156 (1): 1 phytotaxa Article - Plantas Carnívoras villosa - complexo.pdf · PAULO MINATEL GONELLA....

PHYTOTAXA

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2014 Magnolia Press

Phytotaxa 156 (1): 1–40 (2014)

www.mapress.com/phytotaxa/Article

http://dx.doi.org/10.11646/phytotaxa.156.1.1

Exhuming Saint-Hilaire: revision of the Drosera villosa complex (Droseraceae)

supports 200 year-old neglected species concepts

PAULO MINATEL GONELLA1, FERNANDO RIVADAVIA2, PAULO TAKEO SANO3 & ANDREAS

FLEISCHMANN4

1 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: [email protected] 2 1 Daniel Burnham Ct., San Francisco, 94109, USA; e-mail: [email protected] 3 Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, CEP 05508–

900, São Paulo, Brasil; e-mail: [email protected] 4 LMU Munich, Systematic Botany and Mycology, Menzinger Strasse 67, D-80638 Munich, Germany;

e-mail: [email protected]

Abstract

The Drosera villosa complex is here reviewed and includes six species endemic to Brazil: D. villosa, here identified for

the first time as a narrow endemic species native to the neighboring highlands of the Serra Negra and Serra do Ibitipoca,

in southern Minas Gerais state; D. ascendens, rediscovered nearly 200 years after its description, narrowly endemic to

the Diamantina Plateau, central Minas Gerais; D. graomogolensis, endemic to northern Minas Gerais, but here found to

be more widespread than previously reported; D. latifolia, a highly polymorphic and widespread taxon, previously

placed in synonymy of D. villosa and heretofore misidentified as D. ascendens, is here elevated to species rank; and two

new species here described, D. riparia and D. chimaera. Furthermore, two new natural hybrids are reported: D. villosa ×

D. tomentosa var. glabrata and D. latifolia × D. tomentosa. The morphological characters distinguishing these taxa from

each other and from similar species are discussed, together with habitat and ecological information, detailed illustrations

and photographs, distribution maps, and a key to the species of the D. villosa complex is provided.

Key words: Brazil, carnivorous plants, Chapada Diamantina, Espinhaço Range, new species

Resumo

O complexo Drosera villosa é aqui revisado e é composto por seis espécies endêmicas do Brasil: D. villosa, aqui

identificada pela primeira vez como uma espécie endêmica das vizinhas Serra Negra e Serra do Ibitipoca, no sul de

Minas Gerais; D. ascendens, redescoberta após quase 200 anos, micro-endêmica no Planalto de Diamantina, no centro de

Minas Gerais; D. graomogolensis, endêmica do norte de Minas Gerais, porém aqui considerada mais amplamente

distribuída do que reportado anteriormente; D. latifolia, um táxon altamente polimórfico e amplamente distribuído,

anteriormente colocado em sinonímia de D. villosa e até então erroneamente identificado como D. ascendens, é aqui

elevado ao status de espécie; e duas novas espécies que são aqui descritas, D. riparia e D. chimaera. Dois novos híbridos

naturais são reportados: D. villosa × D. tomentosa var. glabrata e D. latifolia × D. tomentosa. As características

morfológicas que distinguem esses táxons uns dos outros e de espécies similares são discutidos, juntamente com

informações sobre habitat e ecologia, ilustrações detalhadas e fotografias, mapas de distribuição e uma chave para as

espécies do complexo D. villosa é apresentada.

Palavras-chave: Brasil, Cadeia do Espinhaço, Chapada Diamantina, espécies novas, plantas carnívoras.

Accepted by Duilio Iamonico: 28 Nov. 2013; published: 3 Jan. 2014 1

Introduction

The cosmopolitan sundews of the genus Drosera Linnaeus (1753: 281) (Droseraceae Salisb.) comprise nearly 200 species, with ca. 100 of these endemic to southwestern Australia (McPherson 2010). After the works of Saint-Hilaire (1824, 1826), the genus Drosera has been hitherto very poorly researched in Brazil, with most recent studies, such as Silva & Giulietti (1997) and the generic treatment in Flora Neotropica (Correa & Silva 2005), lacking substantial field work, relying mostly on herbarium specimens – which are often badly deformed due to the delicate nature of most Drosera taxa. As a result, these and most other previous studies failed to fully describe the diversity of the genus in Brazil, resulting in a small number of species currently registered for the country: only 14 are noted in the online Lista da Flora do Brasil (Silva 2013), the most extensive work on the Brazilian flora since Martius’ Flora Brasiliensis (produced between 1840 and 1906).

Nonetheless, several recent studies suggest that Brazil hosts a much greater diversity of Drosera (Santos 1989, Rivadavia 2003, 2008, 2009, Fleischmann et al. 2007, Rivadavia et al. 2009, Rivadavia & Gonella 2011, Gonella et

al. 2012). We estimate this number to be closer to 30 taxa, of which 18 belong to the so-called “Brazilian-tetraploid clade” (Rivadavia et al. 2003), characterized by a chromosome number of 2n=40, circinate leaf vernation, and flower scapes usually erect at the base.

The species complex of the affinity of D. villosa Saint-Hilaire (1826: 267) belongs to the tetraploid clade, together with D. camporupestris Rivadavia (2003: 85), D. chrysolepis Taubert (1893: 505), D. graminifolia Saint-Hilaire (1826: 269), D. grantsaui Rivadavia (2003: 82), D. quartzicola Rivadavia & Gonella (2011: 34), D.

schwackei (Diels 1906: 89) Rivadavia (2008: 39), D. spiralis Saint-Hilaire (1826: 270) and the four species that make up the D. montana Saint-Hilaire (1826: 260) complex: D. montana, D. tentaculata Rivadavia (2003: 79), D.

tomentosa Saint-Hilaire (1826: 261), and a yet unpublished taxon.The history of the D. villosa complex is summarized below:

— During a long trip through the interior of Brazil in the early 19th century, the French naturalist Auguste de Saint-Hilaire discovered and later published several Drosera species, including D. villosa and D. ascendens

Saint-Hilaire (1826: 268); the former from the Serra Negra, southern Minas Gerais state, and the latter from the Serra de Curumatahy (currently Curimataí), near Diamantina, central Minas Gerais, southeastern Brazil (Saint-Hilaire 1826).

— Eichler (1872) added a new taxon to the group, D. villosa var. latifolia Eichler (in Martius & Eichler 1872: 395), based on material collected by Glaziou at the Serra dos Órgãos, Rio de Janeiro state, southeastern Brazil, differing from type variety by the broader leaves and shorter petioles.

— Diels (1906) considered both D. ascendens and D. villosa var. latifolia as synonyms of D. villosa in his generic revision, and this taxonomic opinion has been followed by all subsequent taxonomic treatments (e.g. Santos 1980, Silva 1994, Silva & Giulietti 1997, Duno de Stefano & Silva 2001, Correa & Silva 2005, Silva 2013). According to the concept of D. villosa proposed by Diels (1906), this species could be recognized by the oblanceolate-linear leaves and fusiform-curved seeds.

— Buxbaum (1924) published D. villosa var. bifurca Buxbaum (1924: 119), differing from the type variety by the bifurcated scapes, and small differences in sepal shape.

— Silva (1997) published a new species, D. graomogolensis Silva (1997: 85), based on material collected at Grão Mogol, northern Minas Gerais state, eastern Brazil. The author considered the species endemic to Grão Mogol and very similar to D. villosa, differing in the seed and style shapes, as well as in the length of the upright stem.

— Rivadavia et al. (2003), Rivadavia (2005), Rivadavia (2008), and Rivadavia (2009) considered D. ascendens

and D. villosa as separate species, but did not mentioned the distinctive characteristics or the respective distributions.

— In 2007 a new taxon belonging to this complex was discovered in northern Minas Gerais, and a further very unusual taxon was uncovered in central Minas Gerais in 2010. These discoveries raised doubts about the true identity of D. ascendens, and eventually led to a surprising reversal in the taxonomic views previously held by the current authors.The aim of this study is to present a detailed morphological and ecological circumscription of the taxa of the D.

villosa complex, including: I. Identify and circumscribe D. villosa, as described by Saint-Hilaire (1826);II. Define the taxa allied to D. villosa based on morphological evidence; III. Evaluate the distribution range of each taxon;

GONELLA ET AL.2 • Phytotaxa 156 (1) © 2014 Magnolia Press

IV. Present a complete botanical description of all taxa recognized as belonging to this complex, including descriptions of eventual new species.

Material & Methods

Extensive field studies were undertaken by the authors between 1990 and 2013 across the known natural range of D. villosa complex populations throughout eastern Brazil. In addition, herbarium specimens at B, BHCB, CESJ, F, GFJP, HB, HRB, M, MBM, MO, NY, OUPR, P, R, RB, SP, SPF, UEC, and US (herbarium acronyms according to Thiers 2011) were personally examined and annotated by at least one of the authors. Type materials held at P, R, and SPF were personally examined under stereo dissecting microscope. Digitized high-resolution images of the type materials held at K, P, and WU 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 2012).

Appendages are divided into trichomes and emergences. To avoid confusion, the glandular vascularized emergences present on the margins and adaxial lamina surface are referred in the text as “tentacles”. Three types of trichomes are found in the species here treated, and the following terminology is used (adapted from Seine & Barthlott 1993): — “eglandular-trichomes”: non-glandular trichomes;— “sessile glands”: short glandular trichomes with inconspicuous stalk, usually 0.03–0.05 mm in diameter;— “glandular trichomes”: glandular trichomes with conspicuous stalk, usually longer than 0.2 mm in length.

Results

Considerable taxonomic confusion has surrounded the name Drosera villosa for over 100 years, since the proposal by Diels (1906) to include D. ascendens and D. villosa var. latifolia in this taxon. Plants matching the type of D.

villosa have remained for the most part unknown since the species’ discovery nearly 200 years ago by Saint-Hilaire. The main reason for this was its restricted geographical distribution, lack of field studies, the paucity of herbarium specimens, and difficulty to access type material from P. Also, the name of the type location, “Serra Negra”, is extensively used for other mountain ranges across Brazil, which helped keep the type locality in obscurity until multiple new collections were made over the past two decades, including at the type location.

As a result, since Diels (1906) most taxonomists and carnivorous plant enthusiasts worldwide have mistaken D. villosa for another member of this complex. These two taxa are so different that one of the authors (FR) even believed to have discovered a new species at Serra do Ibitipoca in 1992, different from what was then thought to represent D. villosa (viz. D. ascendens).

The true nature of D. villosa was first evident to one author (FR) in 1996, when Saint-Hilaire’s types were personally examined at P, and this is here discussed in detail for the first time. Nevertheless, the identity of D.

ascendens was missed for yet another 15 years, during which a widespread taxon in the D. villosa complex temporarily usurped the identity of D. ascendens.

Due to the similarity in shape and indumentum of the leaves, most specimens treated as D. villosa by Correa &

Silva (2005) and previous works of the 20th century were wrongly considered as conspecific with D. ascendens by the current authors. This view was also increasingly adopted by carnivorous plant enthusiasts since the mid 1990’s, the exception being the actual type D. villosa and a handful of specimens from Ibitipoca.However, the discovery in 2010 of unusual plants similar to D. graomogolensis, but with ascending scapes bearing unique indumentum, growing near the type location of D. ascendens raised a series of taxonomic questions regarding the true identity of the latter taxon. Careful re-inspections of photographs obtained from P of the holotype and isotypes of D. ascendens, as well as subsequent field studies in 2011, helped reveal that what was initially believed to be a new species native to the Diamantina Plateau in Minas Gerais state, was in fact the long-neglected D. ascendens as described by Saint-Hilaire (1826).

Phytotaxa 156 (1) © 2014 Magnolia Press • 3REVISION OF THE DROSERA VILLOSA COMPLEX

With the realization that true D. ascendens had apparently never been recollected since its original discovery

by Saint-Hilaire in the early 19th century, came the understanding that all populations more recently recognized as D. ascendens, comprised a separate species that required a name. Eichler’s D. villosa var. latifolia from Rio de Janeiro is the earliest name published for plants belonging to this widespread taxon, distributed along nearly 1500 km from the Santa Catarina-Rio Grande do Sul border in southern Brazil to the Serra do Cabral in central Minas Gerais. Considering its distinctive, well-circumscribed morphology, different from all remaining members of the D. villosa complex, it is here raised to species rank, as D. latifolia (Eichler) Gonella & Rivadavia (see taxonomic treatment).

Furthermore, two new species are here described, both belonging to the D. villosa complex. The first was discovered in 1992 at the Chapada Diamantina by FR and it has since then been studied at several other locations along this mountain range. The second new species was first discovered in 2007, in northern Minas Gerais by the carnivorous plant enthusiast Adilson Peres. Morphologically intermediate between members of the D. villosa and D. montana complexes, and initially thought to constitute a natural hybrid, the latter was first recognized as a new species during field studies in 2010 and 2011.

The D. villosa complex, as redefined here, comprises six species endemic to Brazil (Figures 1−2), which are commonly characterized by semi-erect, oblong to lanceolate leaves, small, rectangular stipules, and a lack of translucent-yellow short-stalked globose trichomes (see Rivadavia & Gonella 2011). Their distribution ranges are presented (Figures 1−2), and the diagnostic morphological characters are compared in Table 1.

