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Phytotaxa 376 (5): 214–222
http://www.mapress.com/j/pt/
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
214 Accepted by Andreas Fleischmann: 9 Nov. 2018; published: 21 Nov. 2018
https://doi.org/10.11646/phytotaxa.376.5.4
Utricularia biceps (Lentibulariaceae), a new carnivorous species endemic to the
campos rupestres of Brazil
PAULO MINATEL GONELLA1,* & PAULO CÉSAR BALEEIRO2
1 Universidade Federal dos Vales do Jequitinhonha e Mucuri, Diamantina, MG, Brazil.
2 National Herbarium of New South Wales, Mrs Macquaries Road, Sydney, NSW 2000, Australia.
* Author for correspondence. E-mail: paulogonella@uol.com.br
Abstract
Utricularia biceps (Lentibulariaceae), a new species belonging to U. sect. Foliosa, is here described and illustrated. This
new species is endemic to the campos rupestres of eastern Brazil, an extremely biodiverse and endangered vegetation. Notes
on phenology, ecology, habitat, and conservation are provided, along with a discussion on the features that distinguish this
species from the other taxa of the genus. The recent discovery of this and many other new species in the Botumirim region,
in northern Minas Gerais, highlight this area as a priority for biodiversity conservation and emphasize the importance of
extensive studies on the flora of the region.
Key-words: carnivorous plants, conservation, Eudicots, Lamiales, taxonomy
Resumo
Utricularia biceps (Lentibulariaceae), uma nova espécie pertencendo a U. sect. Foliosa, é aqui descrita e ilustrada. Essa
nova espécie é endêmica dos campos rupestres do leste do Brasil, uma vegetação extremamente biodiversa e ameaçada.
Notas sobre fenologia, ecologia, habitat e conservação são providos, junto com uma discussão sobre as características
que distinguem essa espécie das demais do gênero. A descoberta recente desta e de outras espécies novas na região de
Botumirim, no norte de Minas Gerais, destacam essa área como prioritária para a conservação da biodiversidade e enfatizam
a necessidade de estudos extensivos da flora da região.
Palavraschave: conservação, Eudicotiledôneas, Lamiales, plantas carnívoras, taxonomia
Introduction
The campos rupestres (rupestrian fields) of eastern Brazil host around 15% of the Brazilian flora in less than 1% of
the national territory with high levels of endemism, representing an important hotspot of biodiversity (Silveira et al.
2016). The campos rupestres’ core area is centered on the Espinhaço Range, a mountain chain that stretches for over
1.200 km from central Minas Gerais (SE Brazil) to northern Bahia (NE Brazil). Acknowledging the great biodiversity
and importance of the natural resources of these highlands, in 2005 its southernmost portion in Minas Gerais was
recognized by UNESCO as a Biosphere Reserve (UNESCO 2018). Despite having been left out of this Reserve, the
northern portion of the Espinhaço Range in Minas Gerais, (placed at the ecotone of three major Brazilian biomes, the
Caatinga, Cerrado and Atlantic Forest) is also extremely rich in biodiversity and natural resources, representing an
important conservation priority.
The acidic and impoverished soils of the campos rupestres are home to one of the greatest biodiversity of carnivorous
plants in the Neotropics, with 21 species of Drosera (Droseraceae), four species of Philcoxia (Plantaginaceae; the
remaining three out of the seven known species of this newly discovered carnivorous genus occur in the Cerrado
vegetation), 13 species of Genlisea, and at least 32 species of Utricularia (Lentibulariaceae) (Gonella 2018; Fleischmann
2012; Scatigna 2018; Flora do Brasil 2020 under construction).
Utricularia is a cosmopolitan genus currently comprising around 250 species, with an important center of diversity
in the Neotropics (Ellison & Adamec 2018). Of the 64 species recorded for Brazil, 16 are endemic to the country
SPECIES OF UTRICULARIA BICEPS (LENTIBULARIACEAE) Phytotaxa 376 (5) © 2018 Magnolia Press • 215
(Flora of Brazil 2020 under construction). The genus presents a wide array of life forms, including aquatic, terrestrial,
lithophytic, epiphytic, and rheophytic species, and can be easily recognized by the presence of bladder-like traps that
capture prey by suction, the utricles (Taylor 1989).
During field expeditions to study the carnivorous flora of the campos rupestres of the northern Espinhaço Range
in Minas Gerais, a remarkable species of Utricularia was discovered, which is here described as new.
