme, bearing 2–7 flowers; bracts caducous,
1.5–2 mm long, oblong-lanceolate, eglandular-pilose; pedi-
cels 1–2.5 mm long, inserted 1.5–7 mm apart, sparsely
eglandular-pilose; sepals 3.5–4 91.5–2.5 mm, united at
basal half of length, elliptic, sparsely eglandular-pilose, apex
obtuse to acute; petals 8–12 95 mm, obovate, light pink to
violet-pink in color; stamens 5, c. 7 mm long, anthers c.
1 mm long, bithecate, yellow; ovary 3-carpellate, fused, ca.
1 mm in diameter, globose, slightly 3-lobed in outline; styles
3, forked at the base, ca. 4 mm long (including stigmata),
style arms curved inwards after anthesis, stigmata bilobed or
trilobed. Fruit a dry dehiscent capsule, ca. 3 mm long,
ellipsoid, 3-valvate; seeds oblong, 0.6–0.7 90.2–0.25 mm,
testa reticulate, black.
Fig. 1 Maps showing the
distribution of Drosera
cendeensis (triangles), D.
condor (stars), and D. peruensis
(circles)
1422 P. M. Gonella et al.
123

Phenology: Flower phenology information is very scarce
for this species, since there are only four known herbarium
collections from two localities, all of which were collected
with flowers and fruits in June and July.
Habitats:Drosera condor was found at elevations
between 2090 and 2500 m. The type collection at the
Achupallas site occurred in vegetation characterized as a
mosaic of sclerophyllous shrubs and herbazales, the latter
defined by Schulenberg and Awbrey (1997) as ‘‘meadows
dominated by dense and species-rich clumps of bromeliads
and orchids, interspersed with an array of other herbs and
shrubs including dwarf palms, the insect-eating Drosera,
and the rare curly grass–fern (Schizaea pusilla).’’
At the Cerro Machinaza site, D. condor grows on
outcrops of bare sandstone bedrock (Fig. 3b), in pockets of
soil over rock, a habitat it shares with a few other
herbaceous plants such as the fern Pterozonium brevifrons
(A.C.Sm.) Lellinger (Adiantaceae). Between the sandstone
Fig. 2 Drosera condor ahabit,
bdetail of the adaxial petiole
surface, cdetail of the abaxial
petiole surface, dstipule with
leaf base attached, epedicel and
calyx, and fseed. Based on
Gentry 80306 (isotypes: MO,
SPF)
A revision of Andean Drosera (Droseraceae) 1423
123

outcrops, the soil is a deep white quartzite sand derived
from the parent bedrock with a low, open vegetation of
shrubs and herbaceous plants not exceeding 1 m in height.
Many of these plants are local endemics known only from
the sandstone crest of the Cordillera del Co
´ndor that have
been described in recent years, such as the shrubs Clethra
concordia D.A.Neill, H.Beltra
´n & Quizhpe (Clethraceae),
Miconia machinazana C.Ulloa & D.A.Neill, Chalybea
brevipedunculata Penneys, C.Ulloa & D.Ferna
´ndez (Me-
lastomataceae), Weinmannia condorensis Z.S.Rogers
(Cunoniaceae), and the terrestrial herb Anthurium achu-
pallense Croat. (Araceae). The vegetation at the Cerro
Machinaza locality also includes shrub species with a
wider distribution such as Purdiaea nutans Planch.
(Clethraceae) and Arcytophyllum ciliolatum Standl. (Rubi-
aceae) (D. Neill personal observation; Fig. 3a).
Distribution area: Peru (Amazonas) and Ecuador (Mor-
ona-Santiago and Zamora-Chinchipe): endemic to the
Cordillera del Co
´ndor, along the border between the two
countries (Fig. 1). Drosera condor grows on the sandstone
mesas (tepui-like formations) at two known localities,
Achupallas at 2090–2500 m elevation and Cerro Machi-
naza (Machinaza Plateau) at 2160–2440 m elevation.