Key to the species of the Drosera villosa complex

1. Petiole longer than one third of total leaf length.................................................................................................................... 2- Petiole shorter than or equal to one third of total leaf length................................................................................................. 52. Plants with base of scape curved (ascending); lower half of scape densely covered by long (up to 1.8 mm) glandular

trichomes decreasing in density and size towards the apex; seeds ellipsoid ...................................................... D. ascendens

- Plants with base of scape erect; lower half of scape glabrescent to densely covered by eglandular trichomes and/or sparse short (up to 0.3 mm long) glandular trichomes, glandular trichomes increasing in density and size towards the apex; seeds fusiform or obconical ............................................................................................................................................................. 3

3. Both surfaces of the petiole densely eglandular-pilose; lamina narrowly oblong to narrowly lanceolate; scapes densely eglandular-pilose on basal third of length .................................................................................................................D. villosa

- Abaxial petiole surface glabrescent to densely eglandular pilose, adaxial surface glabrous to glabrescent; lamina oblong to lanceolate; scapes lacking eglandular trichomes or, if present, then very sparse .................................................................. 4

4. Petiole equal to or longer than the lamina, as well as distinct from it, narrower; seeds obconical ......................... D. riparia

- Petiole shorter than the lamina, grading into it and of near equal width; seeds fusiform .......................................D. latifolia

5. Glandular trichomes uniformly dense along the whole length of scape; styles 4–4.5 mm long (stigmata included) ................................................................................................................................................................................. D. graomogolensis

- Glandular trichomes absent or very sparse at the lower half of scape, increasing in density towards the apex; styles only up to 3 mm long (stigmata included)........................................................................................................................................... 6

6. Basal third of scape densely eglandular-pilose; seeds ovoid up to 0.4 mm long ................................................. D. chimaera

- Basal third of scape glabrous or sparsely eglandular-pilose; seeds fusiform 0.8–1 mm long ................................ D. latifolia

Taxonomic treatment

Drosera villosa A.St.-Hil. (Figures 1, 3, 4)Type:—BRAZIL. Minas Gerais: Crescit in sabuletis humidis montis dicti Serra Negra in provinciae Minas Geraes, haud longè

à finibus provinciae Rio de Janeiro, January−February 1817, Saint-Hilaire C1-75 (holotype P-578095!, isotypes K parte

image!, P-578097!, P-578099!).

Description:—Perennial rosetted herbs, acaulescent or forming short upright stems up to 50 mm tall, sparsely covered by persistent dead leaves. Roots slightly succulent, up to 0.5 mm wide, sparsely branched, densely covered by root hairs. Indumentum consisting of white to red (bronze-colored when old) eglandular trichomes up to 1.8 mm long on leaves, scapes, and sometimes on pedicels, bracts and sepals abaxially; short glandular capitate trichomes 0.05–0.3 mm long, the stalk light red and translucent, the head dark red, present on scapes, pedicels, bracts and


sepals abaxially; minute sessile glands ca. 0.03 mm in diam., red in color, densely covering both leaf surfaces, scapes, pedicels and sepals abaxially. Leaves 17–115 mm long, with circinate vernation, semi-erect, persistent when old, linear-lanceolate, apex obtuse to acute, green to wine-red in color; petiole 6.5–58 mm long, 0.5–1.6(–2) mm wide, linear, transversely elliptic in cross section, densely eglandular-pilose on both surfaces, somewhat denser on the abaxial surface; lamina 10.5–60 mm long, 1.4–4 mm wide, narrowly oblong to narrowly lanceolate, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface densely to sparsely eglandular-pilose, especially towards the base and along the margins; stipules (1.8–)3–6.5 mm long, 1–2.5 mm wide at the base, rectangular, membranaceous, pinkish to red (drying bronze colored), the apical 1/2 divided into several long laciniate segments, the marginal segments longer. Scapes 1–2(–3) per plant, 90–355 mm long (including floriferous part), terete, 0.4–2 mm in diameter at the base, erect base, densely eglandular-pilose on basal third (decreasing in density towards the apex), densely glandular-pilose on the apex (decreasing in size and density towards the base), green to wine-red in color; inflorescence a scorpioid cyme, rarely bifurcating, bearing 2–30 flowers; bracts 1.8–4 mm long, filiform-lanceolate, densely glandular-pilose, often sparsely eglandular-pilose, caducous, and thus often absent on mature scapes; pedicels 2–6 mm long, inserted 4–9 mm apart from each other, densely glandular-pilose and often sparsely eglandular pilose; sepals 5, (2.8–)3.3–5.8 mm long, 1–1.8 mm wide, oblong-lanceolate to lanceolate, united at basal 1/3 of length, densely glandular-pilose, more rarely with few eglandular trichomes; petals 5, 5–8 mm long, 3.5–4.5 mm wide, obovate, pale pink to whitish in color; stamens 5, 3.5–4 mm long, anthers ca. 1 mm long, bithecate, yellow; ovary 3-carpellate, fused, 1–1.2 mm in diameter, globose, slightly 3-lobed in outline; styles 3, forked at the base, 2.5–3 mm long (including stigmata), style arms curving inwards at the end of anthesis, stigmata bilobed to flabellate, pale pink to whitish in color; fruit a dry dehiscent capsule, ellipsoid, 2–2.5 mm long, 3-valvate; seeds fusiform, 0.7–0.8 mm long and ca. 0.2 mm wide, testa reticulate, black.

Illustrations:—Ferrero & Mello-Silva (2011), p. 17, fig. 2A–C.Distribution and ecology:—Drosera villosa is here newly recognized to be a narrow endemic species,

occurring in montane to high-montane habitats at elevations between (1200–)1400–1750 m in the two small highlands of the Serra do Ibitipoca and Serra Negra, ca. 30 km apart from each other. These are located in southern Minas Gerais, near the border with Rio de Janeiro state, and are both part of the larger Serra da Mantiqueira highlands (Fig. 1). At lower elevations (approximately up to 1500 m) D. villosa occurs mostly along the margins of perennial rivers with the moss Sphagnum Linnaeus (1753: 1106), or in cracks of quartzitic rocks (Fig. 4A), occasionally semi-shaded by riparian vegetation. At higher elevations (usually above 1500 m), it grows in seepages or damp areas of “campo rupestre” and “campo de altitude” vegetation, often associated with Sphagnum species (Fig. 4B). Drosera villosa also colonizes disturbed sites such as trail sides, where it may form large populations, sometimes in relatively dry sandy areas that depend mostly on condensation from fog for humidity.

Drosera tomentosa var. glabrata Saint-Hilaire (1826: 262) has been observed growing sympatric with D.

villosa at several sites at the Serra do Ibitipoca, where these two taxa have even been found to form a rare, newly discovered, natural hybrid (Fig. 5A, B). Drosera villosa was also observed growing near D. montana and D.

communis Saint-Hilaire (1824: t. XV(1)) at a few sites at Ibitipoca, and it also co-occurs with a few species of Utricularia Linnaeus (1753: 18) (Lentibulariaceae Rich.) on both mountain ranges, as well as with Genlisea

violacea Saint-Hilaire (1833: 431) (Lentibulariaceae) occasionally at Ibitipoca.Phenology:—Drosera villosa was collected in flower and fruit from October to February, which corresponds

to the wet season. Flowers of this species (as with most Drosera spp., and all taxa treated in this article) last for only one day, opening early on sunny days and usually closing around midday.

Conservation status:—Only a single population of D. villosa is currently known from the Serra Negra, which is the type location and where it was only recently rediscovered and recollected (2006 and 2009), after nearly 200 years. Although numerous populations have been identified at the Parque Estadual da Serra do Ibitipoca since 1987 (with only one previous collection from 1896), this park is a small fragile island of native vegetation surrounded by farms and ranches. Also, the most numerous populations of this species are concentrated along trail margins, where there is less competition for light, but the plants face higher risk of human disturbance. Duno de Stefano & Silva (2001) considered D. villosa as Least Concern, however their classifaction stemmed from a broader circumscription of this species (sensu Diels 1906), including the widespread D. latifolia. Based on its newly recognized narrow endemism, we here propose to reclassify D. villosa as Vulnerable (VU, criteria B2a, biii; IUCN,

2001).


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FIGURE 1. Distribution map of Drosera villosa (triangles) and D. latifolia (circles) in southern and southeastern Brazil.


FIGURE 2. Distribution of Drosera ascendens (triangles), D. chimaera (inverted triangles), D. graomogolensis (circles), and D.

riparia (squares) in eastern Brazil.

Discussion:—Drosera villosa is easily distinguished from other members of the complex by its narrow leaves with elongated petioles often as long as the lamina and always exceeding 50% of the lamina length (petiole/lamina ratio higher than 0.5), narrow petioles that are transversely elliptic in cross section and densely eglandular pilose on both surfaces, leaf lamina narrowly oblong to narrowly lanceolate, basal third of the scape densely eglandular-pilose, relatively small to medium-sized flowers (petals 5–8 mm long) with short styles (2.5–3 mm long), and seeds fusiform 0.7–0.8 × 0.2 mm (Figs. 3, 4). General morphology and seed shape suggest that D. villosa may be most closely related to D. latifolia (Table 1).


FIGURE 3. Drosera villosa. A, habit. B, detail of the indumentum of the base of the scape. C, detail of the indumentum of the apex of

the scape. D, stipule. E, leaf, adaxial surface. F, leaf, abaxial surface. G, cross section of the petiole at mid length. H, calyx. J, petal. K,

stamen. L, gynoecium. M, seed. (Based on Ferrero et al. 02 – drawn by P.M. Gonella.)


FIGURE 4. Drosera villosa. A, habitat, in cracks of sandstone along the margin of a river at Parque Estadual da Serra do Ibitipoca,

Minas Gerais. B, habitat, along trailside in high-montane campo rupestre at Parque Estadual do Ibitipoca, Minas Gerais. C, habit. D,

detail of the center of the rosette, showing the densely eglandular-pilose petioles and emerging flower scape. E, flower. (A–D by P.M.

Gonella; E by A. Fleischmann.)


FIGURE 5. Hybrids. A, C, Drosera villosa × D. tomentosa var. glabrata at the Serra do Ibitipoca, Minas Gerais. B, Drosera latifolia

× D. tomentosa var. tomentosa (bottom two rosettes) and D. latifolia (top rosette) at Couto de Magalhães de Minas, Minas Gerais. D,

E, Drosera latifolia × D. tomentosa var. glabrata at Diamantina, Minas Gerais. (A, C, by Adilson Peres; B by F. Rivadavia; D, E, by

P.M. Gonella.)

Saint-Hilaire collected D. villosa twice at the Serra Negra, as he mentioned in the general generic description of Droseraceae, on page 257 (Saint-Hilaire 1826), and also evident by the existence of two collections of this species made by him with distinct numbers. During his first trip to the Serra Negra, in 1817, he collected the type specimens (Saint-Hilaire C1-75), and then he re-collected this species in 1822 (Saint-Hilaire J-146).

Line drawings and SEM images of the seed presented for D. villosa by Silva & Giulietti (1997) and Correa & Silva (2005) corresponds to D. latifolia. Similarly, the broad distribution presented for D. villosa in these and other studies refer mostly to D. latifolia. A collection of D. villosa reported from Bolivia (Schulenberg & Awbrey 1997) was examined and is here considered to be conspecific with D. montana.


Before naming this species as D. villosa, Saint-Hilaire may have considered naming it as D. montana, as is suggested by the original herbarium label and the handwritten description attached to the holotype with the epithet “montana” crossed out and substituted by the name “villosa”.

Specimens examined:—BRAZIL. Minas Gerais: Município de Bias Fortes, distrito de Ibitipoca, June 1896, Magalhães 1328 (R). Município de Lima Duarte, Parque Florestal da Serra de Ibitipoca, 25 February 1977, Coons

77-275 (VIC), 05 May 1987, Andrade & Souza 960 (BHCB), 20 July 1992, Rivadavia 143 (SPF), 30 October 1995, Rivadavia & Padovese 507 (SPF), 23 November 2001, Rivadavia 1323 (SPF), 24 November 2001, Rivadavia 1325

(SPF), 24 November 2001, Rivadavia 1326 (SPF), 24 November 2001, Rivadavia 1327 (SPF), 26 February 2005, Rivadavia 1954 (SPF); Parque Estadual do Ibitipoca, trilha entre a Gruta do Cruzeiro e a Lombada, 1710 m, 26 January 2010, Ferrero et al. 02 (SPF); próximo à Ponte de Pedra, 15 December 2010, Gonella et al. 362 (SPF); Prainha, 15 December 2010, Gonella et al. 364 (SPF); Pico da Lombada, 16 December 2010, Gonella et al. 366

(SPF); entre a Prainha e a Ponte de Pedra, 19 June 2013, Gonella et al. 618 (SPF). Município de Rio Preto, Serra Negra, Burro de Ouro, 21 January 2006, Abreu et al. 63 (CESJ); topo da Serra do Funil, 1500 m, 27 December 2009, Gonella et al. 252 (SPF). Without attribution to administrative area, Serra Negra, 1822, Saint-Hilaire J-146

(MPU); Serra Negra, January/February 1817, Saint-Hilaire C1-75 (P; type).Specimen examined of the hybrid between D. villosa and D. tomentosa var. glabrata:—BRAZIL. Minas

Gerais: Município de Lima Duarte, Parque Estadual da Serra do Ibitipoca, Pico da Lombada, 16 December 2010,Gonella et al. 367 (SPF).

Drosera ascendens A.St.-Hil. (Figures 2, 6, 7)Type:—BRAZIL. Minas Gerais: Crescit in sabuletis humidis montium dictorum Serra de Curumatahy, parte boreali provinciae

Minas Geraes dicta Distrito dos Diamantes; alt. circiter 3700 ped., September 1817, Saint-Hilaire B1-1988 (holotype P-578094!, isotype P-578098!, B!).