Material & Methods
Field surveys were carried out by the first author in February and September 2011. No preceding herbarium collections
of the taxon are known. Another later specimen collected and provided by Carlos Rohrbacher and deposited at SPF
was also studied. The specimens were examined under a stereo dissecting microscope and the description was based on
fresh, dried and spirit material. Morphological terminology and description structure follow Taylor (1989). Herbarium
abbreviations cited through the text follow Thiers (2018).
Informal conservation status category was assessed by range size (B criterion), following IUCN Standards and
Petitions Subcommittee (2016) recommendations. The extent of occurrence (EOO) and area of occupancy (AOO)
were estimated using GeoCAT (Bachman et al. 2011).
Taxonomic Treatment
Utricularia biceps Gonella & Baleeiro, sp. nov. (Figures 1 and 2)
Type—BRAZIL. Minas Gerais, Botumirim, Rio do Peixe, 10 February 2011, Gonella et al. 376 (holotype SPF!).
Diagnosis—Species most similar to Utricularia tricolor A.St.-Hil., but distinguished by the corolla lower lip with a prominent bilobed
crest with tuberculate texture on the gibbose palate, a spur with apex swollen and bifid, about the same length of the lower lip, the
upper lip of the stigma with ciliate apex, ovoid seeds, and with trap appendages emerging from a flat extension in the apical portion
of the trap mouth.
Description—Medium-sized perennial, terrestrial. Rhizoids few, filiform, with simple short branches, up to 1 cm
long, c. 0.2 mm thick. Stolons few, filiform, sparsely branched, up to 6 cm long, c. 0.2 mm thick. Leaves few, 1–4
rosulate at the base of the peduncle, petiolate, lamina obovate with apex rounded, green to red in color, with numerous
anastomosing nerves, 5–20 × 1.5–4.0 mm. Traps numerous on the stolons, broadly ovoid, on stalk of c. 1.5 mm length,
c. 1 × 1 mm, mouth basal, with a pair of appendages emerging from a flat extension in the apical portion of the trap
mouth, apex of appendages acute, distal end of the stalk with a distinct swelling, inner surfaces of appendages and the
distal end of the stalk densely covered with fine, distally inflexed, gland-tipped hairs. Inflorescence a raceme, erect,
simple, solitary, 12.0–31.5 cm long; peduncle filiform, terete, glabrous, 0.5–0.8 mm thick, deep red in color. Scales,
bracts and bracteoles basifixed and single-nerved. Scales few, ovate-deltoid, with apex acute, 0.6–1.2 mm long. Bracts
ovate-deltoid, with apex acute, 1.0–1.2 × 0.3–0.5 mm. Bracteoles linear-subulate, slightly shorter than and basally
connate for half to 2/3 of length to the bract, with apex acute, 0.8–1 × 0.1–0.2 mm. Flowers 2–7; pedicel ascending,
filiform, terete, 4–15 mm long (longer towards the base of the inflorescence), c. 0.2 mm in diam. Calyx lobes unequal,
upper lobe longer than the lower lobe, glabrous, with few simple parallel nerves, red to greenish-red in color; upper
lobe broadly ovate with apex rounded, strongly convex, 2.0–2.2 × 1.5–1.8 mm, firm in texture with a narrow hyaline
margin; lower lobe convex, ovoid, with apex sub-acute or minutely bifid, 1.7–2.0 × 1.5–1.6 mm. Corolla violet, with
yellow-orange crest at the base of the lower lip, minutely papillose, 8–10 mm long; upper lip oblong-ovate 3–4 × 3
mm, apex rounded; lower lip transversally elliptic, shallowly 3 lobed, with a prominent yellow-orange bilobed crest at
the palate with tuberculate texture, 5.5–7.0 × c. 9 mm; spur cylindrical, narrow at the base and swollen at the mid-apex,
apex bifid, slightly shorter than or equal in length to the lower lip, c. 2 mm in diam., 5–6 mm long. Filaments curved, c.
0.7 mm long, the anther thecae subdistinct, anther c. 0.7 × 0.3 mm. Ovary globose, glandular, 0.7 mm long; style short,
0.3 mm long; stigma bilabiate, lower lip semicircular, 0.5 mm long, with apex ciliate, upper lip truncate, 0.2 mm long.
Capsule globose, to 2 mm in diam., shorter than the calyx lobes, dorsiventrally bivalvate. Seeds ovoid, reticulated in a
helical arrangement relative to the longer axis of the seed, testa cells elongate, 0.35 × 0.20 mm.
Additional specimens examined (paratypes)—BRAZIL. Minas Gerais, Botumirim, Rio do Peixe, 06 September
2011, Gonella et al. 466 (SPF); 18 July 2015, Rohrbacher 15 (SPF).
