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Rodriguésia

Print version ISSN 0370-6583On-line version ISSN 2175-7860

Rodriguésia vol.69 no.3 Rio de Janeiro July/Sept. 2018

http://dx.doi.org/10.1590/2175-7860201869336 

Original Papers

Amazon canga: the unique vegetation of Carajás revealed by the list of seed plants

Nara Furtado de Oliveira Mota1  2 

Mauricio Takashi Coutinho Watanabe1 

Daniela Cristina Zappi1 

Alice Lima Hiura1 

Julieta Pallos1 

Raquel Stauffer Viveros1 

Ana Maria Giulietti1 

Pedro Lage Viana3  4 

1Instituto Tecnológico Vale de Desenvolvimento Sustentável, R. Boaventura da Silva 955, 66055-090, Belém, PA, Brasil.

2Museu Paraense Emílio Goeldi, Coord. Botânica, Prog. Capacitação Institucional, Av. Perimetral 1901, 66077-830, Belém, PA, Brasil.

3Museu Paraense Emílio Goeldi, Coord. Botânica, Av. Perimetral 1901, 66077-830, Belém, PA, Brasil.

Abstract

The Floresta Nacional de Carajás (FLONA of Carajás) and the Parque Nacional dos Campos Ferruginosos (PNCF) are home to canga vegetation where 856 species distributed in 116 families of seed plants were recorded. The richest families were Poaceae (86), Fabaceae (65) and Rubiaceae (46) and the best represented habit was herbaceous. Two genera, 24 species and one subspecies are considered endemic from the canga of the studied area. Within the FLONA of Carajás, the Serra Norte was the best sampled site, and is home for 659 species of seed plants, while 545 species were recorded for the Serra Sul. Approximately 60% of the species documented, including the endemics, have no record within the PNCF. Through the taxonomic list here presented, it was possible to demonstrate considerable distinction between the canga of the Serra dos Carajás and the same formation in the Quadrilátero Ferrífero, in Minas Gerais, with little correspondence between these two lists and the one from Corumbá, Mato Grosso do Sul state. The richness and singularity of the regional flora, which includes many endemisms, associated to the current threats to these environments due to mineral exploration, point to the need for careful and consistent planning to ensure the conservation of the flora of the canga of Carajás.

Key words: Amazon; Angiosperms; endemism; Gymnosperms; floristics; species richness

Introduction

The Amazon covers an area of approximately 6 million square kilometers, comprising predominantly diverse types of rainforest (Pires & Prance 1985). In order to assess the diversity of plants found within this biome, a recent survey recorded over 14 thousand species of seed plants in lowland rainforest (Cardoso et al. 2017). Almost 12 thousand seed plants were recorded in the Brazilian Amazon (BFG 2015), considering all its diverse phytophysionomies that include, alongside the forests, several types of open vegetation. Meanwhile, a relatively low number of species was referred to these open areas, such as campinaranas (1.350 species), cerrado (876), highland rocky fields (683) and rock outcrops (625) (FBO 2020, under construction). These data, however, are underestimated as the Brazilian Amazon is still in need of floristic studies. The sheer size and access difficulties (Zappi et al. 2016) and the relatively recent history of the scientific investigations in the area result in a very low density of collections and, consequently, patchy knowledge of the plant diversity in the region (BFG 2015).

The Serra dos Carajás, in the state of Pará, is home to an open type of vegetation that is mostly formed by herbs and shrubs associated to iron-rich conglomerates that occur on the top of some local ranges. This vegetation, known as canga (Rizzini 1979; Secco & Mesquita 1983; Viana et al. 2016), comprises a peculiar flora with high number of endemic species. Plants growing there are adapted to extreme conditions of acid and nutrient poor soils (Nunes et al. 2015), high concentrations of heavy metals (Schettini et al. 2018), high temperatures and strong seasonality, with a clearly defined dry season (Mota et al. 2015).

The Serra dos Carajás spans between the municipality of São Félix do Xingu to the west and Curionópolis at the east, with the main plateaus found within Parauapebas and Canaã dos Carajás (Viana et al. 2016). Two conservation units include areas of canga within the last two municipalities: the Floresta Nacional de Carajás (FLONA Carajás), a sustainable use preservation area encompassing Serra Norte and Serra Sul, where mining activities occur, and the Parque Nacional dos Campos Ferruginosos (PNCF), a full protection area that includes the Serra do Tarzan and Serra da Bocaina (Zappi 2017).

The Flora of the canga of Carajás (FCC) project (Viana et al. 2016) began in 2015, with the objective of developing a detailed information base for the local flora, contributing to the knowledge of the Amazonian plants. This project culminated with the publication of four fascicles of monographs that provide detailed information regarding 164 families of land plants and 1,108 species that grow in the region of the Serra dos Carajás, including bryophytes, ferns and lycophytes, gymnosperms and angiosperms. The project aimed to monograph only native and or naturalized species found on the canga (sensuMota et al. 2015) from the FLONA Carajás (Serra Norte and Serra Sul) and the PNCF (Serra da Bocaina and Serra do Tarzan (Viana et al. 2016; Zappi 2017). However, some species that occur in the forest and/or on other rock formations (e.g., granitoid inselbergs) were included by some authors in order to increase the usefulness and scope of their monographs and to facilitate the identification of the species from the canga, considering the shortage of similar studies within the Brazilian Amazon (Viana et al. 2016).