Conservation status: Endangered: EN B1ab(iii) ?B2a-
b(iii). The available data suggest that D. condor is endemic
to the Cordillera del Co
´ndor, a remote region that is still
very poorly explored botanically. According to Rodrı
´guez-
Rodrı
´guez et al. (2013), although deforestation and mining
activities have increased in the area over the past decades,
the central areas of the sandstone highlands remain mostly
intact. However, the expansion of large-scale mining
activity in that region poses a major threat to the biodi-
versity of the Cordillera del Co
´ndor, resulting in habitat
loss and environmental degradation. Based on the esti-
mated extent of occurrence (EOO) of 137.8 km
2
, the area
of occupancy (AOO) of 16 km
2
, and the fact that it is
restricted to very specific, narrow habitats that can be
easily degraded or destroyed by human activities, D.
Fig. 3 Drosera condor ahabitat in the Cordillera del Co
´ndor, bdetail of the leaves and cplants growing in cracks of sandstone (Neill & Kajekai
16902). Photos by David Neill
1424 P. M. Gonella et al.
123

condor is considered Endangered, according to the IUCN
Red List (2012) categories and criteria. This new species is
known to occur inside the recently created Ichigkat Muja—
Cordillera del Co
´ndor National Park, in Peru, as well as in
two protected areas in Ecuador: the El Quimi Biological
Reserve and the Cordillera del Co
´ndor Protection Forest.
Notes: The putative closely related taxa to D. condor are
the Andean species D. cendeensis, from northwestern
Venezuela, and D. peruensis, from Peru and Ecuador (see
below). All three species share a somewhat similar leaf
shape and indumentum (considering only the long, erect
leaves of the heterophyllous D. cendeensis) and similar
scape indumentum.
Drosera condor is easily recognized by the stipule
morphology (Fig. 2d), which is unique among all Neotrop-
ical taxa, except for the only distantly related D. meristo-
caulis from Cerro Neblina on the border between
Venezuela and Brazil, which similarly presents two sub-
parallel longitudinal ridges along the whole length of the
abaxial surface of the stipule. This is a curious case of
morphological convergence, since D. meristocaulis
belongs to a distantly related clade of mostly Australian
species, the ‘‘pygmy sundews’’ (Drosera section Bryas-
trum; Rivadavia et al. 2012).
Drosera condor and D. cendeensis share similar leaf
morphology, if one considers only the erect leaves of the
latter species. Besides the characteristic stipule morphol-
ogy, D. condor differs from D. cendeensis by the isomor-
phic leaves (vs. dimorphic leaves in D. cendeensis,a
heterophyllous species which produces smaller rosetted
leaves during the dry dormancy periods), the narrower
lamina up to 1.5 mm wide (vs. 2–3 mm wide in the erect
leaves of D. cendeensis), and the abaxial leaf surface with
denser indumentum (vs. glabrous to glabrescent abaxial
leaf surface).
Although not very similar at a first glance, D. condor
resembles a lax, long-leaved D. peruensis. Both species
share similar leaf and scape indumentum, but D. peruensis
has more compact and dense rosettes with shorter,
narrowly oblong leaves (and further differs from the other
two, pink-flowered Andean species by its white petal
color).
Similarities with D. chrysolepis and D. camporupestris,
both from eastern Brazil, include the overall habit, the
narrowly spatulate leaves, the triangular stipules, and the
similarly sized oblong seeds. Drosera condor further shares
with D. camporupestris the characteristic lax rosette, each
with very few active leaves (usually only 1–3). However,
the two Brazilian species are very distinctive in their larger
stipules (9–16 mm long in D. camporupestris; 8–15 mm in
D. chrysolepis vs. 6.5–10 mm long in D. condor), the acute
leaf apex terminating in a single carnivorous tentacle (vs.
rounded leaf apex) and the dense indumentum of long
eglandular trichomes and translucent-yellow, short-stalked,
globose multicellular glandular trichomes (‘‘TSG tri-
chomes’’; Rivadavia and Gonella 2011) that are found on
both petiole surfaces, the abaxial surface of the lamina, the
scape, pedicels, and sepals.
Additional specimens examined (paratypes): Ecuador.
