Menu
Brazil
home Table of contents navigate_before previous current next navigate_next
home Table of contents
Original Article • Acta Bot. Bras. 38 • 2024 • https://doi.org/10.1590/1677-941X-ABB-2024-0011 linkcopy

Open-access Orchidaceae in the state of Pará, Brazilian Amazon: an updated checklist reveals underestimated species richness

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    Abstract

    We present an updated list of Orchidaceae species and their distribution data in the mesoregions of the state of Pará, Brazil, based on a systematic review of the literature and analysis of herbarium voucher specimens (and their labels) to confirm the occurrences of the species in the state. Our list has 434 species belonging to 98 genera; 209 species are endemic to the Amazon and 44 are restricted to the state of Pará. We also present 45 new records for the state. Catasetum, Epidendrum, Habenaria, and Maxillaria are the genera with the greatest species richness. Orchids were collected in all mesoregions of the state. Still, collections are absent in 21 of the 144 municipalities of the state, indicating the need for surveys and floristic assessments to understand the factors that influence the distribution of the species at the regional level and assist in conservation actions in the state.

    Keywords:
    Brazil; endemisms; flora; list of species; new occurrences; orchids

    Introduction

    Orchidaceae is one of the most diverse botanical families, with 31,372 species belonging to 740 genera, and has a cosmopolitan distribution, occurring mainly in the tropics (POWO, 2024). Among angiosperms, it is one of the most specialized families, with flowers exhibiting complex color patterns and unique shapes that vary according to different reproductive strategies (Fay & Chase, 2009). These characteristics confer high ornamental value, which is why orchids have been cultivated throughout the world for a long time in history and significantly contributed to the ornamental plants market (Hinsley et al., 2018; Tiwari et al., 2024).

    Orchids also stand out in the food industry, mainly due to the extraction of vanilla from fruits of the genus Vanilla Plum. (Baqueiro-Peña & Guerrero-Beltrán, 2017). Several secondary metabolites and biologically active compounds extracted from orchids have antioxidant, contraceptive, and antifungal effects, promising for pharmaceutical purposes (Asseleih et al., 2015; Gantait et al., 2021; Lipińska, 2023). Flowers, pseudobulbs, tubers, roots, and fruits of orchids also constitute relevant non-timber forest products (Hinsley et al., 2018; Anghelescu et al., 2020), however often illegally extracted from individuals in native vegetation remnants (Cruz-Garcia et al., 2015; Fay, 2015; Hinsley et al., 2018; Emeterio-Lara et al., 2021). Furthermore, orchids are ecologically important in the most varied ecosystems, establishing diverse interactions with other plants (e.g. epiphytism), birds and insects (e.g. pollination and dispersal of individuals), fungi and bacteria (symbiotic associations) (Hinsley et al., 2018; Castillo-Pérez et al., 2019; Houlihan et al., 2020; Tubarão & Barberena, 2022).

    Almost 9% of orchid species occur in Brazil and 2,526 are native to the country. The greatest diversity is observed in the country’s tropical forests. Orchidaceae is the largest family in terms of species richness in the Atlantic Forest (1,379 native species) and the second largest in the Amazon (786 native species) (Flora e Funga do Brasil, 2024). In the Brazilian Amazon, the orchid flora is threatened by the loss of habitats caused by increasing deforestation, especially in the state of Pará. A total of 35.2% of native Amazon forest wiped out in Brazil between 1985 and 2021 corresponded to areas deforested in Pará (Gomes et al., 2019; Projeto Mapbiomas, 2022; Silva et al., 2023) and in 2021, the largest rate of deforestation in the country was found in this state (Projeto MapBiomas, 2022). This scenario is thought to impact the climate, leading to, for example, increased temperatures, frequency and intensity of droughts, and reduced local rainfall rates (Silva et al., 2023). In turn, these environmental changes are likely to affect the structure, composition, phenology, pollination, and growth behavior of orchids (Barman & Devadas, 2013; Kumar & Rawat, 2022).

    The first list of orchid species for the state of Pará was presented by Pabst & Dungs (1975), with 77 genera and 249 species. However, research on the state’s orchid flora was only intensified in the 1990s, through floristic-taxonomic studies carried out in some areas, such as, for example, Serra dos Carajás (Silveira et al., 1995; Koch et al., 2018), Serra das Andorinhas (Atzingen et al., 1996), Ilha do Combu (Cardoso et al., 1995; Medeiros & Jardim 2011; Medeiros et al., 2013), Floresta Nacional de Caxiuanã (Koch et al., 2014), and in the municipalities of Abaetetuba (Afonso et al., 2016; Ferreira Filho et al., 2021), Igarapé-Miri (Ferreira Filho & Barberena, 2022) and Capitão Poço (Costa & Barberena, 2023). Recently, new species have been described (e.g., Eltroplectris guimaraesii A.E.S.Rocha & Afonso, Maxillaria luizotavioi Engels & E.C.Smidt, Mormodes ivanaluciae P.C.Cantuária, T.D.S.Medeiros & J.B.F.Silva, Palmorchis triquilhada Ferreira Filho & Barberena and, Vanilla labellopapillata A.K.Koch, Fraga, J.U.Santos & Ilk.-Borg.; Koch et al., 2013; Cantuária et al., 2019; Ferreira Filho & Barberena, 2020; Rocha & Afonso, 2022; Engels & Smidt, 2023), and other species have been recorded for the first time in the state (e.g., Stelis ciliaris Lindl., Uleiorchis ulei (Cogn.) Handro and Vanilla hartii Rolfe; Ferreira et al., 2020; Ferreira Filho & Barberena, 2022).

    In the meantime, other lists of orchid species for Pará have been published. Ilkiu-Borges & Cardoso (1996) documented the occurrence of 90 genera and 383 species, while Barros et al. (2010) listed 106 genera and 405 species. These numbers were later updated by BFG (2015), which recorded 112 genera and 431 species, and Flora e Funga do Brasil (2024), which reported a reduction of around 15% in the number of species. Therefore, 374 species of orchids belonging to 95 genera have been recognized for the state until now (Flora e Funga do Brasil, 2024).

    However, none of the above studies associated every species listed for the state with herbarium materials (vouchers). Furthermore, Pabst & Dungs (1975), Barros et al. (2010), BFG (2015) and Flora e Funga do Brasil (2024) consist of inventories conducted at a national level, allowing us to assume that the botanical collections deposited in herbaria in Pará were not thoroughly analyzed. In turn, although Ilkiu-Borges & Cardoso (1996) had a regional approach, their study was restricted to a list of species, without additional comments. Thus, considering the need to systematize the growing volume of species occurrence data and the numerical inconsistencies in previous lists, the objective of this study was to present an updated list of native Orchidaceae species occurring in Pará along with a diagnosis of specimens in the state and collection gaps.