GONELLA & BALEEIRO
216 • Phytotaxa 376 (5) © 2018 Magnolia Press
FIGURE 1. Utricularia biceps. A, habit. B, utricle, side view. C, bract and bracteoles. D, calyx. E, corolla, front view. F, spur. G, stamen.
H, gynoecium. I, fruit, side view. J, seed. A–H based on the holotype; I–J based on Gonella et al. 466. Illustration by PMG.
Etymology—from the Latin biceps (two-headed), referring to the most remarkable characteristics of the species,
the crest on the palate with two protruded lobes, and also the spur with bifid apex (Figs. 1E,F; 2E–G).
SPECIES OF UTRICULARIA BICEPS (LENTIBULARIACEAE) Phytotaxa 376 (5) © 2018 Magnolia Press • 217
FIGURE 2. Utricularia biceps. A, general view of the habitat at Rio do Peixe, Botumirim. B, inflorescence. C, immature fruit, side
view. D, leaves and inflorescence bases. E, flower, front view. F, flower, side view, highlighting the projected bilobed crest on the palate
with tuberculate texture. G, flower seen from below, highlighting the spur with bifid apex (top center), and dorsal sepal seen from above
(bottom right). All by PMG.
Phenology—The species was collected with flowers in February and July and flowers and fruits in September. In
cultivation, the species flowers in the spring and fall, but no seed production was observed (C. Rohrbacher, pers.com.).
GONELLA & BALEEIRO
218 • Phytotaxa 376 (5) © 2018 Magnolia Press
Distribution and habitat—Utricularia biceps is only known from a single location, where it grows on the
margin of a perennial river on islands of vegetation over quartzitic rocks or on cracks of the same rock, in sandy soil
with organic matter (Fig. 2A, D). It was only observed growing on the north-facing margin of the river, at elevations
around 790 m.
The species is sympatric with the white and the lilac flowered forms of U. amethystina Salzm. ex Saint-Hilaire
& Girard (1838: 870), U. tricolor Saint-Hilaire (1833: 418), U. subulata Linnaeus (1753: 18), U. nana Saint-Hilaire
& Girard (1838: 869), U. nervosa Weber ex Benjamin in Martius (1847: 247), U. gibba Linnaeus (1753: 18), U.
tenuissima Tutin (1934: 334), U. purpureocaerulea Saint-Hilaire & Girard (1838: 869), Genlisea repens Benjamin in
Martius (1847: 254), G. aurea Saint-Hilaire (1833: 429) var. minor (Saint-Hilaire 1833: 430) Fleischmann (2012: 525)
and several Drosera spp. (Droseraceae).
Conservation status—Critically Endangered (CR B2abiii). Utricularia biceps presents an AOO (area of
occurrence) equal to 4 km2, being considered critically endangered based on the IUCN criterion “AOO less than 10
km2, number of locations equal to one and continuing decline of the quality of the habitat”. The occurrence of this
species lies within a recently created conservation unit, the Botumirim State Park (Decreto No 302, 04 July 2018).
However, the area where the species was found is used by tourists as a recreational spot, due to its crystal clear waters
and scenic landscapes (Fig. 2A). Incorrect management of the recreational use of this area could pose a threat through
trampling and pollution of soil and water, potentially having a negative impact on the fragile habitat in the short
term.
Taxonomic notes—Utricularia biceps belongs to Utricularia sect. Foliosa Kamiénsky (1891: 120), characterized
by the traps broadly ovate with a basal mouth and two dorsal appendages, and capsule dorsiventrally bivalvate (Taylor
1989). The species of the section can be further recognized by the bracts and bracteoles proximally connated. The
section traditionally included three species, but U. amethystina is clearly a species complex composed by numerous
taxa, which were tentatively united by Taylor (1989) under that name justified by the presence of intermediate forms
between the morphotypes. A morphometric approach, however, has corroborated the segregation of synonyms and new
species currently under U. amethystina (Baleeiro et al. 2016), and the group is currently under taxonomic revision by
Baleeiro et al. (in prep.). All accepted taxa and synonyms within this complex were analyzed and none represent the
newly described taxon.
Molecular phylogenetic studies suggested the merger of a paraphyletic U. sect. Psyllosperma Taylor (1986: 8)
within a more broadly circumscribed U. sect. Foliosa (Jobson et al. 2003; Müller & Borsch 2005). This hypothesis,
however, was based on analysis with insufficient taxon sampling and also on the position of a misidentified voucher of
U. huntii Taylor (1986: 8), which actually represents a taxon in the U. amethystina complex [Rivadavia-Lopes & Sato
1000 (SPF!)]. Unpublished phylogenetic studies (Baleeiro et al. in prep.), based on a broader taxon sampling, support
a monophyletic U. sect. Psyllosperma, sister to a monophyletic U. sect. Foliosa, a phylogenetic placement we follow
here.
