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Synopsis of Orchidaceae from Fazenda Sete Irmãos: a fragment of Amazon Forest in northwestern Maranhão, Brazil

Abstract

This study provides an inventory of the Orchidaceae in one of the most threatened areas of the Brazilian Amazon, situated in the eastern portion of the Belém Endemism Center. A total of 51 species, distributed in 24 genera, are recorded. Epiphytes represent 86% of the species, the remaining are terricolous (6%), hemiepiphytic (6%) or myco-heterotrophic (2%). The most representative genera were Epidendrum (6 spp.) and Maxillaria (5 spp.). This study cites for the first time five genera (Coryanthes, Sarcoglottis, Stelis, Trichosalpinx and Wullschlaegelia) and ten species for the flora of Maranhão, of which four are new records to the Brazilian Northeast. An identification key, photographs of the new records, taxonomic and ecological comments, as well as geographic distribution are provided. The data reinforce the importance of the forest fragments of the Amazon Forest in Maranhão for the conservation of Orchidaceae which is constantly threatened by deforestation in the Belém Endemism Center.

Key words
Belém Endemism Center; epiphytes; inventory; taxonomy

Resumo

Este estudo apresenta um inventário das Orchidaceae em uma das áreas mais ameaçadas da Amazônia Brasileira, localizada na porção mais oriental do Centro de Endemismo Belém. São registradas 51 espécies distribuídas em 24 gêneros. Epífitas representam 86% das espécies, as demais são terrícolas (6%), hemiepífitas (6%) ou micoheterotrófica (2%). Os gêneros mais representativos foram Epidendrum (6 spp.) e Maxillaria (5 spp.). Este estudo registra pela primeira vez cinco gêneros (Coryanthes, Sarcoglottis, Stelis, Trichosalpinx e Wullschlaegelia) e dez espécies para a flora do Maranhão, dos quais, quatro também são novos registros para a Região Nordeste do Brasil. Uma chave de identificação, pranchas fotográficas dos novos registros e comentários taxonômicos, ecológicos e de distribuição geográfica são apresentados. Os dados evidenciam a importância dos fragmentos de Floresta Amazônica do Maranhão para a proteção das Orchidaceae, constantemente ameaçadas pelo desmatamento no Centro de Endemismo Belém.

Palavras-chave
Centro de Endemismo Belém; epífitas; inventário; taxonomia

Introduction

Orchidaceae Juss. is widely known as one of the most species-rich plant families (Dressler 1981Dressler RL (1981) The orchids: natural history and classification. Harvard University Press, Cambridge. 317p.; Chase et al. 2015Chase MW, Cameron KM, Freudestein JV, Pridgeon AM, Salazar G, Van Den Berg C & Schuiteman A (2015) An updated classification of Orchidaceae. Botanical Journal of the Linnean Society 177: 151-174.). It is especially diverse in the Neotropical region where Brazil stands out with about 220 genera and 2,500 species recorded, of which about 65% are endemic (BFG 2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113., 2018). In northeastern Brazil inventories of Orchidaceae species have been conducted in areas of the states of Bahia (Bastos & Van den Berg 2012Bastos CA & Van den Berg C (2012) Flora da Bahia: Catasetum (Orchidaceae). Sitientibus série Ciências Biológicas 12: 83-89.; Marinho & Azevedo 2014Marinho LC & Azevedo CO (2014) Orchidaceae na Reserva do Poço Escuro, Vitória da Conquista, Bahia, Brasil. Sitientibus série Ciências Biológicas 13: 1-14.; Vieira et al. 2014Vieira TL, Barros F & Roque N (2014) Orchidaceae no Município de Jacobina, estado da Bahia, Brasil. Hoehnea 41: 469-482.), Maranhão (Silva et al. 1999Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393.; Oliveira et al. 2021Oliveira MS, Ferreira AWC, Oliveira HC & Pessoa E (2021) Orchids in the central region of eastern Maranhão, Brazil. Rodriguésia: 72: 1-15.), Paraíba (Almeida et al. 2007Almeida A, Felix WJP, Andrade LA & Feli LP (2007) A família Orchidaceae em inselbergues da Paraíba, Nordeste do Brasil. Biociências 5: 753-755. ; Moreira et al. 2020Moreira LHL, Neto RLS & Barbosa MRV (2020) Flora da Mata do Buraquinho, João Pessoa, Paraíba: Orchidaceae. Rodriguésia 71: 1-11.), Pernambuco (Pessoa & Alves 2012Pessoa E & Alves M (2012) Flora da Usina São José, Igarassu, Pernambuco: Orchidaceae. Rodriguésia 62: 341-356. , 2014, 2015) and Sergipe (Pessoa & Alves 2011Pessoa E & Alves M (2011) Orchidaceae Juss. na Serra de Itabaiana, Sergipe, Brasil. Caatinga 24: 102-114.; Monteiro et al. 2012Monteiro SHN, Silva TC, Santos LAS, Nascimento-Júnior JE & Prata APN (2012) Survey of Orchidaceae from the state of Sergipe, Brazil. Biota Neotropica 12: 167-175.). However, knowledge of the eastern portion of the Amazon forest located in the state of Maranhão is fairly low.

Some recent studies have pointed out new records of Orchidaceae for the state (Ferreira et al. 2017Ferreira AWC, Oliveira MS, Silva EO, Campos DS, Pansarin ER & Guarçoni EAE (2017) Vanilla bahiana Hoehne and Vanilla pompona Schiede (Orchidaceae, Vanilloideae): two new records from Maranhão state, Brazil. Check List 13: 1131-1137. , 2019c; Gomes et al. 2021Gomes GS, Ferreira AWC, Silva MJC, Conceição GM & Pessoa EM (2021) Taxonomic study of Epidendrum L. (Laeliinae-Orchidaceae) in the state of Maranhão, Northeastern Brazil. Rodriguésia: 72: 1-13.; Oliveira et al. 2021Oliveira MS, Ferreira AWC, Oliveira HC & Pessoa E (2021) Orchids in the central region of eastern Maranhão, Brazil. Rodriguésia: 72: 1-15.), indicating that it is still in need of further taxonomic studies, yet it remains neglected by the majority of Brazilian botanists. Currently 115 species and 49 genera of the family are cited to Maranhão (BFG 2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113., 2018), however a large portion of its area remains fairly poorly collected, especially the northwest region where it includes part of the Belém Endemism Center, one of the most threatened portions of the Brazilian Amazon (Almeida & Vieira 2010Almeida AS & Vieira IGV (2010) Centro de Endemismo Belém: status da vegetação remanescente e desafios para a conservação da biodiversidade e restauração ecológica. Estudos Universitários 36: 95-111.).

Thus, this study aims to provide a synopsis of the Orchidaceae species from a fragment of Amazon Forest located in the northwest part of the state. In addition to the inventory, we provide an identification key, photographs of the new records, taxonomic and ecological comments, as well as the geographic distribution of each species. This study will contribute to expanding the knowledge of the orchid flora along the eastern border of the Amazon domain.