Description:—Perennial rosetted herbs, forming upright or decumbent stems up to 125 mm tall, densely covered by persistent dead leaves; rarely plants acaulescent. Roots slightly succulent, up to 2 mm wide, sparsely branched, densely covered by root hairs. Indumentum consisting of white (gold-colored to brown when old) eglandular trichomes 0.8–1.5 mm long, present on the abaxial leaf surface (increasing in size and density towards the base); short to long glandular capitate trichomes 0.2–1.8 mm long, the stalk and head deep red in color, present on entire scape (increasing in size and density towards base), pedicels, abaxial surface of bracts and sepals; minute sessile glands ca. 0.05 mm diameter, red in color, with translucent head, present on both leaf surfaces, scapes, pedicels and sepals abaxially (especially dense on adaxial petiole surface and towards the apex of the inflorescence). Leaves 14–42 mm long, with circinate vernation, semi-erect, persistent when old, lanceolate, apex broadly acute, deep-red to greenish-red in color; petiole 6–17 mm long, 1.5–3 mm wide, narrowing towards the base, transversely narrowly elliptical in cross section, margins revolute, adaxial surface glabrous, abaxial surface densely eglandular-pilose; lamina 8–25 mm long, 2–5 mm wide, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface densely eglandular-pilose; stipules 1.5–4 mm long, 1–1.8 mm wide at the base, rectangular, membranaceous, translucent white in color (drying gold-colored to brown), the apical 2/3 divided into several laciniate segments shortening towards the center. Scapes 1–2 per plant, (90–)110–310 mm long (including floriferous part), terete, 0.5–1.5 mm in diam. at the base, with slight to pronounced ascending curve near base, densely glandular-pilose from base to apex (glandular trichomes increasing in size and density towards the base), deep-red to greenish-red in color; inflorescence a scorpioid cyme, rarely bifurcating, bearing 4–23 flowers; bracts

caducous, extremely rarely present on mature scapes, ca. 4 mm long, linear; pedicels 3–4.5 mm long, inserted 5–11(–14) mm apart from each other, densely glandular-pilose; sepals 5, 4–6 mm long, 1–1.5 mm wide, narrowly ellipsoid to lanceolate, united at basal 1/4 of length, densely glandular-pilose, apex acute to obtuse; petals 5, ca. 8 mm long, 7–8 mm wide, obovate, pale pink in color; stamens 5, ca. 3.5 mm long, anthers ca. 1.5 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. 4 mm long (including stigmata), style arms remaining straight at the end of anthesis, stigmata bilobed or flabellate, pale pink to whitish in color; fruit a dry dehiscent capsule, ca. 2 mm long, ellipsoid, 3-valvate; seeds ellipsoid, ca. 0.5 mm long and ca. 0.2 mm wide, testa reticulate, black.


Distribution and ecology:—Drosera ascendens is a narrowly endemic taxon occurring on the northern part of the Diamantina Plateau, in central Minas Gerais, Brazil (Fig. 2). This species occurs in campo rupestre vegetation, mostly in open fields with sparse grasses around seasonal springs and along the margins of seasonal or perennial rivers in acidic sandy soil or on a thin layer of sandy soil overlying sandstone rock (Fig. 7A). It was found at both a lowland site at ca. 730 m near the village of Inhaí in the municipality of Diamantina, and at a few highland locations at ca. 1300 m inside the Sempre Vivas National Park, in the municipality of Buenópolis, just north of the village of São João da Chapada, where it was observed growing sympatrically with D. camporupestris, D.

communis, D. grantsaui, D. hirtella Saint-Hilaire (1826: 262) var. hirtella, D. montana, D. spiralis, D. tentaculata

and D. tomentosa (both varieties), as well as D. latifolia (totaling 11 taxa). The high alpha-diversity of Droseraceae corroborates the creation and maintenance of this newly created National Park, which comprises the area with the highest concentration of Drosera spp. known in the New World, the Diamantina Plateau, containing a total of 13 taxa, followed by the Serra do Cipó (12 taxa) and the eastern Gran Sabana of southern Venezuela and Guyana (ca. 11 taxa).

Phenology:—Based on the few existing collections, D. ascendens is only known to bloom during the height of the dry season, from July to September.

Conservation status:—Drosera ascendens was only recently rediscovered in 2010, after nearly 200 years of obscurity, and is here newly recognized as a narrowly endemic species. The extent of its distribution can only tentatively be assumed as spanning the northern tip of the Diamantina Plateau – a relatively isolated area ca. 30 km wide by ca. 50 km long. Only four populations were studied by the authors during two short visits, but D. ascendens

is assumed to be common in this region, based on population density and availability of suitable habitats. Although D. ascendens is officially protected within the borders of the Sempre Vivas National Park, our current knowledge of its geographical distribution is insufficient to allow an adequate assessment of its conservation status. It is thus here considered as Data Deficient (DD), according to IUCN Red List criteria (IUCN 2001).

Discussion:—Drosera ascendens is distinguished from all other members of the complex by its lanceolate leaves with broad petioles that are only slightly shorter than the lamina; in combination with a glabrous adaxial petiole surface; an abaxial leaf surface that is densely covered by white eglandular trichomes; an arcuate, curved (ascending) base of the scape; scape, pedicels and sepals densely covered by long (up to 1.8 mm long) red glandular trichomes that increase in size and density towards the base of the scape; large flowers with petals ca. 8 mm long, styles ca. 4 mm long; and ellipsoid seeds (Figs. 6, 7).

Drosera ascendens appears to be most closely related to D. graomogolensis, with which it not only shares a preference for drier sandy habitats, but also characters such as a dense and well-expressed eglandular indumentum on the abaxial leaf surface, scapes with a dense glandular indumentum starting from the base, usually absent bracts, sepals united to 1/5–1/4 of their length (only fused to 1/4–1/2 of their length in the other species of the complex), large petals, and long styles (compared to the remaining species of the complex), small ellipsoid seeds, a contemporary flowering period in the dry season, the habit of commonly forming semi-erect stems covered by columns of dead leaves, and relatively thick roots. However, D. ascendens can be easily distinguished from D.

graomogolensis by its longer petioles, which are 6–17 mm in length, and which have their margins revolute (curved downwards); by scapes always arcuately curved from the base (ascending), and especially by the longer glandular trichomes up to 1.8 mm long present on the inflorescence parts (Table 1).

The dense indumentum of long red glandular trichomes present on the inflorescence (Fig. 7B) is the most distinctive character of D. ascendens, a feature not found in any other New World Drosera species. The study of dried specimens under a stereomicroscope suggests that the glandular trichomes of D. ascendens have a multiseriate stalk, although this is not clear due to difficulty in discerning the cells. Comparatively, other species of the complex have trichomes with easily discernable cells in the stalks, which are assumed to be uni- to biseriate, a character that requires further confirmation by anatomical studies.

Despite the unique morphology of the inflorescence trichomes in D. ascendens, they were surprisingly not highlighted in Saint-Hilaire’s original description (as well as in all subsequent studies), and were described as mere eglandular trichomes. Although the type specimen has lost most of the inflorescence indumentum over time, the unique glandular trichomes are nonetheless still visible today.

The petioles with revolute margins of D. ascendens (Fig. 6H) are also unique within the D. villosa complex. This character is also expressed (to different degrees) in some Neotropical species such as D. grantsaui, D.

meristocaulis Maguire & Wurdack (1957: 332), and D. amazonica Rivadavia, Fleischmann & Vicentini (2009: 13).


FIGURE 6. Drosera ascendens. A, habit. B, detail of the indumentum at the base of the scape. C, detail of the indumentum at the apex

of the scape. D, stipule. E, leaf, adaxial surface. F, leaf, abaxial surface. G, cross section of a young petiole at mid length. H, cross

section of the petiole of a fully developed leaf at mid length. J, calyx. K, petal. L, stamen. M, gynoecium. N, seed. (All based on

Gonella et al. 460 – drawn by P.M. Gonella.)


FIGURE 7. Drosera ascendens. A, habitat at Parque Nacional das Sempre-Vivas, Minas Gerais. B, detail of the indumentum of the

scape. C, rosette seen from above, note the dense indumentum of white eglandular trichomes on the abaxial leaf surface of the young

leaves. D, side view of the rosette, with ascending scape. E, detail of the apex of the inflorescence. F, flower. (A–E by P.M. Gonella; F

by Adilson Peres.)


The lack of bracts in D. ascendens was first pointed out by Saint-Hilaire (1826), who states in the diagnosis: “pedicellis omnibus ebracteatis”. Among ca. 40 inflorescences studied on the herbarium specimens, bracts were only found on 2 inflorescences belonging to different plants, thus supporting Saint-Hilaire’s statement.

Because typical D. ascendens has remained in obscurity for nearly 200 years (it was apparently never collected after Saint-Hilaire), this name was mistakenly used for nearly two decades for specimens of the widespread D.

latifolia (Rivadavia et al. 2003, Rivadavia 2005, 2009), which shares a similar leaf shape and leaf indumentum, and which also occurs on the Diamantina Plateau. Although D. ascendens was observed at the Sempre Vivas National Park growing mostly in dry habitats consisting of sandy soil among sparse grasses, a few individuals were found in wet sandy soil along a stream, only a few meters apart from D. latifolia, which was abundant in such wet habitats along this same stream, growing on islands of vegetation or in cracks of sandstone. Although, no hybrids were observed between D. ascendens and D. latifolia, it is likely that these two species are indeed capable of hybridizing. This is the only know location in Brazil where two taxa of the D. villosa complex co-occur.

During his voyages in Brazil from 1816 to 1822, Saint-Hilaire collected D. ascendens at the Serra de Curumatahy (currently Curimataí) in September 1817, according to the species description. Drosera ascendens

received its epithet from the ascending curve present at the base of the flower scape (Figs 6A, 7D). Although a common character among diploid South American taxa such as D. communis and D. hirtella (as well as many South African sundews), D. ascendens is the only species of the Brazilian tetraploid Drosera clade that consistently presents this character (although slightly ascending scapes may occasionally be found in D. graomogolensis, D.

quartzicola and D. riparia). Eichler (1872) considered D. ascendens to be distinct from D. villosa on the basis of the following characters: leaves densely pilose abaxially, ascending scape, and sub-infundibuliform (funnel shaped) stigmata. However, he made no mention of the obvious differences in scape indumentum.

The holotype of D. ascendens (at P) was at some point labeled by T.R.S. Silva as D. canescens H.B.K. This is an unpublished, invalid name, including wrong authority, and clearly represents a misreading of the original label.

The entries named “D. ascendens” in the phylogenetic study of Rivadavia et al. (2003) as well as in the chromosome study of Rivadavia (2005), and most photographic reports of this species published online all correspond to D. latifolia.

Specimens examined:—BRAZIL. Minas Gerais: Município de Diamantina, distrito de Inhaí, ao norte de Inhaí, 05 September 2011, Gonella et al. 462 (SPF). Município de Buenópolis, Parque Nacional das Sempre Vivas, 04 September 2011, Gonella et al. 454 (SPF), Gonella et al. 455 (SPF), Gonella et al. 460 (SPF), 8 March 2013, Gonella et al. 603 (SPF), Gonella et al. 607 (SPF), Gonella et al. 612 (SPF). Without attribution to administrative area, Serra de Curumatahy, September 1817, Saint-Hilaire B1-1988 (B, P; type).

Drosera graomogolensis T.R.S.Silva (Figures 2, 8, 9, 10B, C)Type:—BRAZIL. Minas Gerais: Grão Mogol, Córrego da Bonita, subida para o Morro Jambeiro, 800 m, 07 September 1990,

Silva et al. CFCR 13506 (holotype SPF!, isotype RB!).

Description:—Perennial rosetted herbs, usually forming short stems up to 50 mm long, often forming stems up to 270 mm long in wetter habitats; stems occasionally branched, densely covered by persistent dead leaves, rarely acaulescent. Roots slightly succulent, up to 2.5 mm wide, sparsely branched, densely covered by root hairs.Indumentum consisting of white eglandular trichomes up to 2 mm long (brown when old) on the abaxial leaf surface (increasing in size and density towards the base), very rarely present on the base of the scape; glandular capitate trichomes 0.1–0.4(–0.5) mm long, the stalk light red and translucent, the head dark red, densely covering the entire scape, pedicels and sepals abaxially; minute sessile glands 0.03–0.05 mm in diam., red in color, with translucent head, densely covering both leaf surfaces, scapes, pedicels, bracts and the abaxial sepal surface. Leaves 15–40(–57) mm long, with circinate vernation, semi-erect, persistent when old, lanceolate to oblong-lanceolate, apex broadly acute to acute (rarely obtuse), green to deep-red in color (more rarely wine colored); petiole 4–7(–12) mm long, 1.5–3.5(–4) mm wide, rectangular, narrowly transversely elliptical in cross section, adaxial surface glabrous, abaxial surface densely eglandular-pilose; lamina 11–35(–45) mm long, 2.2–5 mm wide, lanceolate to oblong-lanceolate, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface densely eglandular-pilose (increasing in size and density towards the base); stipules 2–4.5 mm long, 1.2–3.5 mm wide at the base, rectangular, membranaceous, translucent white to pinkish (drying brown), the apical 1/2–2/3 divided into several laciniae. Scapes 1–3 per plant, 160–380 mm long (including floriferous part), terete, 0.6–2 mm in diameter at the base, with erect to slightly curved base, densely glandular-pilose from base to apex, green to deep-red in color;


inflorescence a scorpioid cyme, rarely bifurcating, bearing 6–24(–30) flowers; bracts caducous, rarely present on mature scapes, ca. 3 mm long, filiform-lanceolate, glandular pilose; pedicels (1.5–)2.5–9(–13) mm long, inserted (2–)6–16(–25) mm apart from each other, densely glandular pilose; sepals 5, 3.6–7 mm long, 1–2 mm wide, oblong-lanceolate to lanceolate, united at basal 1/5–1/4 of length, densely glandular pilose abaxially; petals 5, 7.5–11 mm long, 7.5–9.5 mm wide, obovate, pale pink in color; stamens 5, 4–5 mm long, anthers 1.2–2 mm long, bithecate, yellow; ovary 3-carpellate, fused, 1–2 mm in diameter, globose, smooth to slightly 3-lobed in outline; styles 3, forked at the base, 4–4.5 mm long (including stigmata), style arms remaining straight at the end of anthesis, stigmata unequally bilobed, pale pink to whitish in color; fruit a dry dehiscent capsule, ca. 1.5 mm long, ellipsoid, 3-valvate; seeds oblong-ovoid to ellipsoid, 0.5–0.6 mm long and 0.2–0.3 mm wide, testa reticulate, dark brown.

