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Phytotaxa 576 (1): 029–054
https://www.mapress.com/pt/
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Andreas Fleischmann: 22 Nov. 2022; published: 15 Dec. 2022
https://doi.org/10.11646/phytotaxa.576.1.2
29
Unveiling Utricularia amethystina’s true colours: a taxonomic revision of one of the
largest species complexes (U. sect. Foliosa, Lentibulariaceae)
PAULO C. BALEEIRO1,2,5*, PAULO M. GONELLA3,6, PAULO T. SANO4,7 & RICHARD W. JOBSON2,8
1Department of Biological Science, The University of Queensland, St Lucia, Qld 4076, Australia.
2National Herbarium of New South Wales, Royal Botanic Gardens and Domain Trust, Mrs Macquaries Road, Sydney, NSW 2000,
Australia.
3Universidade Federal de São João del-Rei, Departamento de Ciências Exatas e Biológicas, Campus Sete Lagoas, Rodovia MG-424, km
47, 35701-970, Sete Lagoas, MG, Brazil.
4Department of Botany - Institute of Biosciences, University of São Paulo. Rua do Matão, 277, 05508-090, São Paulo, SP, Brazil.
5
�
uqpbalee@uq.edu.au; https://orcid.org/0000-0002-0484-3166
6
�
pmgonella@gmail.com; https://orcid.org/0000-0001-8332-5326
7
�
ptsano@usp.br; https://orcid.org/0000-0002-1709-1215
8
�
richard.jobson@botanicgardens.nsw.gov.au; https://orcid.org/0000-0002-1822-9634
*Author for correspondence.
Abstract
The genus Utricularia (Lentibulariaceae) is the most diverse and cosmopolitan among carnivorous plants. Utricularia sect.
Foliosa sensu Taylor (1989) is composed of three South American species, U. amethystina, U. tricolor and U. tridentata,
the first species being recognized as highly diverse. In his taxonomic revision of the genus Utricularia, Taylor (1989)
circumscribed the polymorphic U. amethystina comprising 27 synonyms involving a wide variation in corolla size and
colour, morphology of corolla parts, indumentum, and overall plant size. The original publications of the synonyms
provide limited information disencouraging a revision study and therefore inflating U. amethystina as new variations were
discovered. In order to shed light onto the problem, a taxonomic revision, a morphometric and a molecular study showed
evidence of a larger complex than previously thought, and U. amethystina s.str. sister to U. tricolor. A previously published
molecular phylogeny provided strong support for all sampled U. amethystina synonyms and potential new taxa found to
be nested outside the U. amethystina s.str. and U. tricolor clade. Here, we incorporate these new relationships and describe
and comment on the species that comprise U. sect. Foliosa, including the restablishment of six synonyms, the description of
three new species and lectotypification of five names. The current study disentangles U. amethystina complex so that further
studies can approach it in a more realistic way, especially studies using population genetics tools. Our study shows that more
needs to be done and here U. hirtella is presented as a new complex.
Keywords: carnivorous plants, Lamiales, Neotropical biodiversity, new species, taxonomy
Introduction
Plant monographs of classical taxonomy have a great impact on elucidating biodiversity and it is the backbone for
the discovery of new species and disentangling of unrealistic diversity (Grace et al. 2021). The accurate association
of natural populations to their designated type specimens is essential, and it is made by conference of holotypes and/
or isotypes and their original publications, so called protologues. When a taxon carries synonyms more attention is
necessary so that the right populations are selected and therefore the study is optimized showing the best scenario.
The genus Utricularia Linnaeus (1753: 18) (Lentibulariaceae, Lamiales) presents a nearly cosmopolitan
distribution and comprises around 250 species, being one of the two largest carnivorous plant genera together with
the only distantly related Drosera Linnaeus (1753:281) (Droseraceae, Caryophyllales). The genus comprises small to
medium sized herbs that inhabit a great diversity of habitats, from terrestrial to aquatic, from rheophytic to epiphytes.
Utricularia was last revised in the celebrated work of Taylor (1989), who presented a treatment for 214 recognized
species from an initial list of over 600 names, also presenting an infrageneric classification that found strong support
in later molecular works. Among these 214 species, it is fair to state that no other species gave Taylor (1989) a harder
BALEEIRO ET AL.
30 • Phytotaxa 576 (1) © 2022 Magnolia Press
time than U. amethystina Salzm. ex Saint-Hilarie and Girard (1838: 870), from U. sect. Foliosa Kamiénski. In his
monograph, Taylor (1989: 291) states that “Utricularia amethystina is a most ‘difficult’ and excessively polymorphic
species, and although I am far from satisfied with the circumscription here proposed, a long study of a large amount of
material has failed to provide me with anything better”.
As of today, Utricularia is segregated into three subgenera U. subgen. Polypompholyx (Lehmann, 1844: 48)
(Taylor, 1986: 1), U. subgen. Bivalvaria (Kurz, 1874: 54), and U. subgen. Utricularia, and 37 sections differentiated
based mainly on trap morphology (Taylor 1989; Jobson et al. 2018). Within U. subgen. Utricularia, U. sect. Foliosa
(Kamieński, 1891: 120) sensu Taylor (1989) contains the three species, U. amethystina, U. tricolor (Saint-Hilare,
1833: 418) and U. tridentata (Sylveìn, 1909: 28), that are mainly distributed across South America to the Caribbean,
with most of the diversity occuring in the Cerrado domain region of Brazil, and the Guiana Shield (Fig1C) (Baleeiro
et al. 2019). A fourth member of U. sect. Foliosa was recently described, U. biceps Gonella & Baleeiro (2018: 215).
Taylor (1989) noted that U. amethystina, unlike the other two accepted species, was a polymorphic taxon under
which he listed 27 synonyms. Included in his list of U. amethystina synonyms there seemed to be some well-defined
species, although taxonomic limitations such as continuous morphological characters led to the lumping of all synonyms
under a single name (Taylor 1989). These polymorphic taxa contain wide variation of qualitative and quantitative
characters and the whole section was in need of a detailed phylogenetic study (Baleeiro et al. 2016, 2019).
After an extensive systematic revision of synonyms’ protologues, type specimens, and specimens not matching any
of the synonyms, an attempt was made to include populations representing most of these synonyms in a morphometric
study, in order to recognize or not morphologically distinct taxa. The results of Baleeiro et al. (2016) presented
morphometric analyses of floral characters that indicated where gaps in morphological variation supported the
resurection of several synonyms to species status. These morphological divisions were further supported by molecular
phylogenetic results that showed accessions representing the type of U. amethystina (Fig.2) fall into a clade that was
sister to U. tricolor (Baleeiro et al. 2019). The same study also showed that all of the examined accessions representing
synonyms fell into clades outside the U. amethystina-U. tricolor grouping, and several taxa not previously included as
synonyms formed clades supporting their new species status. Therefore, the results indicate that either several species
needed to be uncovered, or U. amethystina s.l. would have to make room for U. tricolor and U. tridentata.
As lumping by far is not the best decision in this case, we here utilize the molecular phylogenetic hypothesis of
Baleeiro et al. (2019) in a taxonomic treatment of Taylor’s U. sect. Foliosa circumscription, with taxa resurrected
from synonymy under U. amethystina and new taxa named, and compared with the morphology of related species. We
include notes on distinguishing characters, distribution and ecology, phenology, and conservation status. In addition, a
taxonomic key is provided for identification with terminology following Taylor (1989).
Material and methods
For this study, approximately 700 herbarium sheets were examined including type specimens of all synonyms (F, K,
M, MO, NY, P, R, SPF, US, W). Other herbaria in Brazil holding important collections from savannah formations
were also examined: ASE, BHCB, CEN, CEPEC, ESA, ESAL, HEPH, HMS, HUEFS, IBGE, INPA, JPB, MBM,
MG, MIRR, NX, UB, UFG, RB, SP, TANG, UEC, UFMT as well as the Argentinian MERL (Acronyms from Thiers,
continuously updated).
Fieldwork was carried out at targeted sites across South America in order to observe plants in their habitats and to
collect samples for the morphometric study (Baleeiro et al. 2016) (250 specimens in alcohol-preserved inflorescences
were used), molecular phylogeny (Baleeiro et al. 2019), along with additional samples for this revision, representing
the wide morphological and geographic range of the complex, these specimens being deposited at SPF.
Measurement of quantitative variables were made using Leica Application Suite software (LAS V. 3.7) with a
stereoscopic light microscope (Leica M125) and camera lucida (DFC 425).
Maps were prepared with DIVA-GIS (Hijmans et al. 2012) with data input as co-ordinates, and displayed as
dots representing geographic distribution and elevation of sites of populations within the regions of Guayana Shield,
Cerrado, Pantanal, and Northern Territory in Brazil.
Conservation status was assessed using IUCN Red List 3.1 Categories and Criteria (IUCN 2012). The Conservation
Assessment Tool described in Bachman et al. (2011) was used to estimate the extent of occurrence (EOO) and area of
occupancy (AOO), for which the standard IUCN cell size of 4 km2 was employed.
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 31
Results
Taxonomic delimitation
This new revision proposes that most of the entities within U. sect. Foliosa under synonymy should be maintained
as synonyms, particularly those under U. tridentata and U. tricolor (Taylor 1989). Regarding U. amethystina, from
the 27 binomials under synonymy (Taylor 1989), seven are here recognized as valid species, while the remaining 20
names remains under synonyms but rearranged across the recognized taxa. After the recent resurrection of U. trinervia
(Benjamin, 1847: 247) to species status by (Guedes et al., 2021) we will not treat this taxon in current study. The species
resurrected in the current study are U. bicolor Saint-Hilaire & Girard (1838: 870), U. damazioi (Beauverd, 1907: 703)
U. lindmanii Sylvén (1909: 26), U. hirtella Saint-Hilaire & Girard (1838: 869), U. roraimensis (Brown, 1901: 54), and
U. velascoensis (Kuntze, 1898: 240). In addition, the four previously recognised species U. amethystina, U. biceps,
U. tridentata and U. tricolor are here maintained at the specific level. Morphotypes that are not currently assigned to
any of these binomials (novel taxa uncovered during the studies of Baleeiro et al. 2016, 2019) are here described at the
specific level (refer to ‘Comments’ within U. sect. Foliosa description below). The new species presented here are U.
lunaris Baleeiro, Gonella & R.W.Jobson (Fig. 3H–I), U. pantaneira Baleeiro, Gonella & R.W.Jobson (Fig. 4L), and U.
chapadensis Baleeiro, Gonella & R.W.Jobson (Fig. 4M–N).
Moreover, Baleeiro et al. (2019) provides evidence of potential new taxa under the phrase names U. sp “Noel
Kempff”, Utricularia sp. “Marahuaka” and U. sp. “Minas Gerais”, in addition to potential of new taxa within the
highly diverse U. hirtella complex (Fig. 4C–D). Unfortunately, the current study does not utilize enough data to
sufficiently address these potential new taxa.
Taxonomic treatment
Utricularia sect. Foliosa Kamieński, Lentibulariaceae preprint from Nat. Pflanzenfam. 120 (1891)
TYPE: Utricularia amethystina Salzm. ex A.St.-Hil. & Girard. (Designated by P. Taylor).
Illustrations: figs.77a,b; 78; 79 P.Taylor (1989); fig.2 Baleeiro et al. 2016, fig.1 Baleeiro et al. 2019.
Description. Terrestrial, affixed sub-aquatic or lithophitic herbs. Rhizoids few to many, simple or ramified branches,
normally present semi-aquatic species; stolons few to many, filiform. Leaves simple at peduncle base, or rosulate,
lamina filiform, spathulate to circular, base attenuate, apex rounded, ascendant or prostate, 1- or multinerve (0–)1–
7; traps globular, stalk basally inserted c. 1 mm long, two dorsal deltoid appendages with trichomes on abaxially
portion and around trap door, chin-like swelling at base of trap present or absent. Inflorescence simple or ramified,
erect, glabrous or papillose near base; scales, bract and bracteoles basifix, entire, glabrous; bract deltoid to broadly
ovate; bracteoles linear to ovate, united at some degree with the bract; calyx lobes similar or unequal, glabrous,
papillose, or with simple or glandular trichomes, margin entire; corolla bilabiate with spur, lower lip violet with yellow
surrounded by white spot at base of, or entirely yellow, or white with yellow spot at base; upper lip ovate, elliptic
or circular, glabrous or papillose, or simple or glandular trichomes; lower lip obovate to transversally elliptical, base
with a prominent bilobed swelling, apex ± entire to deeply 3-lobed. Filament curved to ± straight, theca distinct or
subdistinct; polen oblade to prolate, 6–15 colporate. Ovary globose, glabrous or glandular, style usually shorter than
the ovary, stigma bilabiate, lower lip semicircular with ciliate margin, upper lip reduced. Capsule globose 1.5–4mm
diam., dorsiventrally bivalvate. Seed ovoid to narrowly cylindrical.
Distribution and ecology across the biodiverse regions Cerrado and The Guiana Shield: Utricularia sect.
Foliosa is found in most of the countries in South and Central America, Mexico and possibly USA (Fig.1); however,
it is in the Brazilian Cerrado (including campos rupestres) and the Guiana Shield formation the responsibility for the
striking diversity not only inside this section, but also for the entire genus Utricularia (Taylor 1989). These adjacent
complex regions have been documented highly important due to their high levels of diversity and endemisms of
vascular plants (Giulietti et al. 1997; Kelloff & Funk 2004; Funk et al. 2007). U. sect. Foliosa habitat elevation ranges
from sea level in the Brazilian restingas to 2994 m on Neblina peak in Brazil and countless table mountains in the
pantepui province (e.g. Mt. Roraima and Kukenan tepui ~ 2800 m). Even though a range of different elevation is
seen with one of the newest species from Pantanal wetlands shown in this study, the greatest levels of diversity and
endemism seem to be associated with high elevations and cooler climates in three main areas, such as the campos
rupestres of the quartzitic massifs of the Chapada dos Veadeiros (Souza & Bove 2012) and Espinhaço Range in Brazil
BALEEIRO ET AL.
32 • Phytotaxa 576 (1) © 2022 Magnolia Press
and the pantepui province of the Guiana Shield. These regions have been traditionally reported as showing similar
levels of richness and distribution for other group of plants (Rapini et al. 2008; Rando & Pirani 2011).
The “archipelago” nature provided by the pantepui province with tabletop mountains above the surrounding
rainforest is a unique biogeographic region and harbours an incredible diversity. The evolutionary historic pathways
provided by these so called “islands in the sky” across the landscape is still controvertial, but nonetheless, their old
uplifting age has probably been providing as a barrier and limiting genetic flow, in the flora of the different tabletop
Mountains. The Pantepui is less than one percent in area of the total Guiana Shield. Yet, it harbours about 2450
vascular plant species, with 60% endemic to the Guiana Shield, 40% to the Pantepui and 25% to a single mountain
(Berry & Riina 2005). According to the same authors, the genus Utricularia comes in the list of the richest genera for
the Pantepui flora (21spp.), but we think this number is underestimated as new species of animals (Kok, 2010; Kok et
al., 2015, 2018) and not as much of plants (Barbosa-Silva et al., 2018) have been introduced.