Material and Methods

The state of Maranhão, with an area of 331,983 km², presents vegetation with ecotonal features since it is located in a transition zone among the Amazon Forest, the Cerrado and the Caatinga (Muniz 2006Muniz FH (2006) Estudos da estrutura e distribuição da vegetação no Maranhão. In: Silva AC & Bringel JCM (orgs.). Projeto e ações em biologia e química. Vol. 1. EDUEMA, São Luís. Pp. 99-117.). The northwest portion of the state in dominated by ombrophilous and semi-deciduous forests, while savannahs and palm rich ecosystems locally called “mata dos cocais” are more common in the south and east portions (Abreu 1949Abreu SF (1949) O estado do Maranhão. Boletim Geográfico 7: 743-745.; Froés 1953Froés RL (1953) Limites meridionais e orientais da área de ocorrência da floresta amazônica em território brasileiro. Revista Brasileira de Geografia 15: 3-95.; Ribeiro 1971Ribeiro JR (1971) O Maranhão e seu revestimento florístico. Brasil Florestal 2: 9-20.; Muniz 2004Muniz FH (2004) A vegetação da região de transição entre a Amazônia e o Nordeste: diversidade e estrutura. In: Moura EG (ed.) Agro ambientes de transição entre o Trópico Úmido e o Semiárido do Brasil: atributos, alterações e uso na produção familiar. 2nd ed. Programa de Pós-graduação em Agroecologia/UEMA, São Luís. Pp. 53-69.; Almeida & Vieira 2010Almeida AS & Vieira IGV (2010) Centro de Endemismo Belém: status da vegetação remanescente e desafios para a conservação da biodiversidade e restauração ecológica. Estudos Universitários 36: 95-111.; Celentano et al. 2017Celentano D, Rousseau GX, Muniz FH, Varga, ID, Martinez C, Carneiro MS, Miranda MCV, Barros MNR, Freitas L, Narvaes IS, Adami M, Gomes AR, Rodrigues JC & Martins MB (2017) Towards zero deforestation and forest restoration in the Amazon region of Maranhão state, Brazil. Land Use Policy 68: 692-698. ; Silva et al. 2017Silva JMC, Leal IR & Tabarelli M (2017) Caatinga: the largest tropical dry forest region in South America. Springer International Publishing, Cham. 769p.; Silva-Moraes et al. 2019Silva-Moraes HG, Cordeiro I & Figueiredo N (2019) Flora and floristic affinities of the Cerrados of Maranhão state, Brazil. Edinburg Journal of Botany 76: 1-21.). The study area is in the northwest part of the state, in the municipality of Cândido Mendes, on a private property called Fazenda Sete Irmãos (01º51’37”S, 45º46’10”W) (Fig. 1). The area includes a fragment of about 7,000 ha of Amazon Forest (Almeida & Vieira 2010Almeida AS & Vieira IGV (2010) Centro de Endemismo Belém: status da vegetação remanescente e desafios para a conservação da biodiversidade e restauração ecológica. Estudos Universitários 36: 95-111.), considered one of the largest of the state (Koch & Araújo-Silva 2014Koch AK & Araújo-Silva LE (2014) Primeiro registro de Voyria tenella Guild. ex. Hook. (Gentianaceae) para o estado do Maranhão, Brasil. Biota Amazônica 4: 132-134. ; Celentano et al. 2017Celentano D, Rousseau GX, Muniz FH, Varga, ID, Martinez C, Carneiro MS, Miranda MCV, Barros MNR, Freitas L, Narvaes IS, Adami M, Gomes AR, Rodrigues JC & Martins MB (2017) Towards zero deforestation and forest restoration in the Amazon region of Maranhão state, Brazil. Land Use Policy 68: 692-698. ). According to Köppen the climate type is “Am” (Alvares et al. 2013Alvares CA, Stape JL, Sentelhas PC, Gonçalves JLM & Sparovek G (2013) Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift 22: 711-728. ) with mean temperatures between 26–27 °C and annual precipitation between 2,300–2,500 mm (NuGeo 2016NuGeo (2016) Bacias hidrográficas e climatologia no Maranhão. Universidade Estadual do Maranhão, São Luís. 165p. ).

Figure 1
Geographic position of Fazenda Sete Irmãos. Fieldwork points highlighted.

Field expeditions were conducted monthly between August 2017 and February 2020. The fertile specimens were processed following standard procedures (Fidalgo & Bononi 1984Fidalgo O & Bononi VLR (1984) Manual prático de coleta, herborização e preservação. Instituto de Botânica do estado de São Paulo, São Paulo. 62p.) and deposited at SLUI (Thiers, continuously updated). Sterile specimens were kept under cultivation, awaiting flowering. Taxonomic identification was based upon the specialized literature (Pabst & Dungs 1975Pabst GFJ & Dungs F (1975) Orchidaceae brasilienses. Vol. 1. Kurt Schmersow, Hildesheim. 408p., 1977Pabst GFJ & Dungs F (1977) Orchidaceae brasilienses. Vol. 2. Kurt Schmersow, Hildesheim. 418p.; Batista et al. 2008Batista JAN, Silva JBS & Bianchetti LB (2008) The genus Habenaria in the Brazilian Amazon. Revista Brasileira de Botânica 31: 105-134.; Pessoa & Alves 2011Pessoa E & Alves M (2011) Orchidaceae Juss. na Serra de Itabaiana, Sergipe, Brasil. Caatinga 24: 102-114., 2012, 2014Pessoa E & Alves M (2014) Orchidaceae em afloramentos rochosos do estado de Pernambuco, Brasil. Rodriguésia 65: 717-734., 2015; Pessoa et al. 2015Pessoa E, Barros F & Alves M (2015) Orchidaceae from Viruá National Park, Roraima, Brazilian Amazon. Phytotaxa 192: 61-96. ; Koch et al. 2018Koch AK, Miranda JC & Hall CF (2018) Flora of the cangas of the Serra dos Carajás, Pará, Brazil: Orchidaceae. Rodriguezia 69: 165-188.; Klein & Piedade 2019Klein VP & Piedade MTF (2019) Orchidaceae occurring in white-sand ecosystems of the Uatumã Sustainable Development Reserve in Central Amazon. Phytotaxa 419: 113-148.) in addition to an analysis of type specimens when available on-line. The circumscription of the genera follows the Flora do Brasil 2020 (continuously updated).

The morphological terminology follows Harris & Harris (2001)Harris J & Harris M (2001) Plant identification terminology - an illustrated glossary. 2nd ed. Spring Lake Publishing, Payson. 216p., the data on geographical distribution of the species follow Flora do Brasil 2020 (continuously updated) and Govaerts et al. (2020)Govaerts R, Pfahl J, Campacci MA, Holland-Baptista D, Tigges H, Shaw J, Cribb P, George A, Kreuz K & Wood J (2020) World checklist of Orchidaceae. The board of trustees of the Royal Botanic Gardens, Kew. Available at <http://apps.kew.org/wcsp/>. Access on 9 March 2020.
http://apps.kew.org/wcsp/...
. The map was produced using QGis® 2.18.12 Essen (QGIS Development Team 2020QGIS® Development Team (2020) QGIS Geographic Information System. Open Source Geospatial Foundation. Available at <https://QGIS.osgeo. org>. Access on 20 March 2020.
https://QGIS.osgeo. org...
) under SIRGAS 2000 datum.

Results and Discussion

Orchidaceae is represented in the study area by 51 species distributed in 24 genera. Epiphytes represent 86% of the species (34 spp.), the remaining are terricolous (6%, 3 spp.), hemiepiphytic (6%, 3 spp.), or myco-heterotrophic (2%, one sp.) (Tab. 1; Figs. 2-3). The predominance of epiphytes can be explained by the dense canopy observed in the area which does not allow much light to reach the soil. This result agrees with Dressler (1993)Dressler RL (1993) Phylogeny and classification of the orchid family. Dioscorides Press, Portland. 314p. who argued that at least 2/3 of the orchid species in the tropics are epiphytes. Epidendrum L. with six species (12%) and Maxillaria Ruiz & Pav. with five species (10%) are the most representative genera in the area. These genera have also been found to be representative in other studies conducted in the Amazon domain (Silveira et al. 1995Silveira EC, Cardoso ALR, Ilkiu-Borges AL & Atzingen N (1995) Flora Orquidológica da Serra dos Carajás, estado do Pará. Boletim do Museu Paraense Emílio Goeldi 11: 75-81. ; Pessoa et al. 2015Pessoa E, Barros F & Alves M (2015) Orchidaceae from Viruá National Park, Roraima, Brazilian Amazon. Phytotaxa 192: 61-96. ; Koch et al. 2018Koch AK, Miranda JC & Hall CF (2018) Flora of the cangas of the Serra dos Carajás, Pará, Brazil: Orchidaceae. Rodriguezia 69: 165-188.; Klein & Piedade 2019Klein VP & Piedade MTF (2019) Orchidaceae occurring in white-sand ecosystems of the Uatumã Sustainable Development Reserve in Central Amazon. Phytotaxa 419: 113-148.).

Table 1
Orchidaceae from Fazenda Sete Irmãos, Cândido Mendes, Maranhão. (Habit: E = Epiphyte; HE = Hemiepiphyte; T = Terricolous; MI = Mycoheterotrophic). ** = first record for Maranhão. ## = first registration for the Northeast region of Brazil. ++ = exsiccate to confirm the report by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393..

