Content uploaded by Adriane Maciel de Araújo
Author content
All content in this area was uploaded by Adriane Maciel de Araújo on Jul 20, 2022
Content may be subject to copyright.
LANKESTERIANA 22(2): 123–131. 2022. doi: http://dx.doi.org/10.15517/lank.v22i2.51844
THE DISCOVERY OF SCAPHYGLOTTIS PUNCTULATA (LAELIINAE)
IN THE HIGHLANDS OF BRAZILIAN AMAZONIA WITH A KEY TO THE
SPECIES OF THE REGION
AdriAne MAciel de ArAújo1, FrAncisco FArroñAy2, ricArdo de oliveirA Perdiz1,3,
edlley PessoA4,5 & leAndro GiAcoMin1,6,7
1Programa de Pós-Graduação em Botânica, Instituto Nacional de Pesquisas da Amazônia (INPA),
Manaus, Amazonas 69060-001, Brazil.
2Laboratório de Botânica Amazônica (LABOTAM), Instituto Nacional de Pesquisas da
Amazônia (INPA), Manaus, Amazonas 69060-001, Brazil.
3Luz da Floresta, Boa Vista, Roraima 69306-320, Brazil.
4Laboratório de Estudos Integrados de Plantas, Departamento de Botânica e Ecologia,
Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso 78060-900, Brazil.
5Programa de Pós-Graduação em Biodiversidade, Ambiente e Saúde,
Universidade Estadual do Maranhão, Caxias, Maranhão 65604-290, Brazil.
6Centro de Ciências Exatas e da Natureza - Campus I, Departamento de Sistemática e Ecologia,
Universidade Federal da Paraíba, João Pessoa, Paraíba 58051-090, Brazil.
7 Instituto de Ciências e Tecnologia das Águas & Herbário HSTM,
Universidade Federal do Oeste do Pará, Santarém, Pará 68040-255, Brazil.
8Author for correspondence: adrianne.maciel.a@gmail.com
ORCID of the Authors: AMA , FF , ROP , EP , LG
Received 21 April 2022; accepted for publication 23 June 2022. First published online: 19 July 2022.
Licensed under a Creative Commons Attribution-NonCommercial-No Derivs 3.0 Costa Rica License.
Introduction. Scaphyglottis Poepp. & Endl. ranges
from Mexico to southern Brazil and contains 78 spe-
cies (Dressler 2001, Dressler et al. 2004, Govaerts et
al. 2021, Szlachetko & Kolanowska 2014). Molecu-
lar and morphological studies place the genus in the
Laeliinae (Dressler 2004). The genus may be distin-
guished from the other members of the subtribe by the
overlapping pseudobulbs, usually apical leaves, ow-
ers that are almost always resupinate, and a free col-
umn (Dressler 2001, Szlachetko & Kolanowska 2014).
Brazil has 13 recorded species of Scaphyglot-
tis (Barros et al. 2015, Brazil Flora Group 2022), of
which 11 occur in the North region of the country:
S. bidentata (Lindl.) Dressler, S. boliviensis (Rolfe)
B.R.Adams, S. emarginata (Garay) Dressler, S. fusi-
formis (Griseb.) R.E.Schult., S. graminifolia (Ruiz &
Pav.) Poepp. & Endl., S. imbricata (Lindl.) Dressler, S.
modesta (Rchb.f.) Schltr., S. prolifera (Sw.) Cogn., S.
reexa Lindl., S. sickii Pabst, and S. stellata Lodd. ex
Lindl. (Brazil Flora Group 2022). Of these 11 species,
eight are recorded from the Guiana Shield (GS) (Brazil
Flora Group 2022, Cantuária et al. 2021, Pessoa et al.
2015), a region that, within Brazil, is within the states
of Roraima, Amapá, Amazonas, and Pará (Barbosa-
Silva et al. 2020, Funk et al. 2007).