Illustrations:—Silva (1994), p. 59, fig. 11M–X (as D. villosa var. graomogolensis); Silva (1997), p. 86, fig. 1; Silva & Giulietti (1997), p. 90, fig. 8; Silva (2003), p. 249, fig. 1J–N; Correa & Silva (2005), p. 40, fig. 24F–H.

Distribution and ecology:—Drosera graomogolensis is endemic to the “Northern Mountains Complex” of the Espinhaço Range (terminology following Echternacht et al. 2011) in Minas Gerais state, Brazil (Fig. 2). It was originally published based on a few collections from areas surrounding the town of Grão Mogol. However, extensive fieldwork over the past two decades have revealed that it is in fact more widespread locally, occurring throughout the Serra de Grão Mogol region and also further south in the nearby municipalities of Cristália, Botumirim and Itacambira. Although most collections originate from around the town of Grão Mogol, D.

graomogolensis is probably equally common in the less explored ranges south of that town, and it may also possibly extend further north along the Espinhaço, closer to the border with Bahia.

Drosera graomogolensis occurs over a wide altitudinal range in montane areas, between 700–1420 m elevation. Populations are most common at lower elevations where they are often found in relatively dry habitats of open cerrado and caatinga vegetation, around the edges of seasonal springs in sandy soil with coarse quartz gravel, shallow sand overlying rocks, and in cracks of quartzite, often partially shaded by grasses and short bushes (Fig. 9). This species is more rarely also known to occur along permanent seepages over wet sandstone or in boggy soil with Sphagnum where it may even grow partially submerged. Furthermore, dense populations have been seen growing at one location in wet sandy soil on islands of vegetation along the margins of a large perennial river. A unique lowland population was even observed growing in sandy soil along a vereda – a seepage area dominated by the palm Mauritia flexuosa Linnaeus (1782: 454). At higher elevation, D. graomogolensis becomes less frequent and occurs in humid sandy soil of campo rupestre vegetation.

Although the formation of column-like stems around 3–5 cm in height is a common characteristic of D.graomogolensis, the production of longer stems (Fig. 9C) is usually associated with flooded or semi-shaded habitats, where competition for sunlight is stronger. The same correlation was observed for the inflorescence length, where the longest scapes were found on plants growing in more shaded habitats. In drier open habitats, plants of D. graomogolensis are more compact, producing stunted columns of dead leaves, and shorter inflorescences.

Drosera graomogolensis does not usually occur with other Drosera spp., but has occasionally been observed growing sympatric with D. tomentosa var. tomentosa, D. spiralis, D. grantsaui, D. hirtella var. hirtella, D.

communis, and D. × fontinalis Rivadavia (2009: 121) (the natural hybrid between D. tomentosa and D. grantsaui), as well as several species of Utricularia and Genlisea.

Phenology:—Drosera graomogolensis was collected in flower from July to October, corresponding to the mid dry season and early wet season.

Conservation Status:—Silva & Giulietti (1997) and Correa & Silva (2005) considered D. graomogolensis as endemic to Grão Mogol, incorrectly assigning studied specimens from nearby Botumirim and Itacambira to D.

villosa. Due to this wrongly presumed and restricted occurrence, the species was considered as Vulnerable in the above publications, as well as by Silva (2009) and in the database Biodiversitas (2005). Silva (2009) also wrongly claims that the species has not been collected since 1990, although representative herbarium collections have in fact been continuously made since then, including several locations near Grão Mogol, all of them deposited at SPF with the holotype (see Specimens Examined). Duno de Stefano & Silva (2001) indecisively proposed D. graomogolensis

as “Data Deficient—Least Concern?”. Drosera graomogolensis is here found to be more common and widespread than previously described, occurring at numerous populations approximately along a 70 km stretch of the Northern Mountains Complex, including the Grão Mogol State Park. Thus, with more distribution data available, this taxon is here reclassified as of Least Concern (LC), according to IUCN Red List criteria (IUCN 2001).


http://herbarium.univie.ac.at/database/detail.php?ID=192179

FIGURE 8. Drosera graomogolensis. A, habit. B, detail of the indumentum of the base of the scape. C, detail of the indumentum of

the apex of the scape. D, E, stipule. F, leaf, adaxial surface. G, leaf, abaxial surface. H, cross section of the petiole at the middle. J,

calyx. K, petal. L, stamen. M, gynoecium. N, seed. (A–E, H–M based on Gonella et al. 485; F, N based on Gonella et al. 486 – drawn

by P.M. Gonella.)


FIGURE 9. Drosera graomogolensis. A, habitat, on islands of vegetation in quartzite along the margin of a river at Botumirim, Minas

Gerais. B, habitat, waterlogged area in lowland caatinga at Grão Mogol, Minas Gerais. C, side view of a specimen with developed

stem. D, rosette seen from above, note the dense indumentum of white eglandular trichomes on the abaxial surface of the young

leaves. E, F, flower. (All by P.M. Gonella.)


Discussion:—Drosera graomogolensis is distinguished from other members of the complex by its lanceolate to oblong-lanceolate leaves with very short petioles, a glabrous adaxial petiole surface, an abaxial leaf surface that is densely covered by long white eglandular trichomes, by short uni- to bi-seriate glandular trichomes densely covering the entire scape, relatively large flowers (petals 7.5–11 mm long) with long styles (4–4.5 mm long), ; and small oblong-ovoid to ellipsoid seeds 0.5–0.6 mm long (Figs. 8, 9). It is here considered most closely related to D.

ascendens (Table 1).Silva (1997) cites the shape of the stigma as being one of the main differences between D. graomogolensis

(flabellate) and D. villosa (bilobed), the latter defined including D. latifolia. However, significant variation in stigma shape has been observed within D. villosa and D. latifolia, ranging from bilobed to flabellate. The main floral difference between the taxa in this complex is actually not the stigma shape, but the length of the styles, which in D. graomogolensis and D. ascendens are 4–4.5 mm long, but only up to 3 mm long in D. villosa, D.

latifolia, D. chimaera, and D. riparia. The bimodal style length observed within this complex may be correlated with different reproductive strategies (see below).

Drosera graomogolensis populations recently studied in Botumirim have flower scapes that tend to be slightly ascending at the base, whereas those of all other populations are erect from the base. This appears to represent a mere intraspecific variation, as the plants can be promptly assigned to D. graomogolensis by the other characteristics described above. Wine-red plants were also observed in Botumirim, but it was not possible to evaluate if the difference in pigmentation from the usual deep-red coloration was genetically fixed or ecologically caused.

Although Silva (1997) cites an isotype at RB, the specimen could not be located. While studying the collections at SPF, a duplicate of the type was found and forwarded to RB, to comply with the original protologue.

The larger flowers with longer styles, the scapes that are densely glandular-pilose from the base to apex, and the oblong-ovoid to ellipsoid seeds are characters which segregate D. ascendens and D. graomogolensis from the other four species in the complex. Most Drosera taxa possess flowers that are open for a single day, opening early in the morning on sunny days, and usually closing by early afternoon. Curiously, Drosera flowers do not produce nectar (A.F. pers. obs., Murza & Davis 2003, 2005), and thus only offer pollen as reward to their pollinators. We observed that most Neotropical Drosera species have short styles that bend inwards before the flowers close, bringing the stigmata in contact with the anthers (Fig. 10A), thus accomplishing successful self-pollination and often resulting in abundant fruit and seed set.

Observations in the field and in cultivation have highlighted a peculiarity of D. ascendens and D.

graomogolensis: these two species rarely fructify and produce very few seeds per fruit. This unusually low fertility may be explained by their very distinctive flowers, which have relatively large petals and long styles when compared to other species in the complex, and to most other New World Drosera. The long styles appear to never touch the anthers, remaining extended in the closed flower (Fig. 10B), suggesting a possible shift away from self-pollination and towards cross-fertilization. The unusually large petals may be a compensation to help attract more pollinators, whereas the unusually large thecae (up to 2 mm in D. graomogolensis) may represent a larger source of food in the form of pollen for some pollinators.

Flowers of D. graomogolensis were observed to be visited in the field by Africanized honey bees (a hybrid variety of Apis mellifera; Hymenoptera: Apidae) and also by a native bee species of the large family Halictidae (Hymenoptera; Fig. 10C; Rivadavia 1996). Despite the low seed production observed in D. ascendens and D.

graomogolensis, these species often form large and dense populations, suggesting successful asexual reproduction. Both taxa were observed to produce small plantlets from the roots, the base of the stem, and occasionally even from the leaves in D. graomogolensis growing in exceptionally wet habitats. Though vegetative reproduction is known from many species of Drosera, it is possibly more common in these two taxa, relative to other native Brazilian species.

Prior to the official publication by Silva (1997), D. graomogolensis was considered as a variety of D. villosa by the same author in her Master thesis (Silva 1994), and the name “D. villosa var. graomogolensis” was also applied for that taxon by Rivadavia (1996), a status that was never validly published.

Specimens examined:—BRAZIL. Minas Gerais: Município de Botumirim, Serra da Canastra, 1300 m, 20 December 1994, Rivadavia 334 (SPF); Rio do Peixe, 780 m, 10 February 2011, Gonella et al. 371 (SPF), 06 September 2011, Gonella et al. 478 (SPF), 791 m, 06 September 2011, Rivadavia 2702 (SPF), 796 m, 06 September 2011, Rivadavia 2703 (SPF); estrada entre o povoado de Canta Galo e a Várzea da Estiva, 790 m, 10


February 2011, Gonella et al. 388 (SPF); Várzea da Estiva, 7 March 2013, Gonella et al. 590 (SPF). Município de Grão Mogol, Serra Grao Mogul [sic], north base of mountain, 600-700 m, 16 August 1960, Maguire et al. 49217

(SPF, NY); ca. 16 km west of Grão Mogol, 950 m, 17 February 1969, Irwin et al. 23452 (NY); Estr. Grão Mogol-Cristália, 15 April 1981, Cordeiro et al. CFCR 933 (SPF); Vargem do Quartel, 500-650 m, 09 July 1985, Wanderley et al. 842 (SP); Jambeiro a 7km de Grão Mogol, 05 September 1985, Cavalcanti et al. CFCR 8510

(SPF, SP, RB. BHCB); 5-7 km N de Grão Mogol, 900 m, 01 November 1988, Wanderley & Kral 1432 (SP); Alto do Morro do Jambeiro, Córrego Escurinha, 900 m, 06 September 1990, T.R.S. Silva et al. CFCR 13478 (SPF); Córrego da Bonita, subida para o Morro Jambeiro, 07 September 1990, Silva et al. CFCR 13506 (RB, SPF; type); Morro do Jambeiro, 03 June 1994, Rivadavia 287 (SPF), 700 m, 08 September 1994, Rivadavia 297 (SPF); Vale do córrego Jambeiro, 26 September 1997, Rapini et al. 336 (SPF, SP); estrada para Montes Claros, 700 m, 12 October 2001, Rivadavia 1256 (SPF); Morro do Jambeiro estrada para Montes Claros, 700 m, 12 October 2001, Rivadavia

1264 (SPF); estrada Grão Mogol-Montes Claros, Morro do Jambeiro 715 m, 12 August 2010, Gonella 358 (SPF), 07 September 2011, Gonella et al. 485 (SPF); Grão Mogol, trilha para Cachoeira Véu das Noivas, 11 February 2011, Gonella et al. 382 (SPF); Capim Puba, 07 September 2011, Gonella et al. 486 (SPF). Município de Itacambira, ao sul da cidade, 1250 m, 12 December 1994, Rivadavia 325 (SPF); à esquerda da estrada para Montes Claros, ca. 5 km de Itacambira, 1420 m, 05 March 1997, Rivadavia 615 (SPF); a alguns km de Itacambira pela estrada para Montes Claros, 1350 m, 14 July 1999, Rivadavia & Pinheiro 1142 (SPF), 14 October 2001, Rivadavia

1294 (SPF), 30 June 2002, Rivadavia & Gibson 1367 (SPF).

Drosera latifolia (Eichler) Gonella & Rivadavia, comb. et stat. nov. (Figures 1, 10A, 11, 12, 13).Basionym: Drosera villosa var. latifolia Eichler (1872: 395). Type (lectotype designated by Silva & Giulietti 1997):—BRAZIL. Rio de Janeiro, Serra dos Órgãos, 13 December 1869,

Glaziou 3868 (R!, isolectotypes K image!, P image!).= Drosera villosa var. bifurca Buxbaum (1924: 119), syn. nov.

Type:—BRAZIL. São Paulo, prope Campo Grande inter Santos et urben S. Paulo, 750 m, 1902, M. Wacket s.n. (holotype WU image! Image available at http://herbarium.univie.ac.at/database/detail.php?ID=192179).