According to BFG (2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113., 2018BFG - The Brazil Flora Group (2018) Brazilian Flora 2020: innovation and collaboration to meet Target 1 of the Global Strategy for Plant Conservation (GSPC). Rodriguésia 69: 1513-1527.), five genera are cited for the first time for the Maranhão: Coryanthes Hook., Sarcoglottis C. Presl., Stelis Sw., Trichosalpinx Luer and Wullschlaegelia Rchb.f. Ten species (about 20%) are new records for the state: Campylocentrum pachyrrhizum (Rchb.f.) Rolfe, Coryanthes speciosa Hook., Lockhartia imbricata (Lam.) Hoehne, Notylia microchila Cogn., Ornithocephalus cujeticola Barb. Rodr., Sarcoglottis acaulis (Sm.) Schltr., Scaphyglottis prolifera (R.Br.) Cogn., Stelis paraensis Barb. Rodr., Trichosalpinx egleri Pabst and Wullschlaegelia calcarata Benth. Furthermore, four of the species were collected for the first time in northeastern Brazil: L. imbricata, S. paraensis, T. egleri and W. calcarata. These four species are restricted to the Amazon dominion (Flora do Brasil 2020, continuously updated), and the presence of W. calcarata is even more interesting due to the particular ecology of this species which is highly water-dependent.

Another five species cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. for Maranhão but without testimony vouchers, Campylocentrum micranthum Rolfe., Dichaea picta Rchb.f., Laelia gloriosa (cited as Schomburgkia gloriosa Rchb.f.), Oncidium baueri Lindl and Polystachya concreta (Jacq.) Garay & H.R.Sweet, have their occurrence currently not confirmed to the state in Flora do Brasil 2020 (continuously updated), yet they were collected by us. These do not represent new records, but confirmations of occurrence.

Among the species 30% are widespread in the Neotropics, while 47% are endemic to South America, and only 6% are restricted to Brazil (BFG 2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113., 2018; Govaerts et al. 2020Govaerts R, Pfahl J, Campacci MA, Holland-Baptista D, Tigges H, Shaw J, Cribb P, George A, Kreuz K & Wood J (2020) World checklist of Orchidaceae. The board of trustees of the Royal Botanic Gardens, Kew. Available at <http://apps.kew.org/wcsp/>. Access on 9 March 2020.
http://apps.kew.org/wcsp/...
). The majority of the species (about 75%) were collected along rivers or water courses, these areas work as corridors for epiphytes linking the Cerrado and the Amazon (Pabst & Dungs 1975Pabst GFJ & Dungs F (1975) Orchidaceae brasilienses. Vol. 1. Kurt Schmersow, Hildesheim. 408p.). In the anthropized portion of the Fazenda Sete Irmãos only three species were observed, Catasetum macrocarpum Rich. ex Kunth. and Vanilla palmarum Lindl. growing in Attalea speciosa Mart. (Arecaceae, locally called “Babaçu”) and Sacoila lanceolata (Aubl.) Garay a terricolous species.

The Fazenda Sete Irmãos is one of the last well-preserved localities of Amazon Forest in Maranhão, at least 3,000 ha are primary forest. Our results indicate it is one of the richest areas in Orchidaceae species in northeastern Brazil (Coelho & Amorim 2014Coelho MM & Amorim AM (2014) Floristic composition of the Montane Forest in the Almadina-Barro Preto axis, Southern Bahia, Brazil. Biota Neotropica 14: 1-41.; Pessoa & Alves 2015Pessoa E & Alves M (2015) Synopsis of Orchidaceae from Serra do Urubu: an area of montane forest, Pernambuco state, Brazil. Hoehnea 42: 109-133.) and it is fundamental for conservation of the Belém Endemism Center (Almeida & Vieira 2010Almeida AS & Vieira IGV (2010) Centro de Endemismo Belém: status da vegetação remanescente e desafios para a conservação da biodiversidade e restauração ecológica. Estudos Universitários 36: 95-111.; Celentano et al. 2017Celentano D, Rousseau GX, Muniz FH, Varga, ID, Martinez C, Carneiro MS, Miranda MCV, Barros MNR, Freitas L, Narvaes IS, Adami M, Gomes AR, Rodrigues JC & Martins MB (2017) Towards zero deforestation and forest restoration in the Amazon region of Maranhão state, Brazil. Land Use Policy 68: 692-698. ). The new records of species presented here are just part of several other studies recently undertaken in the Amazon portion of Maranhão that are also expanding distributions of taxa of other families (Guarçoni et al. 2018Guarçoni EAE, Bastian RE, Silva EO & Ferreira AWC (2018) First record of the rare Dyckia racemosa Baker (Bromeliaceae) in Maranhão state (northeastern Brazil), with an update on the species description, geographic distribution, and conservation status. Phytotaxa 349: 273-280., 2020Guarçoni EAE, Saraiva RVC & Ferraz TM (2020) Dyckia maranhensis (Bromeliaceae, Pitcairnioideae), a new species from the Cerrado of Maranhão, Northeastern Brazil. Systematic Botany 45: 47-52. ; Ferreira et al. 2019aFerreira AWC, Lima GP, Silva MJC & Almeida Junior EB (2019a) Expansion of the geographic distribution of Rapatea paludosa Aubl. (Rapateaceae) to the state of Maranhão, Northeastern Brazil Check List 15: 921-925., 2019bFerreira AWC, Oliveira MS, Silva Junior WR, Andrade IM, Coelho MAN, Oliveira HC & Mayo SJ (2019b) New occurrences of small aquatic duckweeds (Araceae, Lemnoideae) in Maranhão state, northeastern Brazil. Check List 15: 1153-1160. , 2019cFerreira AWC, Oliveira MS, Engels ME & Pessoa E (2019c) Found in Brazil again! Expanding the distribution of Maxillaria aureoglobula Christenson (Orchidaceae, Maxillariinae) and a key to the species of Maxillaria sect. Rufescens Christenson from Brazil. Check List 15: 1107-1112. ; Silva et al. 2016Silva EO, Guarçoni EAE, Ferreira AWC, Oliveira MS & Junior CNO (2016) First record of Passiflora pedata L. (Passifloraceae) from Maranhão state, northeastern Brazil. Check List 12: 1-4. ; Koch & Araújo Silva 2014; Scatigna et al. 2019Scatigna AV, Brandão CM, Colletta GD, Teles RM, Cavalcante KSB, Souza VC & Simões AO (2019) Dizygostemon riparius (Plantaginaceae, Gratioleae), a new species from Maranhão, northeastern Brazil. Willdenowia 49: 177-186. ; Silva-Júnior et al. 2020). We highlight the need for a stronger effort of studying this neglected and threatened region of Brazil.

Key for the orchid species from the Fazenda Sete Irmãos

  • 1. Myco-heterotrophic, achlorophyllous herbs...................51. Wullschlaegelia calcarata

  • 1’. Autotrophic, chlorophyllous herbs...................2

    • 2. Leafless plants; roots flat, green...................4. Campylocentrum pachyrrhizum

    • 2’. Leafy plants; roots cylindrical, grayish or white...................3

      • 3. Hemiepiphytes; stem voluble...................4

        • 4. Lip with a multi-ridged callus on the disc...................50. Vanilla pompona

        • 4’. Lip without a callus on the disc...................5

          • 5. Sepals and petals undulate at margin, green...................48. Vanilla mexicana

          • 5’. Sepals and petals flat at margin, yellow...................49. Vanilla palmarum

      • 3’. Epiphytes or terrestrial herbs; stem erect or pendulous, never voluble...................6

        • 6. Inflorescence terminal...................7

          • 7. Terrestrial herbs; leaves convolute...................8

            • 8. Leaves present during flowering; flowers greenish-white.........................................................40. Sarcoglottis acaulis

            • 8’. Leaves absent during flowering; flowers magenta...................39. Sacoila lanceolata

          • 7’. Epiphytes; leaves conduplicate or plicate...................9

            • 9. Leaves plicate...................44. Sobralia macrophylla

            • 9’. Leaves conduplicate...................10

              • 10. Pseudobulbs ovoid, clavate or fusiform...................11

                • 11. Flowers non-resupinate...................12

                  • 12. Lip 3-lobed; flowers greenish-yellow.........................................................36. Polystachya concreta

                  • 12’. Lip entire; flowers white with purple stripes.........................................................37. Prosthechea aemula