Regarding hydrography, the GS region covers
parts of the Amazon, Negro, and Orinoco river basins
(Barbosa-Silva et al. 2020, Berry & Riina 2005). In
Brazil, the region contains elevation areas higher than
Central Amazonia, including mountains and tepuis that
harbor considerable plant species diversity and ende-
mism (Riina et al. 2019). In addition, a series of scien-
AbstrAct. The genus Scaphyglottis Poepp. & Endl. presents 78 species and occurs from Mexico to Brazil.
Scaphyglottis punctulata (Rchb.f.) C.Schweinf. is known from various high elevation locations from Panama
to Bolivia. Here, we report new records in two regions of the Brazil portion of the Guiana Shield. We provide a
detailed description, photographic plates, updated distribution map, ecological and taxonomic comments for S.
punctulata, and an identication key for Scaphyglottis species from the Brazilian portion of the Guiana Shield.
Keywords/PAlAbrAs clAve: América del Sur, epítas, epiphytes, Escudo Guayanés, Guiana Shield, South
America, taxonomía, taxonomy
LANKESTERIANA
124
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
tic expeditions carried out recently to some Brazilian
protected areas within the Guiana Shield found new re-
cords for the Brazilian ora (Barbosa-Silva et al. 2016,
2020, Farroñay et al. 2019, Flores & Rodrigues 2017),
including several species of Orchidaceae (Cantuária et
al. 2020, Pessoa et al. 2013).
Recently, while analyzing specimens deposited in
the herbarium of the Instituto Nacional de Pesquisas da
Amazônia (INPA) coming from oristic surveys, we
registered for the rst time in Brazil the occurrence of
S. punctulata (Rchb.f.) C.Schweinf. Here, we present
these new records, accompanied by an updated de-
scription, taxonomic and ecological comments, distri-
bution, photographic plates, and an identication key
to species of Scaphyglottis recorded in the Brazilian
portion of the GS.
Materials and methods. All cited specimens are de-
posited in the INPA herbarium (acronym according to
Thiers 2022, continuously updated). Specimen iden-
tities were conrmed with the help of various proto-
logues and specialized literature (Reichenbach 1855,
Schweinfurth 1960, Szlachetko & Kolanowska 2014,
Vásquez & Ibisch 2004). Morphological analyzes and
measurements were carried out in dried material only,
and dried owers were hydrated and dissected for
identication. Morphological measurements were per-
formed using ImageJ software (Schneider et al. 2012)
on images of the specimens. The distribution map was
made within the R environment (R Core Team 2021),
using the R packages base (R Core Team 2021), pret-
tymapr (Dunnington 2017), raster (Hijmans 2022),
rgdal (Bivand et al. 2021), rworldmap (South 2011),
and sf (Pebesma 2018). To determine the conservation
status of S. punctulata, we calculated the extent of oc-
currence (EOO) and area of occupancy (AOO) using
the GeoCAT platform (Bachman et al. 2011). Field
photographs were taken with a Nikon D3300 camera
and photographs of dehydrated specimens with a Leica
M205C Stereomicroscope.
Taxonomic TreaTmenT
Scaphyglottis punctulata (Rchb.f.) C.Schweinf., Bo-
tanical Museum Leaets 17(2): 47. 1955. ≡ Ponera
punctulata Rchb.f., Bonplandia 3: 220. 1855. (Fig.
1). TYPE: Venezuela. Geitner s.n. (holotype: W!,
W0109821, photo; isotypes: unknown).
Epiphytic or rupicolous herbs, 17.6–50.0 cm tall.
Pseudobulbs 6.5–20.6 × 0.2 cm, green, overlapping,
cylindrical and elongated. Leaves 2 per pseudobulb,
5.3–14.0 × 0.4–0.5 cm, green, conduplicate, apical,
lanceolate, apex obtuse to slightly emarginate. Inores-
cence 1–2 owers, 22–29 mm, terminal. Floral bracts
5–8 × 1 mm, purple, lanceolate, base attenuated, apex
acute. Flowers 12.7–12.8 mm, resupinate, petals and
sepals cream with 5 purple lines, lip dark purple and
cream; ovary pedicellate 16–19 × 1 mm, purple and
green. Dorsal sepal 8 × 2–3 mm, oblong to ovate, apex
acute. Lateral sepals 7–8 × 3 mm, oblong to elliptic,
apex acute. Petals 6–8 × 2 mm elliptical to oblong,
wider at midportion, apex acute. Lip entire 10–11 × 3–4
mm, obovate, apex emarginate to straight, margin entire
to slightly eroded, reex, concave callus. Column 5–6 ×
1 mm, dark purple and cream, wings curved deltoid at
the apex, anther obovate, 4-celled, stigmatic cavity con-
spicuous, obovate. Pollinarium and Capsules not seen.