    Material and Methods

    Study area

    The state of Pará is located between 2º N - 10º S and 46° - 59° W, in the Amazon, Northern Brazil, and occupies 1,245,871 km² (an area more than twice the size of Ukraine, the second largest country in Europe) (Zappi, 2020; IBGE, 2022a). Politically, the federative unit is divided into six mesoregions with different historical-cultural characteristics: i - Lower Amazonas (15 municipalities, total area of 345,436 km²); ii - Marajó (16 municipalities, 104,139 km²); iii - Metropolitan region of Belém (11 municipalities, 6,890 km²); iv - Northeast Pará (49 municipalities, 83,031 km²); v - Southeast Pará (39 municipalities, 297,651 km²), and vi - Southwest Pará (14 municipalities, 415,788 km²) (IBGE 2022a).

    The Metropolitan Region of Belém, Northeast Pará, and Southeast Pará mesoregions are occupied by more than 1.8 million inhabitants each (IBGE, 2022a). The large population size of these mesoregions is associated with their strategic location close to the state capital (Belém) and the historical context of the National Integration Program and implementation of the Belém-Brasília highway (Cordeiro et al., 2017). On the other hand, the Lower Amazonas, Southwest Pará, and Marajó mesoregions are comparatively less populated and have a greater portion of their territory within protected areas (Cardoso & Souza Jr, 2020; IBGE, 2022a).

    The state’s climate is classified as Tropical Equatorial Hot Humid, according to the Köppen classification (types Af, Am, and Aw), with annual precipitation ranging from 1,900 to 3,000 mm and an average annual temperature of 25-27 ºC (Alvares et al., 2013). The vegetation is heterogeneous, composed of the following types: Dense Ombrophylous Forest (terra-firme forest), Open Ombrophylous Forest (vine forest), Semideciduous Forest (dry forest), Igapó Forest, Várzea Forest, and Campestre Vegetation (including Amazonian savannah, Campinarana, Campo rupestre over canga, and Amazonian restinga) (Zappi, 2020; IBGE, 2022b).

    Data collection

    Data from the labels of exsiccates of Orchidaceae materials collected in the state of Pará were obtained during visits to herbaria in person (FCCM, HCP, HF, IAN, and MG), through a listing request to the curators (HIFPA and HCJS), or through consultations of virtual herbarium databases available on the SpeciesLink (2024), Reflora (2024) and Jabot (2024), allowing the collection of information from specimens deposited in 55 national and international repositories (ALCB, AMES, AMO, BHCB, BM, CAS, CEN, CNMT, EAC, EAFM, EAN, ESA, HAMAB, HATM, HB, HBRA, HEPH, HERBAM, HPL, HRB, HSTM, HUAM, HUEFS, HVASF, IAC, ICN, INPA, IPA, K, LBMBP, MAC, MAR, MBM, MBML, MFS, MICH, MIRR, MO, NHM, NY, P, R, RB, RFA, S, SOF, SP, SPF, TANG, UB, UEC, UFMT, UFP, UPCB, and US; acronyms according to Thiers, 2024).

    A systematic literature review using the Google Scholar, Scopus, and Web of Science databases was carried out to identify floristic-taxonomic studies with vouchers of Orchidaceae species collected in the state of Pará. We used the search terms “Orchidaceae/orchid” and “Pará State/Estado do Pará” appearing in the title, abstract, and keywords sections (see Mengist et al., 2020). Herbaria containing testimonial materials of the species found in the searches were contacted to confirm their occurrence.

    The compiled data was subjected to rigorous analysis and refinement, mainly regarding the identification of materials (especially those with conflicting identifications), accepted names for the species, origin of collections, and information on collectors. For specimens without coordinates of the collection site, coordinates were assigned based on records of the locality or municipality (Magdalena et al., 2018) using the tool GeoLoc (http://splink.cria.org.br/geoloc). The accepted names of the species follow Flora e Funga do Brasil (2024) and International Plant Names Index (IPNI, 2024), except in specific cases of nomenclatural updates (e.g. Barberena, 2021; Smidt et al., 2021; Camelo-Júnior et al., 2022; Sambin & Aucourd, 2023) and species not previously reported for the Brazilian flora, for which we adopted Plants of the World Online (POWO, 2024). Geographic distribution information was obtained from Flora e Funga do Brasil (2024) and POWO (2024). Information about life forms and substrate (concepts adopted according to the Flora e Funga do Brasil 2024) was obtained from the specimen labels.

    In total, 7,178 Orchidaceae exsiccates registered as originating from the state of Pará were initially analyzed. Of this total, the duplicates (13.8%; 990 specimens), specimens that could not be identified at the specific level (11.2%; 804 specimens), and specimens with conflicting occurrence data (2.4%; 161 specimens) were removed, resulting in a final sample of 5,223 exsiccates (72.6%). A voucher was selected for each species, with priority being given to specimens analyzed in person, collected in the most recent years, identified by experts in Orchidaceae, and/or with digital images available in botanical databases (such as Jabot, REFLORA, and SpeciesLink).

    To assess the influence of the proximity to access routes (land or rivers) on the distribution of specimen records, we calculated the distances from the records to the nearest route using QGis v. 3.22. We used a random sampling model to generate a null distribution of distances, which was then compared with the observed records. The Mann-Whitney test, performed with 1000 repetitions, was used to compare the observed distances with the null model using the `dplyr`, `geosphere`, and `DescTools` R packages (Signorell et al., 2017; Hijmans et al., 2017; Wickham et al., 2017; R Core Team, 2024).

    Results

    There are 98 genera and 434 species of Orchidaceae in the state of Pará. The most representative genera are Catasetum Rich. ex Kunth (35 spp. and a natural hybrid), Habenaria Willd. (30 spp.), Maxillaria Ruiz & Pav. (30 spp.), and Epidendrum L. (28 spp.). Thirty-two genera are represented by only one species (Tab. 1).

    Thumbnail
    Table 1.
    List of Orchidaceae species occurring in the state of Pará, Brazilian Amazon. (1) = Species with only one record in the state; (2) = Endemic to the Amazon; (3) = Restricted to the state of Pará; (4) = New record for the state of Pará.

    Of the total species, 209 species are endemic to the Amazon, 44 species are restricted to the state of Pará, and 45 species are new records for the state. Among the new records for Pará, more than half (25 spp.) are also endemic to the Amazon. Of the 434 species of orchids occurring in Pará, 133 are documented with ten or more specimens from Pará deposited in herbaria. Among the species with the highest number of collection records, we highlight Catasetum macrocarpum Rich. ex Kunth, Epidendrum nocturnum Jacq., Maxillaria uncata Lindl., Polystachya concreta (Jacq.) Garay & Sweet, Scaphyglottis stellata Lodd. ex Lindl. and, Sobralia liliastrum Salzm. ex Lindl., with more than 100 specimens each. Eighty-four species were collected only once in the state.