Intensive collecting expeditions to the area carried out during the FCC Project were aimed at ensuring sufficient sampling for the authors to prepare their monographs. Images of the species in nature and associated material, including samples for micromorphology and tissues to extract DNA, destined to form a DNA bank for molecular studies at the Instituto Tecnológico Vale (ITV) were also targetted by the expeditions. The continued collecting activity even after the publication of some of the monographs has brought occasional new records of species to light, and these were included in the present list.

The results obtained by the FCC have contributed considerably to improve the knowledge of the floristic compostition of the canga of Carajás, as the last available compilation for this area (Silva 1991), counted with 231 species found in 144 genera and 57 seed plant families. Nowadays, the seed plants are represented by 120 families and 977 species monographed by a wide network of 131 plant specialists from Brazil and elsewhere. Some of the monographs, however, included species that were not recorded for the canga, and, as some new species records were made after the conclusion of some of the first monographs, it became necessary to prepare this updated compilation of the species that occur on the canga of the Serra dos Carajás.

The present work brings a list of seed plants from the FLONA of Carajás and the PNCF discriminated by area of occurrence, highlighting the endemic taxa found exclusively in the canga of these conservation units. The results allowed for a comparison of the seed plants from the studied area with other canga sites elsewhere in Brazil. This work focuses only on the seed plants while ferns and lycophytes (Salino et al. 2018) and bryophytes (Ilkiu-Borges & Oliveira-da-Silva 2018) were treated similarly in separate papers within this volume.

Material and methods

The study area was delimited at the onset of the FCC Project (Viana et al. 2016), that included the canga of the FLONA of Carajás (Serra Sul: blocks S11A, S11B, S11C, S11D; and Serra Norte: N1, N2, N3, N4, N5, N6, N7, N8) and PNCF (Serra do Tarzan, Serra da Bocaina), within the municipalities of Canaã dos Carajás and Parauapebas (Fig. 1). The plateaus of the study area add up to an area of approximately 120 km2 of canga vegetation calculated using Google Earth software, by delimiting polygones encompassing the rock outcrops that were visible on satellite imagery (Google Earth 2018).

Figure 1 Study area, indicating the canga plateaux sampled. In black, remaining canga areas; in red, suppressed canga areas (representation based on satelite image taken in December 2016, available on Google Earth). In grey, Floresta Nacional de Carajás. In green, Parque Nacional dos Campos Ferruginosos. 

During the years between 2015 and 2017, 30 collecting expeditions were organized to the Serra dos Carajás, covering all months of the year and resulting in 3,533 new plant samples that are deposited at the MG herbarium. This material has been added to the material already available at the start of the project, of around 3,300 specimens from the BHCB herbarium and another 6,000 from MG (Viana et al. 2016). These collections constitute the main source of data consulted by monograph authors. Some authors have also examined specimens from the HCJS, IAN, INPA, NY, RB and UB herbaria.

Collecting records from the canga were compiled from the published monographs - Rodriguésia 2016/67(5), 2017/68(3), 2018/69(1), 2018/69 (current) - including taxonomic information, type of environment and location of the specimens. The reference for the publications consulted for this end is cited on the heading for each plant family in Table 1. These data, together with collections made after the publications and named by specialists were organized as a database from where the floristic list presented here was extracted. This list is restricted to the plants that were recorded in the canga of the FLONA of Carajás and PNCF. Records made after the publications are accompanied by a voucher citation per species. The list excludes species that did not have at least one collection for specific areas of canga.

Table 1 List of species of seed plants occurring in the canga of the Serra dos Carajás. Caption: * nomenclatural update; + problem native; ++ invasive exotic; ® sample of DNA tissue; also occurs in the canga of Corumbá; □ also occurs in the canga of Quadrilátero Ferrífero; ST = Serra do Tarzan; SB = Serra da Bocaina; Ar = tree; Arb = shrub; Sb = subshrub; Li = liana; Er = herb; Par = parasite; Pal = palm; E-Sb = herb or subshrub; Ar-Arb = tree or shrub; Sb-Arb = subshrub or shrub; Vr = open rupestrian vegetation; Ff = Forest Formation; Vh = hydromorfic vegetation; Aa = disturbed areas. Endemic species of the canga of the Serra dos Carajás in bold. Only for the new records, subsequent to the publication of the floras, a voucher will be given after the species name. 