Description:—Perennial rosetted herbs, acaulescent or more rarely with short upright stems up to 100 mm tall, usually sparsely covered with persistent dead leaves, when occurring in semi-aquatic and semi-shaded habitats. Roots slightly succulent, up to 0.5 mm in diameter, sparsely branched, densely covered by root hairs. Indumentum

consisting of white eglandular trichomes (brown when old), up to 1.5 mm long, present on the abaxial leaf surface (rarely on adaxial petiole surface), and occasionally sparsely present on scapes; glandular capitate trichomes 0.1–0.2 mm long, the stalk light red and translucent, the head dark red, present on scapes, pedicels, bracts and sepals abaxially; minute sessile glands ca. 0.03 mm in diameter, red in color, on both leaf surfaces, scapes, pedicels, bracts and sepals abaxially. Leaves (6.5–)10–57(–70) mm long, with circinate vernation, semi-erect, rarely decumbent, persistent when old, lanceolate to oblong, apex acute to obtuse, green, or deep to wine-red in color; petiole (2–)3–22(–30) mm long, 1–3(–4) mm wide, rectangular-linear, transverse narrowly elliptic in cross section, adaxial surface glabrous, rarely with sparse eglandular trichomes, abaxial surface glabrescent to densely eglandular-pilose, trichomes concentrated along the margins; lamina (4.5–)7–35(–45) mm long, 1.5–4(–7) mm wide, lanceolate, oblong, broad to narrowly oblong-lanceolate, rarely oblong-elliptic, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface glabrescent (rarely densely eglandular pilose), trichomes concentrated along the margins; stipules (2.5–)3–6 mm long, 1.5–3 mm wide at the base, rectangular, membranaceous, pinkish (drying brown), the apical 1/2–1/3 divided into several laciniate segments, the marginal ones longer. Scapes 1–2 (up to 6 in branched specimens) per plant, (50–)100–460 mm long (including floriferous part), terete, 0.5–2 mm in diam. at the base, base erect, basal half with sessile to sub-sessile glands only, apical half with glandular trichomes (increasing in size and density towards the apex), more rarely with very sparse eglandular trichomes uniformly distributed from base to apex, green to deep or wine-red in color; inflorescence a scorpioid cyme, often bifurcating, bearing (1–)2–30 flowers; bracts 1.5–2.5 mm long, filiform-lanceolate, often stipulate, glandular pilose, caducous and thus frequently absent on mature scapes; pedicels 1.5–5 mm long, inserted up to 11 mm apart from each other, sparse to densely glandular-pilose, rarely with few eglandular trichomes; sepals 5, (2–)3–6.5 mm long, 1-2 mm wide, oblong-lanceolate, ovate, obovate, or lanceolate, united at basal 1/3–1/5 of length, glabrescent or densely glandular-pilose at the base (decreasing in size and density towards the apex), rarely with few eglandular trichomes, apex acute or more rarely obtuse; petals 5, 5–8.5 mm long, 3–6.5 mm wide, narrowly to


broadly obovate, pale pink in color; stamens 5, 4–4.5 mm long, anthers 0.7–1.2 mm long, bithecate, yellow; ovary

3-carpellate, fused, 1.5–2 mm in diam., globose, 3-lobed in outline; styles 3, forked at the base, 2–3 mm long (including stigmata), style arms curving inwards at the end of anthesis, stigmata flabellate to bilobed, pale pink to whitish in color; fruit a dry dehiscent capsule, 2–3.5 mm long, globose, 3-valvate; seeds fusiform, often falciform, 0.8–1 mm long and 0.2–0.3 mm wide, testa reticulate, dark brown.

FIGURE 10. A, a senescent flower of D. latifolia, with curved styles and stigmata touching the anthers for self-pollination. B,

senescent flower of D. graomogolensis, with straight styles and stigmata not touching the anthers. C, a halictid bee (sweat bee) visiting

a flower of D. graomogolensis at Grão Mogol, Minas Gerais. (A, B by P.M. Gonella; C by F. Rivadavia.)

Illustrations:—Eichler (1872), t. 91, I (as D. villosa); Silva (1994), p. 59, fig. 11A–L (as D. villosa var. villosa); Silva & Giulietti (1997), p. 88, fig. 7 (as D. villosa); Correa & Silva (2005), p. 53, fig. 28A–C (as D.

villosa).


FIGURE 11. Drosera latifolia. A, B, habit. C, D, stipule. E, leaf, adaxial surface. F, leaf, abaxial surface. G, outline of a leaf, showing

the distribution of eglandular trichomes on the petiole adaxial surface. H, J, K, leaf outlines (dotted line indicates the limit between

lamina and petiole). L, M, cross section of the petiole at mid length. (A, C, E, based on Gonella et al. 452; B, D, K, based on Rivadavia

et al. 1083; F, M based on Rivadavia 2515; G, L based on Rivadavia 2137; H based on Rivadavia & Gibson 1379; J based on

Rivadavia et al. 2464 – drawn by P.M. Gonella.)


FIGURE 12. Drosera latifolia. A–D, detail of the base (A, C) and apex (B, D) of the scape. E, F, calyx. G, H, petal. J, stamen. K, L,

gynoecium. M, seed. (A, B, F, G, J, K, M based on Gonella et al. 452; C, D based on Rivadavia 2515; E based on Souza 7207; H based

on cultivated material from São Bernardo do Campo, São Paulo; L based on cultivated material from Aiuruoca, Minas Gerais.)


Distribution and ecology:—Drosera latifolia is endemic to Brazil. This species is the most widespread and common, as well as most variable, taxon of the D. villosa complex, occurring in the states of Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, and Santa Catarina (Fig. 1), in montane to high-montane regions at elevations between 700–2550 m. The distribution of D. latifolia is geographically and ecologically divided between the quartzitic Espinhaço Range in central Minas Gerais and disjunctly along the igneous Serra da Mantiqueira and Serra do Mar in southern and southeastern Brazil.

On the igneous highlands, the original habitat of D. latifolia is the campos de altitude, where it grows on mountain tops in clayey or peaty soil or over a thin layer of soil overlying basaltic rock (Fig. 13B). However, D.

latifolia is also frequently found in disturbed areas at lower elevation in red clayish soils (rarely in sandy soils) along railways and roads – areas that were covered by rainforests until recent history (Fig. 13A, D). At these coastal igneous habitats, D. latifolia has been observed growing sympatric with D. viridis Rivadavia (2003: 87), D.

grantsaui, D. montana, D. brevifolia Pursh (1813: 211), and D. communis, as well as with several species of Utricularia and Genlisea.

On the quartzitic Espinhaço Range, D. latifolia commonly occurs in campo rupestre vegetation, in sandy soil or in cracks on sandstone, most frequently on islands of vegetation along stream margins or seepages (Fig. 13C), sometimes shaded by riparian vegetation. It is also commonly found occupying areas disturbed by mining activities around the city of Diamantina, especially inside gullies. Along the Espinhaço Range, D. latifolia has been observed growing sympatric with D. ascendens, D. camporupestris, D. communis, D. grantsaui, D. spiralis, and D.

tomentosa (both varieties), as well as with numerous species of Utricularia and Genlisea. Although D. latifolia is known to grow on several highlands neighboring those where D. villosa is endemic,

the two species do not appear to co-occur. This is possibly due to ecological differences: the Serra do Ibitipoca and Serra Negra (home to D. villosa) are quarzitic formations, contrasting geologically from the surrounding igneous formations that dominate the sourthern coastal highlands of Brazil (where D. latifolia is abundant). Paradoxically, D. latifolia occurs further north on the quartzitic Espinhaço Range, as mentioned above.

Phenology:—The flowering period of D. latifolia varies widely across its range. While plants from the Serra do Espinhaço usually bloom during the dry season (from June to September, with a second and minor flowering period in the wet season, in February and March), populations found along the coastal highlands will flower during the late dry season and into the rainy season (from September to February).

Conservation status:—Drosera latifolia is widespread and common in southern and southeastern Brazil, occurring in several Environmental Protection areas, and is thus considered of Least Concern (LC), according to IUCN Red List criteria (IUCN 2001).

Discussion:—Drosera latifolia is distinguished from other members of the complex by its lanceolate, oblong, broad to narrowly oblong-lanceolate, or rarely oblong-elliptic leaf lamina, by petioles that are always shorter than the lamina, transversely flat in cross-section and 1–3(–4) mm wide, the adaxial petiole surface glabrous to glabrescent, by erect scapes with glabrous to glabrescent base (eglandular trichomes very sparse, when present), relatively small to medium sized flowers (petals 5–8.5 mm long), and relatively large fusiform seeds 0.8–1 mm long (Figs. 11, 12, 13; Table 1).

The glandular trichomes concentrated towards the apex of the scapes, together with seed shape of D. latifolia, suggest a closer affinity to D. villosa and D. riparia, two taxa from which the former species is readily distinguished by the petioles that are usually much shorter than the lamina (Table 1).

Drosera latifolia is the most widespread species of the D. villosa complex and thus, not surprisingly, also the morphologically most variable. Although some characteristics are very conserved, such as seed shape and distribution of glandular trichomes on the scape, considerable variation can be seen in the overall leaf size, shape, and indumentum; sepal shape; as well as in the relative abundance of eglandular trichomes on the scapes. Nevertheless, the morphological variations observed within D. latifolia appear to form a continuum, with gradual overlap of the cited characters. However, these morphological characters remain stable in cultivation in specimens grown under identical conditions, and thus are certainly not ecologically induced by different habitats and environmental conditions (e.g. elevation or soil). Although there is not enough evidence at the present time to support a taxonomic subclassification of D. latifolia, a few regional morphotypes can be recognized and are described below:


FIGURE 13. Drosera latifolia. A, side view of a flowering rosette of the Serra do Mar morphotype, notice the erect and glabrous base

of the flower scape (Paranapiacaba, São Paulo). B, habit of the type morphotype (at Pico da Caledônia, Nova Friburgo, Rio de

Janeiro). C, Espinhaço morphotype habitat, in cracks of sandstone (at Parque Nacional das Sempre-Vivas, Minas Gerais). D, rosette of

Serra do Mar morphotype seen from above (Paranapiacaba, São Paulo). E, flower of Serra do Mar morphotype (cultivated specimen

from São Bernardo do Campo, São Paulo). F, flower of Espinhaço morphotype (cultivated specimen from Diamantina, Minas Gerais).

(A, C, D, F by P.M. Gonella; B by F.Rivadavia; E by A. Fleischmann.)


Typical morphotype: small to medium-sized, compact rosettes (plants at lower elevation often larger); petioles short (shorter than half the lamina length); lamina oblong to oblong-ellipsoid (Figs. 11J, 13B); leaves usually decumbent (often only the young leaves are semi-erect), densely eglandular pilose abaxially, adaxial petiole surface with sparse eglandular trichomes, apex obtuse; entire scapes with sparse eglandular trichomes.

The geographical distribution of this morphotype extends from the Serra dos Órgãos, in central Rio de Janeiro state, to the Serra da Mantiqueira (border between São Paulo, Rio de Janeiro and Minas Gerais) including the Serra de Itatiaia and Serra do Papagaio. It is also known to occur at the Serra da Araponga, in eastern Minas Gerais, and at the Morro do Forno Grande in Espírito Santo.

Serra do Mar morphotype: medium-sized, compact rosettes; petioles short (usually shorter than half the lamina length); lamina lanceolate or oblong-lanceolate (Figs. 11F; 13A, D); leaves semi-erect, abaxial leaf surface glabrescent to densely eglandular pilose, adaxial petiole surface glabrous (rarely sparsely eglandular-pilose), apex acute to obtuse; scapes usually glandular-pilose only.

This is the most widespread morphotype, present along almost 1000 km of mountain ranges collectively referred to as the Serra do Mar. It occurs from the Serra do Rio do Rastro (near the Santa Catarina—Rio Grande do Sul border) to the Serra da Bocaina (São Paulo—Rio de Janeiro border).

Salesópolis morphotype: medium to large-sized, lax rosettes (up to 12 cm in diameter); petioles of medium length (exceeding half the lamina length); lamina lanceolate; leaves semi-erect, densely eglandular-pilose abaxially (villous), adaxial petiole surface often sparsely eglandular-pilose (Fig. 11G, L), apex acute; entire scapes with sparse eglandular trichomes.

Although located on the Serra do Mar in São Paulo state, plants from this locality differ from the above morphotype by the larger rosettes, with longer petioles and a denser indumentum of eglandular trichomes on the leaves, and also by the presence of eglandular trichomes on the adaxial petiole surface and scape.

Serra do Espinhaço morphotype: medium to large-sized lax rosettes (Fig. 11A, 13C); petioles short to medium in length (shorter than or equalling half the lamina length); lamina narrowly oblong-lanceolate; leaves semi-erect, abaxial surface glabrescent, the eglandular trichomes usually restricted to the leaf margins, extending to the middle length of the lamina, adaxial petiole surface glabrous (Fig. 11E), apex acute to obtuse; scapes always lacking eglandular trichomes (Fig. 12A, B). This morphotype further differs from all other populations of D. latifolia by the flowering period (see Phenology section).

This morphotype is most common on the Serra do Cabral and Diamantina Plateau in northern-central Minas Gerais, but is also known to occur disjunctly on highlands surrounding the towns of Ouro Preto and Mariana in central-southern Minas Gerais. Although close to these two towns, a specimen from the Serra do Caraça (Damazio

2098) is somewhat dubious as no other collections are known from these heavily-botanized highlands. The largest known specimens of D. latifolia were found in this region, at the Serra do Cabral, where plants are often relatively large, with rosettes reaching 15 cm in diameter—although this morphotype is generally smaller on other parts of the Serra do Espinhaço.

Pico do Itambé morphotype: medium-sized, compact rosettes; leaves semi-erect, broadest among all D.

latifolia morphotypes, densely eglandular-pilose abaxially; lamina oblong to lanceolate, up to 7 mm wide, apex acute to obtuse; petioles short (shorter than half of the lamina length), up to 4 mm wide, adaxial surface with sparse eglandular trichomes (Fig. 11H); entire scape covered with sparse eglandular trichomes.