Utricularia biceps grows sympatric with two other species of U. sect. Foliosa: U. tricolor and two distinctive taxa
currently assigned as morphotypes of a variable U. amethystina s.l. (sensu Taylor 1989). Like U. biceps, U. tricolor also
presents transversally elliptic and shallowly 3-lobed corolla lower lobe, but can be easily distinguished by the conical
spur, which is much longer than the lower lip and with a single, acute apex (as in all other species of the section). The
sympatric morphotypes of U. amethystina can be distinguished by the deeply 3-lobed lower lip with a long conical spur
(lilac form) or by the white corolla color, with a very short lower lip, broadly conical spur, and densely glandular-pilose
indumentum on spur and calyx (white form). Furthermore, these two species lack the tuberculate crest on the palate,
having only an inflated palate with papillose texture.
The two projections with tuberculate texture of U. biceps are a novel character described for U. sect. Foliosa.
This is similarly observed in U. geminiloba Benjamin in Martius (1847: 242) and most remarkably in U. nephrophylla
Benjamin in Martius (1847: 247) of U. sect. Orchidioides A.DC. in DeCandolle (1844: 23) (U. sect. Iperua Taylor
(1986: 10) sensu Taylor 1989), which present a swelled bilobed palate, each lobe with a rugose crest.
The upper lip of the stigma with ciliate margin is also a new character for U. sect. Foliosa. Although not cited in
the description, Taylor (1989) illustrates a pilose stigma for U. hispida Lamarck (1791: 50), of U. sect. Psyllosperma,
but in the illustration, the trichomes are distributed along the surface of the upper lip and not exclusively on the margin,
as in U. biceps.
Unlike all other species of U. sect. Foliosa, the two appendages of the trap originate from a flat extension in the
apical portion of the mouth (Fig. 1B), while in all other species the appendages are free from the base, representing
another new character among the taxa of this section.
Seeds of U. biceps are very distinct from those of the other species of U. sect. Foliosa, presenting an ovoid shape
SPECIES OF UTRICULARIA BICEPS (LENTIBULARIACEAE) Phytotaxa 376 (5) © 2018 Magnolia Press • 219
(Fig. 1J), while seeds are narrowly cylindrical in U. tricolor and U. tridentata Sylvén (1909: 28), and obliquely ovoid
to narrowly cylindrical in the U. amethystina complex. It shares with U. amethystina the rows of testa cells markedly
helically arranged relative to the longer axis of the seed. The seeds studied were harvested from unripe fruits from a
herbarium specimen, which could explain the disparate seed shape observed. No seeds were produced in cultivation
(C.Rohrbacher, pers.comm.).
Within U. sect. Foliosa, U. biceps presents the most restricted distribution, being microendemic to the Rio do Peixe
area, in the municipality of Botumirim. The other three taxa included under U. sect. Foliosa in the circumscription
of Taylor (1989) are widely distributed across South America and even Central and North America in the case of U.
amethystina. Utricularia tricolor is widely distributed across South America, from Argentina to Venezuela, whereas
U tridentata is restricted to southeastern South America, in the Brazilian states of Rio Grande do Sul, Santa Catarina,
Paraná, and Rio de Janeiro, as well as Uruguay and Argentina (Taylor 1989; Flora of Brazil 2020 under construction). On
the other hand, in the broad circumscription of Taylor (1989), U. amethystina is widely distributed across tropical and
sub-tropical America, from Florida in the United States to São Paulo, in southeastern Brazil. Such a wide distribution
of U. amethystina, however, may be a result of its broad circumscription, including a wide range of polymorphism
(Taylor 1989). Baleeiro et al. (2016) started to dissect this species complex using a morphometric approach and
showed that some of the taxa synonymized under U. amethystina may be resurrected and even undescribed species are
hidden under this name.
Discussion
Thanks to the recent discovery of a new record of the critically endangered blue-eyed ground-dove (Columbina
cyanopis Pelzeln, 1870; Aves: Columbidae) in the Botumirim region (Bessa et al., in prep.), the area received great
public attention, which pressured the state government to protect the area, and culminated in the recent creation of a
conservation unit at the state level (Decreto No 302, 04 July 2018).