Family - monograph / species - Voucher Habit Habitat Distribution in the study area
Acanthaceae - Reis et al. (2017)
Justicia divergens (Nees) A.S.Reis, A.Gil & C.Kameyama Sb Vr, Ff N1, ST
J. potamogeton Lindau ® Sb Ff N8, S11D
J. riedeliana (Ness) V.A.W.Graham Sb Ff S11D
J. sprucei V.A.W.Graham ® Sb Vr N1, N2, N3, N4, N8
Justicia sp. 1 Sb Vr, Ff S11A, S11D, ST
Justicia sp. 2 Arb Vr, Ff S11A, S11B, S11D
Justicia sp. 3 Arb Vr, Ff N2, N3, N4, N5, S11A, S11B, S11D, ST, SB
Justicia sp. 4 Sb Vr, Ff ST
Justicia sp. 5 Sb Vr S11C, ST
Mendoncia aspera Ruiz & Pav. Li Vr, Ff N1, N3, ST, SB
Ruellia anamariae A.S.Reis, A.Gil & C.Kameyama Arb Vr, Ff N1, N2, N3, N4, N5
R. exserta Wassh. & J.R.I.Wood Li Vr, Ff N1, N2, N3
R. inflata Rich. ® Arb Vr, Ff N1, N3, N4, N5, S11D
R. wurdackii Wassh. ® Sb Vr, Ff S11B, ST
Alismataceae - Hall & Gil (2016)
Helanthium tenellum (Mart. ex Schult.f.) J.G.Sm. ® Er Vh N1, N3, N7, S11A, ST
Sagittaria rhombifolia Cham. Er Vh S11A, S11B, S11C,
Alstroemeriaceae - Koch (2016)
Bomarea edulis (Tussac) Herb. Li Ff N5, S11B, S11D
Amaranthaceae - Senna & Lima (2017)
Alternanthera dentata (Moench) Stuchlik ex R.E.Fr. Sb Vr N1, N5
A. tenella Colla + ® Sb Aa N4
Cyathula achyranthoides (Kunth) Moq. Sb Vr, Ff N4
C. prostrata (L.) Blume Sb Vr, Ff S11D
Anacardiaceae - Hall & Gil (2017)
Anacardium occidentale L. ® Ar Vr N1, N2, N3, S11A, S11B, ST
Tapirira guianensis Aubl. Ar Ff N4, S11D
Thyrsodium spruceanum Benth. Ar Ff N6
Annonaceae - Lobão (2016)
Guateria punctata (Aubl.) R.A.Howard Ar Ff N1, N4, S11D
Xylopia aromatica (Lam.) Mart. ® Ar Vr, Ff N1, N2, N3, N4, N5, N6, S11D, ST, SB
Apocynaceae - Fernandes et al. (2018)
Asclepias curassavica L. ++ ® Er Aa N5, S11D
Aspidosperma brasiliense A.S.S.Pereira Ar Vr, Ff S11C
A. multiflorum A.DC. Ar Vr, Ff S11D
A. subincanum Mart. ex A.DC. Ar Vr, Ff S11D
Blepharodon pictum (Vahl) W.D.Stevens ∆ □ Li Vr N1, S11A, S11B, S11C, S11D
Forsteronia affinis Müll.Arg. ® Li Vr, Ff N1, S11D
Forsteronia sp. Li Vr, Ff S11D
Hemipogon sprucei E.Fourn. ® Sb Vr N1, N3, N4, N5, N6, S11A, S11D
Lacmellea arborescens (Müll.Arg.) Markgr. Ar Vr, Ff N1, SB
Mandevilla scabra (Hoffmanns. ex Roem. & Schult.) K.Schum. ® Li Vr, Ff N1, N2, N3, N4, N5, N6, N7, S11A, S11B, S11C, S11D
M. tenuifolia (J.C.Mikan) Woodson ® Sb Vr N1, N2, N3, N4, N7, N8, S11A
Marsdenia bergii Morillo ® Li Vr, Ff N5, N7, S11A, S11B, S11D
Matelea microphylla Morillo Li Vr N1, N2, N3
Odontadenia nitida (Vahl) Müll.Arg. Li Vr N1, N4, N5, S11D
Prestonia ornata (Hoehne) J.F.Morales Li Vr N1, SB
P. quinquangularis (Jacq.) Spreng. Li Vr, Ff N1, N5
Secondatia densiflora A.DC. ® Li Vr, Ff N1, N3, N4, N5, N6, S11A, S11B, S11C, S11D
Tabernaemontana flavicans Will ex Roem. & Schult. ® Arb Vr, Ff S11B, S11D, SB
T. heterophylla Vahl Arb Vr, Ff S11D, ST
Tabernamonatana cf. linkii A.DC. Arb Vr, Ff S11B, S11D
T. macrocalyx Müll.Arg. ® Arb Vr, Ff N1, N3, N5, S11A, S11B, S11D
Apodanthaceae - Pastore et al. (2018)
Pilostyles blanchetii (Gardner) R.Br. Par Vr N1
Araceae - Coelho (2018)
Anthurium bonplandii G.S.Bunting Er Vr, Ff S11A, S11B, S11D
A. gracile (Rudge) Lindl. Er Vr, Ff S11A, S11C, S11D
A. kunthii Poepp. Er Ff S11B
A. lindmanianum Engl. ® Er Vr, Ff N1, N4, N5, S11B, S11C, S11D
A. sinuatum Benth. ex Schott Er Vr, Ff N3, S11A, S11B, S11C
Anthurium. sp. Er Vr, Ff N7
Dieffenbachia cf. seguine (Jacq.) Schott Er Ff S11D