Zamora-Chinchipe: Paquisha Canto
´n, Cordillera del Co
´n-
dor, Machinaza Plateau, precisely on the Ecuador–Peru
border, 2440 m, 22 Jun 2009, Neill and Kajekai 16902
(MO, QCNE). Morona-Santiago: Gualaquiza Canto
´n, crest
of Cordillera del Co
´ndor, ridge top 15 km ENE of Guala-
quiza, 2500 m, 26 Jul 1993, Gentry 80462 (MO, QCNE–
94752). Peru. Amazonas: Condorcanqui, Cordillera del
Condor, la cima del ‘‘tepui’’ Cerro Machinaza, cabeceras
del Rio Comainas, tributario al oeste del Rio Cenepa,
2160 m, 31 Jul 1994, Beltra´n and Foster 1488 (F, USM).
Drosera cendeensis Tamayo & Croizat. Lilloa 17: 175.
1949.—TYPE: Venezuela. Trujillo/Lara: Pa
´ramo de
Cende
´, 3000 m, 24 Jun 1940, Isabel Venegas Perdomo
s.n. [Tamayo 1430] (holotype: VEN!; isotypes: F!, UC!,
US–1800890!, US–1800021!) (Figs. 1,4,5a–d).
Vernacular names:dı
´tamo, dı
´tamo real, dı
´ctamo de
venado, yerba de Lucas (Duno de Stefano and Riina 2003).
Etymology: Named after the pa
´ramos de Cende
´(now in
the Dinira National Park) in the Venezuelan Andes, where
it was first discovered (Tamayo and Croizat 1949).
Description: Perennial rosetted herbs, acaulescent or
forming very short, inconspicuous stem up to 5 mm long.
Roots terete, dark, sparsely branched. Indumentum of white
eglandular trichomes (bronze colored when old) on scape,
pedicels and sepals and eventually on leaves; minute sessile
glands c. 0.03 mm in diameter on both leaf surfaces, scape,
pedicels, and sepals. Leaves dimorphic, with circinate
vernation, lamina adaxial surface covered with numerous
red, carnivorous, capitate tentacles with radial symme-
try; basal leaves numerous, 10–25 mm long, oblong-
spatulate, decumbent to semierect, glabrous, petioles
7.5–17 91.5–2 mm, lamina 2.5–5 92–2.5 mm, continu-
ous with the petiole, oblong to ellipsoid, apex rounded to
obtuse, green with red tentacles; erect leaves 3–5 per plant,
30–140 mm long, very narrowly spatulate, glabrous or with
very sparse eglandular trichomes in the abaxial surface,
petioles 25–106 90.6–2 mm, linear with slightly broader
base, narrowing toward apex, lamina 5–34 92–3 mm,
oblong to narrowly oblanceolate, apex rounded, red to red-
dish-green; stipules 4–7(–8) 92–3 mm, rectangular, mem-
branaceous, translucent white to reddish in color (drying
gold-colored), the apical third divided in three segments,
the two lateral segments simple and laciniate, the central
A revision of Andean Drosera (Droseraceae) 1425
123

segment broader and shorter, divided into a few fimbriae.
Scape 1 per plant, 150–250 mm long (including florif-
erous part), terete, 1.5 mm in diameter at base, base
erect, sparsely eglandular-pilose at the base, becoming
densely eglandular-pilose toward apex, red in color.
Inflorescence a simple scorpioid cyme, bearing 3–7
flowers; bracts not seen; pedicels 1–4 mm long, sparsely
eglandular-pilose; sepals 6–9 9ca. 1.5 mm, united at
basal 1/3–1/4 of length, lanceolate, densely eglandular-
pilose, apex acute; petals 7–12 mm long, obovate, pink
in color; stamens 5, ca. 5 mm long, anthers ca. 1 mm
long, bithecate, yellow; ovary 3-carpellate; styles 3,
forked at the base, ca. 3 mm long, stigma bilobed; fruits
not seen; seeds not seen [according to Tamayo and
Croizat (1949): seeds broadly ellipsoid, 0.75 mm long,
testa reticulate, black].