Besides its natural resources and scenic beauty, the Botumirim area is also home to a great number of endemic and
endangered plant species described over the last few decades, including Paepalanthus multistelaris Andrino & Sano in
Andrino et al. (2017: 56) (Eriocaulaceae), Philcoxia rhizomatosa Scatigna & V.C.Souza in Scatigna et al. (2015: 276)
(Plantaginaceae), Psyllocarpus scatignae Carmo, Sobrado & R.M.Salas in do Carmo et al. (2018: 587) (Rubiaceae),
Schefflera botumirimensis Fiaschi & Pirani (2005: 117) (Araliaceae), Vellozia armata Mello-Silva (1996: 258)
(Velloziaceae), Habranthus botumirimensis R.S. Oliveira in Oliveira & Sano (2009: 538) (Amaryllidaceae), Cantinoa
nanuzae Harley (2014: 9530) (Lamiaceae), Sauvagesia bryoclada Queiroz-Lima & D.B.O.S.Cardoso in Queiroz-Lima
et al. (2018: 222) (Ochnaceae), Cardiospermum cristobaliae Ferrucci & Urdampilleta (2011: 479) (Sapindaceae), and
Drosera graomogolensis Silva (1997: 85) (Droseraceae). Less charismatic animal species (compared to the dove)
were also recently described from this area, such as the frog Hypsiboas botumirim (Anura: Hylidae; Caramaschi et al.
2009), and the fish Microlepidogaster discus (Siluriformes: Loricariidae; Martins et al. 2014). Despite the description
of so many new species in recent years, the surroundings of Botumirim are still very poorly explored scientifically,
highlighting the urgent need for more extensive studies in the region.
In these unfavorable times for the study and conservation of Brazilian biodiversity, when Brazil’s government
threatens to undermine the country’s research by successive cuts in funding (Fernandes et al. 2017), by passing legislation
that hinders taxonomic studies (Alves et al. 2018; Bockmann et al. 2018), and with an even more uncertain future in
the next administration that seeks to promote agricultural and industrial expansion at the expense of environmental
protections (Tollefson 2018), it is important to recognize the relevance of basic science such as taxonomy for the
conservation of the megadiverse Brazilian flora and fauna. It is also important to acknowledge the involvement of
public society in the protection of sensitive areas for the conservation of flora, fauna and hydric resources, such as with
the Botumirim region.
Throughout its range, the campos rupestres are threatened by the expansion of farming, mainly cattle ranching,
mining, uncontrolled anthropic fires, and biological invasion, especially of African grasses introduced for livestock
feeding. In the northern portion of the Espinhaço Range in Minas Gerais, Eucalyptus plantations are especially
threatening, transforming not only the landscape but destroying the campos rupestres’ topsoil. And last but not least,
the vegetation is further threatened by climate change, which threatens to reduce its suitable area by up to 95% until
the end of the century (Fernandes et al. 2014).
Current conservation units and legislation are far from ideal for protecting the biodiversity of the Espinhaço
GONELLA & BALEEIRO
220 • Phytotaxa 376 (5) © 2018 Magnolia Press
Range (Silva et al. 2008), lacking government support, with most units still not having their territories expropriated,
and/or lacking administrative and physical infrastructure (Echternacht et al. 2011). Given the low resilience of campos
rupestres vegetation and the difficulty in restoration of degraded areas (Le Stradic et al. 2014), urgent protection is a
high priority, including specific legislation to help prevent their decline. In this context, with the lawful creation of the
new conservation unit of Botumirim, it is now important that it be properly implemented, funded, and managed. The
involvement of society is once again crucial, actively supervising the implementation and management of the new park
so that it can effectively protect its unique biodiversity.
Acknowledgements
We would like to thank Fernando Rivadavia, Paulo Sano, Carlos Rohrbacher, Nílber Silva, Jonathan Santos, Rui Alves,
and Adilson Peres for field assistance; Caroline Andrino for aid in gathering literature about the new species from
Botumirim; Carlos Rohrbacher for sharing his observations on cultivated material and providing herbarium specimens
of the new species; Eduardo Gomes for valuable regional data of the recently created Botumirim park; Viviane Jono
and Roberta Figueiredo from SPF; Caroline Andrino, Fernando Rivadavia, Andreas Fleischmann, and Jan Schlauer for
critical comments and suggestions on the manuscript. During fieldwork, PMG received a scholarship from Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and financial support from the Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES).