FIGURE 1. Geographic distribution map of Utricularia sect. Foliosa species. A. Distribution of all 14 species in South and Central
Americas. B. Map of the species distribution in the Espinhaco range and Chapada dos Veadeiros National park. C. Map of the species
distribution at the Guiana Shield.
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 33
The study of Vasconcelos et al. (2020) shows recent and rapid radiations (around 2–5 Myr) of different linneages
on an ancient formation of campos rupestres in Brazil. It points out studies of plant radiations in such old mountains,
unlike what is more recently seen in the literature of lineage age matching mountain uplifting. Another study show
recent radiations (2–4 Myr) rather than old of the endemic lineages of frogs Tepuihyla (of the 7 spp., 4ssp. are known
from a single tepui) in an ancient region this time the pantepui province presented by Salerno et al. (2012).
Notes: Examination of all binomials and morphotypes of U. sect. Foliosa using morphometric analyses (descriptive
and multivariate) (Baleeiro et al. 2016) and molecular phylogenetics (Baleeiro et al. 2019) highlighted the need to
reassess 11 synonymised taxa. Seven of these were considered accepted species prior to Taylor`s taxonomic treatment
of the genus and are here raised to species level (Taylor 1989). A further three non-synonymised taxa included in the
study are here described as new species.
Characters that group the taxa in the section include the basally positioned trap door with two trichome-covered
dorsal appendages, bract and bracteoles basifix, bracteoles usually attached near the base or up as high as the middle
length of the bract, and capsule dehiscence via a dorsiventral slit. Key defining characters between species include the
shape and size of corolla lower and upper lip, calyx, and corolla colour and indumentum, and the pedicel and peduncle
length (Baleeiro et al. 2016). At the between section level defining characters include the small stature of the stamens
that does not differ across the section, although the size of which is not seen across the genus. Pollen size and shape
can vary strongly between species, but also within the same population or on an individual. Therefore, these two latter
characters are not considered useful for differentiation at the specific level. In addition, a morphometric study of seed
size and shape across and between U. sect. Foliosa and U. sect. Psyllosperma (Menezes et al., 2014) shows that there
is insufficient morphological information for differentiation at the specific level and sectional levels.
Taxonomic key Utricularia sect. Foliosa
1. Corolla purple to pale pink .................................................................................................................................................................2
1´. Corolla white or yellow ....................................................................................................................................................................12
2. Calyx with papillae and/or trichomes near margin .............................................................................................................................3
2´. Calyx glabrous ....................................................................................................................................................................................4
3. Corolla lower lip with white and yellow spot, spur x2> lower lip, upper lip ovate x3> than calyx ...........................U. velascoensis
3´. Corolla lower lip with only yellow spot, spur slightly larger than lower lip, upper corolla lip rounded x2> than calyx ... U. bicolor
4. Corolla lower lip base with a bilobed protruded crest with tuberculate texture, spur cylindrical with apex inflated and bilobed ......
...............................................................................................................................................................................................U. biceps
4’. Corolla lower lip base with crest not protruded and smooth texture, spur conical tapering towards the simple apex (eventually also
bilobed) ...............................................................................................................................................................................................5
5. Corolla lower lip entire to shallowly 3-lobed, spur the same length or slightly larger than the lower lip, lower sepal much smaller
than upper sepal, pedicel half the size of upper sepal ........................................................................................................................6
5´. Corolla lower lip deeply 3-lobed, spur usually x2> than lower lip, both sepals similar in size, pedicel at least x2> the size of upper
sepal ....................................................................................................................................................................................................7
6. Inflorescence 240–750 mm long, corolla 10–25 mm long ..................................................................................................U. tricolor
6´. Inflorescence 40–80 mm long, corolla up to 8 mm long ..............................................................................................U. roraimensis
7. Corolla lower lip same length or smaller than upper lip; leaves linear ........................................................................... U. lindmanii
7´. Corolla lower lip much larger than the upper lip, leaves spathulate to almost circular .....................................................................8
8. Inflorescence peduncle robust (ca. 1mm wide), leaves numerous in a rosette ...................................................................................9
8´. Inflorescence peduncle not robust (<0.8mm), leaves absent or usually not forming a rosette ........................................................10
9. Corolla minutely papillose, lower lip shallowly 3-lobed, leaves mostly circular ........................................................... U. tridentata
9`. Corolla glabrous, lower lip distinctively 3-lobed, leaves mostly spathulate ............................................................... U. amethystina
10. Corolla purple with white and yellow spot, spur glabrous, leaves absent or spathulate to almost circular from a short attenuate
base ................................................................................................................................................................................... U. damazioi
10’. Corolla pale pink with white and yellow spot, spur with trichomes or stipitate papillose, leaves obovate to circular or almost
reniform from a short or long attenuate base ....................................................................................................................................11
11. Calyx with papillae and/or trichomes at least near the margin; leaves mostly absent, or obovate, reniform, to almost circular ........
............................................................................................................................................................................................. U. hirtella
11’. Calyx glabrous; leaves circular or reinforms from a long attenuate base ........................................................................... U. lunaris
12. Calyx with papillae and/or trichomes at least near the margin............................................................................................U. hirtella
12´. Calyx glabrous ..................................................................................................................................................................................13
13. Corolla white ............................................................................................................................................................... U. amethystina
13´. Corolla yellow ..................................................................................................................................................................................14
14. Corolla pale yellow up to 10 mm long, traps monomorphic .............................................................................................U. trinervia
14´. Corolla bright yellow 15–25 mm long, traps dimorphic ..................................................................................................................15
15. Straight spur up to x2> than lower lip ........................................................................................................................... U. pantaneira
15´. Curved spur at least x3> than lower lip .......................................................................................................................U. chapadensis
BALEEIRO ET AL.
34 • Phytotaxa 576 (1) © 2022 Magnolia Press
1. Utricularia amethystina Salzm. ex Saint-Hilaire & Girard (1838: 870)—Figs. 2, 3A–C, E–F
Type:—BRAZIL. Bahia: no date, Salzmann s.n. (Holotype: P!; Isotypes: F!, G image!, K!, LE, MEL!, MO!, MPU, P!, R!, W).
≡Calpidisca amethystina (Salzm. ex A.St.-Hil & Girard) Barnhart (1931: 468)
=Utricularia modesta De Candolle (1844: 17)
=Calpidisca modesta (A.DC.) Barnhart (1931: 469)
Type:—BRAZIL: Pará, no date, Poeppig 3017 (Holotype: G image!; Isotypes: B (destroyed; photo, M), LE, P image! (photo, M), W).
=Utricularia obovata Miquel (1847: 144)
Type:—SURINAME: Onoribo, no date, Focke 1051 (Holotype: U3678).
=Utricularia versicolor Benjamin (1847: 488)
Lectotype (designated here):—VENEZUELA: Monegas: Maturin, Otto 946 (P!).
=Utricularia spatulifolia Pilger (1914: 190)
Type:—VENEZUELA: Roraima, no date, Ule 8757 (Lectotype designated here: K!; Isolectotype: F!,G image!).
=Calpidisca standleyae Barnhart (1933: 1234)
=Utricularia standleyae (Barnhart) Rickett (1967: 518)
Type:—USA: Florida: Lee County, “Vicinity of Fort Meyers” Among (c.f.) Palmetos; Standley 406 (Holotype: NY!; Isotype: US!).
=Utricularia williamsii Steyermark (1950: 126)
Type:—HONDURAS: oak-pine forest near Hoya-Grande, Río Yeguare drainage, Williams 10639 (Holotype: F!; Isotypes: BM!, EAP).
=Utricularia turumiquirensis Steyermark (1953: 552)
Type:—VENEZUELA: Sucre, Cerro Turumiquire, Steyermark 62752 (Holotype: F!; Isotype: VEN).
Illustrations:—Taylor (1989), figs. 77a, 77b, showing a mix of specimens from different localities; Baleeiro et al. (2016), figs. 2a,g;
Baleeiro et al. (2019), fig. 1a.
Description:—Terrestrial or affixed sub-aquatic. Small to medium sized annual herb. Rhizoids 1–4, up to 14 mm
long, branches ramified. Stolons numerous. Leaves 1–7, rosulate from peduncle base, lamina normally spathulate to
circular (15–)30–40 × 1.5–8.5 mm; traps numerous on rhizoids and stolons, c. 1 mm long, globular, the trap door basal,
stalk deltoid 0.5mm long. Inflorescence simple or loosely ramified (30–) 40–300 (–400) mm long, peduncle glabrous,
capillary or sometimes thicker near the base, 0.35–1.3 mm diam.; scales basifix, glabrous, ovate, margin entire (3–6) c.
1.1 mm long; bracts and bracteoles unequal, basifix, often connate near base, glabrous without nerves, bract broadly
ovate, apex acute, bracteoles linear, apex obtuse 1.2–2.0 mm long. Flowers 1–9 (usually single flowered); pedicel
erect, filiform (–1) 2–11 (–18) mm long, 0.4 mm diam.; calyx lobes sub-similar, cartaceous, concave, glabrous, with
inconspicuous simple nerves, upper lobe broadly ovate or oblong, apex rounded, entire margin hyaline (1.5–) 2.5–3.0
(–3.9) mm long.; lower lobe ± broadly elliptic, ± entire margin not hyaline, apex rounded or emarginate, 1.3–2.8 mm
long; corolla lilac with single yellow spot bordered by white at base of lower lip, spur white at base with apex lilac,
7–12 mm long; upper lip glabrous, oblong, apex rounded 1.35–4.37 × 1.41–4.13 mm; lower lip broadly obovate,
base forming bilobed swelling, apex shallowly 3-lobed, 1.4–5 × 2.22–6.73 mm; spur conical from the base, apex
obtuse 3.2–7.0 mm long, sparsely glandular; filaments slightly curved, anther thecae 1/3 the size of the filament;
ovary globose, some glands present, style very short, stigma lower lip semicircular, upper lip absent. Capsule globose
2–3mm diam., calyx lobe not embracing the stigma lips. Seeds obliquely ovoid, c. 0.35 mm long 0.15 mm wide.
Distribution and ecology:—Widely distributed across northern South America, from the Brazilian Northeast
coast (i.e. restingas and tabuleiro ecoregions) (Fig.1A,C), through to the Guiana Shield (Brazil, Guyana; French
Guiana; Surinam, Venezuela), Amazonia lowland savannahs (Brazil and Peru), and on Trinidad and Tobago (Taylor
1989). Grows in water-soaked sand or clay substrate. On the Brazilian Northeast coast, it is distributed across the states
of Bahia, from Maraú Peninsula’s restinga and further north; Sergipe, from Itabaiana mountain and coastal restinga;
Paraíba, at Guaribas Biological Reserve; Rio Grande do Norte, Piauí and Maranhão. In North Brazil it is found in
the states of Pará (at Restinga or Mangrove), Amapá, Amazonas, and Roraima (Viruá National Park to Tapequém
Mountain at northwest of Roraima state). In Venezuela it is found at Gran Sabana and some Tepuis, Guyana to French
Guiana; and far west of Amazonia in Peru, a savannah site of Santuario Nacional de las Pampas del Heath. Flowering
all year round, specially from Jun to Oct. Refer to species examined.
Etymology:—From the Greek amethystos meaning violet, referring to the lilac corolla.
Conservation status:—Least Concern. Based on its broad distribution range (EOO = 7,693,931 km2; AOO = 228
km2) and frequent presence in protected/conservation areas (Fig. 1), U. amethystina is here considered Least Concern
under (IUCN, 2012) categories and criteria.
Notes:—After Taylor’s (1989) monograph, U. amethystina was reported to be a polymorphic species. As a
result, the whole U. section Foliosa became a large species complex, with all specimens not fitting the narrowly
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 35
circumscribed U. tricolor and U. tridentata being placed in a broadly circumscribed U. amethystina by exclusion.
Utricularia amethystina s.str., as defined here, is the most common and widespread species of the complex in northern
South America. The common morphotype from the Brazilian Cerrado domain, with deeply 3-lobed lower corolla lip,
is here recognized as U. damazioi. A typical character that differentiates U. amethystina from the others is the lower lip
of corolla being slightly 3-lobed, with half to 2/3 of corolla lower lip base swollen and given the name gibba; and the
leaves usually forming a rosette. For some, U. amethystina s.str. used to be mistaken as a stunted form of U. tricolor,
a similarity that is supported by their closer position in the phylogenetic hypothesis of Baleeiro et al. (2019).
FIGURE 2. Utricularia amethystina. A. Habit showing leaves, stolons, bladder traps and rhizoids, B. Inflorescence, C. Calyx
frontal view, D. Corolla frontal view, E. Corolla rear view, F. Bract and bracteoles, G. Bladder trap, H. Capsule, I. Stamens, J. Stamens
and ovary showing dorsal stigma surface, K. Seeds.
As described by E. Ule (n. 8757) in the original publication of U. spatulifolia from Mt. Roraima’s lower campos
(1600 m) “The species is related to U. roraimensis N.E.Br., but it is distinguished by its larger growth, longer spur etc.”.
However, here we list this name as synonym of U. amethystina s.str. and not U. roraimensis, due to morphological
affinities with the former. Besides that, the collections of Steyemark 59648 (F, NY), 59742 (US), 60154 (US) under
the name U. spatulifolia and original from Ptari Tepui is here designated as U. amethystina. Future molecular studies
of well-sampled populations from across the range might shed light and provide more evidence for elevating these
populations to associated names under synonymy or new names. As mentioned before, studies suggest that the tepuis
from the Guiana Shield harbour high levels of endemicity and that these regions are still poorly explored for many
group of plants.
BALEEIRO ET AL.
36 • Phytotaxa 576 (1) © 2022 Magnolia Press
The USA population collected on 14 Jan 1946 in Florida (L.J.Brass 15873, FSU 469) is described to have pale
yellow flowers, but due to lack of sufficient material and since its natural population hasn’t been re-collected, this
could not be placed under one of the names in the current monograph and the name U. standleyae remains as synonym.
The Mexican population from Guerrero have yellow or white flowers and is found in savanna habitats around 1230m
above sea level (Rendón and Ruiz, 2017). These populations are shown with a star in the map of distribution of species
(Fig.1).
Specimens Examined:—BRAZIL. BAHIA: Camaçari, km 11 da estrada Salvador-Camaçari-Dias d`Avila, em
grande área brejosa a direita, 1 Jul. 2002, Rivadavia 1458 (SPF). Ituberá, encostas de morros com nascentes rodeando
braço de mangue, a direita do km 15 da estrada p/ a praia de Pratigi, 8 Out. 2005, Rivadavia 2109 (SPF). Maraú, ca.