Heteropsis oblongifolia Kunth Er Ff N1, N3, S11D
Philodendron blanchetianum Schott - Vasconcelos 764 ® Er Ff N5
P. carajasense E.G. Gonç. & A.J.Arruda Er Ff N1, N4, N7, S11A, S11D, SB
P. distantilobum K.Krause Er Ff N1, N3
P. solimoesense A.C.Sm. Er Ff N1, N8
P. wullschlaegelii Schott ® Er Vr, Ff N1, N4, S11B, SB
Philodendron sp. Er Ff N1, S11A
Spathiphyllum gardneri Schott Er Ff S11A, S11D
S. humboldtii Schott Er Ff Serra Norte
Wolffia brasiliensis Wedd. Er Ff Serra Sul
Arecaceae - Hiura & Rocha (2018)
Attalea maripa (Aubl.) Mart. Pal Ff N6
A. spectabilis Mart. Pal Ff N1
Mauritia flexuosa L.f. Pal Vh S11C, SB
Mauritiella armata (Mart.) Burret Pal Vh N5, SB
Oenocarpus distichus Mart. Pal Ff N1, N3
Syagrus cocoides Mart. ® Pal Vr, Ff Serra Norte
Aristolochiaceae - Abreu et al. (2018)
Aristolochia rugosa Lam. Li Vr S11D
Asteraceae - Cruz et al. (2016)
Ageratum conyzoides L. + □ Er Aa N3, N4, S11D, SB
Aspilia attenuata (Gardner) Baker ® Arb Vr N1, N4, N5, S11D
Bidens bipinnata L. + Arb Aa S11D
B. pilosa L. + □ Arb Aa S11D, ST
Calea caleoides (DC.) H.Rob. Arb Vr S11D
Cavalcantia glomerata (G.M.Barroso & R.M.King) R.M.King & H.Rob. Er Vr, Ff N1, N2, N3, N4, N5, N6, N8, S11A
C. percymosa R.M.King & H.Rob. ® Er Vr N7, N8, S11A, S11B, S11C, S11D
Chromolaena maximilianii (Schrad. ex DC.) R.M.King & H.Rob. ® Arb Ff N1, ST
Eclipta prostrata (L.) L. + ® Er Aa N3, S11D
Elephantopus mollis Kunt. + Sb Aa S11D
Emilia sonchifolia (L.) DC. + □ Er Aa N4, N5, S11A, S11D, ST, SB
Erechtites hieracifolius (L.) Raf. ex DC. + □ Er Vh, Aa N4, N5, S11D, ST, SB
Hebeclinium macrophyllum (L.) DC. Arb Ff N4
Ichthyothere terminalis (Spreng.) S.F.Blake ® Er Vr, Ff S11A, S11B, S11D, ST, SB
Ichthyothere sp. Arb Ff ST
Lepidaploa arenaria (Mart. ex DC.) H.Rob. ® Arb Vr N1, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST
L. paraensis (H.Rob.) H.Rob. ® Sb Vr N1, N3, N7, S11A, S11B, S11C, S11D, ST, SB
L. remotiflora (Rich.) H.Rob. Arb Ff N7, S11A, S11B, ST, SB
Lessingianthus monocephalus (Gardner) H.Rob. Arb Vr S11D
Mikania divaricata Poepp. Li Ff N1, S11A, S11B
M. psilostachya DC. Li Ff N4
M. micrantha Kunth Li Ff ST
Mikania sp. Li Ff N3
Monogereion carajensis G.M.Barroso & R.M.King ® Er Vr, Ff N1, N2, N3, N4, N5, N7, N8, S11A, S11B, S11C, S11D, ST, SB
Parapiqueria cavalcantei R.M.King & H.Rob. ® Er Vr, Ff N1, S11A, S11C
Pluchea sagittalis (Lam.) Cabrera Er-Sb Vh, Aa N1, N3, S11D
Porophyllum ruderale (Jacq.) Cass. + □ Arb Aa N1, N5, SB
Praxelis asperulacea (Baker) R.M.King & H.Rob. Er Vr S11A, S11B, S11C, S11D, SB
Pterocaulon alopecuroides (Lam.) DC. ® Sb Aa Serra Norte
Riencourtia pedunculosa (Rich.) Pruski ® Er Vr N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
Rolandra fruticosa (L.) Kuntze + ® Arb Ff, Aa N1
Tilesia baccata (L.) Pruski Arb Vr, Ff N1, S11C, ST, SB
Unxia camphorata L.f. Arb Vr, Ff N1, N2, N3, N4, N5, N7, N8, S11D, SB
Vernonanthura cf. brasiliana (L.) H.Rob. Ar Ff N1
Balanophoraceae - Meirelles (2016)
Helosis cayennensis (Sw.) Spreng. ® Par Ff N1, S11B, S11D, ST
Langsdorffia hypogaea Mart. Par Ff N1, N3, ST, SB
Begoniaceae - Kollman (2016)
Begonia guaduensis Kunth Arb Vr, Ff N3, N4
B. humilis Aiton ® Er Vr, Ff N1, N2, N3, N4, N5, S11A, S11B, S11C, S11D, SB
B. saxicola A.DC. Arb Vr, Ff N4, N5, N8, S11A, S11C, S11D
B. wollnyi Herzog Sb Vr, Ff N7, S11A, S11B, S11D
Bignoniaceae - Lohmann et al. (2018)