                • 11’. Flowers resupinate...................13

                  • 13. Peduncle at least four times longer than the rachis.........................................................18. Laelia gloriosa

                  • 13’. Peduncle as long as the rachis, to slightly longer or shorter.........................................................12. Epidendrum purpurascens

              • 10’. Pseudobulbs cylindrical or stem not swollen...................14

                • 14. Leaves 1 on each stem...................15

                  • 15. Dorsal sepal connate with the lateral sepals.........................................................45. Stelis paraensis

                  • 15’. Dorsal sepal free...................16

                    • 16. Lepanthiform sheaths absent; flowers yellow, lateral sepals free...................29. Octomeria grandiflora

                    • 16’. Lepanthiform sheaths present; flowers dark purple, lateral sepals connate...................47. Trichosalpinx egleri

                • 14’. Leaves 2-several...................17

                  • 17. Pseudobulbs superposed; leaves terminal...................18

                    • 18. Sepals connate at base; perianth < 3.0 mm long.........................................................42. Scaphyglottis sickii

                    • 18’. Sepals free; perianth > 5.0 mm long...................19

                      • 19. Flowers pink; lip 3-lobed; column with lateral appendixes...................43. Scaphyglottis stellata

                      • 19’. Flowers white or beige; lip entire; column without lateral appendixes...................41. Scaphyglottis prolifera

                  • 17’. Pseudobulbs not superposed; leaves distributed along the stem......................................20

                    • 20. Column free or adnate at base to the lip...................21

                      • 21. Flowers pink, column adnate at base with the lip...................8. Dimerandra emarginata

                      • 21’. Flowers yellowish-brown, column free...................32. Orleanesia amazonica

                    • 20’. Column completely adnate to the lip...................22

                      • 22. Rachis completely covered by bracts; lip entire...................23

                        • 23. Flowers green, lip suborbicular...................13. Epidendrum rigidum

                        • 23’. Flowers white, lip cordate...................14. Epidendrum strobiliferum

                      • 22’. Rachis exposed; lip obscurely to clearly 3-lobed...................24

                        • 24. Inflorescence short pedunculate (< 1 cm long)...................10. Epidendrum carpophorum

                        • 24’. Inflorescence long pedunculate (at least 1.6 cm long)...................25

                          • 25. Peduncle completely covered by bracts; lip margin entire.........................................................9. Epidendrum anceps

                          • 25’. Peduncle exposed; lip margin denticulate...................11. Epidendrum macrocarpum

        • 6’. Inflorescence lateral...................26

          • 26. Leaves cylindrical...................27

            • 27. Inflorescence uniflorous; flowers white...................25. Maxillaria uncata

            • 27’. Inflorescence multiflorous; flowers yellow with brown spots...................46. Trichocentrum cepula

          • 26’. Leaves flat...................28

            • 28. Leaves equitant, unifacial...................29

              • 29. Leaves much shorter than the stem...................19. Lockhartia imbricata

              • 29’. Leaves as long as or longer than the stem...................30

                • 30. Flowers pink; lip clawed...................20. Macroclinium wullschlaegelianum

                • 30’. Flowers white, green or yellow; lip not clawed...................31

                  • 31. Flowers yellow with brown dots; petal margins entire; lip apex 2-lobed; column wings developed...................15. Erycina pusilla

                  • 31’. Flowers white or greenish; petal margins minutely denticulate; lip apex rounded or cuspidate; column without wings...................32

                    • 32. Sepal apex rounded to obtuse; lip entire.........................................................33. Ornithocephalus cujeticola

                    • 32’. Sepal apex acuminate; lip 3-lobed...................34. Ornithocephalus gladiatus

            • 28’. Leaves not equitant, dorso-ventrally flattened...................33

              • 33. Flowers with a spur...................34

                • 34. Pseudobulb present; leaves 1, terminal...................30. Oeceoclades maculata

                • 34’. Pseudobulb absent; leaves 3-12, distributed along the stem.........................................................3. Campylocentrum micranthum

              • 33’. Flowers without spur...................35

                • 35. Pseudobulb absent...................7. Dichaea picta

                • 35’. Pseudobulb present...................36

                  • 36. Pseudobulb homoblastic; flowers unisexual...................5. Catasetum macrocarpum

                  • 36’. Pseudobulb heteroblastic; flowers bisexual...................37

                    • 37. Inflorescences uniflorous...................38

                      • 38. Peduncle > 6.5 cm long; lip < 0.6 cm long.........................................................24. Maxillaria subrepens

                      • 38’. Peduncle < 4.5 cm long; lip > 1.0 cm long...................30

                        • 39. Rhizome fully fixed in the substrate; flowers yellow.........................................................22. Maxillaria aureoglobula

                        • 39’. Rhizome pendent, fixed in the substrate only at the base; flowers white...................4

                          • 40. Apical leaves 2; lateral lobes of the lip developed.........................................................23. Maxillaria lutescens

                          • 40’. Apical leaf 1; lateral lobes of the lip vestigial.........................................................21 Maxillaria alba

                    • 37’. Inflorescences multiflorous...................41

                      • 41. Flowers non-resupinate...................42

                        • 42. Inflorescence lax; lip hypochile 0.7–0.9 cm long, pair of callus > 0.2 cm long.........................................................16. Gongora nigrita

                        • 42’. Inflorescence congested; lip hypochile 0.5–0.7 cm long, pair of callus < 0.1 cm long.........................................................17. Gongora quinquenervis

                      • 41’. Flowers resupinate...................43

                        • 43. Leaves plicate, main veins abaxially evident...................44

                          • 44. Pseudobulb smooth; lip not divided; column without glands...................35. Peristeria serroniana

                          • 44’. Pseudobulb ribbed; lip divided into epichile, mesochile and hypochile; column with a pair of glands...................6. Coryanthes speciosa

                        • 43’. Leaves conduplicate, veins not evidente...................45

                          • 45. Lip clawed...................46

                            • 46. Lateral sepals completely connate; lip blade ovate...................27. Notylia microchila

                            • 46’. Lateral sepals free at distal half; lip blade lanceolate...................47

                              • 47. Inflorescence 3.0-5.0 cm long; lip blade apex laterally recurved.........................................................28. Notylia yauaperyensis

                              • 47’. Inflorescence 6.0- 22.0 cm long; lip blade apex plane...................26. Notylia aromatica

                          • 45’. Lip not clawed...................48

                            • 48. Flowers magenta, lateral sepals connate...................38. Rodriguezia lanceolata

                            • 48’. Flowers yellow or beige with brown dots or lines, lateral sepals free...................49

                              • 49. Lip adnate to the base of the column, margin erose...................1. Aspasia variegata

                              • 49’. Lip free from the column, margin entire...................50

                                • 50. Flowers at least 17; lateral sepal apex acute; lip 3-lobed.........................................................31. Oncidium baueri

                                • 50’. Flowers at least 5-8; lateral sepal apex caudate; lip entire.........................................................2. Brassia caudata

1. Aspasiavariegata Lindl. Edwards’s Bot. Reg. 22: t. 1907. 1836.

Examined material: trail near the Pirarucu lake, 01°50’45.2”S, 45°47’17.5”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 001 (SLUI 5724); Igarapé Reno, 01º51’48”S, 45º46’02”W, 3.XI.2017, fl., A.W.C. M.J.C. Silva & A.W.C. Ferreira 002 (SLUI 5725).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain, but it is also recorded to Cerrado. It is a species that while sterile can be confused in the area with O. baueri and B. caudata but in flower is recognized by the lip adnate to the base of the column and with an erose margin (vs. free and entire). The local flowering period is between November and January.

2. Brassiacaudata (L.) Lindl., Bot. Reg. 10: t. 832.1825.

Examined material: Igarapé Cumaruzal, 01°50’36”S,45°47’09”W, 3.VII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 005 (SLUI 5726).

Widespread in the Neotropical region, in Brazil it is widely distributed in the Amazon domain. This species is easily recognized by its long caudate sepals, the most similar species in the area is A. variegata from which it can be distinguished by having more flowers per inflorescence (5–8 vs. 1–3). The local flowering period is between May and July.