    Regarding the life forms, 96.3% of the species are classified as herbaceous and 3.7% as climbers (represented by 16 species of the genus Vanilla). With respect to the substrates colonized, some species were restricted to only one substrate, while others occurred on two or three substrates: exclusively epiphytic species predominated (73.3%) followed by exclusively terrestrial (16.8%), hemi-epiphytic (3.7%), rupicolous (1.8%), aquatic (0.2%), epiphytic/rupicolous (1.8%), epiphytic/ terrestrial (0.7%), rupicolous/terrestrial (1.2%), and epiphytic/rupicolous/terrestrial (0.5%).

    Most species are native (432 spp.) (Fig. 1 and 2) and two are naturalized [(Eulophia alta (L.) Fawc. & Rendle and Oeceoclades maculata (Lindl.) Lindl.)]. Seven species not included in our list, but which have been registered in the state, are cultivated: Arundina graminifolia (D.Don) Hochr., Bulbophyllum medusae (Lindl.) Rchb.f., Dendrobium anosmum Lindl., Maxillaria acutifolia Lindl., Papilionanthe teres (Roxb.) Schltr., Renanthera coccinea Lour., and Renanthera storiei Rchb.f.

    Figure 1.
    Species of Orchidaceae from Pará, Brazil. a- Anathallis brevipes (H.Focke) Pridgeon & M.W.Chase; b- Aspasia variegata Lindl.; c - Brassavola martiana Lindl.; d - Brassia caudata Lindl.; e, f - Campylocentrum fasciola (Lindl.) Cogn.; g - Campylocentrum micranthum (Lindl.) Rolfe; h - Catasetum albovirens Barb.Rodr.; i - Catasetum galeritum Rchb.f.; j - Catasetum macrocarpum Rich. ex Kunth; k - Catasetum mojuense A.T.Oliveira & J.B.F.Silva; l - Catasetum roseoalbum (Hook.) Lindl.; m - Dichaea picta Rchb.f.; n - Dimerandra emarginata (G.Mey.) Hoehne; o - Epidendrum carpophorum Barb.Rodr.; p - Epidendrum purpurascens Focke; q - Epidendrum strobiliferum Rchb.f.; r - Epidendrum viviparum Lindl.; s - Erycina pusilla (L.) N.H.Williams & M.W.Chase; and t - Laelia gloriosa (Rchb.f.) L.O.Williams. Photographs by Ana Laura da Silva Luz (a, b, d, g, k, n, q, t); Deivid Lucas de Lima da Costa (f, i); Felipe Fajardo Villela Antolin Barberena (c, e, j, o, r); Glenda Santa Brígida Quadros (p); Mateus Felipe dos Anjos Souza (s); and Sariane Moraes Bentes (h, l, m).

    Figure 2.
    Species of Orchidaceae from Pará, Brazil. a - Macradenia paraensis Barb.Rodr.; b - Maxillaria brasiliensis Brieger & Illg.; c - Maxillaria lutescens Scheidw.; d - Maxillaria subrepens (Rolfe) Schuit. & M.W.Chase; e - Maxillaria uncata Lindl.; f - Notylia aromatica Baker ex Lindl.; g - Notylia peruviana (Schltr.) C.Schweinf.; h - Notylia yauaperyensis Barb.Rodr.; i - Oeceoclades maculata (Lindl.) Lindl.; j - Ornithocephalus gladiatus Hook.; k - Peristeria cerina Lindl.; l - Polystachya caracasana Rchb.f.; m - Polystachya concreta (Jacq.) Garay & Sweet; n - Rodriguezia lanceolata Ruiz & Pav.; o - Scaphyglottis prolifera (Sw.) Cogn.; p - Sobralia macrophylla Rchb.f.; q - Stelis papaquerensis Rchb.f.; r - Trichocentrum cepula (Hoffmanns.) J.M.H.Shaw; s - Trichosalpinx orbicularis (Lindl.) Luer; and t - Vanilla phaeantha Rchb.f. Photographs by Ana Laura da Silva Luz (a, b, c, d, e, h, j, k, o, p, q); Antônio Augusto C. da Silva (g); Felipe Fajardo Villela Antolin Barberena (f, i, l, m, r, s); Glenda Santa Brígida Quadros (t); and Sariane Moraes Bentes (n).

    A total of 134 species mentioned in previous lists as occurring in Pará were excluded in the present study for two reasons: (1) some specimens had been mistakenly identified and did not correspond to the species indicated in the exsiccate label, (2) some materials listed in the literature or mentioned in electronic databases were not found when sought in person in herbaria, as they were possibly lost or were not even deposited in herbaria (Tab. S1 Table S1. Species previously reported for the state of Pará, Brazilian Amazon, and excluded in the present list with respective reasons for non-inclusion. ). Given this, we highlight the possible occurrence in the state of Pará of the following species whose specimen(s) were not located during visits to the herbaria: Anathallis sertularioides (Sw.) Pridgeon & M.W.Chase, Bulbophyllum bracteolatum Lindl., Caluera vulpina Dodson & Determann, Cattleya nobilior Rchb.f., Coryanthes cataniapoensis G.A.Romero & Carnevali, Coryanthes speciosa Hook. var. speciosa, Epidendrum nuriense Carnevali & Hagsater, Macroclinium bragae Campacci & J.B.F.Silva, Macroclinium subroseum Campacci & J.B.F.Silva, Specklinia tribuloides (Sw.) Pridgeon & M.W.Chase, and Trichosalpinx quitensis (Rchb.f.) Luer.

    Orchid collections in the state began in 1849 with the record of Erycina pusilla (L.) N.H.Williams & M.W.Chase by naturalist botanist Richard Spruce. Until 1950, other naturalists such as Jacques Huber, Adolpho Ducke, and João Murça Pires stood out as the main collectors in Pará. From 1976 onwards, there was a significant increase in the number of collectors and collections, which remains on an increasing trend to this day (Tab. S2 Table S2. Number of collectors and collections of Orchidaceae specimens, by period, in the state of Pará, Brazil. ).