8km na estrada Maraú/Ubaitaba, 4 Set. 1999, Carvalho et al. 6768 (NY!, CEPEC!); ibid, 22 Jun. 2015, Baleeiro 384
(SPF); ibid, estrada sentido Itacaré/Maraú, depois da travessia do Rio de Contas, ca. 2 km, restinga arbórea, 7 Ago.
2005, Paixão et al 560 (CEPEC); ibid, estrada de Ubaitaba para a Barra Grande, na bifurcação para Itacaré, 8 Out.
2005, Rivadavia 2115 (SPF); ibid, in restinga area, 3.8km N of the Rio de Contas at Itacaré ferry crossing on road to
Maraú, 05 Aug. 2002, W.W. Thomas et al. 13121 (CEPEC). Mata de São João, brejo a direita no km 94 da rodovia do
coco (BA099), próximo a praia de Massarandupio, 18 Jan. 2003, Rivadavia 1523 (SPF). Santa Luzia, km 31 da estrada
Camaca-Canavieiras, a 10 km de Hermilandia, 26 Mar. 2005, Rivadavia 1971 (SPF). MARANHÃO: Barreirinhas,
restinga alagada próximo a Lagoa Azul, Lençóis Maranhenses, 30 Mar. 2007, Rivadavia 2499 (SPF). PARÁ: Maracanã,
Ilha de Maiandeua, 13 Jun. 1994, M.N. Bastos et al. 1688 (MG). Marapanim, estrada de Maruda para Praia do Crispim,
7 Ago. 2005, Rivadavia 2100 (SPF). Santarém, estrada Santarém-Alter do Chão, em brejo com buritis ao lado direito
da estrada, 4 Mar. 2000, Rivadavia & Pozza 1194 (SPF) . Vigia, km 16 da estrada de Vigia para São Caetano de
Odivelas, 7 Ago. 2005, Rivadavia 2104 (SPF). PARAÍBA: Mamanguape, Reserva Biológica Guaribas, área 1 Capim
Azul, 187 m, 19 ago. 2002, A.C. Sevilha & G. Pereira-Silva 2250 (CEN); ibid. Reserva Biológica Guaribas, área I -
Capim Azul, 22 Jul. 2012, Silva 4 (JPB). RORAIMA: estrada Porto Velho-Cuiabá, BR 364, km 159, margem esquerda,
vegetação rupestre, 40m mais elevada que o resto da área, 9 Fev. 1983, Teixeira et al. 1400 (INPA). Caracaraí, Manaus-
Caracaraí Road, km 130, Igarapé Lages, swampy ground on rocks, 16 May 1974, Prance et al. 21041 (NY!, INPA!).
Parque Nacional Viruá, na grade da PPBIO, L1/N6, campina, 12 Set. 2010, Costa & Barbosa 695 (INPA); ibid, Estrada
Perdida, 22 Jul. 2010, Cavalcati et al 187 (INPA); ibid, campinarana gramíneo-lenhosa, com muitas Malpighiaceae,
Clusiaceae, área alagável com solo arenoso, nas porções mais secas, areia branca, 12 Sep. 2010, El Ottra et al. 65
(INPA); ibid, grade da PPBIO L3-500, 04 Sep. 2012, Baleeiro 231 (SPF); ibid, estrada Perdida, próximo à guarita lado
direito em direção à primeira bueira, 06 Sep. 2012, Baleeiro 249, 250, 256 (SPF). Boa Vista, Zona Urbana, 07 Sep.
2012, Baleeiro 263 (SPF); ibid, beira da estrada em igarapé do lado direito entre Boa Vista e entrada para Tepequém,
08 Sep. 2012, Baleeiro 268 (SPF). Serra do Tepequém, atrás da pousada da Gisele, 09 Sep. 2012, Baleeiro & R. Perdiz
270 (SPF). SERGIPE: Areia Branca, E. Ecológica Serra de Itabaiana, 16 Sep. 1995, Landim et al. 670 (UEFS); ibid, 28
Sep. 2013, Carregosa-Silva & E. Santos 501 e 502 (ASE). Pirambu, povoado samambaia, 09 Sep. 2013, Carregosa-
Silva et al. 500 (ASE).
COLOMBIA. Upper Apaporis basin, Macaya river, Mount Chiribiquete, 24 Jul. 1943, Schultes 5618 (US). Cerro
Castilho, 27 Jul. 1943, Schultes 5668 (US). Rio Kuduyari, Cerro Yaapoboda, 5–6 Oct. 1951, Schults & Cabrera
14378 (US). Rio Negro, San Felipe and vicinity (below confluence of Rio Guainia and Rio Casiquiare), 22 Oct. 1952,
Schultes et al. 18131 (US). Rio Kubiy’u (tributary of Rio Vaupes), Cerro Kanenda, savannahs about 15 miles upstream
from mouth. Quarzite base, 10 Nov. 1952, Schultes & Cabrera 18391 (US). Rio Kuduyari (tributary of Rio Vaupes),
Yapoboda, 18 November 1952, Schultes et al. 18542 (US). Rio Parana Pichuna (tributary of Rio Vaupes), Jun. 1953,
Schultes & Cabrera 19957 (US).
FRENCH GUIANA. Wilhelmina Gebergte, Lucie Rivier, 2 km of the river, 2 km west of Oost Rivier, 225 m, 11
Jul. 1963, Maguire et al. 54135 (US).
GUYANA. Cuyuni-Mazaruni, Imbaimadai, Savannah between Imbaimadai and Partang River crossing, 570 m,
14 Dec. 1989, Gillespie, L.J. & D.R. Smart 2667 (US). Potaro-Siparuni, Kaietuer Falls National park, near airstrip, 500
m, 2 Apr. 1988, Hahn, W., E. Judziewicz, S. Taiwari, & D. Gopaul. 4127 (US). Ttrail from abandoned balata bleeders’
camp at base of Mt. Makarapan to Rupununi river, 20 Apr. 1988, Maas, P.J.M., Koek-N, J., Lall, H., Ter Welle, B.J.H.
& Westra L.Y.Th. 7567 (INPA).
PERU. MADRE DE DIOS: Rio Heath, Santuario nacional de las Pampas del Heath, 15 Jun. 1992, Castilho &
Foster 6913 (F).
SURINAME. East Creek branch of Lisa creek, savana like, sandy soil, 26 Jun. 1998, Lohmann, L.G. & Peckham
H. 153 (MO). Tafelberg (Table Mountain), shallow bogs savannah 1, 3 Aug. 1944, Maguire, Basset 24197 (F); ibid, 5
km above falls, 525 m, 13 Sept. 1944, Basset Maguire 24727 (F). Lucie Rivier, 2 km north of Rivier, 225 m, 11 Jul.
1963, Maguire et al. 54135 (F).
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 37
TRINIDAD AND TOBAGO. 1977–80, Fendler 551 (NY). Long stretch road side, 26 Sept. 1951, Simmonds et al.
4 and 9 (K).
VENEZUELA. ESTADO BOLIVAR: Llanos between Cano Azul and rocky ridge 3 km. east cano Azul, 23 Oct.
1954, Wurdack & Guppy 167 (K). Vicinity of Salto Pacairao, bordering Rio Pacairao, about 3 km east to northeast off
Santa Teresita de Kavanayen, 24 Nov. 1944, Steyemark 60483 (F). by Ven. 10, 8 km N of San Rafael just S of San Juan
de Camoran, gravelly sandy seep in wet savanna, 29 Jul. 1983, Kral & Gonzales 70588 (MO). Piar, Gran Sabana. 25
km N de Kama-meru, Carretera El Dorado-Sta. Elena, 4 Abr. 1985, Holst et al. 2179 (MO). Kanavayen, estrada para
Liworiwo, Salto Aponwao, em campos rupestres ao lado esquerdo da estrada, já próximo de Liworiwo, 20 Jan. 1999,
Rivadavia & Hoogenstrijd 815 (SPF). Luepa, em campos rupestres brejosos cheios de Stegolepis sp., beirando o km
145 da estrada El Dorado - Sta. Elena, 21 Jan. 1999, Rivadavia & Hoogenstrijd 833 (SPF).
2. Utricularia biceps Gonella & Baleeiro (2018: 215)—Fig. 4J
Type:—BRAZIL. Minas Gerais, Botumirim, Rio do Peixe, 10 February 2011, Gonella et al. 376 (holotype: SPF!).
The species is endemic to the northern Espinhaço Range in Minas Gerais, Brazil. For the complete treatment of the species, see Gonella
& Baleeiro (2018).
3. Utricularia bicolor Saint-Hilaire & Girard (1838: 870)—Fig. 3K
Lectotype (designated here):—BRAZIL: Minas Gerais: Serra da Candonga, near Tapanhuacanga (Itapanhoacanga, c. -18.75, -43.41667)
1816–21, Saint Hilaire B1-919 (P603309!; isolectotype P603308 [mixed collection with U. damazioi] image!).
Description:—Terrestrial. Medium to large, perennial herb. Rhizoids numerous, from the base up to 10 mm above of
peduncle base, robust with thin ramified branchlets; stolons few. Leaves 1–2, circular lamina from a long attenuate
5 mm long; traps on rhizoids, c. 1 mm long, simple trichomes around trap door, without chin-like swelling, stalk up
to 1.5 mm long. Inflorescence simple, loosely, 170–360 mm long, peduncle usually papillose near base, 0.5–1.0 mm
long; scales deltoid, 0.7–1.2 mm long; bracts and bracteoles subequal, connate near the base or at middle, glabrous
or papillose, bract broadly ovate, apex acute, bracteoles linear, single nerved, obtuse apex, 1.2–2.0 mm long. Flowers
1–2(–3); pedicel erect, filiform 2.5–4.2 mm long; calyx lobes unequal, cartaceous, concave, papillae and glandular
trichomes present, with inconspicuous simple nerves; upper lobe broadly ovate to almost circular, apex rounded,
entire, slightly hyaline margin, 2.5–3.0 × 2.1–3.5 mm; lower lobe broadly elliptic, entire, margin not hyaline, apex
emarginate, 1.1–1.8 × 2.1–2.5 mm; corolla lilac with a yellow spot at base of lower lip, 12–15 mm long; upper lip
papillose, some glands present, broadly elliptic to circular, rounded apex 2.9–3.9 × 4.6–7.1 mm, twice as long than
calyx; lower lip transversally elliptic, base forming bilobed swelling, apex shallowly 3-lobed, 5.8–7.9 × 7.3–11.0
mm; spur cylindrical from a larger base, apex obtuse, slightly longer than the lower lip, 6.5–8.7 mm long, sparsely
glandular; filaments curved, 1.5 mm long, thecae 1mm long.; orary globose. Capsule globose c. 3 mm long., calyx not
covering the stigma lip. Seeds obliquely ovoid c. 0.10 mm long.
Distribution and ecology:—Utricularia bicolor is endemic to the Espinhaço Range in Minas Gerais, Southeast
Brazil (Fig.1). It is found growing on rocky outcrops or on edges of creeks in campos rupestres. Its patchy and lower in
number nature makes it hard to identify populations’ boundary, a pattern seen through out the genus in different levels
(see comments on U. hirtella). (P.C.Baleeiro, pers. observation).
Etymology:—From the latin bicolor = two colours, refering to the colour of the corolla; lilac with a yellow spot
at the base of the lower lip.
Conservation status:—Endangered—EN B1ab(iii) + B2ab(iii). Utricularia bicolor present a restricted range
(AOO = 24 km2; EOO = 4,325 km2) with all known populations located outside of conservation areas and subject to
threats such as increased fire frequency and intensity, invasive species, and habitat destruction.
Notes:—Initially, PCB treated this plant from the Diamantina area (Minas Gerais) as a new taxon based on an
initial comparison to one of Saint-Hilaire’s type specimens, and it was Included in the morphometric study of Baleeiro
et al. (2016) with the code “DIA” (Fig.4K). However, after examining the other type for Utricularia bicolor at a
different moment, it became clear that these plants from Diamantina corresponded to this taxon. The confusion is due
to a mixture of both U. bicolor and U. damazioi (both species grow sympatrically) in the first type, and it seems that
Saint-Hilaire & Girard (1838) based their description on both species as they describe the upper corolla lip as ovate,
which is found in U. damazioi but not on U. bicolor. Therefore, here we designate as lectotype the specimen without
the mixture. Utricularia bicolor is distinguished from all other species by the presence of slightly 3-lobed lower corolla
lip lacking a white spot, and papillose calyx lobes containing some marginally placed glandular trichomes.
BALEEIRO ET AL.
38 • Phytotaxa 576 (1) © 2022 Magnolia Press
FIGURE 3. A. U. amethystina s.str. Type locality Marau peninsula Bahia, B. U. amethystina s.str. Virua National Park, C. U. amethystina
s.str. white-flowered specimen, D. U. roraimensis, E. U. amethystina s.str. pink morphotype, F. U. amethystina Type locality, G. U.
lindmanii, H–I. U. lunaris, J–N. U. damazioi. J. from Chapada Diamantina, K., M., N. from Chapada dos Veadeiros, L. from Diamantina
Plateau.
Additional specimens Examined:—BRAZIL. MINAS GERAIS: Botumirim, alto da Serra da Canastra, 13 Out.
2001, Rivadavia 1267 (SPF); ibid, extremidade norte, 19 Set. 1992, Mello-Silva et al. 679 (SPF). Diamantina, grassy
meadow and adjacent campo, 15 Jan. 1969, Irwin et al. 22001 (NY); ibid, ponto atrás do Campus 2-UFVJM, beira
de córrego próximo ao ponto de captação de água da Universidade, 9 Jun. 2012, Baleeiro et al. 181 (SPF); ibid, 06
Mar. 2013; Baleeiro et al. 291 (SPF); ibid, voltando da comunidade de Galheiros, lado direito da estrada, campo
com córrego em meio a afloramento rochoso, 06 Mar. 2013, Baleeiro et al. 294 (SPF); ibid, São João da Chapada,
brown sand and campo on steep gravelly slopes, ascending through dense creekside vegetation to c. 75 cm, 26 Mar.
1970, Irwin et al. 28411 (F/NY). Itacambira, alguns km de Itacambira pela estrada para Montes Claros, 13 out. 2001,
Rivadavia 1284 (SPF). Santana do Pirapama, Serra do Cipó (Serra da Lapa), Distrito de São José da Cachoeira, trilha
da Senhorinha, 19 Fev. 2007, Souza et al. 32788 (SPF).
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 39
4. Utricularia chapadensis Baleeiro, Gonella, R.W.Jobson, sp. nov.—Fig. 4L–M
Type:—BRAZIL: Mato Grosso. Municipality of Chapada dos Guimarães, outside the limit of Chapada dos Guimaraes National Park,
beside the track to Cave Aroe Jari, soaked rocky soil. -15.611278, -55.49475 766 m alt, 28 April 2013, Baleeiro & Cardoso 320
(Holotype: SPF!; Isotype: NSW!, UFMT!).
Diagnosis:—U. chapadensis is morphologically distinct for its bright yellow corolla with rounded gibba and long curved spur at least 2x
longer than the lower lip.