3. Campylocentrum micranthum (Lindl.) Rolfe, Orchid Rev. 9: 136. 1903. Fig. 2a

Figure 2
a-h. Orchidaceae of Fazenda Sete Irmãos – a. Campylocentrum micranthum; b. Campylocentrum pachyrrhizum; c. Coryanthes speciosa; d. Dichaea picta; e. Laelia gloriosa; f. Lockhartia imbricata; g. Maxillaria alba; h. Notylia microchila. (a. M.J.C. Silva & A.W.C. Ferreira 006; b. M.J.C. Silva & A.W.C. Ferreira 007; c. M.J.C. Silva, W.R. Silva Junior & A.W.C. Ferreira 012; d. W.R. Silva Junior, M.J.C. Silva & A.W.C. Ferreira 014; e. M.J.C. Silva & A.W.C. Ferreira 055; f. M.J.C. Silva & A.W.C. Ferreira 057; g. W.R. Silva Junior & A.W.C. Ferreira 105; h. M.J.C. Silva & A.W.C. Ferreira 069). Photos: a-h. A.W.C Ferreira.

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.VII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 006 (SLUI 5727).

Widespread in northern South America and the Caribbean, in Brazil it is found in the Amazon and Atlantic Forest domains. This species was cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. to Maranhão, but without a testimony voucher. As it is not listed to the state in BFG (2015, 2018), here its presence is confirmed. It is easily distinguished from the other species of the genus in the area by the presence of leaves (vs. leafless). The local flowering period is between June and July.

4. Campylocentrumpachyrrhizum (Rchb.f.) Rolfe, Orch. Rev.11(128): 246. 1903. Fig. 2b

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.VII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 007 (SLUI 5728).

Widespread in the Neotropical region, in Brazil it is found in the Amazon and Atlantic Forest domains. This study presents its first record to Maranhão. It is easily recognized among the species of the area by being leafless and by the flat, green roots. The local flowering period is between June and August.

5. Catasetum macrocarpum Richard, L. C. & Kunth, C., Syn. Pl. Aequin., I: 331, 1822.

Examined material: near the border with Zé Pedro’s farm, 02°07’47.4”S, 45°48’42.2”W, 15.X.2017, fl., M.J.C. Silva & A.W.C. Ferreira 009 (SLUI 5730).

Widespread in South America, in Brazil it is widely distributed, except in the South region. It is distinguished in the area by its homoblastic pseudobulbs and it is also the only species with unisexual flowers. The local flowering period is between April and November.

6. Coryanthes speciosa Hooker, W.J., Bot. Mag., 58: 3102, 1831. Fig. 2c

Examined material: Igarapé Reno, near the wood bridge, 01°52’35.6”S, 45°45’18.9”W, 13.X.2017, fl., M.J.C. Silva, W.R. Silva Junior & A.W.C. Ferreira 012 (SLUI 5731); 01°52’35.6”S, 45°45’18.9”W, 13.X.2017, fl., M.J.C. Silva, W.R. Silva Júnior & A.W.C. Ferreira 013 (SLUI 5732).

Widespread in northern South America, in Brazil it is widely distributed, except in the South region. This study presents its first record to Maranhão. It is characterized by the lip divided into epichile, mesochile and hypochile and column with a pair of glands. The local flowering period is between September and November.

7. Dichaea picta Rchb.f. W.W.Saunders, Refug. Bot. (Saunders), 2: t. 84, 1872. Fig. 2d

Examined material: Igarapé Cumaruzal, 01°50’55.0”S, 45°45’35.4”W, 14.VII.2017, fl., W.R. Silva Junior, M.J.C. Silva & A.W.C. Ferreira 014 (SLUI 5733).

Widespread in northern South America, in Brazil it is cited only to the states of Amazonas, Pará and Rondônia. This species was cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. to Maranhão, but without a testimony voucher. As it is not listed to the state in BFG (2015, 2018), here its presence is confirmed. It can be confused in the area with a member of Epidendrum due to the cespitose habit and leaves distributed along the stem, but it differs by membranous leaves and 1-flowered inflorescences (vs. coriaceous, multiflorous). The local flowering period is between June and July.

8. Dimerandra emarginata (G.Mey.) Hoehne, Bol. Agric. Estado São Paulo 34: 618. t. 9. 1934.

Examined material: trail to Igarapé Cumaruzal, 01°50’55.0”S, 45°45’35.4”W, 15.VI.2017, fl., W.R. Silva Junior, M.J.C. Silva & A.W.C. Ferreira 015 (SLUI 5734).

Widespread in the Neotropical region, in Brazil it is found in the Amazon and Atlantic Forest domains. It is a species that can be confused in the area with Epidendrum, but is distinguished by the column adnate only at base with the lip (vs. fully adnate). The local flowering period is between May and July.

9. Epidendrum anceps Jacq., Select. Stirp. Amer. Hist. 224 (t. 138). 1763.

Examined material: trail to Igarapé Cumaruzal, 01°50’47.7”S, 45°45’48.7”W, 14.X.2017, fl., M.J.C. Silva & A.W.C. Ferreira 018 (SLUI 5699).

Widespread in the Neotropical region, and widely distributed in Brazil. It can be distinguished from the other species of the genus in the area by its pedunculate inflorescence (at least 1.6 cm long) with peduncle completely covered by bracts, whereas in E. rigidum and E. strobiliferum the rachis is covered by bracts. The local flowering period is between October and November.

10. Epidendrum carpophorum Barb.Rodr., Gen. Sp. Orchid. 2: 148. 1882.

Examined material: Igarapé Cumaruzal, 01º50’48”S, 45º46’02”W, 15.VI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 020 (SLUI 5702); Macaxeira river, 01°52’47”S, 45°47’54”W, 20.II.2020, fl., A.W.C. Ferreira 021 (SLUI 5703).

Widespread in northern South America, in Brazil it is widely distributed, except in the South region. Among the Epidendrum species of the area it can be confused with E. purpurascens due to the deeply 3-lobed white lip, but in E. carpophorum the stem is not swollen (vs. swollen). The local flowering period is between February and August.

11. Epidendrummacrocarpum Rich., Actes Soc. Hist. Nat. Paris 1(1): 112. 1792.

Examined material: Macaxeira river, near the bridge of the old road, 01°52’43”S, 45°45’37”W, 15.VII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 032 (SLUI 5713).

Widespread in northern South America, in Brazil it is found in the Amazon and Atlantic Forest domains. It is distinguished from the other species of the genus in the area by its long inflorescence and red flowers. The local flowering period is between July and September.

12. Epidendrum purpurascens Focke. Tijdschr. Nat. Wetensch. 4: 64-65. 1851.

Examined material: Igarapé Cumaruzal, 01°50’47.7”S, 45°45’48.7”W, 14.X.2017, fl., M.J.C. Silva & A.W.C. Ferreira 036 (SLUI 5716).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain. Among the Epidendrum species of the area it is the only one with pseudobulbs. The local flowering period is between October and February.

13. Epidendrum rigidum Jacq., Enum. Syst. Pl.: 29. 1760.

Examined material: Macaxeira river, 01°52’44.4”S, 45°45’36.2”W, 15.IV.2019, fl., M.J.C. Silva & A.W.C. Ferreira 038 (SLUI 5719).

Widespread in the Neotropical region, it is also widely distributed in Brazil. It can be confused with E. strobiliferum but the stem of this epiphyte is not branched and the flowers are green (vs. branched, white). The local flowering period is between March and Abril.

14. Epidendrum strobiliferum Rchb. f., Ned. Kruidk. Arch. 4: 333. 1859.

Examined material: Igarapé Cumaruzal, 01°50’47”S, 45°48’42”W, 15.VII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 040 (SLUI 5721).

Widespread in the Neotropical region, it is also widely distributed in Brazil. Among the Epidendrum species of the area it is the only one with ramified, pendent stems. The local flowering period is between May and July.

15. Erycina pusilla (L.) N.H.Williams & M.W.Chase. Lindleyana, 16: 136, 2001.

Examined material: Macaxeira, near the border of the Fazenda Sete Irmãos, 01°52’06”S, 45°48’01”W, 29.VI.2019, fl., A.W.C. Ferreira 042 (SLUI 5735).

Widespread in the Neotropical region, in Brazil it is widely distributed, except in the South region. This twig epiphyte with equitant leaves can be confused in the area, if sterile, with a member of Ornithocephalus, but the flowers are fairly distinctive, and it can be recognized by the color (yellow vs. white or greenish), margin of the petals (entire vs. minutely denticulate), and the presence of wings on the column (vs. without wings). The local flowering period is between June and August.