DisTribuTion, habiTaT, anD phenology: Previously re-
ported for Bolivia, Colombia, Panama, Peru, and Ven-
ezuela (Govaerts et al. 2021), and here expanded to
Brazil, Scaphyglottis punctulata occurs in highland re-
gions of northern Brazil, with the rst collection record
in 2001 in the Parque Estadual (PE) Serra do Aracá,
Amazonas state, then later collected in 2019 in the In-
digenous Territory (IT) Raposa Serra do Sol, Roraima
state. At PE Serra do Aracá, the species was recorded
in a plateau at 1200 m, in a submontane evergreen
rainforest with a canopy of ca. 10 m, on white-sand
soil. In the IT Raposa Serra do Sol, the sheet Perdiz
et al. 3619 (INPA) was collected by a waterfall on the
Cotingo River, surrounded by a forest dominated by
Dimorphandra Schott (Fabaceae), with a prevalence
of mosses and other species of epiphytic orchids in an
open submontane rainforest environment. In the same
expedition, Farronãy et al. 1646 (INPA) was found on
a 1500 m plateau, in the same type of forest, with a 25
m high canopy, in organic soil over white clay. Based
on all records available on online platforms, the spe-
cies blooms year-round (CRIA 2022, GBIF 2022).
maTerial examineD: BRASIL. Amazonas: Barcelos,
Parque Estadual Serra do Aracá, 26 August, 2001, .,
A. Vicentini & R.C. Mesquita. 1851. (INPA 214376);
ibid., 22 April, 2014, ., P. H. Labiak et al. 5707. (RB
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
125
de ArAújo et al. — The discovery of Scaphyglottis punctulata in the Brazilian Amazonia
Figure 1. Scaphyglottis punctulata. A–B. Habit. C–D. Flower, ¾ view. E–F. Flower, frontal view. Based on Farroñay 1646
(A, C, D) and Perdiz 3619 (B, E, F). Photographs by F. Farroñay, gure preparation by G. Miranda.
LANKESTERIANA
126
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
Figure 2. Scaphyglottis punctulata. A–D. Dissected perianth showing the morphological variation of the oral pieces, based
on Vicentini 1851 (A), Perdiz 3619 (B), Labiak 5707 (C), and Farroñay 1646 (D). E. Column, pedicellate ovary and
bract, based on Farroñay 1646. Photographs by F. Farroñay (A, B, D, E) and A. M. Araújo (C). Figure preparation by
G. Miranda.
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
127
de ArAújo et al. — The discovery of Scaphyglottis punctulata in the Brazilian Amazonia
Figure 3. Species of Scaphyglottis that occur in the Guiana Shield. A. Scaphyglottis stellata Lodd. ex Lindl. B. Scaphyglot-
tis sickii Pabst. C. Scaphyglottis prolifera (R.Br.) Cogn. D. Scaphyglottis fusiformis (Griseb.) R.E.Schultes. E. Scaphy-
glottis bidentata (Lindl.) Dressler. F. Scaphyglottis reexa Lindl. Photographs by F. Farroñay (A, D), A. M. Araújo (B,
E), L. P. Félix (C), and R. G. Barbosa-Silva (F). Figure preparation by G. Miranda.
LANKESTERIANA
128
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
Figure 4. Geographical distribution of Scaphyglottis punctulata. Sites of old records are indicated by black circles. New
record sites are indicated by red triangles. Map preparation by R. O. Perdiz.