Illustrations:—fig. 2K (Baleeiro et al. 2016).
Description:—Terrestrial. Small to medium sized annual herb. Rhizoids not observed. Stolons leaf like up to 20 mm
long. Leaves dimorphic, at peduncle base, spathulate, 2–4(–6) mm long, multinerved, some nerves converging the
base, or linear 1–3, single nerve up to 10 mm long.; traps dimorphic, both types globose, larger up to 2 mm long,
smaller c. 0.5 mm long. Inflorescence simple, lax, (5–)70–100(–250) mm long, peduncle glabrous; scales chartaceous,
rhombic, deltoid or ovate-elliptic, entire rounded apex 0.35–0.4(–0.6) mm long; bract and bracteoles connate above
the middle, up to 1 mm long. Flowers not racemose, 1(–3); calyx lobes unequal, chartaceous, glabrous, inconspicuous
nerves simple, upper lobe broadly ovate, culpulate, apex rounded, margin not hyaline, 1.9–3.0 x 1.6–2.4 mm wide;
lower lobe shorter, broadly elliptic, not hyaline, apex emarginate, 1.1–2.2 × 1.2–2.3 mm; corolla bright yellow, 14–25
mm long, upper lip with sparse glandular trichomes on adaxial surface, ovate or ovate deltoid, apex rounded, 2x> than
calyx lobes, 2.9–4.9 × 2.3–4.8 mm, lower lip transversally elliptic, base gibbous, apex 3-lobed, 4.4–7.0 × 4.7–8.4 mm,
spur conical usually curved backwards, 8.5–12.7 mm long 2–3x longer than the lower lip; filaments curved, 0.8–1 mm
long, theca similar. Ovary globose. Capsule globose 3 mm long; stigma lip not covered by calyx lobe. Seeds obliquely
ovoid c. 0.35 mm long.
Distribution and ecology:—This species occurs in the southwest of Goias and southeast Mato Grosso, Central–
West Brazil. Flowering from March to June, fruiting from May to June.
Etymology:—The specific epithet refers to the type locality, Chapada dos Guimarães National Park.
Conservation status:—Endangered—EN B2ab(i, ii, iii). Utricularia chapadensis presents a relatively broad range
(EOO = 78,500 km2), but is known from few locations (AOO = 28 km2) in populations that are severely fragmented in
a region that suffers from intense habitat destruction, intensification of anthropogenic fires, eutrophication of wetlands,
and invasive species. Therefore, we recommend that the species be assessed as Endangered, following IUCN (2012)
categories and criteria. While U. chapadensis.
Notes:—While Utricularia chapadensis (Fig. 4L,M) most closely resembles U. pantaneira, due to the yellow
corolla, the molecular phylogeny of Baleeiro et al. (2019) places it as sister to U. lindmanii from Tocantins.
U. pantaneira differs from U. chapadensis by having a round gibba, short lower lobes and a spur at least 2x longer
than the flower lip. The habitat is also distinct from that U. chapadensis, found in in small patches on rocky outcrops of
sandy-soaked soils. Besides that, U. chapadensis can be easily confused with members of U. sect. Setiscapella as seen
in Coelho et al. (2017) identified as U. pusilla Vahl (1804: 202). The easiest way to distinguish both sections is by the
basifix bract with two bracteoles connected in the base in U. sect. Foliosa, opposed to the peltate and lacks bracteoles
in U. sect. Setiscapella.
Additional specimens Examined:—BRAZIL. GOIÁS: Jataí, entrando a direita no km 268 da BR158 para
Estância, seguindo 3km por estrada de terra até morro com encosta brejosa a direita, 2 Mai. 1999, Rivadavia & Sato
1025 (SPF). Portelândia, Alto da Pedra Aparada, alguns km ao norte da cidade pela estrada para Ponte Branca, 5 Mai.
1999, Rivadavia & Sato 1078 (SPF); ibid, 25 Jun. 2013, Baleeiro & Carneiro 350 (SPF). MATO GROSSO: Chapada
dos Guimarães; córrego próximo a caverna Aroe Jari (estrada velha), solo encharcado, 5˚36’33.8’’, 55˚29’03.7’’, 28
Abr. 2013, Baleeiro & Cardoso 317 (SPF). Chapada dos Guimarães, ao lado da trilha para a caverna Aroe Jari (trilha
nova), limite com o Parque Nacional da Chapada dos Guimarães, 28 Abr. 2013, Baleeiro & Cardoso 320 (SPF).
Jaciara, Fazenda Usina, 21 Mai. 2009, Ramos 3 (UFMT).
5. Utricularia damazioi Beauverd (1907: 703)—Fig. 3J–N
Type:—BRAZIL: Minas Gerais: Itaculumi, c. 70 km SE of Belo Horizonte, 2 January 1905, Damazio s.n. (holotype: G image!).
=Utricularia dawsonii Steyermark (1958: 20)
Type:—BRAZIL: Goiás: Chapada dos Veadeiros, 1 May 1956, Dawson 14770 (Holotype: R!; Isotype: F!, LAM).
Illustrations:—(fig.7 p. 703, Beauverd (1907); figs. 77a3, 77 b2, Taylor (1989); Baleeiro et al. (2016), figs. 2 D,E.
Description:—Terrestrial. Small to medium sized, annual herb. Rhizoids 2–4 or absent, simple and ramified, 8–14
mm long. Stolons numerous (3–12) up to 6 mm long. Leaves usually absent or up to 3 from peduncle base; lamina
BALEEIRO ET AL.
40 • Phytotaxa 576 (1) © 2022 Magnolia Press
spathulate to almost circular, multinerved, 2.70–4.85 × 0.8–2.5 mm; traps numerous on stolons and rhizoids 1.2 mm
long, stalk c. 0.9 mm long. Inflorescence simple or ramified, lax, 80–280 mm long., peduncle glabrous or with few
sessile trichomes near base, 0.26–0.70 mm thick. Scales basifix, glabrous, ovate, margin entire c. 1 mm long. Bract
and bracteoles unequal, basifix, connate slightly above the middle, glabrous, without nerves; bract broadly ovate, apex
acute 1 mm long; bracteoles linear, slightly shorter than bracts. Flowers usually 1(–2); pedicel erect, filiform (4.5–
)7.0–15.0(–18.0) mm long; calyx lobes unequal, cartaceous, concave, glabrous, with inconspicuous simple nerves,
upper lobe cupuliform, broadly ovate, apex rounded, margin hyaline, 1.4–2.8 × 1.28–2.40 mm; lower lobe broadly
elliptic, margin not hyaline, apex rounded or truncate-emarginate, 1.2–2.2 × 1.2–2.2 mm; corolla lilac with yellow
spot bordered by white at base of lower lip; upper lip with sparse minute glandular trichomes, oblong, ovate or elliptic,
rounded apex, x2.5 larger than the upper calyx lobe, 2.5–4.8 x 2.0–4.7 mm; lower lip broadly obovate, base forming
bilobed swelling, apex deeply 3-lobed, lateral lobes larger, oblong with apex rounded or truncate, 4.7–8.6 x 5.7–14
mm; spur cylindrical, ligulate, from a larger base, apex rounded, 6.1–11.6 mm long; filaments slightly curved, anther
thecae ±equal length as filaments; ovary globose, style very short. Capsule globose c. 2mm long. Seeds obliquely
ovoid, c. 0.30 mm long.
Distribution and ecology:—Endemic to Campos rupestres in the Cerrado domain restricted to the states of Bahia,
Goiás and Minas Gerais (Fig. 1A, B). Sandy substrate in grassland, and often sympatric with U. hirtella. Flowers from
December to July.
Etymology:—The epithet honours Brazilian botanist Leonidas Bothelo Damazio (1854–1922), who was born in
Salvador and taught physics, chemistry, zoology, and botany at Ouro Preto University, Brazil.
Conservation status:—Least Concern. The species is widespread in the Cerrado domain (EOO = 464.906 km2;
AOO = 188 km2), and abundant in protected areas such as the National Parks of Veadeiros, Sempre Vivas, Serra da
Canastra, and Serra do Cipó.
Notes:—Utricularia damazioi (Fig.3J–N) has been seen as the “real” U. amethystina, since Taylor’s (1989)
revision, probably for being the most common and collected species, especially in Central Brazil. The revision showed
that U. damazioi matched the populations in the Brazilian Cerrado and U. amethystina s.str. from the Brazilian Northeast
coast and savannas from north of the Amazon and the Guiana Shield. Later on, morphometric and phylogenetic results
confirmed the initial hypothesis (Baleeiro et al. 2016, 2019). The terrestrial U. damazioi differs from the semi-aquatic
U. amethystina by the presence of a large, deeply 3-lobed lower corolla lip vs. slightly 3-lobed, an upper lip x2–2.5
larger than upper calyx lobe vs. x1 larger, and cylindrical-ligulate vs. conical spur. Great variation is observed between
the populations of U. damazioi, with the specimens from the Espinhaço Range having a more round lower lip lobe
unlike the obovate to oblong lobes from specimens collected at Chapada dos Veadeiros (Souza and Bove, 2012). For
this reason, further studies involving high throughput technologies of genome skimming with specimens across the
range would uncover further variation.
Additional specimens examined:—BRAZIL. DISTRITO FEDERAL: Brasília, Jardim Botânico, campo sujo
com murundus, 10 Abr. 2010, Dias & Pereira 719 (CEN); ibid, 26 Mar. 2012; Baleeiro et al. 169 (SPF). GOIÁS: Alto
Paraiso de Goiás, Fazenda São Bento, 23 Fev. 1991, D. Alvarenga et al. 764 (IBGE). Parque Nacional dos Veadeiros,
em campos rupestres da trilha para a cachoeira do córrego Santana, 1 Jan. 1993, Rivadavia 185 (SPF); ibid, entrando
+- no km 190 da estrada Brasília-Alto Paraíso, 14 Abr. 1995, Rivadavia & Ogassavara 380 (SPF); ibid, Povoado São
Jorge, campos rupestres, 10 Abr. 1995, Rivadavia et al. 364 (SPF); ibid, 11 Abr. 1995; Rivadavia et al. 369 (SPF; ibid,
em campos rupestres no lado norte do km 196 da BR 118 (estrada A. Paraíso-Teresina), 23 Jun. 2007, Rivadavia &
Batista 2620 (SPF); ibid, campo úmido inclinado próximo ao córrego dos Ingleses, 2 Junho 2009, Baleeiro et al. 11 e
18 (R); ibid, próximo a Sete Lagoas, 3 Jun. 2009, Baleeiro et al. 24 (R); ibid, caminho pelo morro calda da Baleia em
direção a teta de moça, 20 Mar. 2012, Baleeiro et al. 118 (SPF); ibid, campo rupestre do lado esquerdo da estrada São
Jorge—Alto Paraíso de Goiás, 22 Mar. 2012, Baleeiro et al. 133 (SPF); ibid, estrada Alto Paraiso-Teresina de Goiás, 23
Mar. 2012, Baleeiro et al. 137 (SPF); ibid, 23 Mar. 2012, Baleeiro et al. 138 (SPF); ibid, 27 km de Cavalcante, 23 Mar.
2012, Baleeiro et al. 139 (SPF); ibid, campo rupestre logo depois da torre que fica depois do Cruzeiro, 24 Mar. 2012,
Baleeiro et al. 157 (SPF); ibid, 24 Mar. 2012, Baleeiro et al.158 (SPF); ibid, 24 Mar. 2012, Baleeiro et al. 159 (SPF).
Corumbá de Goiás, nascente sobre morro entre a cach. do Corumbá e Cocalzinho de Goiás, 1 Jul. 2007, Rivadavia
& Batista 2672 (SPF). Cristalina, a 5km da cidade pela BR050 para Brasília, entrando a 2km por pequena estrada de
terra, em campos rupestres a direita, 30 Abr. 1999, Rivadavia & Sato 1089 (SPF). Pirenópolis, nascente brejosa com
buritis próximo aos três Picos à direita da estrada vinda de Pirenópolis, 26 Abr. 1999, Rivadavia & Sato 930 (SPF).
Teresina de Goiás, ponto a 18.3 km de Teresina em direção a Alto Paraíso, 23 Mar. 2012, Baleeiro et al. 144 (SPF).
MINAS GERAIS: Conceição do Mato Dentro, Serra do Cipó, Campo Redondo, caminho para nascentes do ribeirão
do campo, acessível por estrada de terra à direita da MG-10, a 7,5km da bifurcação do Mato Dentro, 17 Mai. 2008,
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 41
Gonella et al. 111 (SPF). Congonhas do Norte, Serra as Mangabeira, no alto da segunda crista de serra a partir do ponto
onde a estrada termina num riacho e se torna trilha, após última casa de fazenda, 26 Fev. 1997, Rivadavia & Pinheiro
561 (SPF). Serra da Carapina (Serra Talhada), norte da Serra do Cipó, 2 Mar. 1998, Rapini et al. 519 (SPF). Couto
de Magalhaes, campos rupestres c. 40km leste de Diamantina pela estrada que passa pela Gruta do Salitre e povoado
de Extração, 12 Mai. 2007, Rivadavia 2531 (SPF). Diamantina, campos rupestres ao sul da Cidade, fina camada de
areia e matéria orgânica em nascentes sobre rochas sedimentares, 24 Jul. 2008, Gonella et al. 168 (SPF); ibid, estrada
Diamantina-Conselheiro Mata, Km 185, inselberg a direita da estrada, 28 Jan. 1986, Menezes et al. CFCR 9351 (SPF);
ibid, km 606 BR 259, estrada para Gouveia, 28 Fev. 1998, Forzza et al. 652 (SPF); ibid, em campos rupestres de serra
ao leste da cidade, 23 Fev. 1992, Rivadavia et al. 73 (SPF); ibid, cachoeira Sentinela no caminho para Biribiri, 12 Jul.
1999, Rivadavia et al. 1223 (SPF); ibid, 12 km a partir do trevo saindo de Diamantina em direção a Universidade,
lado esquerdo da BR, Parque Nacional Biribiri, 10 Jun. 2012, Baleeiro et al. 190 e 193 (SPF); ibid, estrada de terra
sentido comunidade de Garelhos-Diamantina, lado direito da estrada, campo com córrego em meio a afloramento
rochoso, 6 Mar. 2013, Baleeiro et al. 295 (SPF); ibid, BR 367 entre Gouveia e Diamantina, 9 Mar. 2013, Baleeiro et
al. 309 (SPF); ibid, em frente ao entroncamento para Conselheiro Mata, 9 Mar. 2013, Baleeiro et al. 313 (SPF); ibid,
Parque Estadual do Biribiri, cachoeira Sentinela, lado direito sentido contrário das águas, na beira da trilha, 9 Mar.