Although the Pico do Itambé is located in the central part of the Serra do Espinhaço in Minas Gerais, plants from this mountain occur at higher elevation and are morphologically rather different from other nearby D. latifolia

populations found around the towns of Diamantina and Serro. The Itambé morphotype is somewhat reminiscent of D. graomogolensis, from which it is readily distinguished by the type of scape indumentum and geographical distribution.

Serra do Caparaó morphotype: small compact rosettes (Fig. 11B); petioles short (shorter than half of the lamina length); lamina lanceolate; leaves decumbent or semi-erect, usually 1–2 cm long, densely eglandular-pilose abaxially, adaxial petiole surface with sparse eglandular trichomes, apex acute; entire scape with sparse eglandular trichomes (Fig. 12C, D).

The Serra do Caparaó is located on the border between Minas Gerais and Espirito Santo states, near the northeastern limit of the D. latifolia range along the igneous highlands. Plants from this mountain range represent the smallest sized morphotype of D. latifolia (Fig. 11B, D, K) as well as the one that occurs at highest elevation (up to 2550 m).


In general, specimens of D. latifolia at higher elevations (above ca. 1500 m), growing lithophytically, and in very exposed habitats, conditions especially common at the Serra dos Órgãos (Rio de Janeiro state), Serra da Bocaina (border between São Paulo and Rio de Janeiro states), and Serra do Caparaó (border between Minas Gerais and Espírito Santo states), tend to be dwarfed, with relatively small compact rosettes, the leaves often decumbent and as small as 1–2 cm long, and shorter inflorescences (Fig. 11B), possibly a result of the more stressful habitats, where they are susceptible to higher water stress as result of stronger winds and insolation. Such dwarfed plants also tend to possess an oblong-elliptic lamina (Fig. 11J, K) instead of the more common lanceolate or oblong-lanceolate shape, and may be confused with D. tomentosa var. glabrata. However, the latter is distinguished by its obovate to oblong-obovate leaves with snap-tentacles on the obtuse leaf apex, as well as by having ovoid to ellipsoid seeds.

Leaf eglandular indumentum varies considerably across the range of D. latifolia and between different habitats. Plants from many regions, especially from the Pico do Itambé and Salesópolis, occasionally to frequently have sparse eglandular trichomes on the adaxial petiole surface (usually concentrated near the base of the lamina), a densely eglandular-pilose (often villous) abaxial leaf surface, and sparse eglandular trichomes uniformly distributed along the scape. However, these characters may vary even within populations according to the habitats occupied, with plants from wetter and shadier habitats being usually less pilose. Plants from the Espinhaço Range (excluding the Pico do Itambé) and from some parts of Paraná state are the least pilose in the species range, with petioles always glabrous adaxially, leaves with glabrescent abaxial surface (eglandular trichomes concentrated along the margins) and scapes completely lacking eglandular trichomes.

Despite the wide variation observed within D. latifolia, all specimens can still be promptly distinguished from D. villosa by the short and broad petioles that are flat in cross section (vs. transversely elliptic), and by the glabrous to glabrescent base of the scape (vs. densely eglandular-pilose) (Table 1).

A mixed collection from Couto de Magalhães de Minas (Rivadavia 2524) contains specimens of both D.

latifolia and the newly discovered hybrid between D. latifolia and D. tomentosa var. tomentosa (the two species were growing sympatrically). A similar natural hybrid was also discovered at two populations around the town of Diamantina, but it is uncertain whether these represent D. latifolia crossed with D. tomentosa var. tomentosa or with D. tomentosa var. glabrata, which was also observed growing sympatrically (Rivadavia & Gibson 1372; Rivadavia & Pinheiro 1114; Gonella et al. 272) (Fig. 5C–E).

Although Saint-Hilaire’s expeditions in the early 19th century brought him to several areas in southeastern Brazil where the widespread D. latifolia occurs, it is very odd that he did not collect and describe this species. However, he may have in fact encountered this species near Itambé in central Minas Gerais, as appears evident from the holotype of D. tomentosa which contains a single robust scape that appears to represent the Pico do Itambé morphotype of D. latifolia.

Although Eichler (1872) recognized D. villosa var. latifolia as distinct from the type variety, he included under his concept of D. villosa var. villosa several specimens that we here identify as D. latifolia, revealing that Eichler did not fully comprehend these two taxa, possibly due to the low number of specimens available at the time. Although the epithet “latifolia” means “broad leaved”, the difference in the lamina width between this species and D. villosa is not diagnostic, with the exception of a few specimens (the lamina width ranges from 1.5–4(–7) mm in D. latifolia and from 1.4–4 mm in D. villosa). However, significant differences can be observed in the petiole width, which ranges between 0.5–1.6(–2) mm in D. villosa and between 1.5–3.5(–4) mm in D. latifolia.

The description of D. villosa var. bifurca was based on a specimen from the coastal highlands of São Paulo with forked inflorescences (Buxbaum 1924). However, this character is neither unique nor uncommon in D.

latifolia (with which the former is here considered to be conspecific), or even among most Drosera species, as later observed by the same author (Buxbaum 1928). Buxbaum (1924) also cites a notch at the apex of the sepals as a diagnostic character for D. villosa var. bifurca, but this is a common character observed in all D. latifolia

morphotypes. For these reasons D. villosa var. bifurca is considered a synonym of D. latifolia.Specimens examined:—BRAZIL. Without attribution to state or administrative area, 22 December 1826,

Burchell 3655 (K parte – mixed with the isotype of D. villosa); without date, Ule 2379 (US). Minas Gerais: Município de Alto Caparaó, Serra do Caparaó, December 1995, Leoni 3141 (GFJP); beira do rio José Pedro, 01 April 1989, Krieger et al. 23587 (BHCB); beirando a trilha do Pico da Bandeira, 31 October 1995, Rivadavia &

Padovese 508 (SPF); nascente próx. à trilha do P.da Bandeira, 19 February 1996, Rivadavia et al. 1083 (SPF); Vale Encantado, 13 June 1993, Rivadavia 218 (SPF), 27 February 2005, Rivadavia 1959 (SPF). Município de Araponga, Serra da Araponga, Faz.Neblina, December 1994, Leoni 2735 (GFJP), 17 February 1996, Rivadavia et al. 515


(SPF). Município de Buenópolis, Parque Nacional das Sempre Vivas, 04 September 2011, Gonella et al. 452

(SPF), 8 March 2013, Gonella et al. 609 (SPF). Município de Couto de Magalhães, ca. 30km ao leste de Diamantina, 12 May 2007, Rivadavia 2524 (mixed with D. tomentosa var. tomentosa hybrids) (SPF); ca. 40km ao leste de Diamantina, 12 May 2007, Rivadavia 2528 (SPF). Município de Diamantina, cachoeira da Sentinela, 02 December 1991, Braga & Chaves 423 (BHCB), 12 July 1999, Rivadavia et al. 1120 (SPF), 10 August 2010, Gonella et al. 348 (SPF); próximo a Sopa, 28 February 1997, Rivadavia & Pinheiro 584 (SPF); estrada para Tapetes Arraiolos, , 31 July 2002, Rivadavia & Gibson 1368 (SPF), 31 July 2002, Rivadavia & Gibson 1371 (SPF); Reserva do Pau-de-Fruta, 21 September 2000, Stehmann et al. 2699 (BHCB); São João da Chapada, estrada para Macacos, 04 September 2011, Gonella et al. 437 (SPF), 8 March 2013, Gonella et al. 616 (SPF). Município de Itamonte, Parque Nacional de Itatiaia, 19 November 2005, Rivadavia et al. 2133 (SPF). Município de Joaquim Felício, Serra do Cabral, 02 September 1985, Pirani et al. 8192 (SPF); Brejo Grande, 03 July 2003, Rivadavia &

Menezes 1655 (SPF); a 23.3km de J.Felício, 04 July 2003, Rivadavia 1673 (SPF). Município de Mariana, passagem de Mariana, 06 March 1977, J. Ruiz s.n. (OUPR 23804); Parque Estadual do Itacolomi, Serrinha, 04 September 2009, E.S. Cândido et al. 200 (OUPR). Município de Ouro Preto, Itacolomi, 28 December 1950, Macedo 2797

(MO, NY), without date, anonymous (OUPR-4072). Município de Santa Rita (de Jacutinga), Serra do Pacau, Bom Jardim, 04 June 1989, Grandi & Braga 17341 (BHCB). Município de São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 18 November 1999, Lombardi 3485 (BHCB); ribeirão das Éguas, 28 June 2003, Rivadavia & Deco

1606 (SPF), 28 June 2003, Rivadavia & Deco 1614 (SPF), 28 June 2003, Rivadavia & Deco 1622 (SPF). Município de Santo Antônio do Itambé, Pico do Itambé, 09 July 1995, Rivadavia & Pinheiro 458 (SPF), 01 August 2002, Rivadavia & Gibson 1379 (SPF), 01 August 2002, Rivadavia & Gibson 1381 (SPF), 01 August 2002, Rivadavia & Gibson 1383 (SPF), 01 August 2002, Rivadavia & Gibson 1392 (SPF), 01 August 2002, Rivadavia &

Gibson 1396 (SPF). Município de Serro, Milho Verde, cachoeira dos Carijós, 22 February 2000, Salino & Morais

5087 (BHCB), 05 April 2003, Rivadavia & Neves 1562 (SPF). Without attribution to administrative area, Serra do Caraça, 1912, Damazio 2098 (mixed with D. tomentosa) (R). Espírito Santo: Município de Castelo, Morro Forno Grande, 12 August 1948, Brade 19249 (RB). Without attibution to adminstrative area: Serra da Caparaó, 25 November 1929, Mexia 9025 (NY); Pico da Bandeira, Serra do Caparaó, 03 March 1959, Irwin 2804 (VIC, NY, US); Serra do Caparaó, próx. a região de Três Lagos, 25 September 1941, Brade 17023 (RB); Topo da Pedra Azul, 19 November 1975, Kautsky 517 (HB). Rio de Janeiro: Município de Nova Friburgo, Pico da Caledônia, 20 January 2007, Rivadavia et al. 2464 (SPF). Município de Petrópolis. Maria Comprida, March 1944, Góes &

Constantino 351 (R); Parque Nacional da Serra dos Órgãos, entre a Pedra do Açu e a Pedra do Sino, 11 October 1993, Rivadavia 244 (SPF); APA Petrópolis – Serra da Maria Comprida, percurso Palmares – Boi Coroado, 18 November 2009, R. Borges et al. 1019 (HUEFS, RB, SPF). Município de Santa Maria Madalena, Campos de Mocotó, November 1933, Santos-Lima 219 (MO, R); Parque Estadual do Desengano, Pedra do Desengano, 30 June 1989, Martinelli 13399 et al. (RB). Município de Teresópolis, Serra dos Órgãos, 13 December 1869, Glaziou

3868 (R; type), 11 December 1891, Ule 2379 (R), December 1960, Vidal 76 (R), 21 July 1970, Barcia 120 (R); Campo das Antas, 04 April 1947, Lutz s.n. (R); Pedra do Sino, December 1952, Vidal 5776 (R); beira da estrada para Itaipava, km 27, 28 October 1962, Abentroth 2391 (HB, NY); 5km SW of Teresópolis, 28 November 1986, Webester et al. 25860 (R); estrada Petrópolis-Teresópolis, 25 February 1996, Rivadavia & Mullins 551 (SPF), 19 November 2000, Rivadavia 1219 (SPF); Parque Nacional da Serra dos Órgãos, 13 November 2005, Rivadavia et

al. 2127 (SPF); estrada Rio – S.Órgãos, without date, Schwacke 77 (R). Without attribution to administrative area, Serra de Itatiaia, February 1894, Ule 49 (R); S.de Itatiaia, 29 May 1902, Dusai 628 (R, US); Pedra Assentada, March 1907, Brade 15580 (B, RB); Planalto do Itatiaia, Várzea das Flores, 24 February 1921, Lobo 33 (R); km 6–6.5, 08 December 1970, Barcia 142 (R); without date, Moura 1887 (R); without date, anonymous (R-63544). Without attribution to administrative area, Mount des Orgnes (Serra dos Órgãos), 12 October 1867, Glaziou 1605

(P). São Paulo: Município de Biritiba Mirim, Estação Biológica de Boracéia, 24 November 1983, Custódio Filho

1925 (SP), 29 December 1983, Custódio Filho 2165 (SP). Município de Campos do Jordão, São José dos Alpes, 17 December 1966, Mattos 14729 (SP), 17 December 1966, Mattos 14729 (SP); estrada para o Pico Itapeva, 12 January 1977, Occhioni 8001 (RFA), 14 April 1999, Rivadavia & Ract 1094 (SPF), 14 April 1999, Rivadavia et al.

1181 (SPF), 25 January 2003, Rivadavia & Read 1530 (SPF). Município de Cananéia, Ilha do Cardoso, 11 January 1977, Sheplerm & Vital 11217 (UEC). Município de Itararé, Fazenda Ibiti, 30 October 1993, Souza 4521 (ESA), 13 November 1994, Souza et al. 7207 (ESA); Rio Verde, 24 April 1995, Rivadavia 415 (SPF). Município de Mogi das Cruzes, morro do Garrafão, 07 March 1999, Rivadavia et al. 836 (SPF). Município de Ribeirão Pires, estr. velha p/ Santos, 20 April 1991, Rivadavia & Demets 7b (SPF), 11 January 1992, Rivadavia & Cardoso 54 (SPF); estrada


Caminho do Mar, 09 October 1999, Rivadavia et al. 1172 (SPF), 06 February 2006, Rivadavia & Périgo 2142

(SPF). Município de Salesópolis, estr. p/ Caraguatatuba, 23 July 1991, Rivadavia et al. 15 (SPF), 9 December 2000, Rivadavia et al. 1223 (SPF), 25 January 2003, Rivadavia & Read 1534 (SPF), 22 January 2006, Rivadavia &

Périgo 2134 (SPF), 22 January 2006, Rivadavia & Périgo 2136 (SPF), 22 January 2006, Rivadavia & Périgo 2137

(SPF). Município de Santo André, estr. Ribeirão Pires – Campo Grande, 11 December 1971, Windisch s.n. (HB); Distrito de de Paranapiacaba, 06 February 2006, Rivadavia & Périgo 2139 (SPF), 06 February 2006, Rivadavia &

Périgo 2141 (SPF) estr. velha para Paranapiacaba, 20 April 1991, Rivadavia & Demets 7a (SPF), 17 December 1992, Rivadavia 155 (SPF); estrada para a reserva biológica, 11 January 1992, Rivadavia & Cardoso 50 (SPF); estrada que sai por trás da cidade, 09 October 1999, Rivadavia et al. 1176 (SPF), 26 January 2003, Rivadavia et al.