601971). Roraima: Uiramutã, Terra Indígena Rapo-
sa Serra do Sol, 6 December, 2019, ., R.O. Perdiz et
al. 3619 (INPA 290578); ibid., 13 December, 2019, F.
Farroñay et al. 1646 (INPA 290579).
conservaTion sTaTus: Scaphyglottis punctulata is dis-
tributed in central and northern South America, with a
calculated EOO of 4,882,458.740 km², and an AOO of
360,000 km². Although AOO < 500 km², the number of
populations exceeds 10 (condition “a”); therefore, the
species is considered Least Concern (LC) according to
the criterea of IUCN (2012).
Taxonomic noTes: Based solely on vegetative charac-
ters, S. boliviensis is the Brazilian species morphologi-
cally closest to S. punctulata, as it also has elongated
pseudobulbs and a stalked ovary. However, S. punctu-
lata is distinguished by having 1–2 owers (vs. 1–4
owers at S. boliviensis), with an entire oblanceolate
lip (vs. trilobed, obovate). Among Brazilian species,
the owers of S. punctulata most closely resemble
those of S. fusiformis, differing by having one leaf non
conduplicate, a longer lip spathulate (1.1 vs. 0.8 cm),
and in the different coloration of the perianth (greenish
cream to purple dark vs. yellowish white).
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
129
de ArAújo et al. — The discovery of Scaphyglottis punctulata in the Brazilian Amazonia
The rst Brazilian collection of S. punctulata dates
from almost 21 years ago but was only detected after
reviewing the Scaphyglottis samples deposited at INPA
herbarium. In Amazonia, it has been relatively common
to describe new species (e.g., Damasco et al. 2019, Prata
et al. 2016) and to document new records of plants (e.g.,
Holanda et al. 2015) based on samples sitting in her-
baria for long periods. Such phenomena are reported by
Bebber et al. (2010) on a global scale but are certainly
more common in hyperdiverse tropical forests, where
access routes are still scarce. We stress that a more com-
prehensive understanding of owering plant diversity
and distribution is crucial for any conservation initia-
tive. Our results highlight the importance of new expe-
ditions to Amazonian highlands, where new records of
plants and new discoveries are expected. Policies and
funding initiatives to facilitate such expeditions would
most certainly expand the likely biased (Hopkins 2007,
2019), currently known distribution of Amazonian plant
species. It is also important to emphasize that taxonomic
studies that unravel such novelties depend on continu-
ous investment, not only for training qualied human
resources but enabling their subsequent presence in
Amazonian institutions, which currently lack adequate
and desired numbers of those professionals.
acknowleDgemenTs. A. M. Araújo thanks the Coorde-
nação de Aperfeiçoamento de Pessoal de Nível Superior
– Brasil (CAPES) – Finance Code 001 for the masters fel-
lowship, Fundo Brasileiro para a Biodiversidade (FUN-
BIO) and Humanize for the 07/2020 Conservando o Futuro
grant, the Fundação de Amparo a Pesquisa do Estado do
Amazonas (FAPEAM) - PAPAC/2019, the RB herbarium
for loans the specimens, the curator of the INPA herbarium
Dr. Michael J. G. Hopkins and botanical technician Maria-
na Mesquita for accessing the INPA herbarium vouchers,
B.S. Gustavo Miranda Montealegre for editing the photos,
MSc. Lourdes Falen Horna for her support, Dr. Leonardo
P. Félix and MSc. Rafael Barbosa-Silva for the photos, and
Dr. Alberto Vicentini for his support in this study. Adrian
Barnett is greatly acknowledged for the help with the Eng-
lish version. L. L. Giacomin received nancial support from
UFOPA/ICTA/Pipex 01/2021 and CNPq (422191/2021-3).