2013, Baleeiro et al. 314 (SPF). Grão Mogol, Alto do morro Papo de Ema, campo rupestre com afloramentos de rocha
quartzitica, 15 Jun. 1990, Pirani et al. CFCR 13048 (SPF); ibid, margem de córrego na saída da cidade, na estrada para
o rio Ventania, 25 Fev. 1986, Semir et al. CFCR 9651 (SPF); ibid, trilha da Tropa, 2 Jun. 1994, Rivadavia 272 (SPF);
ibid, margem de córrego na saída da cidade, na estrada para o rio Ventania, 25 Feb. 1986, Mello-Silva et al. 9607 (SPF);
ibid, campo rupestre, em direção nordeste da Cidade, 22 May 1982, Giulietti et al. 3429 (SPF); ibid, subida da trilha
da Tropa, à esquerda do riacho ribeirão, 27 Jun. 1988, Zappi et al. CFCR12016 (SPF). Itacambira, estrada para Montes
Claros, 9 Jan. 1986, Pirani et al. CFCR 9160 (SPF). Jaboticatubas, campos rupestres dentro da fazenda da Serra do
Cipó, 26 Fev. 1992, Rivadavia 86 (SPF). Moeda, Serra da Moeda, em campos rupestres ao lado esquerdo da estrada
para Moeda, a 1,8km da BR-040, 12 Mar. 2002, Rivadavia 1343 (SPF); ibid, Serra da Moeda, em campos rupestres ao
lado esquerdo da estrada para Moeda, a 1,8km da BR-040, 26 Abr. 2002, Rivadavia 1351 (SPF). Nova Lima, Serra da
Mutuca, near Vargem de Ouro Podre, 11 Mar. 1945, Williams & Assis 6208 (US). Rio Vermelho, Serra da Torre, 11 Jul.
1999, Rivadavia et al. 1109 (SPF). Santa Barbara, Parque Natural do Caraça, ao lado da trilha para o Campo de Fora,
3 Mar. 1992, Rivadavia 113 (SPF); ibid, Parque Natural do Caraça, ao lado da trilha para a gruta do Padre Caio, 2 Mar.
1992, Rivadavia 109 (SPF). Santana do Pirapama, faz. Inhame (Serra Mineira) Serra do cipó, 22 Mar. 1982, Pirani
et al. CFSC 8091 (SPF). Santana do Riacho, Parque Nacional Serra do Cipó, lado direito da BR, pouco antes da casa
do parque, 8 Jun. 2012, Baleeiro et al. 179 (SPF); ibid, Trilha do Travessão, córrego da cachoeira próximo ao final da
trilha, 4 Jul. 2012, Baleeiro et al. 201 (SPF); ibid, trilha para cachoeira Capivara, 5 Jul. 2012, Baleeiro et al. 203 (SPF);
ibid, ao longo da rodovia Belo Horizonte - Conceição do Mato dentro, 22 Apr. 1982, Amaral et al. CFSC 8.400 (SPF);
ibid, Serra do Cipó (Serra da Lapa), Distrito de São Jose da Cachoeira, trilha da senhorinha, campo Rupestre, 19 Fev.
2007, Souza et al. 32788 (SPF); ibid, Serra do Cipó, nascentes descendo de afloramentos rochosos perto de estátua do
Juquinha, do outro lado da estrada, 15 Mai. 2008, Gonella et al. 89 (SPF); ibid, Serra do Cipó, trilha para as velózias
gigantes, em nascentes nas margens da trilha, 15 Mai. 2008, Gonella et al. 77 (SPF); ibid, Serra do Cipó, na beira de
riacho quase seco, do lado direito do km 108 da MG-10, sentido Conceição do Mato Dentro, 16 Mai. 2008, Gonella
et al. 105 (SPF); ibid, Serra do Cipó, km 125 da rodovia BH-Conceição do Mato Dentro, elevação frente ao Juquinha,
26 Apr. 1991, Pirani et al. CFSC12252 (SPF). Santo Antônio do Itambé, descida do Morro do Pico do Itambé, 6 Abr.
1982, Rossi et al. CFCR 3094 (SPF). São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, trilha para o alto da
chapada, em campo úmido cortando pela trilha, solo arenoso úmido, 5 Feb. 2009, Gonella et al. 191 (SPF). São Roque
de Minas, Parque Nacional Canastra, caminho do Reito de Pedra, 19 Mar. 1998, Sano et al. 954 (SPF); ibid, Parque
Nacional Canastra, campos rupestres beirando a estrada alguns km após a nascente do rio São Francisco, 3 Apr. 1999,
Rivadavia & Peixoto 893 (SPF). São Tome das Letras, em campos rupestres de serra ao norte da cidade, 20 Apr. 1992,
Rivadavia 139 (SPF). Serro, campos rupestres à esquerda da estrada de Milho Verde para Diamantina, km 22.5, 6 Abr.
2003, Rivadavia 1575 (SPF).
BALEEIRO ET AL.
42 • Phytotaxa 576 (1) © 2022 Magnolia Press
6. Utricularia hirtella Saint-Hilaire & Girard (1838: 869)—Fig. 4A–D
Type:—BRAZIL: Minas Gerais: in terra humida propè urbem S. Joao del Rei; Saint Hilaire B2-2392 (Holotype: P image! (photo, M)).
=Utricularia genliseoides Benjamin (1847:250)
Type:—BRAZIL: Minas Gerais, Tejuco; Martius s.n. (Holotype: M image!).
=Utricularia spatulata Splitg. ex de Vriese (1848: 324)
Type:—SURINAM: Pará; Splitgerber 674 (Holotype: L image!).
=Utricularia adenantha Standley (1936: 366)
Type:—BELIZE: All Pines, in swampy places, at sea level, growing in sandy soil; 2 Sept, W.A. Schipp S-89 (Holotype: F!; Isotype: BM!,
BRH, G image!, K!, MICH, MO!, NY!, S image!, Z).
=Utricularia kaieteurensis Steyermark (1948: 658)
Type:—GUYANA: Kaieteur Savanna, Maguire & Fanshawe 23204 (Holotype: F!; Isotype: K!, NY!, VEN).
=Utricularia bolivarana Steyermark (1953: 538)
Type:—VENEZUELA: Bolivar: Gran Sabana, Steyermark 59191 (Holotype: F!; Isotype: VEN).
Illustrations:—Baleeiro et al. (2016), fig 2L,N.
Description:—Terrestrial. Small to medium sized annual herb. Rhizoids absent or 2–4, simple ramification, 3–7 mm
long. Stolons few to numerous (10) up to 6 mm long. Leaves absent or up to 7, from peduncle base, lamina obovate,
reniform to almost circular, multinerved; 2 mm long; traps on stolons, globose, 0.5–2.0 mm long, stalked 0.5–3.0
mm long. Inflorescence simple or loosely branched, 30–150 mm long, peduncle glabrous 0.19–0.42 mm thick; scales
basifix, glabrous, ovate, margin entire 0.5–0.7 mm long, bract and bracteoles basifixed, connate at about the middle,
glabrous, without or 1-nerved, bract ovate apex acuminate obtuse, 0.6–1.0 mm long, bracteoles narrowly ovate, shorter
or the same level than the bracts. Flowers 1–3 pedicel erect, filiform, glabrous or glandular trichomes next to calyx,
1–9 mm long; calyx lobes unequal, chartaceous, concave, glandular trichomes present, with inconspicuous simple
nerves, upper lobe ovate or ovate-elliptic, apex rounded, margin hyaline, entire 1.1–2.1 mm long; lower lobe ovate-
oblong, margin not hyaline, apex crenate or emarginate, 0.8–1.6 mm long; corolla white, cream, or light pink, with
yellow spot at base of lower lip, surfaces hirsute (densely covered by trichomes); upper lip oblong towards the base,
ovate towards the rounded apex; lower lip transversally elliptic, base slightly gibbous, apex slightly 3-lobed, lateral
and middle lobes rounded; spur conical, apex obtuse, x2 > the length of lower lip, trichomes cover much surface
2.2–5.1 mm long; filaments slightly curved, anther thecae same width as filaments; ovary globose, surface covered
with glandular or simple trichomes. Capsule globose 1.0–1.5 mm long., not embracing the style lips. Seeds obliquely
ovoid to narrowly cylindrical, 0.2–0.5 × 0.15 mm.
Distribution and ecology:—This group is found in Belize, Brazil, British Guyana, French Guyana, Honduras,
Guatemala, Venezuela and Surinam. In Brazil, in the states of Bahia, Goiás, Mato Grosso, Minas Gerais, Roraima, and
Tocantins. Although its distribution ranges from Central to South America and across many biogeographical barriers
(Fig. 1A–C), it is mostly hard to measure population size, and the number of specimens is low, similar nature found in
U.trinervia. Specimens grow in sandy, wet, poor soils of mostly high in elevation rocky outcrops and grasslands in the
Brazilian and Guiana Shield Savannas and close to sea level savannas of the Amazon so-called campinaranas. Flowers
from March to September.
Etymology:—The species epithet refers to the indumentum of simple and glandular trichomes covering the
surface of the calyx and corolla.
Conservation status:—Least Concern. Utricularia hirtella s.l. (Fig.4A–D) as here defined, is considered to be
a species complex with a widespread distribution (EOO = 7,237,840 km2; AOO = 160 km2), found within several
protected areas. If this complex taxon is further delimited in the future, the additional taxa may require reevaluation of
threat status.
Notes:—Although an attempt was made to sample populations representing each of the currently accepted taxa
or synonyms of published names (Baleeiro et al. 2019), none of the accessions in the U. hirtella clade represents the
type locality so instead morphologically similar specimens were included (Fig.4A,B). The closest or possibly the type
location is presented in the study of da Silva et al., (2011) of Serra de Sao Jose and a photo is provided (Fig. 3A, Silva
et al. 2011). In addition, access to populations across many northern regions of South America is limited (i.e., the
Tepuis, Venezuela), and further collections are required to examine this species complex of species. In summary, U.
hirtella is differentiated from other related taxa by the presence of papillae and/or trichomes on its calyx, and a usually
white corolla. At the same time, Baleeiro et al. (2019) study showed U. hirtella is a multi-lineage independent from U.
amethystina s.str. it also revealed a morphological diversity that requires further investigation.
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 43
FIGURE 4. A–D. U. hirtella s.l., E–G. U. velascoensis, H. U. tridentata, I. U. tricolor, J. U. biceps, K. U. bicolor, L. U. chapadensis,
M–N. U. pantaneira, O. U. trinervia. Images C. Rivadavia, D. Rohrbacher, E–G. by Marcos Cardoso, J., K. Gonella.
Additional specimens examined:—BRAZIL. AMAZONAS: Manaus-Caracaraí, km 130, Igarapé Lages, 9 Mai.
1974, Prance et al. 21041 (NY). GOIÁS, Alto Paraiso de Goiás, campo gallery forest and adjacent wet campo (brejo),
c. 20km N. of Alto Paraiso, 19 Mar. 1971, Irwin et al. 32716 (NY); ibid, Parque Nacional Veadeiros, entrando +- no
km 190 da estrada Brasília-Alto Paraíso, 11 Abr. 1995, Rivadavia & Ogassavara 370 (SPF); ibid, 14 Abr. 1995,
Rivadavia & Ogassavara 379 (SPF); ibid, Parque Nacional Veadeiros, campo úmido inclinado próximo ao córrego dos
Ingleses, 2 Jun. 2009, Baleeiro et al. 13 (R); ibid, estrada, entre Alto Paraíso de Goiás e Teresina de Goiás (GO-118),
km 196, 5 Jun. 2009, Baleeiro et al. 43 (R); ibid, 20 Mar. 2012, caminho pelo morro Cauda da Baleia em direção à
Teta de Moça, Baleeiro et al. 119 (R); ibid, 11 km de Alto Paraíso de Goiás em direção a São Jorge, lado esquerdo da
pista, ambiente com gramíneas beirando córrego pequeno, 23 Mar. 2012, Baleeiro et al. 149 (SPF); ibid, 24 Mar 2012,
campo rupestre 27.8 km de Alto Paraíso em direção a Teresina, Baleeiro et al. 158 (SPF). Cavalcante, ponto a 20 km
de Cavalcante, estrada em direção ao povoado de Kalunga, 23 Mar. 2012, Baleeiro et al. 141 (SPF). Mineiros, encosta
brejosa de morro à direita do km11 da estrada para o Parque Nacional das Emas, 3 Mai. 1999, Rivadavia & Sato
1050 (SPF). Pirenópolis, nascente brejosa com buritis à direita da estrada para Três Picos, 22 Abr. 1999, Rivadavia
& Sato 919 (SPF). Posse, campo limpo úmido envolta de um buritizal, 30 Jul 2012, Baleeiro et al. 208 (SPF). Santo
Antônio do Descoberto, nascente brejosa com buritis à esquerda da BR-070 pouco antes da divisa com o DF, 23 Abr.
1999, Rivadavia & Sato 933 (SPF). MATO GROSSO: Barra do Garças, próximo a Cachoeira da Bateia, no complexo
BALEEIRO ET AL.
44 • Phytotaxa 576 (1) © 2022 Magnolia Press
do Roncador, aproximadamente 40km a leste da cidade, 9 Nov. 2007, Cardoso 105 (SPF). Chapada dos Guimarães,
Parque Nacional da Chapada dos Guimarães, campo antes da entrada da cidade de Chapada dos Guimarães à esquerda,
11 Jun. 2011, Baleeiro & Cardoso 99 (SPF). MINAS GERAIS: Buenópolis, estrada que sobe ao sul da cidade para
alto da Serra do Cabral em direção ao Brejo Grande, 3 Jul. 2003, Rivadavia & Menezes 1647 (SPF). Diamantina, em
campos rupestres próximos a cachoeira da Toca, 25 Fev. 1992, Rivadavia 78 (SPF); ibid, em campos rupestres ca.
4km ao sudeste da cidade pela estrada para a Gruta do Salitre, 13 Mai. 2007, Rivadavia 2548 (SPF); ibid, Biribiri,
córrego/cachoeira Sentinela, 9 Jun. 2012, Baleeiro et al. 184 (SPF); ibid, Parque Nacional Sempre Vivas, 8 Mar 2013,
Baleeiro et al. 304 (SPF); ibid, Parque Estadual do Biribiri, cachoeira Sentinela, lado esquerdo sentido contrário das
águas, na beira da trilha, 9 Mar. 2013, Baleeiro et al. 315 (SPF). Grão Mogol, Trilha da tropa, 2 Jun. 1994, Rivadavia
280 (SPF). Jaboticatubas, campos rupestres dentro da Fazenda da Serra do Cipó, 26 Fev. 1992, Rivadavia 95 (SPF).
São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, descendo o Ribeirão das Águas pela margem esquerda,
entre as cachoeiras Deitada e Sempre-Viva, 28 Jun. 2003, Rivadavia & Deco 1627 (SPF). Serro, Distrito de milho
verde, 5 Abr. 2003, Rivadavia & Neves 1551 (SPF); ibid, campos rupestres à esquerda da estrada de Milho Verde para
Diamantina, Km 22.5, 6 Abr. 2003, Rivadavia 1573 (SPF). SÃO PAULO: Botucatu, Distrito de Vitoriana, Porto Said,
estrada Marciano Zacarias, em nascente brejosa na encosta de morro descendo da estrada, 20 Mar. 1999, Rivadavia et
al. 853 (SPF). TOCANTINS: Mateiros, Cachoeira das Velhas, APA do Jalapão, 14 Jun. 2006, Rivadavia 2220 (SPF).