Fig. 4 Drosera cendeensis
ahabit depicting a fertile
specimen with basal and erect
leaves, bhabit depicting a
specimen at rosette stage only
with basal leaves, cstipule of an
erect leaf with base of leaf
attached, ddetail of the
indumentum on the abaxial
petiole surface, ebasal leaf with
stipule attached and fpedicel
and calyx. a,c–ebased on Riina
et al. 722;bbased on van der
Werff and Rivero 7997;fbased
on Tamayo 1430 (isotype UC)
1426 P. M. Gonella et al.
123

Phenology:Drosera cendeensis was collected with erect
leaves and flowers between May and August, while spec-
imens collected in November presented only basal leaves
and no inflorescences. Also, in field studies conducted in
September 2006 (Bricen
˜o2009), only immature plants with
basal leaves were observed.
The species has a dormancy period in the dry season (R.
Duno de Stefano personal comm.), during which shorter
and more spatulate-shaped leaves are present (Fig. 4b, e;
quite similar to the distantly related D. stenopetala Hook.f.
from New Zealand). Interestingly, the unrelated
carnivorous plant Pinguicula elongata Benj. (Lentibulari-
aceae, Lamiales), endemic of high Andean pa
´ramos of
Colombia and Venezuela, also has dimorphic leaves, which
enable it to form dormant, bulb-like resting buds during the
two annual dry seasons (Ferna
´ndez-Pe
´rez 1964; Beck et al.
2008; S. Vieira personal comm.).
Habitats:Drosera cendeensis grows in montane forest and
subpa´ramo vegetation, where it is usually found associated
with riparian vegetation, or open wet areas in peat-rich soils
often covered by carpets of Sphagnum L. (Fig. 5a, b).
Fig. 5 a–dDrosera cendeensis at Dinira National Park, Venezuela:
ahabitat at La Lajita, bgroup of individuals at the rosette stage
growing among Sphagnum sp., ccollected specimens (Alvarado et al.
3377) presenting early emerging erect leaves, note the circinate leaf
vernation, dcollected fertile specimens (Riina et al. 722). e–gD.
peruensis ecollected specimen from Cerro Plateado (Neill et al.
17459), forming a column of dead leaves, fplant covered by dew, at
Sira Mountain, Peru (Graham 5751) and gflower (Graham 6038). a,
dby Ricarda Riina; bby Re
´gulo Bricen
˜o; cby Hipo
´lito Alvarado;
eby David Neill; f,gby James G. Graham
A revision of Andean Drosera (Droseraceae) 1427
123

Distribution area: Endemic to the easternmost part of the
Venezuelan Andes, in the states of Trujillo and Lara
(Fig. 1). Drosera cendeensis has been collected in the
pa
´ramos de Cende
´, Las Rosas, Los Nepes and Jabo
´n, at
elevations ranging between 2300 and 3200 m. A previous
report from the Guaramacal National Park (Duno de Ste-
fano 1995) turned out to be an error, as no Drosera species
is recorded from this area (Dorr et al. 2000).
Conservation status: Critically Endangered: CR
A2d ?B1b(v)c(iv). Drosera cendeensis is locally considered
to be medicinal, reportedly used as an antispasmodic for
pertussis, a tonic, and a vasodilator (Duno de Stefano and
Riina 2003). The species is collected in large quantities and
sold in local markets in Trujillo and Lara, which explains its
increasing rarity (Bricen
˜o2009). Also, the flowering period
coincides with the time when plants are collected for
medicinal use (24 June—Day of Saint Juan, Bricen
˜o2009),
which directly affects the number of mature individuals and
jeopardizes the reproduction of the species. Given its
restricted occurrence (EOO of 20 km
2
) associated with
overexploitation, the species is here signaled as Critically
Endangered. Drosera cendeensis is found inside the Dinira
National Park, but according to Duno de Stefano and Riina
(2003), that is not enough to ensure its protection.