    Collections are distributed across all mesoregions of the state, but disproportionately (Fig. 3-4). Most collections come from the mesoregions Lower Amazonas, Northeast, Southeast, and Southwest Pará (more than 660 collections in each), mainly from the municipalities of Parauapebas (with 516 collections), Marabá (390), Abaetetuba (305), and Vitória do Xingu (293) (Fig. 3). In the Metropolitan Region of Belém and Marajó, there were fewer collections (465 and 256 collections, respectively), with 11 of the 27 municipalities presenting less than 10 collections each. Furthermore, no collections of Orchidaceae were found in 21 municipalities, namely: Curuá, Placas (Lower Amazonas); Bonito, Mãe do Rio, Nova Esperança do Piriá, Nova Timboteua, Oeiras do Pará, Primavera, São Domingos do Capim, São João da Ponta, Tomé-Açu (Northeast Pará); Brejo Grande do Araguaia, Dom Eliseu, Palestina do Pará, Paragominas, Piçarra, Rio Maria, Sapucaia, Ulianópolis (Southeast Pará); Bagre, and Santa Cruz do Ariri (Marajó). The Mann-Whitney test (p-value < 0.05) indicated a bias in the collection records relative to access routes (roads and rivers). Approximately 9% and 42% of the species occurred exclusively inside and outside protected areas, respectively.

    Figure 3.
    Distribution of records of Orchidaceae specimens in the state of Pará, Brazilian Amazon. Mesoregions: Lower Amazonas (LA), Metropolitan Region of Belém (MB), Marajó (M), Northeast Pará (NP), Southeast Pará (SP), and Southwest Pará (SWP).

    Figure 4.
    Heatmap of Orchidaceae specimen records in the state of Pará, Brazilian Amazon. The colors of the circles represent the density of records. Mesoregions: Lower Amazonas (LA), Metropolitan Region of Belém (MB), Marajó (M), Northeast Pará (NP), Southeast Pará (SP), and Southwest Pará (SWP).

    Discussion

    The richness of orchids in Pará corresponds to around 17% of the family's richness in Brazil. However, this study showed that the richness of Orchidaceae in the state of Pará was underestimated, as we recognized an increase of almost 16% in the number of species, going from 374 (Flora e Funga do Brasil, 2024) to 434 species. Although the majority of the species (389) were registered in previous checklists for Pará (e.g. Ilkiu-Borges & Cardoso, 1996; Barros, 2010; BFG, 2015; Flora e Funga do Brasil, 2024), 45 species constitute new occurrence records for the state. These numbers are significant and highlight the importance of frequent and careful updating of regional floristic lists.

    Regarding the lists of Orchidaceae from the state of Pará provided by Ilkiu-Borges & Cardoso (1996), Barros et al. (2010), BFG (2015), and Flora e Funga do Brasil (2024), we detected notable numerical inconsistencies, as we were not able to confirm the occurrence of, respectively, 55, 83, 84, and 53 of the species cited in these studies due to the absence of testimonial materials (vouchers) in the databases consulted. It was, therefore, not possible to confirm the occurrence of a total of 87 species in the state (see Table S1 Table S1. Species previously reported for the state of Pará, Brazilian Amazon, and excluded in the present list with respective reasons for non-inclusion. - list of excluded species).

    About 11% of the Orchidaceae specimens from Pará could not be identified at specific level because they were sterile, with only fruits, or damaged, and less frequently due to the impossibility of accessing these materials (e.g., materials loaned to other researchers and temporary suspension of in-person consultations to herbarium collections) and due to the overload of the staff of the herbaria to carry out the automation and digitization of collection images (Carranza-Rojas et al., 2018). For orchids, including those from Pará, these limitations may result in additional gaps in taxonomic studies, mainly because they hinder the recognition of some species or natural hybrids, or new occurrence records.

    Regarding the life forms and substrates, the majority of species were epiphytes, which calls attention to their ability to explore vertical strata in trees and successful colonization in epiphytic niches, mainly due to the presence of highly specialized roots, giving them the ability to explore varied microhabitats, such as the canopy, and increasing their chances of establishment and survival (Zhang et al., 2017).

    Among the Orchidaceae species that occur in the state of Pará, 84 have only one record and can be provisionally considered rare (Işik, 2011). The low frequency of records may be related to the lack of researchers dedicated to studying the local flora or to the difficulties of collecting in more remote or difficult-to-access locations in the state (Fig. 4) or strata in the forest to reach epiphytic species (Quaresma et al., 2022; Luz et al., 2023), structural changes in habitats (Trudgill, 2022), or even the taxonomic complexity of some genera and the impossibility of analyzing sterile materials, making the collection and recognition of species difficult (Hinsley et al., 2018). Furthermore, among the species rarely collected are those recently described in the state (e.g. Maxillaria luizotavioi, Mormodes ivanaluciae, and Palmorchis triquilhada) or species known only from the type material. Collections need to be conducted continuously to identify, confirm, or rediscover rare species (Barberena et al., 2020). Among the rare species, it is possible that Catasetum juruenenseHoehne, Catasetum rolfeanum Mansf. and Microchilus mystacinus (Rchb.f.) E.C.Smidt & M.W.Chase have been suppressed regionally because the areas where they were collected (originally forest or grassland formations) were converted into temporary crops, mining, or pasture lands (Mapbiomas, 2021; SpeciesLink, 2024). Habenaria, Epidendrum and Catasetum are the genera with the greatest species richness and the most collected in Pará, which can be associated with their wide distribution in the Neotropics (Batista et al., 2011; Milet-Pinheiro & Gerlach, 2017; Arista et al., 2023). The high specific richness of these three genera was also observed in other floristic surveys carried out in the Amazon (Klein & Piedade, 2019; Ferreira Filho & Barberena, 2022) and in other regions of Brazil (Barros et al., 2018; Pessoa et al., 2022). The greatest abundance of collections of E. nocturnum, S. liliastrum, P. concreta, and M. uncata may also be related to the wide geographic distribution of these species, which may be associated with ease of reproduction (diversification of pollinators) and adaptation in different phytophysiognomies (Kolanowska et al., 2020; Kolanowska et al., 2022; Arista et al., 2023).

    Collections were concentrated in municipalities with easier access (land or river) (Fig. 4), greater demographic density, or in areas close to higher education institutions, which had specialists in Orchidaceae who were active collectors, such as E.A.L Afonso, N.V. Atzingen, and R.L. Ferreira Filho, whose collections were mainly concentrated in the municipalities of Abaetetuba, Marabá, and Parauapebas. Higher collection effort is also related to surveys in conservation units, where 9% of the species occurred exclusively (e.g. Floresta Nacional de Carajás; Koch et al., 2018) and areas influenced by hydroelectric plants (Carneiro-Silva et al., 2015). However, we highlight the absence of collections in 21 municipalities in the state (Fig. 3, Fig. 4), which may be related to the distance from research centers or a lower proportion of preserved native forests resulting from high rates of deforestation for logging [as, for example, in the municipalities of Dom Eliseu, Nova Esperança do Piriá, Paragominas and Tomé-Açu (Cardoso & Souza Jr, 2020)] that cause a cascade effect, culminating in the loss of epiphytic orchids (Sáyago et al., 2013).