Amphilophium mansoanum (DC.) L.G.Lohmann Li Vr, Ff N8, S11A, S11B
A. rodriguesii (A.H. Gentry) L.G.Lohmann Li Vr, Ff N3, N4
Anemopaegma carajasense A.H.Gentry ex Firetti-Leggieri & L.G.Lohmann Arb Vr, Ff N1, N3, N4, N6, S11A, S11B, S11C, S11D
A. longipetiolatum Sprague Li Vr N1, N3, S11D
Bignonia corymbosa (Vent.) L.G.Lohmann Li Vr, Ff N1, N3, N4, N5, S11A, S11D, SB
Fridericia cinnamomea (DC.) L.G.Lohmann Li Ff N4, S11B
F. craterophora (DC.) L.G.Lohmann Arb Vr, Ff N3, N4
F. tuberculata (DC.) L.G.Lohmann Li Vr N7
Jacaranda ulei Bureau & K.Schum. Arb Vr, Ff N1, S11D, SB
Lundia densiflora DC. Li Vr N5
Pachyptera incarnata (Aubl.) J.N.C.Francisco & L.G.Lohmann Li Vr, Ff N2
Pleonotoma melioides (S.Moore) A.H.Gentry ® Li Vr, Ff S11D, ST
P. orientalis Sandwith Li Vr, Ff N1, N3, N8, S11A, S11B, S11C, S11D
Bixaceae - Fernandes-Junior & Gil (2017)
Cochlospermum orinocense (Kunth) Steud. Ar Ff N3, S11A, S11D
Bromeliaceae - Monteiro & Forzza (2016)
Aechmea angustifolia Poepp. & Endl. Er Vr N1, S11D
A. bromeliifolia (Rudge) Baker Er Vr, Ff S11D
A. castelnavii Baker ® Er Vr, Ff N1, S11A, S11B, S11D
A. mertensii (G.Mey.) Schult. & Schult.f. ® Er Ff S11B
A. tocantina Baker Er Ff N5, S11C, S11D
Ananas ananassoides (Baker) L.B.Sm. ® Er Vr, Ff N1, N7, S11B, S11C, S11D
Bromelia eitenorum L.B.Sm. Er Ff Serra Sul
Bromelia aff. lagopus Mez Er Ff N1
Dyckia duckei L.B.Sm. ® Er Vr N1, N2, N3, N4, N5, N6, S11A, S11C, S11D, SB
Pitcairnia burchellii Mez Er Vr N1, N4, SB
P. torresiana L.B.Sm. Er Ff N1, N4, N7, S11D
Tillandsia paraensis Mez Er Vr, Ff N4, S11A, S11C, S11D
T. streptocarpa Baker Er Vr, Ff S11A, S11B, S11C, S11D
Burmanniaceae - Giulietti (2016a)
Burmannia capitata (Walter ex J.F.Gmel.) Mart. ® Er Vh N2, N3, N7, S11A, S11B, ST
B. flava Mart. ® Er Vh N1, N2, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST
Burseraceae - Hiura & Watanabe (2017)
Protium pilosissimum Engl. ® Ar Ff N4, S11D
Cabombaceae - Lima & Gil (2016)
Cabomba furcata Schult. & Schult.f. Er Vh N1, N5, S11A, S11B, S11C, ST
C. haynesii Wiersema Er Vh N1, N4
Cactaceae - Zappi & Taylor (2017)
Cereus hexagonus (L.) Mill. ® Arb Vr N1, N4, N7, S11A, S11B, S11C, S11D, SB
Epiphyllum phyllanthus (L.) Haw. ® Sb Ff N1, N8, S11D
Calophyllaceae - Marinho & Amorim (2016a)
Calophyllum brasiliense Cambess. ® Ar Vh, Ff S11A, S11B
Cannabaceae - Viana & Gil (2018)
Trema micrantha (L.) Blume ® Ar Ff N5
Caryocaraceae - Nunes & Gil (2016)
Caryocar villosum (Aubl.) Pers. Ar Ff N5
Caryophyllaceae - Lovo & Devechi (2018)
Drymaria cordata (L.) Willd. ex Roem. & Schult. + Er Aa N3
Celastraceae - Lombardi & Biral (2016)
Anthodon decussatum Ruiz & Pav. Li Ff N1, S11C, S11D, ST
Cheiloclinium hippocrateoides (Peyr.) A.C.Sm. Li Ff Serra Norte
Maytenus guyanensis Klotzsch ex Reissek Arb Ff N1, N3, S11D
Tontelea laxiflora (Benth.) A.C.Sm. Li Ff N4
Chrysobalanaceae - Sothers & Prance (2018)
Hirtella hispidula Miq. Ar Ff N6
H. pilosissima Mart. & Zucc. ® Ar Ff S11A, S11B, S11C, ST
H. racemosa Lam. Ar Ff N1, N2, S11C
Leptobalanus octandrus (Hoffmanns. ex Roem. & Schult.) Sothers & Prance Ar Ff S11C
Moquilea egleri (Prance) Sothers & Prance ® Ar Ff N1, N3, S11A, S11B
Cleomaceae - Soares-Neto (2017)
Melidiscus giganteus (L.) Raf. Arb Ff SB
Tarenaya spinosa (Jacq.) Raf. + Sb Aa N5
Clusiaceae - Alencar & Marinho (2017)
Clusia nemorosa G.Mey Ar Ff S11A, S11B, S11C, S11D
C. panapanari (Aubl.) Choisy ® Ar-Arb Ff N4, N5, ST