Notes: In discordance with Tamayo and Croizat (1949)
and Duno de Stefano (1995), who describe the indumen-
tum of the scapes and sepals as being exclusively glandu-
lar-pilose, all fertile specimens examined (including the
type specimens) presented an exclusively eglandular-pilose
indumentum (Fig. 4a, f).
The type specimen of D. cendeensis was originally
collected by Isabel Venegas Perdomo, but deposited under
Tamayo’s collection number 1430.
Additional specimens examined: Venezuela. Lara:
Mora
´n, Pa
´ramo de los Nepes, en sabanas a ma
´sde
2000 mts, Nov 1937, Tamayo 325 (US, VEN), Lajas del
Dı
´tamo, un sector hacia la base de la vertiente oriental del
Pa
´ramo de Cende
´, 2900 m, 10 Jun 1971, Ruiz-Tera´n and
Lo´ pez-Figueiras 2026 (MERF, US), pica que va desde
Buenos Aires a Paramo Las Rosas, 2300–2600 m, 15 Nov
1984, van der Werff and Rivero 7997 (MO, PORT). Parque
Nacional Dinira, ladera del Pa
´ramo de Jabo
´n, vertiente
hacia el Tocuyo, sector ‘‘Los Charquitos,’’ 2800–2900 m,
15 Aug 1999, Riina et al. 704 (VEN), ladera vertiente hacia
el Tocuyo, sector ‘‘La Lajita,’’ 2700 m, 15 Aug 1999, Riina
et al. 722 (M, NY, VEN), turberas del Pa
´ramo Cende
´,
serrania de Barbacoas, 2715 m, s.d.,Alvarado et al. 3377
(UCOB [image!]). Trujillo: Pa
´ramo de las Rosas, 3200 m,
Oct 1912, Jahn 166 (US); Carache, inmediaciones del
Paramo Cende
´, 3000 m, 21 May 1987, Rivero 1185 (MO,
PORT).
Drosera peruensis T.Silva & M.D.Correa. Novon 12(4):
543. 2002.—TYPE: Peru. Pasco: Oxapampa, Cordillera
Yanachaga, Cerro Pajonal, ‘‘chacos’’ 12 km SE of Oxa-
pampa, 2700–2800 m, 7 Oct 1982, Foster 9066 (holotype:
PMA [image!]; isotypes: F!, HUEFS!, MO!, US!)
(Figs. 5e–g, 6).
Etymology: Named after the country of Peru, where it
was first discovered and to which it was believed to be
endemic (Silva and Correa 2002).
Description: Perennial rosetted herbs, forming upright
columns densely covered by the persistent dead leaves, up
to 50(–80) mm long, rarely branching. Roots terete, dark,
sparsely branched. Indumentum of white eglandular tri-
chomes (bronze colored when old) on abaxial surfaces of
petiole and lamina, on scape, pedicels, and sepals; minute
sessile glands c. 0.03 mm in diameter on both leaf surfaces,
scape, pedicels and sepals. Leaves 7–14(–20–28) mm long,
with circinate vernation, semierect, decumbent and per-
sistent when old, narrowly oblong, apex rounded, petiole
green to yellowish-green becoming red toward the apex,
lamina red; petioles 4.5–8(–14–19) 90.35–0.7(–1.2) mm,
with slightly broader base, narrowing toward apex, margins
revolute, adaxial surface glabrous, abaxial surface densely
eglandular-pilose; lamina 2.5–6(–9) 90.5–1(–1.8) mm,
oblong to narrowly oblanceolate, adaxial surface covered
with numerous red, carnivorous, capitate tentacles with
radial symmetry, abaxial surface densely eglandular-pilose;
stipules 5–7 91.5–3 mm, triangular, membranaceous,
white to reddish in color (drying gold-colored), the apical
half divided into three segments, the marginal segments
narrower, longer, and entire, the central segment broader,
shorter and divided into several laciniae. Scape 1 per plant,
50–150(–215) mm long (including floriferous part), terete,
0.6–10 mm in diameter at base, base erect, sparsely
eglandular-pilose from base to apex, red in color; inflo-
rescence a simple scorpioid cyme, bearing 1–7(–10)
flowers; bracts caducous, 1–2 mm long, linear-lanceolate,
eglandular-pilose; pedicels 1–4(–10) mm long, inserted
1–5(–11) mm apart, sparsely eglandular-pilose; sepals
2–5 91.7–2.5 mm, united at basal 1/3–1/4 of length,
ovate to broadly ovate, densely eglandular-pilose, apex
obtuse to rounded (acute); petals 6–7 mm long, obovate,
white or tinged light pink; stamens 5, ca. 3.5 mm long,
anthers ca. 1 mm long, bithecate, yellow; ovary
3-carpellate, fused, ca. 1.5 mm in diameter, globose,
slightly 3-lobed in outline; styles 3, forked at the base, ca.