16. Gongoranigrita Lindl. Lindley, J., Edwards’s Bot. Reg., 25: 59, 1839.

Examined material: Macaxeira river, 01°52’44.4”S, 45°45’36.2”W, 15.X.2017, fl. M.J.C. Silva & A.W.C. Ferreira 048 (SLUI 5736); trail on Igarapé Reno, 01°50’55”S, 45°45’35.4”W, 3.XI.2017, fl. M.J.C. Silva & A.W.C. Ferreira 049 (SLUI 5737).

Distributed in Guyana, Suriname and Brazil, where it is found in the Amazon and Atlantic Forest domains, it has been cited only to the states of Amazonas, Maranhão, Pará and Pernambuco. It is similar to G. quinquenervis but differs by the perianth length (4‒5 cm long vs. 2.5‒4 cm long). The local flowering period is between October and November.

17. Gongoraquinquenervis Ruiz & Pav. Syst. Veg. Fl. Peruv. Chil. 1: 227. 1798.

Examined material: Macaxeira river, 01°52’44.4”S, 45°45’36.2”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 050 (SLUI 5738); Fazenda Sete Irmãos, 02°52’44.4”S, 45°45’36.2”W, 3.XI.2018, fl., M.J.C. Silva & A.W.C. Ferreira 051 (SLUI 5739).

Distributed in Colombia, Ecuador, Peru and Brazil, where it is found in the Amazon and Atlantic Forest domains. As mentioned above, it is similar to G. nigrita, but can also be distinguished by the length of lip callus (< 0.1 cm long vs. > 0.2 cm long). The local flowering period is between October and November.

18. Laeliagloriosa (Rchb.f.) L.O. Williams. Darwiniana 5: 76. 1941. Fig. 2e

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 055 (SLUI 5740); Macaxeira river, 01°52’44.4”S, 45°45’36.2”W, 5.V.2018, fl., A.W.C. Ferreira 056 (SLUI 5741).

Widespread in northern South America, in Brazil it is widely distributed. This species was cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. to Maranhão, but without a testimony voucher. As it is not listed to the state in BFG (2015, 2018), here its presence is confirmed. Among the species of the area, young individuals of this species can be confused with E. purpurascens due to the heteroblastic fusiform pseudobulbs, but these species are easily distinguished by the length of the peduncle (> 20 cm long vs. < 8 cm long) and color of the flowers (sepals and petals brown, lip pale pink vs. completely white). The local flowering period is between April and May.

19. Lockhartia imbricata (Lam.) Hoehne. Arq. Bot. Estado São Paulo 2: 139. 1952. Fig. 2f

Examined material: reforestation branch, trail of the igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 16.VI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 057 (SLUI 5742).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain. This study presents its first record to Maranhão and also to northeastern Brazil. Among the twig epiphytes with unifacial leaves of the area, it is the only one with an elongated stem. The local flowering period is between May and July.

20. Macroclinium wullschlaegelianum (Focke) Dodson. Icon. Pl. Trop. 10: t. 939. 1984.

Examined material: Igarapé Cumaruzal, 01°50’40.4”S, 45°46’01”W2, 14.X.2019, fl., M.J.C. Silva & A.W.C. Ferreira 058 (SLUI 5743).

Widespread in northern South America, the species has also been recorded to Belize, in Brazil it is widely distributed in the Amazon and Cerrado domains. This twig epiphyte is the only one among the species with unifacial leaves that produces pink flowers with a clawed lip. The local flowering period is between October and December.

21. Maxillaria alba (Hook.) Lindl. Gen. Sp. Orchid. Pl., 143, 1832. Fig. 2g

Examined material: Igarapé affluent of Macaxeira river, near the border of Fazenda Sete Irmãos, 01°52’08”S, 45°48’03”W, 29.VI.2019, fl., W.R. Silva Junior & A.W.C. Ferreira 105 (SLUI 5744).

Widespread in the Neotropical region, in Brazil it is widely distributed in the Amazon and Cerrado domains. It can be confused in the area with M. lutescens due to the pendent habit, but differs by having 1 apical leaf (vs. 2) and vestigial lateral lobes of the lip (vs. developed). The local flowering period is between May and June.

22. Maxillaria aureoglobula Christenson, Orchids (West Palm Beach) 71: 125 (2002).

Examined material: Igarapé Cumaruzal, 01°50’50”S, 45°46’10”W, 9.III.2019, fl., A.W.C. Ferreira 100 (MAR 11538).

It is distributed in Costa Rica, Venezuela, Colombia and Brazil, where it is cited only to the states of Maranhão and Mato Grosso. Among the species of Maxillaria in the area, it can be confused with young individuals of M. alba or M. lutescens, but these two species have a pendent habit while M. aureoglobula has its rhizome fully fixed in the substrate. The local flowering period is between February and April.

23. Maxillaria lutescens Scheidw. F Allg. Gartenzeitung, 7: 145, 1839.

Examined material: near the Pirarucu lake and the old headquarter, 01°50’38.1”S, 45°46’07.7”W, 1.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 059 (SLUI 5745); trail to igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 2.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 060 (SLUI 5746).

Widespread in the Neotropical region, and also in Brazil. It is similar to M. alba, but the leaves are wider (up to 2.5 cm wider vs. < 1 cm wider), other features used to distinguish these two species are in the comments of M. alba. The local flowering period is between January and April.

24. Maxillaria subrepens (Rolfe) Schuit. & M.W.Chase. Phytotaxa, 225: 73, 2015.

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 6.VII.2018, fl., M.J.C. Silva & A.W.C. Ferreira 061 (SLUI 5747); 01°49’34.6”S, 45°46’09.8”W, 7.VII.2018, fl., M.J.C. Silva & A.W.C. Ferreira 062 (SLUI 5748).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon and Atlantic Forest domains. It is easily distinguished among the other Maxillaria of the area by its longer peduncle (> 6.5 cm long vs. < 2.0 cm long) and shorter lip (< 0.6 cm long vs. > 1.0 cm long). This species was formerly included under Trigonidium, considered part of Maxillaria s.l. today. The local flowering period is between June and November.

25. Maxillaria uncata Lindl. Edwards’s Bot. Reg., 23: t. 1986, 1837.

Examined material: Igarapé Cumaruzal, 01°50’46.3”S, 45°47’17.5”W, 15.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 063 (SLUI 5749); 01°50’46.3”S, 45°47’17.5”W, 3.XI.2018, fl., M.J.C. Silva & A.W.C. Ferreira 064 (SLUI 5750).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain. It can be distinguished from the other Maxillaria of the area by its cylindrical leaves. The local flowering period is between November and May.

26. Notylia aromatica Barker ex Lindl. Edwards’s Botanical Register 27: Misc. 40. 1841.

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 065 (SLUI 5751); 01°50’37”S, 45°47’10”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 066 (SLUI 5752).

Distributed in French Guyana, Guyana, Suriname, Venezuela and Brazil, where it is widely distributed in the Amazon domain. It is similar to N. yauaperyensis but differs by its longer inflorescences (6–22 cm long vs. 3–5 cm long) and number of flowers (15–70 vs. 8–22). The local flowering period is between September and December.

27. Notylia microchila Cogn., Fl. Bras. (Martius) 3(6): 123 (1904). Fig. 2h

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 069 (SLUI 5753); igarapé affluent do Macaxeira river, near the border of Fazenda Sete Irmãos, 01°52’07.3”S, 45°48’02.2”W, 27.XI.2019, fl., W.R. Silva Junior & A.W.C. Ferreira 070 (SLUI 5754).

Distributed in Ecuador, Guyana, Peru, Venezuela and Brazil, where it is cited only to the states of Bahia, Espírito Santo, Mato Grosso, Pará and Pernambuco. This study presents its first record to Maranhão. This species is easily distinguished from the other Notylia of the area by its lateral sepals completely connate (vs. free at distal half) and lip blade ovate (vs. lanceolate). The local flowering period is between November and December.

28. Notylia yauaperyensis Barb. Rodr. Vellosia (ed. 2) 1: 131. 1891.

Examined material: Igarapé affluent of Macaxeira river, near the border of Fazenda Sete Irmãos, 01°52’07.3”S, 45°48’02.2”W, 27.XI.2019, fl., W.R. Silva Junior & A.W.C. Ferreira 071 (SLUI 5755).