Literature Cited
Alves, R.J.V., Weksler, M., Oliveira, J.A., Buckup, P.A., Pombal, J.P. Jr, Santana, H.R.G., Peracchi, A.L., Kellner, A.W.A., Aleixo, A.,
Bonino, A.R.L., Almeida, A.M.P., Albernaz, A.L., Ribas, C.C., Zilberberg, C., Grelle, C.E.V., Rocha, C.F.D.D., Lamas, C.J.E.,
Haddad, C.F.B., Bonvicino, C.R., Prado, C.P.A., Lima, D.O., Rossa-Feres, D.C., Santos, F.R.D., Salimena, F.R.G., Perini, F.A.,
Bockmann, F.A., Franco, F.L., Giudice, G.M.L.D., Colli, G.R., Vieira, I.C.G., Marinho-Filho, J, Werneck, J.M.C.F., Santos, J.A.D.D.,
Nascimento, J.L.D., Nessimian, J.L., Cordeiro, J.L.P., Claro, K.D., Salles, L.O., Casatti, L., Py-Danie, L.H.R., Silveira, L.F., Toledo,
L.F., Oliveira, L.F., Malabarba, L.R., Silva, M.D.D., Couri, M.S., Martins, M.R.C., Tavares, M.D.S., Sobral, M.E.G., Vieira, M.V.,
Oliveira, M.L.A., Pinna, M.C.C., Hopkins, M.J.G., Solé, M., Menezes, N.A., Passos, P., D’Andrea, P.S., Pinto, P.C.E.A., Viana, P.L.,
Toledo, P.M., Reis, R.E.D., Vilela, R., Bastos, R.P., Collevatti, R.G., Silva, R.C., Fisher, S.C. & Caramaschi, U. (2018) Brazilian
legislation on genetic heritage harms Biodiversity Convention goals and threatens basic biology research and education. Anais da
Academia Brasileira de Ciências 90 (2): 1279–1284.
https://doi.org/10.1590/0001-3765201820180460
Andrino, C.O., Trovó, M. & Sano, P.T. (2016) A field full of green stars: Paepalanthus multistellaris (Eriocaulaceae), a new species from
Minas Gerais, Brazil. Phytotaxa 278 (1): 55–61.
https://doi.org/10.11646/phytotaxa.278.1.6
Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting red list threat assessments with GeoCAT: Geospatial
conservation assessment tool. ZooKeys 150: 117–126.
https://doi.org/10.3897/zookeys.150.2109
Baleeiro, P.C., Jobson, R. & Sano, P.T. (2016) Morphometric approach to address taxonomic problems: the case of Utricularia sect.
Foliosa (Lentibulariaceae). Journal of Systematics and Evolution 54 (2): 175–186.
https://doi.org/10.1111/jse.12186
Bockmann, F.A., Rodrigues, M.T., Kohsldorf, T., Straker, L.C., Grant, T., de Pinna, M.C.C., Mantelatto, F.L.M., Datovo, A., Pombal, J.P.
Jr., McNamara, J.C., de Almeida, E.A.B., Klein, W., Hsiou, A.S., Groppo, M.E., Castro, R.M.C. & de Souza Amorim, D. (2018)
Brazil’s government attacks biodiversity. Science 360 (6391): 865.
https://10.1126/science.aat7540
Caramaschi, U., Cruz, C.A.G. & Nascimento, L.B. (2009) A new species of Hipsiboas of the H. polytaenius clade from southwestern
Brazil (Anura: Hylidae). South American Journal of Herpetology 4 (3): 210–216.
De Candolle, A.P. (1844) Prodromus Systematis Naturalis Regni Vegetabilis, vol. 8. Fortin, Masson et Sociorum, Paris, 684 pp.
Decreto No 302 (2018) Avalable from: http://jornal.iof.mg.gov.br/xmlui/handle/123456789/204095 (accessed 20 November 2018)
do Carmo, J.A.M., Sobrado, S.V., Salas, R.M. & Simões, A.O. (2018) Two new threatened species of Psyllocarpus (Rubiaceae;
Spermacoceae) from eastern Brazil. Systematic Botany 43 (2): 579–590.
SPECIES OF UTRICULARIA BICEPS (LENTIBULARIACEAE) Phytotaxa 376 (5) © 2018 Magnolia Press • 221
https://doi.org/10.1600/036364418X697300
Echternacht, L., Trovó, M., Oliveira, C.T. & Pirani, J.R. (2011) Areas of endemism in the Espinhaço Range in Minas Gerais, Brazil. Flora
206: 782–791.
https://doi.org/10.1016/j.flora.2011.04.003
Ellison, A.M. & Adamec, L. Carnivorous plants: physiology, ecology, and evolution. UOP Oxford, Oxford, 560 pp.