1538 (SPF); Estação Ecológica do Alto da Serra, 20 October 1917, Hoehne & Gehrt 756 (ESA, NY). Município de São Bernardo do Campo, rodovia dos Imigrantes, 12 December 1992, Rivadavia 153 (SPF), 19 August 2007, Gonella & Rivadavia 27 (SPF), 19 August 2007, Gonella & Rivadavia 28 (SPF); velha estrada para Santos, 17 December 1992, Rivadavia 157 (SPF); rodovia Caminho do Mar, 19 August 2007, Gonella & Rivadavia 31 (SPF); estrada ligando Caminho do Mar a Paranapiacaba, 19 August 2007, Gonella & Rivadavia 33 (SPF). Município de São José do Barreiro, Serra da Bocaina, 22 October 2002, Rivadavia & Morais 1416 (SPF), 22 October 2002, Rivadavia & Morais 1420 (SPF). Município de São Paulo, Pico do Jaraguá, 17 March 1907, Usteri 18391 (SP), 8 December 1912, Brade 6534 (SP); Distrito de Campo Grande, Estação Biológica, 23 February 1929, Smith 1977

(NY, US); Jardim Botânico de São Paulo, 25 November 1939, Handro 43851 (SP); Distrito de Parelheiros, ao SE de Parelheiros, 11 February 2006, Rivadavia & Miranda 2149 (SPF), 2151 (SPF), 2153 (SPF); Campo Grande, Serra do Mar, November 1913, Brade 7258 (R). Município de Silveiras, Serra da Bocaina, source of Paraíba River, March 1951, Segadas-Vianna 2693 (R, US). Without attribution to administrative area, Alto da Serra do Mar, antiga S.Paulo – Santos, 14 October 1961, Pabst & Pereira 5773 (HB); Estação Biológica, 20 October 1917, Hoehne 756 (SP); Alto da Serra de S.Paulo, estrada do Vergueiro, 23 December 1920, Hoehne 4700 (SP). Without locality, without date, Diogo 600 (R); Riedel 62538 (R parte – mixed with D. brevifolia). Paraná: Município de Arapoti, Rio das Cinzas, 08 September 1960, Hatschbach 7232 (MBM), 12 October 1968, Hatschbach 20016

(MBM). Município de Balsa Nova, seguindo estrada de terra após São Luiz do Purunã, 2 November 2010, Alves 6

(SPF). Município de Campina Grande do Sul, Serra do Capivari Grande, 15 January 1969, Hatschbach & Koczicki

20750 (MBM, NY); Serra Ibitiraquire, 23 January 1970, Hatschbach 23402 (MBM). Município de Campo Largo, fazenda São José, 2 October 2011, Alves 15 (SPF). Município de Castro, canyon do Guartelá, 22 April 1995, Rivadavia et al. 405 (SPF); km 21 da estr. Castro–P.Grossa, 24 November 1950, Vidal & Araújo s.n. (R). Município de Guaratuba, Serra de Araçatuba, 23 November 1996, Santos et al. 273 (NY). Município de Jaguariaíva, estr. de ferro, 06 November 1928, Hoehne s.n. (SPF); Recanto da Prainha, 20 November 1995, Cervi

et al. 6040 (BHCB); rio Jaguariaíva, 13 November 1974, Hatschbach 35440 (MBM). Município de Palmeira, Rio das Pombas, 30 December 1953, Hatschbach 3386 (MBM, US); Rio dos Papagaios, 28 November 1986, Hatschbach & Souza 50830 (MBM), 21 April 1995, Rivadavia et al. 395 (SPF), 12 October 2010, Alves 1 (SPF); depois do Rio dos Papagaios, 2 November 2010, Alves 8 (SPF); Recanto dos Papagaios, 12 January 2011, Alves 12

(SPF); Rio das Pombas, 20 December 1953, Hatschbach 3386 (MBM), 25 November 2006, Rivadavia &

Rohrbacher 2446 (SPF); Serra de São Luiz, 07 January 1958, Emygdio & Angely 1534 (R). Município de Ponta Grossa, 1874, anonymous (R). Município de Sengés, estr. Ponta Grossa-Sengés, rio Funil, 24 April 1995, Rivadavia 412 (SPF). Without attribution to administrative area: Serra de São Luis, 07 January 1958, Mello-Filho

1374 (R); a 50km de Curitiba, 07 January 1958, Emygdio & Angely 1534 (R). Dubious location: Aguadas, November 1916, S.Brito 77 (R). Santa Catarina: Município de Campo Alegre, Morro Iquererim, 10 October 1956, Smith & Klein 8529 (R, NY, US); Serra do Quiriri, 29 December 1998, Silva et al 2771 (MBM). Município de Garuva, Serra do Quiriri, 19 November 2006, Rivadavia & Rohrbacher 2439 (SPF). Município de Grão Pará, Serra do Corvo Branco, 28 April 2007, Rivadavia 2515 (SPF). Município de Lauro Müller, Serra do Rio do Rastro, 24 October 1958, Reitz & Klein 4075 (B, HRB NY, US). Município de São Francisco do Sul, Morro do Campo Alegre, 07 October 1960, Reitz & Klein 10057 (US), 04 November 1960, Reitz & Klein 10354 (US).

Specimens examined of the hybrid between D. latifolia and D. tomentosa:—BRAZIL. Minas Gerais: Município de Couto de Magalhães, ca. 30km ao leste de Diamantina, 12 May 2007, Rivadavia 2524 [mixed collection with D. latifolia] (SPF). Município de Diamantina, estrada para Tapetes Arraiolos, 12 July 1999, Rivadavia

& Pinheiro 1114 (SPF), 31 July 2002, Rivadavia & Gibson 1372 (SPF), 20 April 2010, Gonella et al. 272 (SPF).


Drosera riparia Rivadavia & Gonella, sp. nov. (Figures 2, 14, 15)Type:—BRAZIL. Bahia: Miguel Calmon, Parque Estadual das Sete Passagens, Cachoeira do Coração, 850 m, 03 August 2012,

Gonella et al. 543 (holotype SPF!, isotypes ALCB!, HUEFS!, MO!, P!, RB!).Diagnosis:—Species similar to Drosera latifolia, but differing by the long petioles that are equal in length or longer than the

lamina, glabrescent scapes with very short glandular trichomes, more delicate inflorescences, usually bearing fewer flowers, petals only up to 5 mm long, and obconical seeds.

Description:—Perennial rosetted herbs, acaulescent or more rarely with decumbent stems up to 45(–105) mm tall, sparsely covered by persistent dead leaves. Roots slightly succulent, up to 0.5 mm wide, sparsely branched, densely covered by root hairs. Indumentum consisting of white eglandular trichomes up to 1.5 mm long (brown when old), on the abaxial leaf surface, on adaxial petiole surface, scape, bracts, and sepals abaxially; glandular capitate trichomes 0.03–0.05 mm long, the stalk light red and translucent, the head dark red, present on scapes, bracts, pedicels, and sepals abaxially; minute sessile glands 0.02–0.03 mm in diameter, red in color, on both leaf surfaces, scapes, pedicels and sepals abaxially. Leaves 11–55 mm long, with circinate vernation, semi-erect, persistent when old, narrowly spatulate, apex obtuse, green to pinkish-red in color; petioles 5–28 mm long, 1–2 mm wide, narrowly ellipsoid in cross section, adaxial surface glabrous (or very rarely sparsely eglandular-pilose), abaxial surface sparsely (rarely densely) eglandular-pilose; lamina 5–27 mm long, 2–4 mm wide, oblong to narrowly oblong, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface glabrous to sparsely eglandular-pilose (specially towards the base and along the margins); stipules 2.5–7 mm long, 1–3 mm wide at the base, rectangular, membranaceous, pinkish to red (drying brown), the apical 1/2–1/3 divided into 3 long laciniate segments, the central wider and shorter. Scapes 1–2 per plant, (70–)115–200 mm long (including floriferous part), terete, thin, up to 1.7 mm diam. at the base, base erect (rarely slightly ascending), glabrous, becoming sparsely glandular-pilose towards apex, rarely with very sparse eglandular trichomes uniformly distributed along entire length, green to pinkish-red in color; inflorescence a scorpioid cyme, rarely bifurcating, bearing (1–)2–9(–13) flowers; bracts 1–1.5 mm long, filiform-lanceolate, caducous and thus usually absent in mature scapes, with sub-sessile glands, the basal bracts usually covered with few eglandular trichomes; pedicels 1.5–3 mm long, inserted 3–14 mm apart from each other, sparsely glandular-pilose; sepals 5, 3–4 mm long, 1–2 mm wide, ovate to elliptical, united at basal 1/3–1/2 of length, glabrescent, glandular-pilose, more rarely with few eglandular trichomes along the margins; petals 5, 4–5 mm long, 2–3 mm wide, obovate, pale pink in color; stamens 5, 2–2.5 mm long, anthers ca. 0.5 mm long, bithecate, yellow; ovary 3-carpellate, fused, 1–1.3 mm in diameter, globose, slightly 3-lobed in outline; styles 3, forked at the base, ca. 2 mm long (including stigmata), style arms curving inwards at the end of anthesis, stigmata flabellate, pale pink to whitish in color; fruit a dry dehiscent capsule, 1.5–2 mm long, ellipsoid, 3-valvate; seeds obconical, 0.7–0.9 mm long and 0.20–0.25 mm wide, testa reticulate, light to dark brown.

Distribution and ecology:—Drosera riparia is endemic to the Chapada Diamantina highlands of central Bahia state, Brazil (Fig. 2). This new species occurs in montane to high-montane regions at elevations between 850–1850 m, from the Serra das Almas in the municipality of Rio de Contas in the South, to Lençóis in the North, with a disjunct occurrence on the Serra de Jacobina, ca. 170 km northeast of Lençóis.

Drosera riparia is most commonly found along the margins of perennial rivers semi-shaded by surrounding rocks and trees, near waterfalls and along canyons, often associated with Sphagnum, but has also been observed growing in sandy soil in open seepages (Fig. 15A, B). It has been found growing sympatrically with D. communis, D. tomentosa var. tomentosa, and also occurs with other carnivorous species such as Genlisea aurea Saint-Hilaire (1833: 429) as well as several species of Utricularia.

Phenology:—Drosera riparia was collected in flower in January, March, April, May, July, August, and October, suggesting that it does not have any specific flowering period and will more or less flower throughout the year.

Etymology:—The epithet “riparia” refers to the habitat where this species is most commonly found, along river margins (Fig. 15D, E).

Conservation status:—Drosera riparia is widespread along the Chapada Diamantina highlands. Although seemingly rare, this is most likely an artifact resulting from insufficient field studies in that region. Fortunately most known populations are protected within a national park (Chapada Diamantina), a state park (Parque das Sete Passagens, Miguel Calmon), and a municipal park (Mucugê). Because only a few locations are currently known, D.

riparia is here considered Vulnerable (VU, criterion B2a, biii; IUCN, 2001).


FIGURE 14. Drosera riparia. A, habit. B, detail of the indumentum at the base of the scape. C, detail of the indumentum at the apex

of the scape. D, stipule. E, leaf, adaxial surface. F, leaf, abaxial surface. G, cross section of the petiole at mid length. H, calyx. J, petal.

K, stamen. L, gynoecium. M, seed. (All based on the holotype – drawn by P.M. Gonella.)


FIGURE 15. Drosera riparia. A, B, habitat at Mucugê, Bahia. C, flower. D, E, habit, note the long petiole compared to the lamina. (A,

B, D by J. Santos; C by A. Fleischmann; E by P.M. Gonella.)

Discussion:—Drosera riparia is distinguished from all other members of the complex by its leaves with oblong to narrowly oblong lamina, the petioles longer or as long as the lamina with the adaxial surface usually glabrous, the abaxial leaf surface glabrescent (very rarely densely eglandular-pilose on abaxial petiole surface),


scape, pedicels and sepals glabrescent, with very short capitate glandular trichomes (0.03–0.05 mm in length), inflorescences few-flowered [(1–)2–9(–13) flowers each], bearing small flowers (petals only up to 5 mm long) with short styles (ca. 2 mm long), and obconical seeds 0.7–0.9 mm long (Figs. 14, 15).

Although the relatively long petioles (compared to the lamina) may suggest a closer affinity with D. villosa, D.

riparia seems to be more closely related to D. latifolia, with which it shares the usually glabrous adaxial petiole surface, glabrescent abaxial leaf surface, and the glabrous or glabrescent lower one third of the scape. These characters suggest a closer affinity between D. riparia and the Espinhaço morphotype of D. latifolia, which is geographically closest among the morphotypes of the latter species (Table 1).