R. O. Perdiz was funded by a doctoral fellowship from the
Conselho Nacional de Desenvolvimento Cientíco e Tecno-
lógico (CNPq) (no. 142243/2015-9). The scientic expedi-
tion to Serra do Sol, located in the Monte Roraima National
Park, was nanced by the MMA’s Amazon Protected Areas
Program (ARPA), and was promoted and coordinated by
the Chico Mendes Institute for Biodiversity Conservation
(ICMBio) in partnership with the Ingarikó People’s Coun-
cil and collaboration of the National Indian Foundation. We
thank the Ingarikó people for warmly welcoming us to their
land - Wakî pe man.
iDenTiFicaTion key For ScaphyglottiS species known From The brazilian porTion oF The guiana shielD
1. Leaves cylindrical S. reexa
1a. Leaves at 2
2. One leaf per pseudobulb S. fusiformis
2a. Two leaves per pseudobulb 3
3. Lip clearly trilobed S. stellata
3a. Lip entire 4
4. Sepals ≤ 5 mm length 5
5. Lips lilac, obovate, petals ≥ 4 mm length S. graminifolia
5a. Lips white-cream, elliptical or spatulate, petals < 4 mm in length 6
6. Inorescence a raceme, lip ≥ 2 mm, apex acute S. sickii
6a. Inorescence a fascicle, lip ≤ 4 mm apex obtuse S. prolifera
4a. Sepals > 5 mm in length 7
7. Lip with emarginate apex, purple and cream S. punctulata
7a. Lip with apex acute or acuminate, red 8
8. Pseudobulbs fusiform, lips with yellow callus S. imbricata
8a. Pseudobulbs cylindrical, lips with brown callus S. bidentata
liTeraTure ciTeD
Bachman, S., Moat, J., Hill, A. W., de la Torre, J. & Scott, B. (2011). “Supporting Red List threat assessments with Geo-
CAT: geospatial conservation assessment tool.” In: V. Smith & L. Penev (Eds.), e-Infrastructures for data publishing
in biodiversity science. ZooKeys, 150, 117–126. doi: http://dx.doi.org/10.3897/zookeys.150.2109
LANKESTERIANA
130
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
Barbosa-Silva, R. G., Bueno, M. L., Labiak, P., Coelho, M. A. N., Martinelli, G. & Forzza, R. (2020). The Pantepui in the
Brazilian Amazon: Vascular Flora of Serra Do Aracá, a Cradle of Diversity, Richness and Endemism. The Botanical
Review, 86, 359–375. doi: https://doi.org/10.1007/s12229-020-09235-x
Barbosa-Silva, R. G., Labiak, P. H., Gil, A. S. B., Goldenberg, R., Michelangeli, F. A., Martinelli, G., Coelho, M. A. N.,
Zappi, D. C. & Forzza, R. C. (2016). Over the hills and far away: new plant records for the Guayana Shield in Brazil.
Brittonia, 68, 397–408.
Barros, F. de., Vinhos, F., Rodrigues, V. T., Barberena, F. F. V. A., Fraga, C. N., Pessoa, E. M., Forster, W., Menini Neto,
L., Furtado, S. G., Nardy, C., Azevedo, C. O. & Guimarães, L. R. S. (2015). Orchidaceae in Lista de Espécies da Flora
do Brasil. Jardim Botânico do Rio de Janeiro. Retrieved from http://oradobrasil.jbrj.gov.br/jabot/oradobrasil/FB179
[Accessed on 23/02/2022].
Bebber, D. P., Carine, M. A., Wood, J. R. I., Wortley, A. H., Harris, D. J., Prance, G. T., Davidse, G., Paige, J., Pennington,
T. D., Robson, N. K. B. & Scotland, R. W. (2010). Herbaria are a major frontier for species discovery. Proceedings of
the National Academy of Sciences of the United States of America, 107(51), 22169–22171.
Berry, P. E. & Riina, R. (2005). Insights into the diversity of the Pantepui Flora and the biogeographic complexity of the
Guayana Shield. In: I, Friis. & H. Balslev (Eds.), Plant Diversity and Complexity Patterns: Local, Regional, and Global
Dimensions (pp. 145–167). Copenhagen: Biologiske Skrifter 55.
Bivand, R., Keitt, T. & Rowlingson, B. (2021). Rgdal: Bindings for the “Geospatial” Data Abstraction Library. Retrieved
from https://CRAN.R-project.org/package=rgdal [Accessed on 06/04/2022].