Distributed in French Guyana, Peru, Venezuela and Brazil, where it is widely distributed in the Amazon domain. It can be confused in the area with N. aromatica, but it differs by the morphological features cited in the comment above. The local flowering period is between November and December.

29. Octomeriagrandiflora Lindl. Edwards’s Bot. Reg. 28 (Misc.): 64. 1842.

Examined material: Igarapé Cumaruzal, 01°50’36”S, 45°47’09”W, 15.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 072 (SLUI 5756).

Distributed in Bolivia, Colombia, Ecuador, Peru, and Brazil, where it is widely distributed. Its most similar species in the area is T. egleri, but these species are easily distinguished by the color the flowers (dark purple vs. yellow) and the fusion of the lateral sepals (free vs. connate). The local flowering period is between January and May.

30. Oeceoclades maculata (Lindl.) Lindl., Gen. Sp. Orchid. Pl.: 237. 1833.

Examined material: trail to igarapé Cumaruzal, 01°50’37”S, 45°47’10”W, 15.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 073 (SLUI 5757).

Widely distributed in the Neotropics and tropical Africa, as well as in Brazil. It is the only terrestrial species in the area with lateral inflorescences and flowers with a distinct spur. The local flowering period is between February and May.

31. Oncidium baueri Lindl., A.C., Fl. Bras. (Martius), 3(6): t. 85, 1906. Fig. 3a

Figure 3
a-h. Orchidaceae of Fazenda Sete Irmãos – a. Oncidium baueri; b. Ornithocephalus cujeticola; c. Polystachya concreta; d. Sarcoglottis acaulis; e. Scaphyglottis prolifera; f. Stelis paraensis; g. Trichosalpinx egleri; h. Wullschlaegelia calcarata. (a. A.W.C. Ferreira 075; b. M.J.C. Silva & A.W.C. Ferreira 077; c. M.J.C. Silva & A.W.C. Ferreira 082; d. A.W.C. Ferreira 088; e. M.J.C. Silva & A.W.C. Ferreira 089; f. M.J.C. Silva & A.W.C. Ferreira 095; g. M.J.C. Silva & A.W.C. Ferreira 098; h. W.R. Silva Junior & A.W.C. Ferreira 103). Photos: a-h. A.W.C Ferreira.

Examined material: Macaxeira river, 01°52’11.0”S, 45°47’59.8”W, 21.XII.2019, A.W.C. Ferreira 075 (SLUI 5758).

Widespread in northern South America, the species has also been recorded to Costa Rica, in Brazil it is widely distributed in the Amazon and Atlantic Forest domains. This species was cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. to Maranhão, but without a testimony voucher. As it is not listed to the state in BFG (2015, 2018), here its presence is confirmed. Its flowers are somewhat similar to E. pusilla, but the presence of pseudobulbs (vs. absent) and the bi-facial leaves (vs. unifacial) distinguish these two. The local flowering period is between November and April.

32. Orleanesia amazonica Barb.Rodr., Genera et Species Orchidearum Novarum, 1, 1877.

Examined material: Macaxeira river, 01°52’15.1”S, 45°47’58.5”W, 20.II.2020, fl., A.W.C. Ferreira 076 (SLUI 5759).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain. It can be confused in the area with a member of Epidendrum but differs by its column totally free (vs. adnate to the lip claw). The local flowering period is between September and February.

33. Ornithocephaluscujeticola Barb. Rodr., Gen. Sp. Orch. Nov. 1:133.1877. Fig. 3b

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 11.VIII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 077 (SLUI 5760); Macaxeira river, 01°52’37.6”S, 45°47’09”W, 12.VIII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 078 (SLUI 5761).

Endemic to Brazil where it is cited only to the states of Amazonas, Ceará, Mato Grosso, Pará and Rondônia. This study presents its first record to Maranhão. In the area it is similar to O. gladiatus Hook. but differs by the sepals rounded to obtuse at apex (vs. acuminate) and the lip entire (vs. 3-lobed). The local flowering period is between June and August.

34. Ornithocephalus gladiatus Hook., Exot. Fl. 2:t. 127. 1824.

Examined material: trail to igarapé Cumaruzal, 01°50’40.4”S, 45°’01”W, 13.IX.2017, fl., M.J.C. Silva & A.W.C. Ferreira 079 (SLUI 5762); Igarapé Reno, 01°51’47.4”S, 45°45’03”W, 14.IX.2017, fl., M.J.C. Silva & A.W.C. Ferreira 080 (SLUI 5763).

Widespread in northern South America, in Brazil it is widely distributed, except in the South and Southeast regions. As mentioned before, it is similar to O. cujeticola; the features used to distinguish them are described in the comment above. The local flowering period is between June and September.

35. Peristeria serroniana (Barb. Rodr) Garay, Arch. Jard. Bot. Rio de Janeiro 13: 47 (1954).

Examined material: Macaxeira river, 01°52’11.0”S, 45°47’59.8”W, 20.II.2019, fl., M.J.C. Silva & A.W.C. Ferreira 081 (SLUI 5764).

Endemic to Brazil, where it is cited only to the states of Pará and Maranhão, in the Amazon domain. It can be confused with C. speciosa due to the heteroblastic pseudobulbs and plicate leaves but is easily distinguished by its smooth pseudobulb (vs. ribbed). The local flowering period is between January and February.

36. Polystachya concreta (Jacq.) Garay & H.R.Sweet, Orquideología, 9: 206, 1974. Fig. 3c

Examined material: trilha para o igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 4.V.2018, fl., M.J.C. Silva & A.W.C. Ferreira 082 (SLUI 5765).

Widespread in the Neotropical region, in Brazil it is cited only to the states of Amapá, Goiás, Mato Grosso and Rondônia, in the Amazon and Cerrado domains. This species was cited by Silva et al. (1999)Silva MFF, Silva JBF & Feiler JM (1999) Orchidaceas do estado do Maranhão, Brasil. Acta Amazonica 29: 381-393. to Maranhão, but without a testimony voucher. As it is not listed to the state in BFG (2015, 2018), here its presence is confirmed. In the area, the only other species with swollen stems, terminal inflorescence and non-resupinate flowers is P. aemula, but it is easily distinguished by its 3-lobed lip (vs. entire). The local flowering period is between February and May.

37. Prosthechea aemula (Lindl.) W.E.Higgins, Phytologia 82: 376 (1997 publ. 1998).

Examined material: near the pirarucu lake, 01°50’46.3”S, 45°47’17.5”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 083 (SLUI 5766); 01°48’46.3”S, 45°50’19.5”W, 4.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 084 (SLUI 5767).

Widespread in northern South America, it has also been recorded to Panama, in Brazil it is widely distributed. Young specimens of this species can be confused in the area with E. purpurascens, due to the fusiform heteroblastic pseudobulbs, but it is easily distinguished by the entire lip (vs. 3-lobed). Specimens of P. aemula has been identified as P. fragrans, but this second species is not found in the state. The local flowering period is between October and April.

38. Rodriguezialanceolata Ruiz & Pav. Syst. Veg. Fl. Peruv. Chil. 1: 219. 1798.

Examined material: Igarapé Cumaruzal, 01°50’45.2”S, 45°47’17.5”W, 3.II.2018, fl., M.J.C. Silva & A.W.C. Ferreira 085 (SLUI 5768); 01°48’45.2”S, 45°50’19.5”W, 10.III.2019, fl., M.J.C. Silva & A.W.C. Ferreira 086 (SLUI 5769).

Widespread in northern South America, it has also been recorded to Panama, in Brazil it is widely distributed in the Amazon domain, but is also cited to surrounding Cerrado. Among the species with heteroblastic pseudobulbs, lateral inflorescences and lip not clawed in the area it is the only one with purple flowers and lateral sepals connate. The local flowering period is between January and March.

39. Sacoila lanceolata (Aubl.) Garay, Bot. Mus. Leafl. 28: 352. 1980.

Examined material: campo na trilha para o Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 087 (SLUI 5770).

Widespread in the Neotropical region, and also in Brazil. Before flowering it can be confused with S. acaulis, but all the leaves shed before the flowering period (vs. leaves present during flowering). The local flowering period is between October and January.