Clusia aff. weddelliana Planch. & Triana ® Ar-Arb Ff N1, N4, S11A, S11B, S11C, S11D
Garcinia madruno (Kunth) Hammel Ar Ff S11A, S11B
Symphonia globulifera L.f. Ar Ff S11A, S11B, S11C
Combretaceae - Praia (2017)
Combretum laxum Jacq. ® Li Vr, Ff N1, SB
Commelinaceae - Aona et al. (2016)
Commelina benghalensis L. ++ □ Er Aa S11D
C. erecta L. ∆ □ Er Vh, Ff S11D
C. obliqua Vahl Er Vh, Ff S11D
C. rufipes var. glabrata Seub. Er Ff S11A
Dichorisandra hexandra (Aubl.) C.B.Clarke ® Er Vr, Vh, Ff N4, N5, S11A, S11C, S11D, SB
D. villosula Mart. ex Schult & Schult.f. Er Ff N5
Tipogandra diuretica (Mart.) Handlos Er Vh N5
Connaraceae - Pastore & Vasconcelos (2017)
Connarus perrotteii (DC.) Planch. Ar Ff N2, N4, S11B, S11C
Rourea doniana Baker ® Li Ff N1, N5, SB
R. ligulata Baker ® Li Vr, Ff N1
Convolvulaceae - Simão-Bianchini et al. (2016)
Aniseia martinicensis (Jacq.) Choisy * Li Vr,Vh, Ff N5, SB
Cuscuta insquamata Yunck. ® Par Vr, Vh, Ff N1, N2, N3, N4, N5, N6, N7, SB
Evolvulus filipes Mart. ® ∆ □ Er Vr, Vh N5, N7, ST, SB
E. lithospermoides Mart. ∆ □ Er Vr N1, S11A
Ipomoea asplundii O'Donell ® Li Vr, Ff N3, N6, N7, ST
I. cavalcantei D.F.Austin ® Li Vr, Ff N1, N2, N3, N4, N5
I. decora Meisn. Li Vr, Ff N4, S11A, S11B, S11C, S11D, ST
I. goyazensis Gardner ® Li Vr, Ff N1, N5, S11D, ST
I. hederifolia L. Li Vr,Ff, Aa N5
I. marabensis D.F.Austin & Secco ® Li Vr, Ff N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
I. maurandioides Meisn.* ® Li Vr N1, N2, N3, N4, N5, N7, S11A, S11B, S11C, S11D, ST
I. minutiflora (M.Martens & Galeotti) House Li Vr, Ff S11D
I. procumbens Mart. ex Choisy - Zappi 3510 + Li Aa S11A, S11B
I. ramosissima (Poir.) Choisy Li Vr, Ff N4, N5, SB
I. reticulata O'Donell - Sperling 5680 Li Vr, Ff N5
I. setifera Poir. - Zappi 3561+ ® Li Aa S11D
Jacquemontia tamnifolia (L.) Griseb. ® Li Vr, Ff N1, N3, N4, N5, N6, N7, S11A, S11B, S11C, S11D, ST
Distimake macrocalyx (Ruiz & Pav.) A.R.Simões & Staples * Li Vr, Ff N4, N5, ST, SB
Operculina hamiltonii (G.Don) D.F.Austin & Staples Li Ff N7, S11D
Turbina cordata (Choisy) D.F.Austin & Staples Li Ff N3
Cordiaceae - Watanabe et al. (2017)
Cordia exaltata Lam. ® Arb Ff N1, N4, N5, SB
C. nodosa Lam. Arb Ff N1, S11D, SB
Varronia multispicata (Cham.) Borhidi + Arb Aa N4
Costaceae - André & Sousa (2018)
Costus lasius Loes. Er Ff N1, S11D
C. scaber Ruiz & Pav. ® Er Ff N5, S11D
C. spiralis (Jacq.) Roscoe Er Ff N7, S11D
Cucurbitaceae - Gomes-Klein et al. (2016)
Cayaponia tayuya (Vell.) Cogn. Li Vr, Ff N4, SB
Gurania bignoniacea (Poepp. & Endl.) C.Jeffrey ® Li Ff N1, N3, S11D
G. sinuata (Benth.) Cogn. Li Ff N1, N2, N6, S11A, ST
G. subumbellata (Miq.) Cogn. ® Li Ff ST
Helmontia cardiophylla Harms Li Ff N5, S11B, ST, SB
Melothria pendula L. Li Vh N1
Cyperaceae - Nunes et al. (2016)
Becquerelia cymosa Brongn. Er Vr,Vh N5
Bulbostylis cangae C.S.Nunes & A.Gil *® Er Vr N1, S11A, ST
B. conifera (Kunth) C.B.Clarke ® ∆ □ Er Vr N1, N2, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
B. lagoensis (Boeckeler) Prata & M.G.López Er Vr,Vh N4, N5, S11A, S11B, S11C, S11D, ST, SB
Bulbostylis sp. Er Vh S11D
Cyperus aggregatus (Willd.) Endl. +∆ □ Er Vh,Ff, Aa N1, N5, S11B, S11C, S11D, SB
C. distans L.f. Er Vr, Ff N3, N4
C. haspan L. ® Er Vr,Vh N1, N2, N3, N4, N5, N8, S11B, S11C, S11D, ST, SB
C. hortensis (Salzm. ex Schltdl.) Dorr* Er Vr N1, N5, S11D
C. laxus Lam. ∆ □ Er Vr N1, S11A, S11B, S11D, ST, SB