Brazil Flora Group. (2022). Brazilian Flora 2020 project - Projeto Flora do Brasil 2020. Version 393.328. Instituto de Pes-
quisas Jardim Botanico do Rio de Janeiro. Dataset/Checklist. doi:10.15468/1mtkaw
Cantuária, P. C., Medeiros, T. D. S., Sobrinho, M. S., Silva, R. B. L., Costa Neto, S. V., Araújo, C. B., Furtado, M. F. M.,
Krahl, A. H., Soares, A. C. S., Farias, A. L. F., Potyguara, P. M. B., Almeida, S. S. M. S. & Farias, J. E. S. (2021). Rique-
za Vegetal da Bacia Hidrográca do Igarapé da Fortaleza, AP-010, Amapá, Brasil. Research, Society and Development,
10(13), e503101321290. doi: 10.33448/rsd-v10i13.21290
Cantuária, P. C., Silva, J. B. F., Farias, J. E. S., Silva, D. A. S. & Medeiros, T. D.S. (2020). Uma nova espécie de Bulbophyllum
Thouars (Orchidaceae Juss.) para a Amazônia Brasileira. Biota Amazônia, 10(3), 36–38. doi: 10.18561/2179-5746/bio-
taamazonia. v10n3p36-38.
CRIA (Centro de Referência e Informação Ambiental). (2022). speciesLink - simple search. Retrieved from https://species-
link.net/ [Accessed on 23/02/2022].
Damasco, G., Daly, D. C., Vicentini, A. & Fine, P. V. A. (2019). Reestablishment of Protium cordatum (Burseraceae) based
on integrative taxonomy. Taxon, 68(2), 1–13. doi: 10.1002/tax.12022.
Dressler, R. L. (2001). Scaphyglottis. In: A. M. Pridgeon, P. J. Cribb, M. W. Chase & F. N. Rasmussen (Eds.), Genera Or-
chidacearum: 4: Epidendroideae (Part 1) (pp. 310–313). Oxford: Oxford University Press.
Dressler, R. L., Whitten, M. & Williams, N. H. (2004). Phylogenetic relationships of Scaphyglottis and related genera
(Laeliinae: Orchidaceae) based on nrDNA ITS sequence data. Brittonia, 56(1), 58–66. doi: 10.1663/0007-196X(2004)
056[0058:PROSAR]2.0.CO;2.
Dunnington, D. (2017). Prettymapr: Scale Bar, North Arrow, and Pretty Margins in R. Retrieved from https://CRAN.R-
project.org/package=prettymapr [Accessed on 06/04/2022].
Farroñay, F., Perdiz, R. O., Prata, E. M. B. & Vicentini, A. (2019). Notes on morphology and distribution of Acman-
thera (A. Juss.) Griseb. (Malpighiaceae), an endemic genus from Brazil. Phytotaxa, 415(4), 199–207. doi: https://doi.
org/10.11646/phytotaxa.415.4.4
Flores, A. S. & Rodrigues, R. S. (2017). First occurrence of Dioclea ruddiae Maxwell (Leguminosae) in Brazil. Hoehnea,
44(4), 607–610. doi: http://dx.doi.org/10.1590/2236-8906-30/2017
Funk, V. A., Berry, P., Alexander, S., Hollowell, T. H. & Kelloff, C. L. (2007). Checklist of the plants of the Guiana Shield
(Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana) (pp. 1–300). Washington: National
Museum of Natural History.
GBIF (2022). Global Biodiversity Information Facility. Retrieved from https://www.gbif.org [Access on 23/02/2022].
Govaerts, R., Dranseld, J., Zona, S., Hodel, D. R. & Henderson, A. (2021). World Checklist of Orchidaceae. Facilitated by
the Royal Botanic Gardens, Kew. Retrieved from http://wcsp.science.kew.org/ [Accessed on 26/10/2021].
Hijmans, R. J. (2022). Raster: Geographic Data Analysis and Modeling. Retrieved from https://CRAN.R-project.org/
package=raster [Accessed on 06/04/2022].