Ponte Alta, Cachoeira da fumaça, estrada entre Rio da Conceição e Ponte Alta, 17 Jul. 2006, Rivadavia 2257 (SPF).
RORAIMA: Tepequém Pré-Tepui, região da Cachoeira do Barata, área de campo aberto entre a casa do Norton e a
mata, 9 Set. 2012, Baleeiro 275 (SPF).
VENEZUELA. BOLIVAR: Gran Sabana, beira da estrada, lado esquerdo sentido Caracas, 11 Set. 2012, Baleeiro
282 (SPF).
7. Utricularia lindmanii Sylvén (1909: 26)—Fig. 3G
Type:—BRAZIL. Mato Grosso: Tangará da Serra, Serra do Itapirapuan, 29 Apr. 1894, Lindman A2935 (holotype: S image!).
Illustrations:—(Sylvén, 1909), fig. 1–3 taf.2.
Description:—Terrestrial. Small to medium sized annual herb. Rhizoids numerous, c. seven simple ramifications.
Stolons few. Leaves 1–7(–10) from peduncle base, filiform, up to 20 mm long, 1 mm wide, traps on rhizoids or stolons,
stalked basal entrance, 2 mm long. Inflorescence simple or loosely branched, 60–180 mm long, peduncle glabrous 0.7
mm thick, scales basifix, glabrous, deltoid or elliptic 0.6 mm long, bract and bracteoles unequal, connate at the middle,
glabrous, three nerves, bract broadly ovate, apex obtuse 1 mm long., bracteoles ovate 0.8 mm long. Flowers 1–3,
pedicel erect, filiform (–5)8–20 mm long, calyx lobes subequal, chartaceous, concave, glabrous, with inconspicuous
simple nerves, upper lobe ovate, apex rounded, entire margin hyaline 1.9–3.3 × 1.4–2.1 mm wide, lower lobe broadly
elliptic, entire margin not hyaline, apex rounded, 1.6–3.2 × 1.6–2.5 mm wide, corolla lilac with a yellow spot bordered
by white margin at base of lower lip, upper lip with few sessile glands, oblong or oblong-ovate, rounded apex 2–4.5
× 2.6–4.0 mm wide, lower lip transversally elliptic, base forming bilobed swelling, apex 3-lobed, 2.4–5.3 × 4.6–8.9
mm wide, spur conical from a larger base, apex obtuse, longer than the lower lip 3.4–6.7 mm long, filaments slightly
curved, anther thecae shorter than the filaments, ovary globose, style very short, stigma lower lip semicircular with
glandular trichomes, upper lip absent. Capsule globose, 2 mm long., calyx lobes covering stigma lips. Seeds obliquely
ovoid c. 0.30 mm long.
Distribution and ecology:—Found mostly in the east and southeast of the state of Tocantins, Brazil (Fig.1 A).
However, the type specimen was collected in the state of Mato Grosso at a location that has since been cleared of native
vegetation and converted to farmland. Grows in moist clay substrate within grassland with other species of Utricularia
and Genlisea. Flowers from June to August. Further fieldwork may uncover populations at under-explored locations
such as Jalapão, west Tocantins and Goiás, and Northeast of Mato Grosso.
Etymology:—The epithet honours Carl Axel Magnus Lindman (1856–1928), a Swedish naturalist who travelled
to Brazil from 1892–1894.
Conservation status:—Vulnerable—VU B1ab(iii) + B2ab(iii). Utricularia lindmanii presents restricted range
(EOO = 5,988 km2, AOO = 28 km2) and occurs in a region severely affected by habitat destruction and interference
with conversion of land use for agricultural purposes. The species is likely extinct at the type location and the new
records from Tocantins fall outside the Jalapão State Park.
Notes:—The holotype specimen is from a savannah habitat in central Mato Grosso. The attempt to sample from
the type population failed, as large portions of the natural land have been lost to agriculture, cattle stations and due
to the constant urban expansion in the past decades. Therefore, the populations from Tocantins are here denominated
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 45
U. lindmanii (Fig. 3G) and not a new taxa after examining the illustration introduced by the original publication
(protologue) for U. lindmanii and type material. Morphologically, U. lindmanii resembles U. amethystina s.str., but
distinguishes by linear vs. spathulate/rotund leaves, upper and lower corolla lobes about the same length vs. lower
corolla lobe larger than upper. Besides, molecular data (Baleeiro et al. 2019) shows no affinities with U. amethystina
s.str., and rather being in a clade sister to U. chapadensis.
Additional specimens examined:—BRAZIL. TOCANTINS: Ponte Alta, estrada sentido Rio da Conceição-Ponte
Alta, lado direito da estrada quase chegando em Ponte Alta, 5 Ago. 2012, Baleeiro et al. 222 (SPF). Rio da Conceição,
TO-476, estrada Dianópolis—Rio da Conceição, a 45 km do trevo, entrada para estrada de terra ca. de 6km, Cachoeira
Cavalo Queimado, 7 Jun. 2009, Baleeiro et al. 63 (R). Taguatinga, beira da rodovia, sentido Alto Paraíso de Goiás, 7
Ago. 2012, Baleeiro et al. 227 (SPF).
8. Utricularia lunaris Baleeiro, Gonella, R.W.Jobson, sp. nov.—Fig. 3H–I
Type:—BRAZIL. GOIÁS: Alto Paraíso de Goiás, Vale da Lua, external border of Veadeiros National Park on the way to São Jorge village.
905m, 21 March 2012, -14.185944, -47.788917 Baleeiro et al. 127 (Holotype: SPF, Isotype: NSW, UFG).
Diagnosis:—U. lunaris presents pale pink to almost white corolla with pale purple and a yellow spot on the lobed swelling lower lip base;
it distinguishes itself from others for its circular, or reniform lamina leaves.
Illustrations:—Baleeiro et al. (2016), fig.2H; Baleeiro et al. (2019).
Description:—Terrestrial. Small perennial herb. Rhizoids 2–5, simple branches, up to 8 mm long. Stolons 3–7, up
to 15 mm long. Leaves 1–5 at peduncle base or stolon, lamina circular or reniform, attenuate towards what should
be a petiole, 3 × 3.5 mm, traps sub sessile on stolons, ovoid, basal entrance, with two short dorsal appendages and
reduced number of trichomes on lower surface of appendages, 1 mm long. Inflorescence simple (50.5–)70–150 mm
long, peduncle glabrous, scales membranous, ovate, apex obtuse or acuminate 0.4–0.8 mm long, bract and bracteoles
connate at base, membranous, bract ovate, apex obtuse or acuminate, bracteoles narrowly oblong-ovate, 0.45 mm long.
Flower 1, pedicel glabrous 2.4–13 mm long, calyx lobes sub-similar, membranous, concave, glabrous, some papilla or
simple trichomes near upper margin, nerves inconspicuous, simple, upper lobe broadly elliptic, apex rounded, margin
not hyaline 1.3–2.1 × 0.9–1.5 mm, lower lobe broadly elliptic, margin entire, not hyaline, emarginate, 0.5–1.7 ×
0.8–1.5 mm, corolla pale pink to almost white with a yellow spot and pale purple and white on a lobed swelling lower
lip base, upper lip papillose with sparse glandular trichomes, ovate, apex round, up to x3> than calyx lobes, 1.6–2.9 ×
1.4–2.6 mm, lower lip transversally elliptic, base lobed swelling, apex 3-lobed, 3.1–5.0 × 3.5–7.5 mm, spur ligulate,
stipitate papillose 3.5–7.5 mm long. Filaments slightly curved, forming a protuberance in the top, 0.8 mm long, thecae
similar, 0.6 mm long. Ovary and Capsule not observed.
Distribution and ecology:—Endemic to a small area on a private land called Vale da Lua, c. 4 km from the border
of Chapada dos Veadeiros National Park, Goiás, Brazil. It grows on sandy substrate in rocky habitats (Fig. 3I) on edges
of creeks in an intriguing `moon like` formation within gallery forest vegetation, surrounded by grasslands. Flowers
during the whole year.
Etymology:—The specific epithet refers to the type locality Vale da Lua, Chapada dos Veadeiros, Brazil.
Conservation status:—Critically Endangered—CR B2ab(iii). Utricularia lunaris is known from a single location
(AOO = 4km2) in a private area with waterfalls and natural pools that attract thousands of visitors yearly. The species
is vulnerable to trampling, invasive species and pollution, potentially reducing the quality of the habitat in the short
term.
Notes:—Results from both morphometric and molecular phylogenic studies (Baleeiro et al. 2016, 2019) support
Utricularia lunaris as a new species, sister to Utricularia damazioi. Utricularia lunaris has a much smaller pale pink
corolla, and its leaves are circular or reniform with a long pedicel reminding that of U. tricolor.
Additional specimens examined:—BRAZIL.GOIÁS: Alto Paraiso de Goiás, Vale da Lua, 4 Fev. 1993, Rivadavia
200 (SPF); ibid, 11 Abr. 1995, Rivadavia & Ogassavara 367 (SPF); ibid, 21 Mar. 2012, Baleeiro et al. 127 (SPF).
9. Utricularia pantaneira Baleeiro, Gonella, R.W.Jobson, sp. nov.—Fig. 4N
Type:—BRAZIL. Mato Grosso: Santo Antônio do Leverger, 15˚36’40.6’’S, 55˚29’41.1’’W, 766m alt., 6 May 2013, Baleeiro & Cardoso
325 (Holotype: SPF, Isotype: NSW, UFMT).
Diagnosis:—U. pantaneira is morphologically close to U. chapadensis due to corolla colour and size, but it is distinguished by its broader
lower lip corolla lobes and spur cylindrical from a larger base. In addition, molecular data support this new species as sister to U.
lindmanii, which does not share morphological similarities with U. pantaneira.
BALEEIRO ET AL.
46 • Phytotaxa 576 (1) © 2022 Magnolia Press
Description:—Terrestrial or affixed sub-aquatic. Small to medium sized perennial herb, rhizoids absent. Stolons leaf
like. Leaves dimorphic, at peduncle base spathulate to almost circular near the apex 2–6, forming a rosette up to 12 mm
long, parallel nerves from the apex to petiole base, or linear 1–3, single nerve, up to 25 mm long. Traps on the linear
leaves, dimorphic, type 1 globoid, sub-sessile, appendages deltoid, shorter trichomes under the appendages and around
the trap door, up to 2.5 mm long., type 2 ovoid and much smaller 0.33 mm long, entrance lateral, chin-like welling and
reduced appendages, stalk the same of trap length, capillary. Inflorescence simple, loosely, (130–)160–280 mm long,
peduncle glabrous 0.5–1.0 mm thick, scales chartaceous, ovate or ovate-elliptic, apex of upper-most scales 3-lobed 0.7
mm long, lower scales with entire rounded apex 0.4–0.6 mm long, bract and bracteoles connate above the middle, up
to 1 mm long. Flowers 1–6, calyx lobes subsimilar, chartaceous, concave (culpulate), glabrous, inconspicuous simple
nerves, upper lobe broadly ovate, apex rounded, margin not hyaline, 1.9–3.0 × 6.0–2.5 mm, lower lobe broadly ovate,
not hyaline, apex rounded, 1.2–2.2 × 1.6–2.4 mm, corolla yellow, 14–22 mm long, upper lip with sparse glandular
trichomes, ovate or broadly ovate, apex rounded, up to x2.5> than calyx lobes, 2.6–4.6 × 2.1–4.4 mm, lower lip
transversally elliptic, base gibbous, apex slightly 3-lobed, 4.8–7.0 × 6.6–10.8 mm, spur cylindrical from a larger base,
minutely papillose 8.0–10.5 mm long, filaments curved, 0.9–1.2 mm long, thecae distinct 0.75–0.80 mm long, ovary
globose, glabrous, 0.5 mm long, style short, sparse trichomes, stigma lower lip semicircular, border ciliate, upper lip
inconspicuous. Capsule globose 3 mm long, stigma covered by calyx at maturity. Seeds obliquely ovoid c. 0.35 mm
long.
Distribution and ecology:—Endemic to flooded regions of Mato Grosso and Mato Grosso do Sul, Centra–West
Brazil, in the Pantanal wetlands. Flowers during the wet season from January to May.
Etymology:—The epithet is a name in apposition of the Portuguese demonym for an inhabitant of the Pantanal,
pantaneira.
Conservation status:—Vulnerable—VU D2. Utricularia pantaneira is known for a few populations from the
east margin of Pantanal Wetland, a region that suffers from intense human interference, such as the record burning of
2020 (Pivello et al., 2021). Although intensifying sampling efforts in the region could uncover further populations of
the taxon, the increasing pressure on natural habitats in the region could lead the species to a more severe category of
threat in the short term.
Additional specimens Examined:—BRAZIL. MATO GROSSO: Santo Antônio do Leverger, região das chácaras
do Aricá, 22 Fev. 1994, Rivadavia & Cardoso 248 (SPF). MATO GROSSO DO SUL: Fazenda Santa Rosa, Paiaguas,
Pantanal, 24 Maio 1986, Pott, A. 2227 (UFMT).
10. Utricularia roraimensis Brown (1901: 54)—Fig.3–D
Type:—VENEZUELA: Mt. Roraima summit, autumn 1898, McConnell & Quelch 685 (Holotype: K!).
≡Calpidisca roraimensis Gleason (1929: 405)
=Utricularia tepuiana Steyermark (1953: 550)
Type:—VENEZUELA: Bolivar: Ptari-tepui, 29 October 1944, Steyermark 59517 (Holotype: F!, Isotype: VEN).
Illustrations:—(Brown, 1901), Plate 11: figs1–4.
Description:—Terrestrial. Small sized perennial herb. Rhizoids not observed. Stolons few. Leaves spathulate or circular
from an attenuate base 4–15 mm long, traps 1 mm long. Inflorescence simple, lax, 40–80 mm long, peduncle glabrous,
0.7 mm thick, bract and bracteoles similar, connate at middle, bract, bracteoles ovate obtuse apex, 1 mm long. Flowers
usually 1–2, pedicel erect, filiform, (2–)3–8 mm long, calyx lobes unequal, vinaceous, chartaceous, concave, glabrous,
with inconspicuous simple nerves, upper calyx lobe elliptic or elliptic ovate, cupulate, 2 mm long, lower calyx lip
broadly elliptic, apex truncate emarginate, 1 mm long, corolla lilac with a yellow spot bordered by white at base of
lower lip, upper lip ovate, rounded or obtuse apex, 3–5 × 2–3 mm, x2> than the calyx, glandular trichomes present,
lower lip transversally elliptic, base forming a glandular bilobed swelling which extends for half of lower lip, apex
slightly 3-lobed, 3 × 4–5 mm, spur conical or ovate, apex rounded, curved upwards, slightly bigger or smaller than
lower corolla lip, c. 3 mm long, filaments slightly straight, anther theca distinct, ovary globose, style very short, stigma
lower lip entire truncate, upper lip teeth like. Capsule not observed.