    More than 48% of the Orchidaceae species recorded in our study are endemic to the Amazon, including the majority of the new records for the state. However, only 50 of the 434 species have been assessed for risk of extinction in Brazil (CNCFLORA, 2024; Flora e Funga do Brasil, 2024), corroborating the low proportion of species evaluated at national level (7% of Orchidaceae species in Brazil; Brasil, 2022) and internationally (3% of Orchidaceae species in the world; Hinsley et al., 2018). It is also noteworthy that 200 orchid species endemic to the Amazon have not been assessed for risk of extinction in Brazil. These results indicate the need for new assessments of the conservation status of species, mainly of endemic and rare species in the Amazon, especially considering their occurrence, population dynamics, commercial demand (at national or state levels), and illegal extraction. Such assessments need to be addressed in future studies, aiming at the identification and conservation of species at a regional or broader level.

    Acknowledgments

    We thank Ana Kelly Koch, Anna Luiza Ilkiu-Borges, André Cardoso, Edlley Pessoa, Emerson Pansarin, João Batista da Silva, Miguel Oliveira and Tiago Vieira for their contributions to the identifications; Helena Joseane Souza of IAN for providing photos of requested materials; and the teams of FCCM, HCP, HF, IAN, and MG for their receptivity and assistance during the consultations in person.