LANKESTERIANA 22(2). 2022. © Universidad de Costa Rica, 2022.
131
de ArAújo et al. — The discovery of Scaphyglottis punctulata in the Brazilian Amazonia
Holanda, A. S. S., Zartman, C. E., Hopkins, M. J. G., Valsko, J. J. & Krahl, A. H. (2015). First occurrence of Schistostemon
(Urb.) Cuatrec. (Humiriaceae) in states of Roraima and Pará, Brazil. Check List, 11(1), 1–3. doi: 10.15560/11.1.1520.
Hopkins, M. J. G. (2007). Modelling the known and unknown plant biodiversity of the Amazon Basin. Journal of Biogeog-
raphy, 34(8), 1400–1411. doi: 10.1111/j.1365-2699.2007.01737. x.
Hopkins, M. J. G. (2019) Are we close to knowing the plant diversity of the Amazon? Anais da Academia Brasileira de
Ciências, 91(3), e20190396. doi: 10.1590/0001-3765201920190396.
IUCN. (2012). IUCN red list categories and criteria: Version 3.1. Second edition. Gland, Switzerland and Cambridge, UK.
Pebesma, E. (2018). “Simple Features for R: Standardized Support for Spatial Vector Data.” The R Journal, 10(1), 439–46.
doi: 10.32614/RJ-2018-009.
Pessoa, E., de Barros, F. & Alves, M. (2013). Novelties in Orchidaceae from the Brazilian Amazon. Check List, 9(4),
823–825. doi: 10.15560/9.4.823.
Pessoa, E., de Barros, F. & Alves, M. (2015). Orchidaceae from Viruá National Park, Roraima, Brazilian Amazon. Phyto-
taxa, 192(2), 61–96. doi: 10.11646/phytotaxa.192.2.1.
Prata, E. M. B., Carvalho, R. & Vicentini, A. (2016). Pagamea spruceana (Rubiaceae, Gaertnereae), a new species from
ooded white-sand forests in the Upper Rio Negro region, Brazil. Phytotaxa, 269(3), 186–192. doi: 10.11646/phyto-
taxa.269.3.2.
R Core Team. (2021). R: A language and environment for statistical computing. R Foundation for Statistical Computing,
Vienna, Austria. Retrieved from https://www.R-project.org/ [Accessed on 06/04/2022].
Reichenbach, H. G. (1855). Symbolae Orchidaceae: Ponera punctulata. Bonplandia, 3(15–16), 220–221.
Riina, R., Berry, P. E., Huber, O. & Michelangeli, F. A. (2019). Vascular plants and Bryophytes. In: V. Rull, O. Huber, C.
Señaris & T. Vegas-Vilarrúbia (Eds.), Biodiversity of Pantepui: The Pristine Lost World of the Neotropical Guiana
Highlands (pp. 121–147). Cambridge: Academic Press.
Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. (2012). NIH Image to ImageJ: 25 years of image analysis. Nature Meth-
ods, 9(7), 671–675. doi:10.1038/nmeth.2089
Schweinfurth, C. (1960). Orchidaceae, Orchids of Peru. Fieldiana, Botany, 30(3), 548–549.
South, A. (2011). “Rworldmap: A New R Package for Mapping Global Data.” The R Journal, 3(1), 35–43.
Szlachetko, D. L. & Kolanowska, M. (2014). Two new species of Scaphyglottis (Orchidaceae, Epidendroideae) from Co-
lombia. Polish Botanical Journal, 59(1), 1–5.
Thiers, B. [continuously updated]. (2022). Index Herbariorum: A global directory of public herbaria and associated staff.
New York Botanical Garden’s Virtual Herbarium. Retrieved from http://sweetgum.nybg.org/science/ih/ [Accessed on
10/03/2022].
Vásquez C., R. & Ibisch, P. L. (2004). Orquídeas de Bolivia: Diversidad y estado de conservación (pp. 317–330). Santa
Cruz de la Sierra: Editorial FAN.
LANKESTERIANA