Distribution and ecology:—Endemic to the Venezuelan tepuis, states Amazonas and Bolivar, on the top of tepuis
Mount Roraima, Cerro Atabapo region, Meseta de Jaua, Chimanta Massif, Ptari tepui, Tramen tepui and Kukenan
tepui. The populations are found at elevation c. 2270m on rocky areas with white sand substrate on top of the table
mountains or white sand savannahs at lower elevations (140 m) like Cerro Atabapo (Fig. 1A, C). It may also occur in
Brazil and Guyana. Apparently, in flower year around.
Etymology:—The species epithet refers to Mt. Roraima, the type specimen locality.
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 47
Conservation status:—Least concern (EOO = 34,560 km2; AOO = 32 km2). Even though geographically
restricted to the Mount Roraima ( which receives considerable environmental pressure from visitors) and from a few
other tepui tops, this species is not considered under threat due to its remoteness. (Fig.1 A,C).
Notes:—Utricularia roraimensis (Fig.3D) is distinct from U. amethystina s.str. on the grounds of its divergent
morphology and geographic isolation. U. roraimensis possess reduced corolla spur that is usually flattened and curved
upwards, glandular trichomes on the upper corolla lip and spur, and vinaceous scape and calyx lobes. Known only from
unique habitats of high elevation on the top of table mountains (tepuis).
Additional speciemens examined:—BRAZIL/VENEZUELA/GUIANA. Bolivar, Roraima summit, Sep. 1974,
Warren s/n (K). Dist. Rocio (ahora Sinfontes): cumbre del Tramen-Tepui, en la porción más noroccidental del macizo
del Ilu-(Uru) Tepui, 23 Jan. 1985, Hubber, O. 10.064 (K). Rocio, Kukenan (Mataui)—Tepui, cumbre meridional,
no collection date, Hubber, O. 10.554 (K). VENEZUELA. AMAZONAS: Atabapo, Cerro Marauca-Huha, cumbre
altiplanicie no arbolada, no collection date, Steyermark et al. 125936 (K).
11. Utricularia tricolor Saint-Hilaire (1833: 418)—Fig. 4–I
Type:—BRAZIL. RIO DE JANEIRO: Lieux humides & sablonneux voisins de la mer près S. Joao de Barra, 1816–21, St.Hilaire B2-161
(holotype: P! (photo, M), Isotype: MPU, P).
=Utricularia fontana Saint-Hilaire & Girard (1838: 870)
Type:—BRAZIL. SÃO PAULO: in quodam fonte prope Paulopolium, Saint-Hilaire C1-1147 (Holotype: P image!, Isotype: P image!).
=Utricularia gomezii De Candolle (1844: 13)
Type:—BRAZIL. RIO DE JANEIRO: Gomez s.n. (Holotype: G, Isotype: K!, P image!).
=Utricularia globulariifolia Martius ex Benjamin (1847: 241)
Type:—BRAZIL: RIO DE JANEIRO: Gardner 590 (Holotype: BM!, Isotype: G!, K!, NY!, P image!, S, W).
=Utricularia fusiformis Warming (1874: 9)
Type:—BRAZIL. MINAS GERAIS: Lagoa Santa, Warming s.n. (Holotype: C image! (photo, M)).
=Utricularia globulariifolia var. caudata Sylvén (1909: 23)
Type:—BRAZIL. MINAS GERAIS: Caldas, Mosén 4349 (Holotype: C image!).
Illustrations:—Taylor (1989), fig. 79; Baleeiro et al. (2016), fig. 2m; Baleeiro et al. (2019), fig. 1c.
Description:—Terrestrial, or affixed sub-aquatic. Small to medium sized, annual herb. Rhizoids up to 90 mm long.
Leaves usually 1(–3) from peduncle base, lamina obovate to circular, 5–55(–65) mm long, traps numerous on rhizoids
and stolons, broadly ovoid, 1–2 mm long. Inflorescence simple (240–)300–750 mm long, peduncle glabrous, 0.5–
1.4mm long, scales basifix, glabrous, deltoid, apex acute, margin entire 1.3–1.5 mm long, bract ovate to deltoid, apex
acute c. 1.5 mm long, bracteoles smaller or similar length, lanceolate, connate near the base, or sometimes middle,
glabrous. Flowers 1–8, pedicel (1–)2–4 mm long, calyx lobes unequal, coriaceous, usually glabrous or papillose near
margin, rarely with sparse sessile glands, inconspicuous simple nerves, upper lobe ovate-oblong or broadly elliptic
to circular, concave, apex rounded, margin entire, hyaline 2.8–3.1 × 2.3–2.7 mm, lower lobe broadly elliptic, entire,
margin not hyaline, apex crenate or emarginate, 1.5–2.0 × 2.5–3.1 mm, corolla lilac with yellow spot with white
margin at base of lower lip, 10–25 mm long, with papillae and few glandular trichomes, upper lip broadly ovate, apex
rounded, equal to about twice the length of calyx lobes, 2.0–4.3 mm long, lower lip transversally elliptic or broadly
obovate, base forming a bilobed swelling, apex entire or shallowly 3-lobed, 6–9 × 8–10 mm, spur slightly conical,
apex obtuse, equal or slightly larger than the lower lip, 7.2–8.4 mm long, filaments curved, anther thecae sub-distinct,
ovary globose with some glands on the outside wall. Capsule globose up to 4 mm diam. Seeds narrowly cylindrical c.
1 mm long, 0.15 mm wide.
Distribution and ecology:—Argentina, Brazil, Colombia, Uruguay, and Venezuela (Fig. 1). In Brazil, in the states
of Bahia, Espirito Santo, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do
Sul, Santa Catarina and São Paulo. Grasslands bordering creeks, lakes, gallery forests, veredas (a phytophysiognomy
composed of the palm Mauritia flexuosa L.f.), restingas, and campo rupestre. Flowering August to May.
Etymology:—The epithet tricolor refers to the three colors of the corolla, lilac with yellow and white at base of
the lower lip.
Conservation status:—Least Concern. Based on its wide distribution (EOO = 8,284,102 km2, AOO = 132 km2)
and frequent presence in protected/conservation areas, U. tricolor is here considered Least Concern, in accordance
with Cross et al. (2020).
Notes:—Utricularia tricolor (Fig.4I) is easily distinguished from all other U. sect. Foliosa species by its robust
specimens with tall inflorescence scapes and large flowers. The most similar species is U. bicolor (previously a
BALEEIRO ET AL.
48 • Phytotaxa 576 (1) © 2022 Magnolia Press
synonym of U. amethystina), which can be mistaken for U. tricolor from the Espinhaço Range in Minas Gerais,
where both taxa are sympatric, and specimens of U. tricolor are often smaller. In some specimens, it is possible to
observe a few trichomes and papillae near the calyx margin, one of the defining characters of U. bicolor. However,
U. tricolor has a larger corolla with lower lip apex almost entirely or shallowly 3-lobed and glabrous calyx, except in
some populations collected from the Espinhaço Range. At this site it is notable that U. bicolor does not possess a white
margin surrounding the yellow spot at the base of lower lip, as does U. tricolor. Nevertheless, the material examined
during this study suggests hybridisation between these two taxa. It is worth mentioning that these two species are not
phylogenetically close, with U. tricolor sister to U. amethystina s.str., and U. bicolor sister to a clade containing the U.
hirtella complex (Baleeiro et al. 2019).
Additional specimens Examined:—ARGENTINA. Corrientes, Dep. San Tome, 4km E de Ruta Nac. n.14,
caminho a colônia Garabi, 3 Dic. 1970, Krapovickas s/n (RB 176498).
BRAZIL. BAHIA: Abaíra, Catolés, subindo riacho que desce da serra do Guarda-mor, 26 Jul. 1995, Rivadavia
479 (SPF); ibid, em campos rupestres beirando a trilha para o campo da Mutuca, Serra da Mesa, Vila de Catolés, 15
Jul. 2005, Rivadavia et al. 2020 (SPF). Mucugê, Serra de São Pedro, 17 Dez. 1984, Lewis et al. CFCR 7078 (SPF).
Piatã, campos rupestres ao lado da capelinha na trilha subindo a Serra do Santana, 13 Jul. 2005, Rivadavia et al. 2001
(SPF). Rio de Contas, campos rupestres próximo a trilha para o alto do Pico do Itobira, Serra da Mesa, 18 Jul. 2005,
Rivadavia et al. 2077 (SPF). GOIÁS: Alto Paraiso de Goiás, Parque Nacional dos Veadeiros, córrego dos Ingleses,
19 Jun. 2013, Baleeiro & Carneiro 335 (SPF); ibid, Parque Nacional dos Veadeiros, cachoeiras Almecegas I e II, 21
Jun. 2013, Baleeiro & Carneiro 341 & 344 (SPF); ibid, Chapada dos Veadeiros, km 200 da estrada Brasília—Alto
Paraíso, 13 Abr. 1995, Rivadavia & R.C. Ogassavara 373 (SPF). Barro Alto, GO-080, 6 Jan. 2001, Rivadavia 1229
(SPF). Cristalina, BR-060. RPPN Linda Serra dos Topázios, 9 Jun. 2009, Baleeiro et al. 72 (R). Mineiros, estrada
para o Parque Nacional das Emas, 3 Mai. 1999, Rivadavia & R.K. Sano 1041 (SPF). Rio Verde, BR 060, 16 Jan.
2004, C.P. Bove et al. 1332 (R). DISTRITO FEDERAL: Brasília, Reserva Ecológica do IBGE, brejo entre o Guará
e o setor da indústria, 21 Fev. 1981, F. Chagas & Silva 390 (IBGE). Gama, estrada para o Santuário Ecológico, 26
Abr. 1999, Rivadavia & R.K. Sano 944 (SPF). MATO GROSSO DO SUL: Rio Verde do Mato Grosso, estrada para
Morro do Padre, vereda, 3 Nov. 20112, A.V. Scatigna et al. 281 (UEC). MINAS GERAIS: Chapada Gaúcha, nascente
brejosa com buritis no lado E do Parque Nacional Grande Sertão Veredas, 28 Abr. 1999, Rivadavia & R.K. Sato 960
(SPF). Diamantina, Parque Nacional Sempre Vivas, 8 Mar. 2013, Baleeiro et al. 307 (SPF); ibid, campo em frente ao
entroncamento para Conselheiro Mata, 9 Mar. 2013, Baleeiro et al. 310 (SPF). Jaboticatubas, Serra do Cipó, 11 Set.
1994, Rivadavia 307 (SPF). Prata, c.2.5km ao noroeste da cidade, brejo beirando a BR497, 28 Jul. 2007, Gonella et
al. 19 (SPF). Rio Vermelho, Pedra Menina, Serra do Ambrósio, Espigão do Meio Faz. Vargem do Anjo, 8 Set. 1986,
Mello-Silva et al. CFCR 10208 (SPF). Santana do Riacho, Serra do Cipó, em nascente brejosa antes da bifurcação
da BR10 para Conceição do Mato Dentro e Morro do Pilar, 12 Nov. 2007, Gonella et al. 43 (SPF); ibid, trilha para
cachoeira Capivara, 5 Jul. 2012, Baleeiro et al. 204 (SPF). São Roque de Minas, 13 Mar. 1990, Souza, H.C. s/n
(BHCB); ibid, caminho do Abrigo de Pedra, Parque Nacional Serra da Canastra, 22 Dez. 1991, Rivadavia 42 (SPF).
Uberaba, em brejo c.30km ao noroeste da cidade, no lado leste da MG 190, 27 Jul. 2007, Gonella et al. 8 (SPF).
PARAÍBA: Mamanguape, 1 Jan. 1900, Felix 2104 (JPB). PARANÁ: Palmeira, Rio das Pombas, beirando o km 522 da
BR376, 25 Nov. 2006, Rivadavia & Rohrbacher 2454 (SPF). RIO DE JANEIRO: Cabo Frio, entre Lagoa de Araruama
e Praia de Massambaba, brejo dos Espinhos, 20 Dez. 1982, fl., Pedrosa, D.S. et al. 742 (GUA). Carapebus, praia entre
Lagoa Paulista e Lagoa de Carapebus, C.P.R. Batista 35 et al. (R). Saquarema, Praia do Sossego, C.B. Moreira 53
et al. (R). Teresópolis, descendo a estrada Petrópolis-Teresópolis (BR 495) em direção a Teresópolis, 19 Nov. 2000,
Rivadavia 1220 (SPF). SÃO PAULO: Bocaina, 10 Fev 1870, Glaziou 8218 (R). Botucatu, c.1km pela estrada de ferro
da FEPASA a partir da rodovia Marechal Rondon, em nascentes brejosas em encosta de morro acima dos trilhos, 21
Mar 1999, Rivadavia et al. 860 (SPF). Cananéia, Parque Estadual Ilha do Cardoso, 9 dez. 2002, Foster et al. 978
(SPF). Itararé, descendo trilho de trem em direção a Gruta Santa, próximo à divisa SP/PR, 25 Jan. 1993, Rivadavia
167 (SPF). Ribeirão Pires, Velha estrada para Santos, próximo ao início da decida da Serra, 11 Jan. 1992, Rivadavia
& Cardoso 53 (SPF). São Jose do Barreiro, Parque Nacional da Bocaina, cachoeira de Santo Izidro, 22 Out. 2002,
Rivadavia & Morais 1414 (SPF). Vargem Grande do Sul, brejo na estrada para Casa Branca, antes do Rio Verdinho, 14
Nov. 1997, J.H.A. Dutilh & W. Marcondes-Ferreira s/n (UEC 94951). SANTA CATARINA: Itapiruba, sul da cidade,
a alguns km da BR 101, 7 Set. 2004, Rivadavia 1861 (SPF). Tocantins, Parque Estadual do Jalapão, Brejo do Porco
Podre, local de colheita do capim dourado, 19 Jun. 2002, Barbosa Cavalcanti 3005 (CEN).
COLOMBIA. SANTANDER: Charala, corregimento de Virolin: em zonas pantanosas cerca de Virolin, 21 Mar.
2008, Vieira, S. 4 (SPF).
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 49
VENEZUELA. ESTADO AMAZONAS: Cerro Duidia, pico ao extremo NW do maciço, vizinho a aldeia indígena
de Culebra, 5 Jan. 2003, Rivadavia & Hoogenstrijd 1515 (SPF).
12. Utricularia tridentata Sylvén (1909: 28)—Fig. 4H
Lectotype (designated by Taylor 1989):—BRAZIL: Rio Grande do Sul, S. Pedro do Rio Grande do Sul (S). Lindman A735 (lectotype: S
image!).
=Utricularia ternata Sylvén (1909: 29)
Lectotype (designated here):—BRAZIL: Rio Grande do Sul: Canoas near Porto Alegre, Malme I: 310 (S image!).