    References

    • Afonso EAL, Koch AK, Costa JM. 2016. Flora preliminar de Orchidaceae no município de Abaetetuba, Pará, Brasil. Biota Amazônia 6: 107-118. doi: 10.18561/2179-5746/biotaamazonia.v6n1p107-118.
      » https://doi.org/10.18561/2179-5746/biotaamazonia.v6n1p107-118.
    • Alvares CA, Stape JL, Sentelhas PC, Gonçalves JLM, Sparovek G. 2013. Koppen’s climate classification map for Brazil. Meteorologische Zeitschrift 22: 711-728. doi: 10.1127/0941-2948/2013/0507.
      » https://doi.org/10.1127/0941-2948/2013/0507.
    • Anghelescu NEDG, Bygrave A, Georgescu ML, Petra AS, Toma F. 2020. A history of orchids: A history of discovery, lust and wealth. Horticulture 64: 519-530. doi: 10.13140/RG.2.2.31997.38880.
      » https://doi.org/10.13140/RG.2.2.31997.38880.
    • Arista JP, Hágsater E, Santiago E et al 2023. New and noteworthy species of the genus Epidendrum (Orchidaceae, Laeliinae) from the the Área de Conservación Privada La Pampa del Burro, Amazonas, Peru. PhytoKeys 227: 43-87. doi: 10.3897/phytokeys.227.101907.
      » https://doi.org/10.3897/phytokeys.227.101907.
    • Asseleih LMC, García RAM, Sageth JY. 2015. Ethnobotany, pharmacology and chemistry of medicinal orchids from Veracruz. Journal of Agricultural Science and Technology A 5: 745-754. doi: 10.17265/2161-6256/2015.09.006.
      » https://doi.org/10.17265/2161-6256/2015.09.006.
    • Atzingen N Von, Cardoso ALR, Ilkiu-Borges AL. 1996. Flora orquidológica na Serra das Andorinhas, São Geraldo do Araguaia - PA. Boletim do Museu Paraense Emílio Goeldi 12: 59-74.
    • Baqueiro-Peña I, Guerrero-Beltrán JA. 2017. Vanilla (Vanilla planifolia Andr.), its residues and other industrial by-products for recovering high value flavor molecules: A review. Journal of Applied Research on Medicinal and Aromatic Plants 6: 1-9. doi: 10.1016/j.jarmap.2016.10.003.
      » https://doi.org/10.1016/j.jarmap.2016.10.003.
    • Barberena FFVA, Costa DLL, Rocha Jr JAL. 2020. Re-discovery of Catasetum mojuense (Orchidaceae: Catasetinae), a poorly-known Amazonian species. Neotropical Biology and Conservation 15: 447-452. doi: 10.3897/neotropical.15.e54142.
      » https://doi.org/10.3897/neotropical.15.e54142.
    • Barberena FFVA. 2021. Taxonomic notes on Catasetum roseoalbum (Orchidaceae: Epidendroideae): Reaffirming Catasetum ciliatum as a synonym. Phytotaxa 529: 171-173. doi: 10.11646/phytotaxa.529.1.13.
      » https://doi.org/10.11646/phytotaxa.529.1.13.
    • Barman D, Devadas R. 2013. Climate change on orchid population and conservation strategies: A review. Journal of Crop and Weed 9: 1-12.
    • Barros F, Hall CF, Paiva Neto VB, Batista JAN. 2018. Check-list das Orchidaceae do estado de Mato Grosso do Sul, Brasil. Iheringia 73: 287-296. doi: 10.21826/2446-8231201873s287.
      » https://doi.org/10.21826/2446-8231201873s287.
    • Barros F, Vinhos F, Rodrigues VT, Barberena FFVA, Fraga CN. 2010. Orchidaceae. In: Forzza RC et al (org.). Catálogo de plantas e fungos do Brasil. Instituto de Pesquisas Rio de Janeiro, Jardim Botânico do Rio de Janeiro. vol. II, p. 1344-1426.
    • Batista JAN, Bianchetti LDB, González-Tamayo R, Figueroa XMC, Cribb PJ. 2011. A synopsis of new world Habenaria (Orchidaceae) I. Harvard Papers in Botany 16: 1-47. doi: 10.3100/025.016.0101.
      » https://doi.org/10.3100/025.016.0101.
    • BFG - The Brazil Flora Group. 2015. Growing knowledge: An overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113. doi: 10.1590/2175-7860201566411.
      » https://doi.org/10.1590/2175-7860201566411.
    • Brasil. Ministério do Meio Ambiente. 2022. Portaria n. 148, de 7 de junho de 2022. Brasília, DF, Oficial da União.
    • Camelo- Júnior AE, Ferreira AWC, Andrade IM et al 2022. Species delimitation in the Trichocentrum cepula (Oncidiinae, Orchidaceae) complex: A multidisciplinary approach. Systematics and Biodiversity 20: 1-18. doi: 10.1080/14772000.2022.2099478.
      » https://doi.org/10.1080/14772000.2022.2099478.
    • Cantuária PC, Medeiros TDS, Negrão MF, Silva JBF, Silva e Silva BM, Lima e Silva RB. 2019. Nova espécie de Mormodes Lindl. para a Amazônia Brasileira. Biota Amazônia 9: 40-42. doi: 10.18561/2179-5746/biotaamazonia.v9n2p40-42.
      » https://doi.org/10.18561/2179-5746/biotaamazonia.v9n2p40-42.
    • Cardoso ALR, Ilkiu-Borges AL, Suemitsu C. 1995. Flora orquidológica da Ilha do Combu, Acará - Pará. Boletim do Museu Paraense Emílio Goeldi 11: 231-238.
    • Cardoso D, Souza Jr C. 2020. Sistema de Monitoramento da Exploração Madeireira (Simex): Estado do Pará 2017-2018. Belém, Imazon.
    • Carneiro-Silva MQ, Koch AK, Viana PL, Ilkiu-Borges AL. 2015. Oncidiinae (Orchidaceae) on the great curve of the Xingu River, Pará state, Brazil. Brazilian Journal of Biology 75: S222-S237. doi: 10.1590/1519-6984.01014BM.
      » https://doi.org/10.1590/1519-6984.01014BM.
    • Carranza-Rojas J, Joly A, Goëau H, Mata-Montero E, Bonnet P. 2018. Automated Identification of Herbarium Specimens at Different Taxonomic Levels. In: Joly A, Vrochidis S, Karatzas K, Karppinen A, Bonnet P (eds.). Multimedia Tools and Applications for Environmental & Biodiversity Informatics. Springer, Cham.
    • Castillo-Pérez LJ, Martínez-Soto D, Maldonado-Miranda JJ et al 2019. The endemic orchids of Mexico: A review. Biologia 74: 1-13. doi: 10.2478/s11756-018-0147-x.
      » https://doi.org/10.2478/s11756-018-0147-x.
    • CNCFLORA. 2024. Centro Nacional de Conservação da Flora. https://cncflora.jbrj.gov.br/portal 13 Aug. 2024.
      » https://cncflora.jbrj.gov.br/portal
    • Cordeiro IMCC, Rangel-Vasconcelos LGT, Schwartz G, Oliveira FA. 2017. Nordeste Paraense: panorama geral e uso sustentável das florestas secundárias. Belém, Embrapa Amazônia Oriental.
    • Costa DLL, Barberena FFVA. 2023. Orchidaceae in Reserva São Geraldo Majela, Belém Endemism Center, Pará State, Brazilian Amazon. Hoehnea 50: e502023. doi: 10.1590/2236-8906e502023.
      » https://doi.org/10.1590/2236-8906e502023.
    • Cruz-Garcia G, Lagunez-Rivera L, Chavez-Angeles MG, Solano-Gomez R. 2015. The wild orchid trade in a Mexican local market: Diversity and economics. Economic Botany 69: 291-305. doi: 10.1007/s12231-015-9321-z.
      » https://doi.org/10.1007/s12231-015-9321-z.
    • Emeterio-Lara A, Garcia-Franco JG, Hernandez M, Toledo-Hernández V, Valencia Díaz S, Alejandro F-P. 2021. Is pseudobulb harvest a sustainable management strategy in wild orchid populations? An experiment with Laelia autumnalis Forest Ecology and Management 491: 119205. doi: 10.1016/j.foreco.2021.119205.
      » https://doi.org/10.1016/j.foreco.2021.119205.
    • Engels ME, Smidt EC. 2023. Maxillaria luizotavioi (Orchidaceae: Maxillariinae): A new species from the Oriental Brazilian Amazon. Nordic Journal of Botany 2023: e03815. doi: 10.1111/njb.03815.
      » https://doi.org/10.1111/njb.03815.
    • Fay MF, Chase MW. 2009. Orchid biology: From Linnaeus via Darwin to the 21st century. Annals of Botany 104: 359-364. doi: 10.1093/aob/mcp190.