Fernandes, G.W., Barbosa, N.P.U., Negreiros, D. & Paglia, A.P. (2014) Challenges for the conservation of vanishing megadiverse rupestrian
grasslands. Natureza & Conservação 12 (2): 162–165.
https://doi.org/10.1016/j.ncon.2014.08.003
Fernandes, G.W., Vale, M.M., Overbeck, G.E., Bustamante, M.M.C., Grelle, C.E.V., Bergallo, H.G., Magnusson, W.E., Akama, A., Alves,
S.S., Amorim, A., Araújo, J., Barros, C.F., Bravo, F., Carim, M.J.V., Cerqueira, R., Collevatti, R.G., Colli, G.R., da Cunha, C.N.,
D’Andrea, P.S., Dianese, J.C., Diniz, S., Estrela, P.C., Fernandes, M.R.M., Fontana, C.S., Giacomin, L.L., Gusmão, L.F.P., Juncá,
F.A., Lins-e-Silva, A.C.B., Lopes, C.R.A.S., Lorini, M.L., de Queiroz, L.P., Malabarba, L.R., Marimon, B.S., Marimon Jr., B.H.,
Marques, M.C.M., Martinelli, B.M., Martins, M.B., de Medeiros, H.F., Menin, M., de Morais, P.B., Muniz, F.H., Neckel-Oliveira,
S., de Oliveira, J.A., Oliveira, R.P., Pedroni, F., Penha, J., Podgaiski, L.R., Rodrigues, D.J., Scariot, A., Silveira, L.F., Silveria, M.,
Tomas, W.M., Vital, M.J.S. & Pillar, V.D. (2017) Dismantling Brazil’s science threatens global biodiversity heritage. Perspectives in
Ecology and Conservation 15: 239–243.
https://doi.org/10.1016/j.pecon.2017.07.004
Ferrucci, M.S. & Urdampilleta, J.D. (2011) Cardiospermum cristobaliae (Sapindaceae, Paullinieae), una nueva especie de Minas Gerais,
Brasil. Brittonia 63 (4): 478–483.
https://doi.org/10.1007/s12228-011-9203-3
Fiaschi, P. & Pirani, J.R. (2005) New species of Schefflera (Araliaceae) from the Espinhaço Range in Minas Gerais, Brazil. Novon 15 (1):
117-122.
Fleischmann, A. (2012) Monograph of the genus Genlisea. Redfern Natural History, Poole, 727 pp.
Flora do Brasil 2020 under construction (2018) Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/
(accessed 10 July 2018)
Gonella, P.M. (2018) Droseraceae. In: Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available from: http://
floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB7420 (accessed 10 July 2018)
Harley, R.M. (2014) Cantinoa nanuzae, a new species of Lamiaceae from the Serra do Espinhaço, Minas Gerais, Brazil. Kew Bulletin
69: 9530.
https://doi.org/10.1007/s12225-014-9530-0
IUCN Standards and Petitions Subcommittee (2016) Guidelines for using the IUCN red list categories and criteria, version 12. Prepared
by the Standards and Petitions Subcommittee. Gland, Switzerland and Cambridge, UK: IUCN Council.
Jobson, R.W., Playford, J., Cameron, K.M. & Albert, V.A. (2003) Molecular phylogenetics of Lentibulariaceae inferred from plastid rps16
intron and trnL-F DNA sequences: implications for character evolution and biogeography. Systematic Botany 28: 157–171.
doi: https://doi.org/10.1043/0363-6445-28.1.157
Kamiénski, F. (1891) Lentibulariaceae. In: Engler, A. & Prantl, K.A.E. (Eds.) Die natürlichen Pflanzenfamilien, vol. IV, 3b. W. Engelmann,
Leipzig.
Lamarck, J.B.A.P.M. (1791) Tableau encyclopedique et methodique des trois règnes de la nature: Botanique, vol. 1. Chez Pancoucke,
Paris, 200 pp.
Le Stradic, S., Buisson, E. & Fernandes, G.W. (2014) Restoration of Neotropical grasslands degraded by quarrying using hay transfer.
Applied Vegetation Science 17: 482–492.
https://doi.org/10.1111/avsc.12074
Linnaeus, C. (1753) Species Plantarum, vol. 1. Impensis Laurentii Salvii, Holmiae, 572 pp.
Martins, F.O., Rosa, A.C. & Langeani, F. (2014) Microlepidogaster discus, a new species of Hypoptopomatinae (Siluriformes: Loricariidae)
from the rio Jequitinhonha basin, southeastern Brazil. Zootaxa 3873 (5): 560–570.
https://doi.org/10.11646/zootaxa.3873.5.6
Martius, C.F.P. von (1847) Flora Brasiliensis 10. Munich, Frid. Fleischer, 338 pp.+59 plates.