=Utricularia ostenii Hicken (1912: 180)
Lectotype (designated here):—URUGUAY: Carrasco near Montevideo, Osten 5230 (IS 2130, Isolectotype: IS 2129, MVFA image!).
=Utricularia rubra Larrañaga (1923: 15)
Lectotype (designated here):—URUGUAY: 22 Dec 1812, presumably Larrañaga s.n. (MVFA image!).
Illustrations:—Taylor (1989), fig. 78; Baleeiro et al. (2017), fig.2m.
Description:—Terrestrial. Medium sized annual herb. Rhizoids few, short branches, papillose, stolons few, c. 0.5
mm thick. Leaves 1–6, rosulate from peduncle base, lamina circular or broadly obovate from an attenuate base, 5–10
mm long, traps broadly ovoid, on rhizoid and stolon, 1.0–1.5 mm long. Inflorescence simple, lax, rarely branched,
65–160(–230) mm long, peduncle glabrous 1 mm thick, scales basifixed, glabrous, ovate, up to 1 mm long, bract and
bracteoles connate attached near middle, glabrous, 3 nerved or none, bracts ovate or deltoid, apex acute, bracteoles
linear, smaller than the bracts. Flowers 1–3(–6), pedicel erect, filiform c. 5 mm long, calyx lobes unequal, cartaceous,
concave, glabrous, with inconspicuous simple nerves, upper lobe ovate-oblong, apex rounded, entire margin hyaline 3
mm long, lower lobe broadly to transversally elliptic, margin entire, apex rounded or emarginate 2.8 mm long., corolla
lilac with yellow spot bordered by white margin at base of lower lip, minutely papillose with few glandular trichomes
7.5–10.0 mm long, upper lobe ovate, apex rounded, same or slightly longer than the sepal, lower lip transversally
elliptic or broadly obovate, base forming a 2–3 lobed swelling, apex 3–lobed, 4 × 10 mm, spur narrowly conical, apex
obtuse, slightly longer than the lower lip, filaments slightly curved c. 1.5 mm long., anther theca sub-distinct, ovary
globose, glandular, style short, stigma lower-lip semicircular, upper-lip absent. Capsule globose 2.5 mm diam., calyx
not covering the stigma lips. Seeds narrowly cylindrical, 0.5–0.6 × 0.1–0.15 mm.
Distribution and ecology:—Southern South America, Argentina, Uruguay and Brazil (Fig.1). In Brazil, the
species is recorded in the southern states of Paraná, Santa Catarina and Rio Grande do Sul, and disjunctly at the
higher mountains of Mantiqueira Range, at Itatiaia National Park in Rio de Janeiro, and Caparaó National Park at the
Minas Gerais-Espírito Santo border. Occurring on wet substrate in high altitude grassland (campo de altitude) habitats.
Flowering throughout the year.
Etymology:—From the Latin tridentata = having three teeth. Refers to the margin of the bract and the bracteoles. In
the past, it was common to describe the bract `tridentatis`, considering both bract and bracteoles as a single structure.
Conservation status:—Least Concern. Even though U. tridentata is restricted to southern regions of South
America (EOO = 181,082 km2, AOO = 40 km2), it commonly occurs within conservation areas. Following IUCN
(2012) categories categories and criteria, it is to be considered Least Concern. The species is listed in the Brazilian Red
List as Vulnerable (CNCFlora, 2021).
Notes:—Utricularia tridentata (Fig. 4H) most closely resembles U. amethystina and U. lindmanii, with few
morphological characters that differentiate between them. Geographically, their distributions do not overlap, and U.
tridentata is found in wetlands at elevations between 1000 and 2000 m (Taylor 1989). The corolla is minutely papillose
in U. tridentata vs. glabrous in U. amethystina and U. lindmanii. The leaves in U. tridentata are usually circular,
whereas in U. amethystina they are spathulate, and filiform in U. lindmanii.
Additional specimens examined:—BRAZIL. MINAS GERAIS: Alto Caparaó, Rivadavia et al. 525 (SPF), Serra
do Cipó, 15.VI.1935, H.L.M. Barreto 1066 (RB). PARANÁ: Palmeira, Rio das Pombas, beirando o km522 da BR376,
25 Nov. 2006, Rivadavia & Rohrbacher 2455 (SPF). RIO GRANDE DO SUL: Cambará do Sul, Parque Nacional
dos Aparados da Serra, Canion de Itaimbezinho, 23 Jul. 2000, Rivadavia & Valente 1207 (SPF). RIO DE JANEIRO:
Itatiaia, estrada Nova, km 15, Mai. 1950, Brade, A.C. 20329 (RB). Teresópolis, Serra dos Órgãos, 12 Jul. 1891, Ule
s.n. (R). SANTA CATARINA: Guaruva, em campos de altitude no alto da serra do Quiriri, Nov. 2006, Rivadavia
& Rohrbacher 2437 (SPF). Grão Pará, início da descida da Serra do Corvo Branco, rodovia SC439, 28 Abr. 2007,
Rivadavia 2516 (SPF). Rancho Queimado, Chapadão Serra Boa Vista, 27 Abr. 2007, Rivadavia 2510 (SPF). São
Joaquim, Dist. Bom Jardim, margin of falls and campo, east of Bom Jardim, 2 Jan. 1965, Smith & Reitzz 14190 (US).
Boggy brook, Santa Barbara, 3 Jan. 1965, Smith & Reitz 14199 (US). Bog, Source of Rio Capivaras, Serra do Oratorio,
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10km east of Bom Jardim da Serra (Cambajuva), 16 January 1957, Smith & Reitz 1013 (US). Urubici, km8 da estrada
para o Morro da Igreja, 28 Abr. 2007, Rivadavia 2514 (SPF).
URUGUAY. Near the mouth of the Arroyo Solis Chico, Dep. Canelones, 4 December 1955, Pedersen 3644 (US).
Montevideo, Jan. 1871, Gilbert 1089 (K 533964), Carrasco, 20 Nov. 2010, Ex HERB. CORN. OSTEN 5337 (US).
13. Utricularia trinervia Benjamin (1847: 247)—Figs.1A,B, 4O
Type:—BRAZIL: São Paulo, Guaratingueta, Martius s.n. (Holotype: M!).
=Utricularia selloi Werber ex Benjamin (1847: 247)
Type:—BRAZIL: Southern Brazil, Sellow 147 (Holotype: photo M!).
=Utricularia punctifolia Benjamin (1847: 492)
Type:—VENEZUELA: Merida, Moritz 1133 (Holotype: BM!).
Utricularia trinervia (Fig. 4O) is not treated in detail here. A complete treatment can be found at (Guedes et al.,
2021).
14. Utricularia velascoensis Kuntze (1898: 240)—Fig. 4E–G
Type:—BOLIVIA, Velasco, July 1892, Kuntze, O. s.n., (Holotype: NY!, Isotype: B (destroyed, photo, M).
Description:—Terrestrial or affixed sub-aquatic. Small to medium sized perenial herb. Leaves not frequent, spathulate
to circular, cuneate base, no nerves, upper surface densely pustulate (5–)7–20 mm long, traps 1 mm long. Inflorescence
mostly simple, glabrous, 80–290(–330) mm long, scales ovate acute or acuminate apex 0.5–0.7 mm long, bract ovate,
acuminate obtuse apex, 0.8–1.0 mm long, bracteoles same size of bract or smaller, 0.8mm. Flowers 1–6, pedicel erect
0.7–1.7, calyx lobes unequal, cartaceous, concave, papillose, sparse simple trichomes mainly near the border, upper
sepal broadly ovate with rounded-ciliate apex, hyaline margin, culpulate, 2.2 × 1.6 mm, lower sepal broadly ovate 1.5
× 1.8 mm, corolla lilac with the lower lip base spotted white and yellow on the swelling apex, 5–15 mm long, upper
lip ovate-elliptic apex rounded, x3 > than the upper calyx lobe, lower lip transversally elliptic to broadly obovate, base
forming a ball like bilobed swelling c. 7 × 10 mm, filaments curved in the base blackish 1.2 mm long (dried specimen),
thecae c. 0.9 mm long, ovary globose, style short, blackish, lower lip stigma circular ciliate, upper lip absent. Capsule
not seen.
Distribution and ecology:—Bolivia and Brazil. In Brazil, in the states of Amazonas, Mato Grosso, and Pará. East
Bolivia at Velasco municipality, and Noel Kempff National Park. Grows within swampy grassland west of the State
of Mato Grosso and Campinas in the Amazon, with Utricularia costata Taylor (1986: 7), U. pusilla and Drosera spp.
Flowers from February to June.
Etymology:—The species epithet refers to the type locality of Velasco province, eastern Bolivia.
Conservation status:—Least Concern. Utricularia velascoensis presents a wide distribution range (EOO = 888,164
km2, AOO = 44 km2) and is recorded from the Noel Kempff National Park in Bolivia. Nevertheless, the populations
from Brazil are all found outside protected areas and are vulnerable to severe human impact from agriculture related
threats, such as eutrophication of wetlands, increased fire regimes, and habitat destruction. Therefore, we recommend
that the species be listed as Endangered in the country according to criterion B2ab(iii) of IUCN (2012), given its
restricted occurrence in the country (AOO = 28 km2) and listed threats.
Notes:—Although U. velascoensis (Fig.4E–G) brings some morphological similarities with U. damazioi, it can
be distinguished by its sepals possessing numerous papillae and simple trichomes mostly in the edges. Its purple
corolla is covered in minute simple trichomes that shine when dried.
Additional specimens examined:—BOLIVIA. SANTA CRUZ: Velasco, Campamento Las Torres, margen de
Rio Itenez (Guapore), frontera com Mato Grosso, lado noroeste del Serrania Huanchaca, 24km sur de Flor de Oro,
mais ou menos 50km norte de Rio Verde, pampa mucho humeda (non inundada), al base del cerro, 24 May. 1991,
Marielos & Foster 207 (F!, SCZ). Velasco, Parque Noel Kempff M. Campamento La Torre, con diferentes tipos de
formaciones desde pampa inundada, 16 Jun. 1994, Quevedo et al. 2579 (MO). Parque Nacional Noel Kempff Mercado,
Campamento Las Gamas, con ecotonos de Pampa, cerrado, bosque de galeria y bosque humedo, 25 Feb. 1997, Guardia
& Surubi 222 (MO). DEPT. LA PAZ: Parque Nacional Madidi, entre Queara y Mojos, campamento Tocoaque, bosque
de yungas montano inferior pluvial y vegetacion secundaria, 1 Mar. 2008, Fuentes & Quisbert 12211 (MO). Nuflo de
Chavez, Est. Santa Maria, 7km S of Concepcion, 5 Apr. 1987, Killeen 2438 (NY).
BRAZIL. AMAZONAS: Manicoré, BR 230, 150km ao L de Humaitá e 18 km ao S pela Rod. Do Estanho, 15 Abr.
1985, Cid Ferreira 5522 (NY). Novo Aripuanã, BR 230, rodovia trans-amazônica entre os kms 350 e 400 ao leste de
U. SECT. FOLIOSA, LENTIBULARIACEAE Phytotaxa 576 (1) © 2022 Magnolia Press • 51
Humaitá, 30 Abr. 1985, Cid Ferreira 5896 (NY). Baixo Rio Negro, 25 Abr. 1911, Ducke s/n (F). MATO GROSSO:
Aguá Boa, Xavantina-Cachimbo road, 60km from Xavantina, wet campo, between cerrado and galllery forest, 6 June
1966, Hunt & Ramos 5826 (NY). Juruena, Rio Juruena, cachoeira São Simão, 21 Mai. 1977, Rosa & Santos 1952
(F). Porto Espiridião, Rodovia MT-265, 28 Jan. 2014, Cardoso 132 (SPF). PARÁ, Itaituba, Serra do Cachimbo, base
aérea, estrada Santarém-Cuiabá, BR 163, Km 794. Campina, solo arenoso, solo úmido, 25 Abr. 1983, Amaral et al. 950
(NY).
Incertae Sedis
Utricularia chamissonis Weber ex Benj., Fl. Bras. 10: 241 (1847). Type not indicated.
Utricularia globulariifolia var. minor Merl ex Luetzelb., Estud. Bot. Nordéste 3: 221 (1926). Nom. nud. Type not found.
Utricularia monantha Benj. Fl. Bras. 10: 244 (1847)
Type:—BRAZIL: southern Brazil, Sellow s.n., (B (destroyed)).
This name could be placed under U. amethystina s.str. or U. tricolor. Taylor (1989) placed it under U. amethytina, but according to its
protologue the lower lip is entire, characteristic seen on U. tricolor flowers.
Utricularia rotundifolia Merl ex Luetzelb., Est. Bot. Nordéste 3: 221 (1926). Nom. nud. Type not found.
Utricularia stolonifera Benj. In Martius, Fl. Bras. 10:245 (1847)
Type:—BRAZIL: R. Amazon, Poeppig 2909 (not located), Bahia Sellow 1422 (not located)
The name was described based on the specimen Poeppig 2909. Edward Frederich Poeppig had his collection at LE (main) NZ, University
of Leipzig but was destroyed during the great world war in 1946. The specimen may be U. amethystina s.str. considering where it was
collected. Poeppig was around the Amazon basin in the state of Para as seen in his collection number 3017 (https://www.jacq.org/
detail.php?ID=509663) (isotypes of U. modesta, seen by Taylor, collected at Colares island and deposited at Austria herbarium W).
Further studies
In the molecular phylogenetic study of Baleeiro et al. (2019), accessions representing the phrase name Utricularia
sp. ‘Minas Gerais’ were found to be sister to U. tridentata s.str., from the South of Brazil. Despite the taxon forming
a strongly supported monophyly, it was not described as a new species in the current study, with further sampling
required to differentiate it from U. tridentata. Baleeiro et al. (2019) also show that U. hirtella is a species complex.
A further study addressing the nested entities will not only disentangle the complex but may also shed light on the
distribution of the genus across the Amazon rainforest. Lastly, we encourage future studies to address other species
complexes within the genus Utricularia from across South and Central Americas.
Acknowledgements
This study was possible due to the grants given to the first author PCB (CNPq 131442/2009-0, FAPESP 2013/02729-
8, and CAPES Ciência sem fronteiras 0242/13-6). We also would like to thank Ricardo Perdiz for the support during
field trips to Viruá National Park, Tapequem Hills and Bela Vista surroundings, Fernando Rivadavia for support during
field trip and discussions related to taxa diversity and distribution, the national parks managers for the logistics and
national and international herbarium managers (Overseas - F, K, M, MERL, MO, NY, P, US, W, Brasil - ASE, BHCB,
CEN, CEPEC, ESA, ESAL, HEPH, HMS, HUEFS, IBGE, INPA, JPB, MBM, MG, MIRR, NX, UB, UFG, R, RB, SP,
TANG, UEC, UFMT).
BALEEIRO ET AL.
52 • Phytotaxa 576 (1) © 2022 Magnolia Press
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