      » https://doi.org/10.1093/aob/mcp190.
    • Fay MF. 2015. Undocumented trade in species of Orchidaceae: Examples from Asia, the eastern Mediterranean region and Africa. PC22 Inf 6. https://cites.org/sites/default/files/eng/com/pc/22/Inf/E-PC22-Inf-06.pdf 02 Jan. 2023.
      » https://cites.org/sites/default/files/eng/com/pc/22/Inf/E-PC22-Inf-06.pdf
    • Ferreira AWC, Franken EP, Pansarin ER. 2020. Confirmation of the presence of Vanilla hartii Rolfe (Orchidaceae, Vanilloideae) in Brazil. Check List 16: 951-956. doi: 0.15560/16.4.951.
      » https://doi.org/0.15560/16.4.951.
    • Ferreira Filho RL, Barberena FFVA. 2020. Palmorchis triquilhada sp. nov. (Orchidaceae; Neottieae) from the Brazilian Amazon. Nordic Journal of Botany 38: e02740. doi: 10.1111/njb.02740.
      » https://doi.org/10.1111/njb.02740.
    • Ferreira Filho RL, Barberena FFVA. 2022. Orchidaceae of the municipality of Igarapé-Miri, Pará, Brazilian Amazon. Rodriguésia 73: e01432021. doi: 10.1590/2175-7860202273103.
      » https://doi.org/10.1590/2175-7860202273103.
    • Ferreira Filho RL, Barberena FFVA, Costa JM. 2021. Orchidaceae in floodplains of the islands of Abaetetuba, Amazonian Brazil: A flora threatened by intensive management for açaí palm (Euterpe oleracea). Brittonia 73: 1-24. doi: 10.1007/s12228-020-09647-4.
      » https://doi.org/10.1007/s12228-020-09647-4.
    • Flora e Funga do Brasil. 2024. Jardim Botânico do Rio de Janeiro. https://floradobrasil.jbrj.gov.br/ 13 Aug. 2024.
      » https://floradobrasil.jbrj.gov.br/
    • Gantait S, Das A, Mitra M, Chen J. 2021. Secondary metabolites in orchids: Biosynthesis, medicinal uses, and biotechnology. South African Journal of Botany 139: 338-351. doi: 10.1016/j.sajb.2021.03.015.
      » https://doi.org/10.1016/j.sajb.2021.03.015.
    • Gomes VHF, Vieira ICG, Salomão RP, Steege H. 2019. Amazonian tree species threatened by deforestation and climate change. Nature Climate Change 9: 547-553. doi: 10.1038/s41558-019-0500-2.
      » https://doi.org/10.1038/s41558-019-0500-2.
    • Hinsley A, Boer H, Fay M et al 2018. A review of the trade in orchids and its implications for conservation. Botanical Journal of the Linnean Society 186: 435-455. doi: 10.1093/botlinnean/box083.
      » https://doi.org/10.1093/botlinnean/box083.
    • Hijmans RJ, Karney C, Williams E, Vennes C. 2017. Geosphere: Spherical Trigonometry. R package documentation. https://cran.r-project.org/web/packages/geosphere/geosphere.pdf 29 Jul. 2024.
      » https://cran.r-project.org/web/packages/geosphere/geosphere.pdf
    • Houlihan PR, Lucky A, Owen M, Emmel TC. 2020. Ants tend ghost orchids: Patrolling of Dendrophylax lindenii (Orchidaceae) by Crematogaster ashmeadi in Florida. Florida Entomologist 103: 369-375. doi: 10.1653/024.103.0309.
      » https://doi.org/10.1653/024.103.0309.
    • IBGE - Instituto Brasileiro de Geografia e Estatística. 2022a. Prévia da população calculada com base nos resultados do Censo Demográfico 2022 até 25 de dezembro de 2022. https://ftp.ibge.gov.br/Censos/Censo_Demografico_2022/Previa_da_Populacao/PA_POP2022.pdf 10 Jul. 2023.
      » https://ftp.ibge.gov.br/Censos/Censo_Demografico_2022/Previa_da_Populacao/PA_POP2022.pdf
    • IBGE - Instituto Brasileiro de Geografia e Estatística. 2022b. Mapa da vegetação brasileira. Banco de Informações Ambientais (BDiA - web). https://bdiaweb.ibge.gov.br/#/consulta/vegetacao 30 May 2023.
      » https://bdiaweb.ibge.gov.br/#/consulta/vegetacao
    • IPNI. 2024. International Plant Names Index. https://www.ipni.org/ 01 Mar. 2024.
      » https://www.ipni.org/
    • Ilkiu-Borges AL, Cardoso ALR. 1996. Flora orquidológica do estado do Pará. Boletim do Museu Paraense Emílio Goeldi 12: 183-205.
    • Işik K. 2011. Rare and endemic species: Why are they prone to extinction? Turkish Journal of Botany 35: 411-417. doi: 10.3906/bot-1012-90.
      » https://doi.org/10.3906/bot-1012-90.
    • Jabot. 2024. Sistema de herbários do Jardim Botânico do Rio de Janeiro, v. 3. https://rb.jbrj.gov.br/v3/consulta.php 12 Aug. 2024.
      » https://rb.jbrj.gov.br/v3/consulta.php
    • Klein VP, Piedade MTF. 2019. Orchidaceae occurring in white-sand ecosystems of the Uatumã sustainable development reserve in central Amazon. Phytotaxa 419: 113-148. doi: 10.11646/phytotaxa.419.2.1.
      » https://doi.org/10.11646/phytotaxa.419.2.1.
    • Koch AK, Fraga CN, Santos JUM, Ilkiu-Borges AL. 2013. Taxonomic notes on Vanilla (Orchidaceae) in the Brazilian Amazon, and the description of a new species. Systematic Botany 38: 975-981. doi: 10.1600/036364413X674706.
      » https://doi.org/10.1600/036364413X674706.
    • Koch AK, Miranda JC, Hall CF. 2018. Flora das cangas da Serra dos Carajás, Pará, Brasil: Orchidaceae. Rodriguésia 69: 165-188. doi: 10.1590/2175-7860201869115.
      » https://doi.org/10.1590/2175-7860201869115.
    • Koch AK, Santos JUM, Ilkiu-Borges AL. 2014. Synopsis of the holoepiphyte and hemiepiphyte Orchidaceae of Floresta Nacional de Caxiuanã, Pará State, Brazil. Hoehnea 41: 129-148. doi: 10.1590/S2236-89062014000100012.
      » https://doi.org/10.1590/S2236-89062014000100012.
    • Kolanowska M, Rewicz A, Baranow P. 2020. Ecological niche modeling of the pantropical orchid Polystachya concreta (Orchidaceae) and its response to climate change. Scientific Reports 10: 14801. doi: 10.1038/s41598-020-71732-1.
      » https://doi.org/10.1038/s41598-020-71732-1.
    • Kolanowska M, Tsiftsis S, Dudek M, Konowalik K, Baranow P. 2022. Niche conservatism and evolution of climatic tolerance in the Neotropical orchid genera Sobralia and Brasolia (Orchidaceae). Scientific Reports 12: 13936. doi: 10.1038/s41598-022-18218-4.
      » https://doi.org/10.1038/s41598-022-18218-4.
    • Kumar D, Rawat S. 2022. Modeling the effect of climate change on the distribution of threatened medicinal orchid Satyrium nepalense D. Don in India. Environmental Science and Pollution Research 29: 72431-72444. doi: 10.1007/s11356-022-20412-w.
      » https://doi.org/10.1007/s11356-022-20412-w.
    • Lipińska MM, Haliński LP, Gołębiowski M, Kowalkowska AK. 2023. Active compounds with medicinal potential found in Maxillariinae Benth. (Orchidaceae Juss.) representatives - a review. International Journal of Molecular Sciences 24: 739. doi: 10.3390/ijms24010739.
      » https://doi.org/10.3390/ijms24010739.
    • Luz ALS, Costa AAS, Moreira CR, Barberena FFVA. 2023. Vascular epiphytes in the Amazon: main gaps, limitations and perspectives for studies on the subject. Acta Botanica Brasilica 37: e20220311. doi: 10.1590/1677-941X-ABB-2022-0311.
      » https://doi.org/10.1590/1677-941X-ABB-2022-0311.
    • Magdalena UR, Silva LAE, Lima RO et al 2018. A new methodology for the retrieval and evaluation of geographic coordinates within databases of scientific plant collections. Applied Geography 96: 11-15. doi: 10.1016/j.apgeog.2018.05.002.
      » https://doi.org/10.1016/j.apgeog.2018.05.002.