Mello-Silva, R. (1996) Two new species of Vellozia (Velloziaceae) from Minas Gerais, Brazil. Botanical Journal of the Linnean Society
120: 257–263.
https://doi.org/10.1111/j.1095-8339.1996.tb00775.x
Müller, K. & Borsch, T. (2005) Phylogenetics of Utricularia (Lentibulariaceae) and molecular evolution of the trnK intron in a lineage
with high substitutional rates. Plant Systematics and Evolution 250: 39–67.
https://doi.org/10.1007/s00606-004-0224-1
Oliveira, R.S. & Sano, P.T. (2009) Two new species of Habranthus (Amaryllidaceae) from the Espinhaço Range, Brazil. Kew Bulletin 64:
GONELLA & BALEEIRO
222 • Phytotaxa 376 (5) © 2018 Magnolia Press
537–541.
https://doi.org/10.1007/s12225-009-9144-0
Queiroz-Lima, A., Amorim, A.M.A. & Cardoso, D.B.O.S. (2018) Two more elegant species of the neglected Sauvagesia elegantissima
complex (Ochnaceae). Systematic Botany 43 (1): 221–230.
https://doi.org/10.1600/036364418X697076
Saint-Hilaire, A. (1833) Voyage dans le district des diamans et sur le littoral du Brésil, volume 2. Librairie-Gide, Paris, 465 pp.
Saint-Hilarie, A. & Girard, F. (1838) Monographie des Primulacées et des Lentibulariées du Brésil méridional. Comptes Rendus
Hebdomadaires des Séances de l’Academie des Sciences, Series. D: Sciences Naturelles 7: 868–870.
Scatigna, A.V. (2018) Philcoxia. In: Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available from: http://
floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB603264 (accessed 10 July 2018)
Scatigna, A.V., Souza, V.C., Pereira, C.G., Sartori, M.A. & Simões, A.O. (2015) Philcoxia rhizomatosa (Gratioleae, Plantaginaceae): a
new carnivorous species from Minas Gerais, Brazil. Phytotaxa 226 (3): 275–280.
https://doi.org/10.11646/phytotaxa.226.3.8
Silva, J.A., Machado, R.B., Azevedo, A.A., Drumind, G.M., Fonseca, R.L., Goulart, M.F., Moraes Jr., E.A., Martins, C.S. & Ramos Neto,
M.B. (2008) Identificacão de áreas insubstituíveis para a conservacão da Cadeia do Espinhaço, estados de Minas Gerais e Bahia,
Brasil. Megadiversidade 4: 248–270.
Silva, T.R.S. (1997) Drosera graomogolensis (Droseraceae), a new species from the campos rupestres of Minas Gerais, Brazil. Novon 7:
85–87.
https://doi.org/10.2307/3392080
Silveira, F.A.O., Negreiros, D., Barbosa, N.P.U., Buisson, E., Carmo, F.F., Carstensen, D.W., Conceição, A.A., Cornelissen, T.G.,
Echternacht, L., Fernandes, G.W., Garcia, Q.S., Guerra, T.J., Jacobi, C.M., Lemos-Filho, J.P., Le Stradic, S., Morellato, L.P.C., Neves,
F.S., Oliveira, R.S., Schaefer, C.E., Viana, P.L. & Lambers, H. (2016) Ecology and evolution of plant diversity in the endangered
campo rupestre: a neglected conservation priority. Plant Soil 403: 129–152.
https://doi.org/10.1007/s11104-015-2637-8
Sylvén, N.O.V. (1909) Die Genliseen und Utricularien des Regnell’schen Herbariums. Arkiv für Botanik 8 (6): 1–48.
Taylor, P. (1986) New taxa in Utricularia (Lentibulariaceae). Kew Bulletin 41: 1–18.
https://doi.org/10.2307/4103020
Taylor, P. (1989) The genus Utricularia: a taxonomic monograph. Kew Bulletin Additional Series XIV. Royal Botanic Gardens, Kew.
Thiers, B. (2018) [Continuously updated] Index Herbariorum: A global directory of public herbaria and associated staff. New York
Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih/ (accessed 20 November 2018)
Tollefson, J. (2018) Brazil’s presidential election could savage its science. Nature 562: 171–172.
https://doi.org/10.1038/d41586-018-06917-w
Tutin, T.G. (1934) New species from British Guiana. Journal of Botany, British and Foreign 72 (864): 333–341.
UNESCO - United Nations Educational, Scientific and Cultural Organization (2018) Available from: www.unesco.org/new/en/natural-
sciences/environment/ecological-sciences/biosphere-reserves/ (accessed 10 October 2018)