3 mm long (including stigmata), stigmata bilobed or
1428 P. M. Gonella et al.
123

flabellate; fruit a dry dehiscent capsule, 3 mm long,
ellipsoid, 3-valvate; seeds ellipsoid, ca. 0.7 90.2 mm
wide, testa reticulate, black.
Phenology:Drosera peruensis was collected with flowers
and fruits in January, August, September, and October,
which suggests somewhat continuous flowering throughout
the year.
Habitats:Drosera peruensis grows in quartzitic sandy
soils or in cracks of sandstone in humid areas of dwarf
forest (shrubland). At the Cerro Plateado locality in
Ecuador, D. peruensis occurs in open, herbaceous vegeta-
tion on white sandy soil derived from sandstone, together
with other low herbaceous plants such as Gentianella
saxifragoides (Kunth) Fabris (Gentianaceae), Lysipomia
sparrei Jeppesen (Campanulaceae), and Pinguicula
Fig. 6 Drosera peruensis
ahabit depicting a specimen
from Sira Mountains, Peru,
bhabit depicting a specimen
from the type location at
Oxapampa, Peru, cstipule with
leaf base attached, dleaf with
stipule attached, edetail of the
indumentum of the abaxial
petiole surface, fpedicel and
calyx. abased on Graham 5751;
bbased on Foster 9066 (isotype
F); c–fbased on Graham 6038
A revision of Andean Drosera (Droseraceae) 1429
123

calyptrata Kunth (Lentibulariaceae) (D. Neill personal
observation).
Distribution area: Peru (Cajamarca, Pasco, Ucayali) and
Ecuador (Zamora-Chinchipe). Drosera peruensis exhibits a
wide, but apparently disjunct distribution along the sub-
Andean cordilleras in central and northwestern Peru and
southern Ecuador (Fig. 1). Silva and Correa (2002)
described the species as being endemic to the Cordillera
Yanachaga, in Central Peru (Pasco), but the new records
presented here expand its range ca. 150 km further to the
northeast on the Sira Mountains (Ucayali) and ca. 750 km
to the northwest on the Cerro Picorana (Cajamarca, NW
Peru) and on a southern portion of the Cordillera del
Co
´ndor, the Cerro Plateado (S Ecuador, close to the border
with Peru), ca. 85 km south from the closest known
occurrence of D. condor.
Consevation status: Endangered: EN B2ab(iii, iv). Dro-
sera peruensis was previously considered as Critically
Endangered (Duno de Stefano and Silva 2001)andEndan-
gered (Leo
´netal.2006) based on its restricted occurrence
and the state of conservation of the type location. However,
the new records presented here expand its distribution con-
siderably, to relatively well-conserved areas. Nevertheless,
the conservation status for D. peruensis is here maintained
as Endangered, meeting the criteria B2a, b(iii, iv). The
species has an estimated AOO of 20 km
2
, occurring in
narrow and highly fragmented habitats. The conservation
status is further aggravated by the known reduction in the
population size, at least at the type location, where it is
apparently extinct, although the habitat seems to not have
suffered from human interference (F. Rivadavia and S.
Vieira personal observation). In both countries, D. peruensis
occurs in protected areas: the Sira Comunal Reserve in Peru
and the Cerro Plateado Biological Reserve in Ecuador.