A few herbarium specimens of D. riparia were identified as D. communis by T.S. Silva, as well as in a picture in Silva (1994), possibly because of the long and distinctly separated petiole of the species. Drosera communis is easily distinguished by its orbicular to oblanceolate leaf lamina, a delicate, filiform scape which strongly arcuately ascending at the base, narrow sepals with large glandular trichomes, and rectangular-fusiform seeds.

Additional specimens examined (paratypes)—BRAZIL. Bahia: Município de Abaíra, Campo da Mutuca, 1400-1500 m, 21 March 1992, Stannard, Silva & Ganev H 52757 (SPF); Catolés, subindo riacho que desce da Serra do Guarda-mor, 1550 m, 26 July 1995, Rivadavia 476 (SPF); trilha para o Campo da Mutuca, Serra da Mesa, vila de Catolés, 1460 m, 15 July 2005, Rivadavia et al. 2018 (SPF). Município de Lençóis, Parque Nacional da Chapada Diamantina, Cachoeira do Sossego, 02 January 1993, Rivadavia 162 (SPF).

Município de Miguel Calmon, Piemonte da Chapada, 1139 m, 17 June 2006, J.S. Santos et al. 144 (ALCB); Parque de Sete Passagens, trilha cachoeira do Coração, 11 March 2006, V.J. Santos et al. 463 (HUEFS), 03 August 2012,Gonella et al. 540 (SPF). Município de Mucugê, Parque Municipal de Mucugê, trilha para a Cachoeira das Andorinhas, 13 May 2007, Roque et al. 1597 (ALCB); Cachoeira das Andorinhas, 1000 m, 06 October 2010, J.F.S.

Santos 1 (ALCB). Município de Piatã, Chapada Diamantina, trilha para a Cachoeira do Patrício, 1429 m, 10 October 2009, Roque et al. 2347 (ALCB). Município de Rio de Contas, trilha subindo para o Pico das Almas, 1630 m, 17 July 2005, Rivadavia et al. 2048 (SPF).

Drosera chimaera Gonella & Rivadavia, sp. nov. (Figures 2, 16, 17)Type:—BRAZIL. Minas Gerais, Cristália, em afloramento de arenito ao lado de banco de areia na margem direita do Rio

Itacambiruçu, 695 m a.s.l., 07 September 2011, Gonella et al. 500 (holotype SPF!, isotypes BHCB!, P!, RB!).Diagnosis:—Species morphologically intermediate between D. latifolia (Eichler) Gonella & Rivadavia and D. tomentosa

A.St.-Hil. var. tomentosa, differing from the former by the scape densely eglandular-pilose at the basal half and by the smaller ovoid seeds, and from the latter by the longer narrowly to broadly oblong-lanceolate semi-erect leaves.

Description:—Perennial rosetted herbs, rarely forming short columns up to 15 mm tall of persistent dead leaves. Roots slightly succulent, up to 0.6 mm wide, sparsely branched, densely covered by root hairs. Indumentum

consisting of white (brown when old) eglandular trichomes 1.5–2 mm long on the abaxial leaf surface and entire scape; capitate glandular trichomes up to 0.1 mm long, the stalk light red and translucent, the head dark red, present on the apex of scapes, pedicels and sepals abaxially; sparse minute sessile glands, ca. 0.03 mm in diameter, red in color, present on both leaf surfaces, scapes, pedicels and sepals abaxially. Leaves 10–30 mm long, with circinate vernation, semi-erect, decumbent when old, narrowly to broadly oblong-lanceolate, apex sub-acute to obtuse; petiole 1.5–7 mm long, 0.7–2.5 mm wide, reddish-green to red in color, transversely narrowly elliptical in cross section, adaxial surface glabrous, abaxial surface sparsely eglandular pilose to glabrescent; lamina 8.5–23 mm long, 2–5 mm wide, lanceolate to oblong-lanceolate, reddish-green to red in color, adaxial surface covered with numerous red, carnivorous, capitate tentacles, abaxial surface sparsely eglandular-pilose to glabrescent, trichomes concentrated along the margins; stipules 2.5–3 mm long, 1.5–2.5 mm wide at the base, rectangular, membranaceous, whitish to pinkish (drying brown), the apical 1/2 divided into several laciniae shorter towards the center. Scapes 1–3 per plant, (55–)90–270 mm long (including floriferous part), terete, 0.3–0.6(–1.3) mm in diam. at the base, erect from the base, densely eglandular-pilose on the basal third, trichomes decreasing in size and density towards the apex, and becoming very sparse to absent on the apical third, basal half with sparse sub-sessile glands, apical half glandular-pilose (increasing in size and density towards apex), red in color; inflorescence a scorpioid cyme, often bifurcating (rarely multiple branching), bearing (2–)4–19(–34) flowers; bracts 1.5–2 mm long, filiform to lanceolate, densely glandular pilose, often sparsely eglandular pilose abaxially, occasionally absent; pedicels 2–3.5 mm long, inserted 2–16 mm apart from each other, densely glandular pilose; sepals 5, 3–5


mm long, 0.6–1.8 mm wide, oblong-lanceolate to lanceolate, united to the basal 1/3 of their length, densely glandular pilose abaxially; petals 5, 4–6 mm long, 3–4 mm wide, obovate, pale pink in color; stamens 5, 2–3 mm long, anthers 0.5–1 mm long, bithecate, yellow; ovary 3-carpellate, fused, globose, 3-lobed in outline, ca. 1 mm in diameter; styles 3, forked at the base, 2–2.3 mm long (including stigmata), style arms curving inwards at the end of anthesis, stigmata flabellate or bilobed, pale pink to dark pink in color; fruit a dry dehiscent capsule, 2 mm long, ellipsoid, 3-valvate; seeds ovoid, ca. 0.4 mm long and ca. 0.2 mm wide, testa reticulate, black.

Distribution and ecology:—Drosera chimaera is narrowly endemic to the Serra de Grão Mogol in northern Minas Gerais, Brazil. It is thus far only known from three populations in the municipalities of Grão Mogol and Cristália (Fig. 2). Drosera chimaera occurs in lowland habitats of montane areas at elevations around 700 m. The type location is a dry south-facing rocky area on the margin of the perennial Itacambiruçu River, which separates the municipalities of Grão Mogol and Cristália (Fig. 17A). A second population was studied further upriver along a small tributary semi-shaded by riparian vegetation, growing in humid sandy soil on islands of vegetation or in a thin layer of soil over sandstone, sympatric with several species of Utricularia. The third population was not studied by any of the authors in the field, but according to herbarium data, it is also located along the margins of a river, in a habitat apparently very similar to that of the type population.

Phenology:—Drosera chimaera was collected with flowers and fruit in July, September and October, suggesting that the flowering period of this species is concentrated in the dry season, but extends into the early wet season.

Etymology:—Rosettes of D. chimaera resemble small specimens of D. latifolia, whereas the inflorescences and seed resemble those of D. tomentosa var. tomentosa. The combination of characters from different species brings to mind the legendary creature from Greek mythology called Chimaera, which was composed of parts from different animals.

Conservation status:—Because the authors only recently became aware of D. chimaera as a distinct taxon, not enough data is currently available to determine its geographical distribution, nor whether it is under any threat. It is thus here considered as Data Deficient (DD), according to IUCN Red List criteria (IUCN 2001).

Discussion:—Drosera chimaera is distinguished from related species by its lanceolate to narrowly oblong lamina, petioles much shorter than the lamina, with glabrous adaxial petiole surface, abaxial leaf surface sparsely eglandular-pilose, scape erect and densely eglandular-pilose (especially on basal third), often bifurcating, bearing small flowers (petals up to 6 mm long), and small ovoid seeds 0.4 × 0.2 mm (Figs. 16, 17).

Drosera chimaera is morphologically intermediate between members of the D. villosa and D. montana

complexes, sharing various characteristics from both groups, which may suggest a hybrid origin between species from these complexes. However, neither of the two D. chimaera populations studied are sympatric with any other Drosera species, excluding the possibility that this taxon is merely a hybrid between the regionally common D.

tomentosa and D. graomogolensis, although the possibility of a hybrid origin for D. chimaera is not excluded. Also, D. chimaera produces an abundance of fertile seeds, which breed true. All other know natural hybrids between New World Drosera species are sterile or produce few fertile seeds, as does D. × fontinalis, for instance.

Out of all D. villosa complex members, D. latifolia appears to be the most closely related to D. chimaera, sharing a similar leaf shape and indumentum. However, D. chimaera is distinguished by its dense indumentum of eglandular trichomes at the basal half of the inflorescence (decreasing in size and density towards the apex), by its ovoid seeds, and by its smaller leaf rosettes (Table 1). Drosera chimaera is morphologically also very similar to D.

tomentosa var. tomentosa, sharing the densely eglandular-pilose scape and seed shape, but the former species is easily distinguished by its larger semi-erect lanceolate to narrowly oblong leaves (decumbent and oblong-obovate in D. tomentosa var. tomentosa).

A hybrid origin from a cross between D. latifolia and D. tomentosa is also excluded by direct comparison with the observed natural hybrids between these two species (compare with Fig. 5B, D & E), which exhibit very low fertility and poor seed production (usually aborted ovules), the seeds elliptic in shape (intermediate between the fusiform seeds of D. latifolia and ovoid seeds of D. tomentosa).

The specimen Harley et al. 25067 (SPF) was considered by Rivadavia (2009) to be D. × fontinalis (a natural hybrid of D. tomentosa and D. grantsaui), due to the semi-erect lanceolate leaves and the base of scape with eglandular trichomes. However, this specimen is here excluded from D. × fontinalis and considered conspecific with D. chimaera. The hybrid D. × fontinalis can be easily distinguished from D. chimaera by its oblong to oblanceolate leaf lamina, flower scapes sparsely eglandular pilose at the base (often even glabrous), and by its obconic seeds up to 0.9 mm in length.


FIGURE 16. Drosera chimaera. A, habit. B–C, detail of the indumentum of the base (B) and apex (C) of the scape. D, stipule. E, leaf,

adaxial surface. F, leaf, abaxial surface. G, calyx. H, petal. J, stamen. K, gynoecium. L, seed. (All based on the holotype – drawn by

P.M. Gonella.)


FIGURE 17. Drosera chimaera. A, habitat at the type location near Cristália, Minas Gerais. B, flower. C, rosette seem from above,

with juvenile inflorescence. D, detail of the base of the inflorescence, with dense indumentum of eglandular trichomes. (All by P.M.

Gonella.)

Additional specimens examined (paratypes):—BRAZIL. Minas Gerais: Município de Cristália, margem

direita do Rio Itacambiruçu (type location), 12 August 2010, Gonella 359 (SPF, BHCB). Município de Grão

Mogol, estrada para Francisco Sá, 15 October 1988, Harley et al. 25067 (SPF); Capim Puba, 13 August 2010,

Gonella 360 (SPF), 07 September 2011, Gonella et al. 489 (SPF).


Conclusions

Detailed analysis of herbarium specimens, together with field surveys throughout the range of the D. villosa

complex, uncovered a much higher diversity than presented in previous works. Each of the six species accepted here as members of this complex can be recognized by a unique combination of morphological characters, of which the most relevant are: leaf shape and indumentum, scape indumentum, petal and style length, seed shape, as well as geographical distribution (Table 1).

The variable and widespread D. latifolia is essential to the understanding of speciation within this complex. Studies focusing on the phylogeography of this species are necessary to better comprehend its variability and whether the morphotypes deserve sub-specific classification. Also, further studies are necessary to uncover the phylogenetic relationships within the D. villosa complex and between this group and the closely related D.

grantsaui and the D. montana complex, the latter being the current focus of a revisional study (Rivadavia et al, in prep.).

Based on the morphological data presented here, hypotheses can be drawn to explain the relationships between the species of the D. villosa complex. Drosera ascendens and D. graomogolensis are deemed close, based on the large flowers with long styles, low self-fertility, as well as the glandular trichomes densely distributed along the whole scape, and the smaller slightly oblong-ovoid seeds. The remaining species of this complex share with D.

tomentosa the small flowers and style morphology, as well as the absence of glandular trichomes on the basal third of the scape. Drosera latifolia, D. villosa and D. riparia have in common the long and somewhat fusiform seeds. Finally, while leaf morphology supports that D. chimaera is allied to the species of the D. villosa complex, seed shape and scape indumentum place it closer to D. tomentosa, suggesting a basal position for D. chimaera, a hypothesis to be corroborated by future molecular studies.

Acknowledgements

We would like to thank Adilson Peres for discovering D. chimaera, rediscovering D. ascendens, sharing photos, location information, and also for providing flowers of cultivated specimens for measurements; Thiago De Roure Bandeira de Mello for providing essential information about the Sempre Vivas National Park and probable D.

ascendens habitats, and to Daniel Borges (PNSV) for the aid to access such locations; Laurent Legendre for providing pictures of type material at P; Jonathan Santos for discovering a new population of D. riparia, helping with measurements of this species, and providing photos used in this work; Carlos Rohrbacher for his skill and dedication in cultivating carnivorous plants, including material used in the present study; Rolf Grantsau for preliminary morphological observations and drawings; Wilson Ganev and Joás Brandão for helping to discover D.

riparia; Josef Mullins, Robert Gibson, Lúcio Leoni, Igor Lins, Vitor Miranda, Linilson Padovese, Mauro Peixoto, Adilson Peres, Fábio Pinheiro, Ed Read, Carlos Rorhbacher, Jonathan Santos, Nílber Silva, Anderson Alves, André Vito Scatigna, Gustavo Shimizu, and many others who have helped with field studies of the D. villosa complex over the past 23 years; and finally to ICMBio for providing the permits for fieldwork. We also thank two anonymous reviewers for useful comments on the manuscript. This work is part of PMG’s M.Sc. thesis and was supported financially by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq). We also thank CNPq for a grant to PTS.

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