C. luzulae (L.) Rottb. ex Retz. ® Er Vr,Vh N2, N5, ST
C. polystachyos Rottb.* Er Vh N3, N4, S11C
C. sesquiflorus (Torr.) Mattf. & Kük * ® ∆ □ Er Vr, Vh S11C, S11D
C. sphacelatus Rottb. + Er Vr, Aa N1, N3, N4, N5, S11A, S11B
C. subsquarrosus (Muhl.) Bauters * Er Vr N3
C. surinamensis Rottb. +∆ Er Vr, Aa N1, S11D
Cyperus sp. 1 Er Vr S11D
Cyperus sp. 2 Er Vr ST
Eleocharis acutangula (Roxb.) Schult. Er Vh N1, N4, S11B, S11D, SB
E. ayacuchensis S.González & Reznicek Er Vh N1, N2, N4, N5, S11D, ST, SB
E. endounifascis Hinchliff & Roalson Er Vh N1, S11B, S11C, SB
E. flavescens (Poir.) Urb. Er Vh N1, N4, N5, S11A, S11B, S11C, S11D, ST, SB
E. geniculata (L.) Roem. & Schult. Er Vh N5
E. pedrovianae C.S.Nunes, R.Trevis. & A.Gil ® Er Vh N1, N3, N4, N7, N8, S11A, S11C, S11D, ST, SB
E. plicarhachis (Griseb.) Svenson Er Vh N1, N3
Fimbristylis dichotoma (L.) Vahl Er Vr N1, ST
F. quinquangularis (Vahl) Kunth * Er Vr N4, S11D, ST
Hypolytrum paraense M.Alves & W.W.Thomas ® Er Ff N1, N2, N3, N5, S11D
Lagenocarpus verticillatus (Spreng.) T.Koyama & Maguire Er Vr S11A, S11B, S11C
Rhynchospora acanthoma A.C.Araújo & Longhi-Wagner Er Vr N1, N3, N4, N5, N6, N7, N8, SB
R. barbata (Vahl) Kunth ® Er Vr, Vh N1, N2, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST, SB
R. candida (Nees) Boeckeler ® Er Vh N3, N5, N6
R. corymbosa (L.) Britton Er Vr, Vh N2, N4, N8, S11C, S11D
R. divaricata (Ham.) M.T.Strong Er Vr N1, N2, N3, N4, N5, N7, S11A
R. eximia (Nees) Boeckeler Er Vr, Vh N3, N4, N5, N7, ST, SB
R. filiformis Vahl Er Vr, Vh N1, N2, N3, N4, N5, N7, S11A, S11B, S11C, S11D, ST, SB
R. holoschoenoides (Rich.) Herter Er Vr, Vh N1, N3, N4, N6, S11A, S11B, S11C, S11D, ST, SB
R. pubera (Vahl) Boeckeler Er Vr, Aa N1, N2, N3, N4, N5, S11A, S11B, S11C, S11D, ST, SB
R. rugosa (Vahl) Gale Er Vh N1, N3, S11D
Rhynchospora sp. 1 Er Vr, Vh ST, SB
Rhynchospora sp. 2 Er Vr, Vh N1, N2, N3, N4, N5, N6, S11A, S11C, S11D, ST, SB
Scleria cyperina Willd. ex Kunth Er Vr, Vh S11A, S11B
S. microcarpa Nees ex Kunth Er Vh N1, N3, N5, N8, S11A, S11D, ST, SB
S. secans (L.) Urb. ® Er Vr, Ff N6, S11A, S11B, S11D, SB
S. verticillata Muhl. ex Willd. Er Vr, Vh N2, N3, S11B, ST, SB
Dilleniaceae - Zappi (2018a)
Davilla cearensis Huber Li Vr, Ff N1, N3
D. rugosa Poir. Li Vr, Ff N2
Doliocarpus brevipedicellatus Garcke Li Ff N2, S11A
Dioscoreaceae - Fraga & Sousa (2018)
Dioscorea pilosiuscula Bertero ex Spreng. Li Ff N1
D. piperifolia Humb. & Bonpl. ex Willd. Li Vr, Ff N1, N8
D. planistipulosa Uline ex R.Kunth. Li Vr S11C
Droseraceae - Mota (2017a)
Drosera sessilifolia A.St.-Hil. Er Vh S11D
Eriocaulaceae - Watanabe et al. (2017)
Eriocaulon carajense Moldenke ® Er Vr, Vh N1, N3, N4, N6, S11A, S11D, SB
E. cinereum R.Br. ® Er Vh N1, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
E. tenuifolium Klotzsch ex Körn. Er Vh S11A
E. aff. setaceum L. ® Er Vh N1, N3, N4, N6, N8, S11A, S11B, S11C, S11D, ST
Paepalanthus fasciculoides Hensold ® Er Vr, Ff N1, N4, N5, N6, N8, S11A, S11B, S11C, S11D, SB
Syngonanthus caulescens (Poir.) Ruhland ® Er Vh S11D,
S. discretifolius (Moldenke) M.T.C.Watan. ® Er Vh N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, SB, ST
S. heteropeplus (Körn.) Ruhland ® Er Vh N1, S11A, S11D
S. simplex (Miq.) Ruhland ® Er Vh N5
S. aff. saxicola (Koern.) Trovó & Stützel ® Er Vh N1
Erythroxylaceae - Costa-Lima & Loiola (2018)