Notes: Silva and Correa (2002) state that the species
resembles the Venezuelan D. arenicola Steyerm., due to
the superficially similar oblong leaves (further similarity
comes from the white petal color; Fig. 5g). Drosera
arenicola is distinguished by the geniculate leaf vernation
(vs. circinate), by the entirely red leaves (vs. bicolored), by
the oblong-obovate lamina (vs. oblong to narrowly
oblanceolate), the shorter and rectangular stipules up to
5 mm long (vs. longer triangular stipules, 5–7 mm long),
by the shorter scapes [up to 55 mm long vs. 50–105(–215)
mm long], and the exclusively glandular-pilose scapes and
sepals (vs. exclusively eglandular-pilose).
The compact, globose rosettes with narrowly oblong, semi-
upright, bicolored leaves of D. peruensis are superficially
reminiscent of those of the Australasian D. neocaledonica
Raym.-Hamet (species endemic to the island of New Caledo-
nia), but such similarities may not reflect a closer phylogenetic
affinity, as D. peruensis is likely to be a derived member in the
SouthAmericancladeofDrosera (with triangular stipules and
circinate leaf vernation), whereas D. neocaledonica (rectangu-
lar stipules and geniculate leaf vernation) branches earlier in the
generic phylogeny of Rivadavia et al. (2003).
Plants of D. peruensis from the Sira Mountains (central
Peru) are much larger when compared to specimens from
other known populations (compare Figs. 6a with b, 5e with
f), usually twice the size. It is currently unknown to us if
this size difference is a result of genetic differences, or just
the effect of environmental conditions (e.g., elevation or
soil), since the Sira Mountains are at lower elevation,
around 2000 m (vs. ca. 2800 m for other D. peruensis
populations).
Additional specimens examined: Ecuador. Zamora-
Chinchipe: Nangaritza, Cordillera del Co
´ndor, Reserva
Biolo
´gica Cerro Plateado, 2790 m, 24 Aug 2012, Neill
et al. 17459 (ECUAMZ, MO). Peru. Cajamarca: San
Ignacio, San Jose
´de Lourdes, cerro Picorana, 2830 m, 17
Aug 1998, Campos et al. 5540 (HUEFS, MO, USM),
campamento Zural, base del Cerro Picorana, 2200 m, 23
Jan 1999, Diaz et al. 10466 (HUEFS, MO). Ucayali: Cor-
onel Portillo, Iparia, Alturas del Cerro Ariapo, Reserva
Comunal el Sira, 2000 m, 16 Dec 2009, Graham 5751 (F),
20 Sep 2010, Graham 6038 (F, SPF).
Acknowledgments We would like to thank Ricarda Riina (Real
Jardı
´n Bota
´nico, Madrid), Hipo
´lito Alvarado A
´lvarez (UCOB), and
Re
´gulo Bricen
˜o for information and photos of D. cendeensis from
Dinira National Park, Venezuela; Jose
´Ramo
´n Grande Allende
(Universidad Central de Venezuela, Caracas) for photos of D. cen-
deensis specimens from MERF and VEN and information about
possible specimens at MER and MERC; Nidia Cuello (Universidad
Nacional Experimental de los Llanos Occidentales, Venezuela) for
photos of specimens at PORT; James G. Graham (Field Museum,
Chicago) for the photos of D. peruensis from Sira Mountains, Peru;
the curators and staff of the visited herbaria; and two anonymous
reviewers for valuable suggestions to the manuscript. DAN thanks the
Ministry of Environment of Ecuador for the research permits enabling
fieldwork in the Cordillera del Co
´ndor. This work is part of PMG’s
Ph.D. thesis, financially supported by Conselho Nacional de Desen-
volvimento Cientı
´fico e Tecnolo
´gico (CNPq – Proc. 140135/2013-8)
and Coordenac¸a
˜o de Aperfeic¸oamento de Pessoal de Nı
´vel Superior
(CAPES – PDSE/Proc. 99999.010612/2014-09). PTS acknowledges
CNPq Research Grant (Proc. 310437/2015-6).
Compliance with ethical standards
Conflict of interest The authors declare that they have no conflict of
interest.
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