    • Medeiros TDS, Jardim MAG. 2011. Distribuição vertical de orquídeas epífitas na Área de Proteção Ambiental (APA) Ilha do Combu, Belém, Pará, Brasil. Revista Brasileira de Biociências 9: 33-38.
    • Medeiros TDS, Jardim MAG, Quaresma AC. 2013. Composição Florística e Estrutura da Comunidade de Orquídeas Epífitas na APA Ilha do Combu, Pará, Brasil. Biota Amazônia 3: 23-33. doi: 10.18561/2179-5746/biotaamazonia.v3n3p23-33.
      » https://doi.org/10.18561/2179-5746/biotaamazonia.v3n3p23-33
    • Mengist W, Soromessa T, Legese G. 2020. Method for conducting systematic literature review and meta-analysis for environmental science research. MethodsX 7: 100777. doi: 10.1016/j.mex.2019.100777.
      » https://doi.org/10.1016/j.mex.2019.100777.
    • Milet-Pinheiro P, Gerlach G. 2017. Biology of the Neotropical orchid genus Catasetum: A historical review on floral scent chemistry and pollinators. Perspectives in Plant Ecology, Evolution and Systematics 27: 23-34. doi: 10.1016/j.ppees.2017.05.004.
      » https://doi.org/10.1016/j.ppees.2017.05.004.
    • Pabst GFJ & Dungs F. 1975. Orchidaceae Brasilienses. Vol. 1. Hildesheim, Brucke-Verlag Kurt Schmersow.
    • Pessoa E, Silva MJC, Oliveira MS, Silva Junior WR, Ferreira AWC. 2022. An updated checklist reveals strong incongruities with previous studies: Insights after revisiting a regional orchid list. Acta Botanica Brasilica 36: e2020abb0487. doi: 10.1590/0102-33062020abb0487.
      » https://doi.org/10.1590/0102-33062020abb0487.
    • POWO - Plants of the World Online. 2024. Orchidaceae. Facilitated by the Royal Botanic Gardens, Kew. https://www.plantsoftheworldonline.org/ 12 Aug. 2024.
      » https://www.plantsoftheworldonline.org/
    • Projeto MapBiomas. 2022. Relatório Anual de Desmatamento 2021. São Paulo, Brasil.
    • Quaresma A, Zartman CE, Piedade MTF et al 2022. The Amazon epiphyte network: A first glimpse into Continental-Scale patterns of Amazonian vascular epiphyte assemblages. Frontiers in Forests and Global Change 5: 828759. doi: 10.3389/ffgc.2022.828759.
      » https://doi.org/10.3389/ffgc.2022.828759.
    • R Core Team. 2024. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ 29 Jul. 2024.
      » https://www.R-project.org/
    • Reflora. 2024. Herbário Virtual Reflora. https://reflora.jbrj.gov.br/reflora/herbarioVirtual/ 12 Aug. 2024.
      » https://reflora.jbrj.gov.br/reflora/herbarioVirtual/
    • Rocha AES, Afonso EL. 2022. Eltroplectris guimaraesii (Spiranthinae, Orchidoideae, Orchidaceae), nova espécie para a Amazônia brasileira. Boletim do Museu Paraense Emílio Goeldi 17: 283-288. doi: 10.46357/bcnaturais.v17i2.212.
      » https://doi.org/10.46357/bcnaturais.v17i2.212.
    • Sambin A, Aucourd M. 2023. Anathallis & Lankesteriana (Orchidaceae, Pleurothallidinae) de Guyane. Révision taxinomique avec quatre nouvelles espèces et une nouvelle occurrence. Richardiana 7: 1-77.
    • Sáyago R, Lopezaraiza-Mikel M, Quesada M, Álvarez-Añorve MY, Cascante-Marín A, Bastida JM. 2013. Evaluating factors that predict the structure of a commensalistic epiphyte - phorophyte network. Proceedings of the Royal Society 280: 20122821. doi: 10.1098/rspb.2012.2821.
      » https://doi.org/10.1098/rspb.2012.2821.
    • Signorell A, Aho K, Alfons A et al 2017. DescTools: Tools for Descriptive Statistics. R package documentation. https://cran.r-project.org/web/packages/DescTools/DescTools.pdf 29 Jul. 2024.
      » https://cran.r-project.org/web/packages/DescTools/DescTools.pdf
    • Silva RM, Lopes AG, Santos CAG. 2023. Deforestation and fires in the Brazilian Amazon from 2001 to 2020: Impacts on rainfall variability and land surface temperature. Journal of Environmental Management 326: 116664. doi: 10.1016/j.jenvman.2022.116664.
      » https://doi.org/10.1016/j.jenvman.2022.116664.
    • Silveira EC, Cardoso ALR, Iilkiu-Borges AL, Atzingen N. 1995. Flora Orquidológica da Serra dos Carajás, Estado do Pará. Boletim do Museu Paraense Emilio Goeldi 11: 75-87.
    • Smidt EC, Salazar GA, Mauad AVSR et al 2021. An Indomalesian origin in the Miocene for the diphyletic New World jewel orchids (Goodyerinae, Orchidoideae): Molecular dating and biogeographic analyses document non-monophyly of the Neotropical genera. Botanical Journal of the Linnean Society 197: 322-349. doi: 10.1093/botlinnean/boab028.
      » https://doi.org/10.1093/botlinnean/boab028.
    • SpeciesLink. 2024. Rede specieslink. https://specieslink.net/search/ 12 Aug. 2024.
      » https://specieslink.net/search/
    • Thiers. 2024. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. https://sweetgum.nybg.rg/science/ih/ 12 Aug. 2024.
      » https://sweetgum.nybg.rg/science/ih/
    • Tiwari P, Sharma A, Bose SK, Park K-I. 2024. Advances in Orchid Biology: Biotechnological Achievements, Translational Success, and Commercial Outcomes. Horticulturae 10: 152. doi: 10.3390/horticulturae10020152.
      » https://doi.org/10.3390/horticulturae10020152.
    • Trudgill D. 2022. Orchids in Scotland: Changes in their occurrence since 1950. British & Irish Botany 4: 34-41. doi: 10.33928/bib.2022.04.034.
      » https://doi.org/10.33928/bib.2022.04.034.
    • Tubarão FN, Barberena FFVA. 2022. Psarocolius decumanus (Icteridae) acts as a seedling disperser of Gomesa recurva (Orchidaceae) in the Brazilian Atlantic Forest. Austral Ecology 47: 1136-1139. doi: 10.1111/aec.13194.
      » https://doi.org/10.1111/aec.13194
    • Wickham H. 2017. Tidy Data. Journal of Statistical Software 59: 1-23. doi: 10.18637/jss.v059.i10.
      » https://doi.org/10.18637/jss.v059.i10.
    • Zappi DC. 2020. Tipos de vegetação da Amazônia com ênfase no Pará. In: Imperatriz-Fonseca VL, Alves DA (orgs.). Abelhas sem ferrão do Pará: a partir das expedições científicas de João M. F. Camargo. Belém, Instituto Tecnológico Vale. p. 17-31.
    • Zhang GQ, Liu KW, Li Z et al 2017. The Apostasia genome and the evolution of orchids. Nature 549: 379-383. doi: 10.1038/nature23897.
      » https://doi.org/10.1038/nature23897.

    Supplementary Data

    Table S1. Species previously reported for the state of Pará, Brazilian Amazon, and excluded in the present list with respective reasons for non-inclusion. Table S2. Number of collectors and collections of Orchidaceae specimens, by period, in the state of Pará, Brazil.

    Edited by

    • Editor Chef:
      Thais Almeida
    • Associate Editor:
      Pedro Viana

    Publication Dates

    • Publication in this collection
      20 Dec 2024
    • Date of issue
      2024

    History

    • Received
      14 Jan 2024
    • Accepted
      14 Oct 2024
    Creative Common - by 4.0
    This is an open-access article distributed under the terms of the Creative Commons Attribution License
    vertical_align_top file_download show_chart
    location_on
    Sociedade Botânica do Brasil SCLN 307 - Bloco B - Sala 218 - Ed. Constrol Center Asa Norte CEP: 70746-520 Brasília/DF. - Alta Floresta - MT - Brazil
    E-mail: acta@botanica.org.br
    rss_feed Stay informed of issues for this journal through your RSS reader
    Accessibility / Report Error