40. Sarcoglottis acaulis (Sm.) Schltr. Fl. Bras. (Martius), 3(4): t. 50, 1895. Fig. 3d

Examined material: near the Pirarucu lake, 01°51’51.5”S, 45°45’56.5”W, 10.IX.2019, fl., A.W.C. Ferreira 088 (SLUI 5771).

Widespread in northern South America and Central America, in Brazil it is widely distributed. This study presents its first record to Maranhão. As mentioned before, it can be confused with S. lanceolata, but its greenish-white flowers are fairly distinctive for lacking a spur (vs. flowers magenta, spur present). The local flowering period is between August and September.

41. Scaphyglottis prolifera (R.Br.) Cogn., Fl. Bras. 3(5): 15. 1898. Fig. 3e

Examined material: trail to igarapé Cumaruzal, 01°51’11.4”S, 45°47’34.6”W, 11.VIII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 089 (SLUI 5772); 01°51’11.4”S, 45°47’34.6”W, 11.VIII.2017, fl., M.J.C. Silva & A.W.C. Ferreira 090 (SLUI 5773).

Widespread in the Neotropical region, in Brazil it is widely distributed, except in the South region. This study presents its first record to Maranhão. It is similar to S. stellata but differs by the white to beige flowers (vs. pink) and column without lateral appendixes (vs. with lateral appendixes). The local flowering period is between July and August.

42. Scaphyglottis sickii Pabst, Orquídea (Rio de Janeiro) 18: 7. 1956.

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 15.VI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 091 (SLUI 5774); 01°50’36.6”S, 45°47’09.8”W, 15.VI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 092 (SLUI 5775).

Widespread in northern South America, in Brazil it is found in the Amazon and Atlantic Forest domains. This species differs from the other Scaphyglottis of the area by its sepals connate at base (vs. free) and shorter perianth (< 3.0 mm long vs. > 5.0 mm long). The local flowering period is between May and July.

43. Scaphyglottis stellata Lodd. ex Lindl., Edwards Bot. Reg. 25: 44. 1839.

Examined material: Igarapé Reno, 01°51’11”S, 45°47’34”W, 17.IX.2017, fl., M.J.C. Silva & A.W.C. Ferreira 093 (SLUI 5776).

Widespread in northern South America and Central America, in Brazil it is widely distributed in the Amazon domain, but is also cited to surrounding Cerrado. In the area it can be confused with S. prolifera but can be distinguished by its 3-lobed lip (vs. entire). The local flowering period is between August and September.

44. Sobraliamacrophylla Rchb. f., Bot. Zeitung (Berlin) 10: 713 (1852).

Examined material: Igarapé Cumaruzal, 01°50’36.6”S, 45°47’09.8”W, 15.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 094 (SLUI 5777).

Widespread in northern South America and Central America, in Brazil it is cited only to the states of Amapá, Amazonas, Maranhão, and Pará in the Amazon Domain. It is the only species of the area with the combination of plicate leaves and stems not swollen. The local flowering period is between March and April.

45. Stelisparaensis Barb.Rodr. Gen. Sp. Orchid. ii. 88. 1881. Fig. 3f

Examined material: Igarapé Cumaruzal, 01°50’58.4”S, 45°45’30.7”W, 15.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 095 (SLUI 5778); Macaxeira river, 01°52’58.4”S, 45°47’30.7”W, 16.IV.2017, fl., M.J.C. Silva & A.W.C. Ferreira 096 (SLUI 5779).

Endemic to Brazil, where it is cited only to the states of Mato Grosso, Pará and Rondônia in the Amazon domain. This study presents its first record to Maranhão and also to northeastern Brazil. It can be confused in the area with S. egleri and O. grandiflora but differs by its lateral and dorsal sepals fused (vs. dorsal sepal free). The local flowering period is between February and April.

46. Trichocentrum cepula (Hoffmanns.) J.M.H.Shaw. Orchid Rev. 120(1297, Suppl.): 16 (2012).

Examined material: Igarapé Reno, 01°51’11.4”S, 45°47’34.6”W, 3.XI.2017, fl., M.J.C. Silva & A.W.C. Ferreira 097 (SLUI 5780).

Distributed in Argentina, Bolivia, Paraguay and Brazil, where it is widely distributed in the Cerrado and Caatinga domains. The other species with cylindrical leaves in the area is M. uncata from which it differs by the multiflorous inflorescences (vs. 1-flowered). The local flowering period is between October and January.

47. Trichosalpinx egleri (Pabst) Luer, Phytologia 54(5): 395. 1983. Fig. 3g

Examined material: Igarapé Reno, 01°51’07.5”S, 45°47’36.9”W, 21.IV.2018, fl., M.J.C. Silva & A.W.C. Ferreira 098 (SLUI 5781).

Widespread in northern South America, in Brazil it is widely distributed in the Amazon domain. This study presents its first record to Maranhão and also to northeastern Brazil. It can be confused in the area with O. grandiflora and S. paraensis but can be easily recognized by its lepanthiform sheaths. The local flowering period is between January and April.

48. Vanilla mexicana Mill., Gard. Dict. ed. 8: n.º 1 (1768).

Examined material: Macaxeira river, 01°52’59”S, 45°47’31”W, 12.VIII.2017, fr., M.J.C. Silva & A.W.C. Ferreira 099 (SLUI 5782).

Widespread in the Neotropical region, in Brazil it is also widely distributed, except in the South region. It can be confused in the area with V. palmarum but differs by its sepals and petals undulate at margin, green (vs. flat, yellow). The local flowering period is between July and August.

49. Vanilla palmarum (Salzm. ex Lindl.) Lindl., Gen. Sp. Orchid. Pl.: 436. 1832.

Examined material: near the Igarapé Reno, 01°51’11.4”S, 45°47’34.6”W, 3.XI.2017, fr., M.J.C. Silva & A.W.C. Ferreira 101 (SLUI 5783).

Widespread in northern South America, in Brazil it is widely distributed, except in the South and Southeast regions. In the area it can be confused with V. mexicana but it grows exclusively on palm species (vs. other phorophytes). The local flowering period is between September and January.

50. Vanilla pompona Schiede, C.J.W., Linnaea, 4: 573-574, 1829.

Examined material: branch near to Carvoaria project, 01°51’48”S, 45°46’02”W, 10.IX.2019, A.W.C. Ferreira 102 (SLUI 5784).

Widespread in the Neotropical region, in Brazil it is widely distributed, except in the South region. It is easily distinguished from the other species of Vanilla in the area by its lip with a multi-ridged callus on the disc (vs. callus absent). The local flowering period is between August and October.

51. Wullschlaegelia calcarata Benth., J. Linn. Soc., Bot. 18: 342 (1881). Fig. 3h

Examined material: trail near to Pirarucu lake, 01°51’51.5”S, 45°45’56.5”W, 26.X.2019, fl. and fr., W.R. Silva Junior & A.W.C. Ferreira 103 (SLUI 5785); 01°51’51.5”S, 45°45’56.5”W, 26.X.2019, fl. and fr., W.R. Silva Junior & A.W.C. Ferreira 104 (SLUI 5786).

Widespread in the Neotropical region, in Brazil it is widely distributed in the Amazon domain. This study presents its first record to Maranhão and also to northeastern Brazil. It is the only myco-heterotrophic herb in the area, recognized by achlorophyllous stems and scale-like leaves. The plants arise from the soil in the area between March and July, and local flowering period is between October and November.

Acknowledgements

We thank Adão Pontarollo and family, for allowing our research in the area; Francisco Ribeiro and the Laboratory of Bee Studies of the Federal University of Maranhão, for assistance during fieldwork; the herbarium SLUI (herbarium Rosa Mochel, Universidade Estadual do Maranhão), for the assistance on the deposit of the specimens; the FAPEMA, by the funding of A.W.C. Ferreira (Edital Universal, 9033/2015, 009033/2015); and CNPQ, by the funding of Edlley Pessoa (Edital Universal, 407513/2018-3).

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Edited by

Area Editor: Dr. Luiz Menini Neto

Publication Dates

  • Publication in this collection
    11 Apr 2022
  • Date of issue
    2022

History

  • Received
    24 Nov 2020
  • Accepted
    04 May 2021
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