Erythroxylum carajasense (Plowman) Costa-Lima * ® Arb Vr, Ff N1, N2, N3, N4, N5, N6, N8, SB
E. citrifolium A.St.-Hil. Arb Vr, Ff N1, N2, N4, N5, S11A, S11B, S11C, S11D
E. leptoneurum O.E.Schulz Arb Ff S11C, S11D
E. mucronatum Benth. Arb Ff N4, N5
E. nelson-rosae Plowman ® Arb Vr N1, N2, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST, SB
E. rufum Cav. Ar-Arb Vr, Ff N7, S11B
E. squamatum Sw. ® Arb Vr, Ff N4, N5, S11C
E. subracemosum Turcz. Arb Vr, Ff S11D
Euphorbiaceae - Costa et al. (2018)
Alchornea discolor Poepp. ® Ar Vr, Ff N1, N5, S11D, ST
Aparisthmium cordatum (A.Juss.) Baill. Ar Ff N1, N5, N6, ST
Astraea paulina Didr. Er Vr,Ff, Aa N1, N3, N4, S11A, S11B, S11C, S11D
Croton aff. subferrugineus Müll.Arg. ® Arb Vr N1, N2, N4, N5, N7, N8, S11A, S11B, S11C, ST
C. cajucara Benth. Ar-Arb Vr, Ff N5
Croton sp. Arb Vr N1, N2, N3, S11A, S11B, S11C, S11D, ST, SB
Mabea angustifolia Spruce ex Benth. ® Ar-Arb Ff N1, N5, S11D
Manihot marajoara Chermont de Miranda ex Huber emend. Secco & Costa Arb Ff S11B, S11D, ST
M. quinquepartita Huber ex D.J.Rogers & Appan Arb Vr, Ff N1, N3, N4, N5, S11C, ST
M. tristis subsp. surumuensis (Ule) Rogers & Appan ® Arb Ff S11D, ST
Sapium argutum (Müll.Arg.) Huber Ar-Arb Vr N1, N4
Fabaceae - Mattos et al. (2018)
Abrus melanospermus subsp. tenuiflorus (Spruce ex Benth.) D.Harder ® Li Vr, Ff N1, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, SB
Aeschynomene americana Hard glandulosa (Poir.) Rudd Sb Ff, Aa N1, N5, S11D
A. filosa Mart. Sb Vh SB
A. sensitiva var. hispidula (Kunth) Rudd Sb Vh N4, N7, S11A, S11D
Aeschynomene sp. Sb Vr, Vh N1, N2, N3, N4, N5, N7, N8, S11A, S11B, S11D, ST
Anadenanthera peregrina (L.) Speg. Ar Ff N1
Ancistrotropis peduncularis (Kunth) A.Delgado ® Li Vr, Ff N1, N5, N8, S11A, S11B, S11C
Bauhinia longicuspis Benth. Ar-Arb Vr, Ff N8, S11B
B. longipedicellata Ducke Ar-Arb Vr, Ff N1, N3
B. pulchella Benth. ® Arb Vr, Ff N1, N3, N4, N5, N6, S11C, S11D, ST, SB
Calopogonium mucunoides Desv. + Li Aa N1, N5, S11D
Camptosema ellipticum (Desv.) Burkart ® Li Vr, Ff N1, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST, SB
C. carajasense Cavalcante ® Li Vr, Vh, Ff N1, N4, N5, S11A, S11B, SB
Centrosema grazielae V.P.Barbosa Li Vr, Vh, Ff N1, N3, N5, S11B
Chamaecrista desvauxii var. langsdorfii (Kunth ex Vogel) H.S.Irwin & Barneby Arb Vr N1, S11D, SB
C. desvauxii var. mollissima (Benth.) H.S.Irwin & Barneby Arb Vr S11A, S11B, S11C, S11D, ST
C. diphylla (L.) Greene Er Vr, Ff N1, N4, N5
C. flexuosa L. Arb Vr, Vh N1, N2, N3, N4, N5, N8, S11A, S11D
C. nictitans subsp. patellaria (DC. ex Collad.) H.S.Irwin & Barneby + □ Arb Ff, Aa N4, N5
C. rotundifolia (Pers.) Greene + □ Arb Aa N5
C. trichopoda (Benth.) Britton & Rose ex Britton & Killip Er Vr, Vh N1, N3, N4, N8, S11B
Chloroleucon acacioides (Ducke) Barneby & J.W.Grimes Ar-Arb Vr, Ff Serra Norte, Serra Sul
Clitoria falcata Lam. ® Li Vr, Vh, Ff N1, N3, N4, N5, N6, N7, S11A, S11B, S11C, S11D, SB
Copaifera martii Hayne ® Ar Ff N2, N4, N5, N8, S11A, S11B, ST
Crotalaria maypurensis Kunth ® Sb Vr, Vh, Ff N1, N2, N4, N5, N6, N8, S11A, S11B, S11C, S11D
Dalbergia spruceana Benth. Ar Ff S11C, SB
D. subcymosa Ducke Li Ff N3
Deguelia amazonica Killip ® Li Vh, Ff N3, N8, ST
Desmodium barbatum (L.) Benth. + Er Vr, Ff N1, N5
D. incanum (Sw.) DC. + □ Sb Aa N1, N5
D. triflorum (L.) DC. + Sb Aa S11D
Dimorphandra gardneriana Tul. Ar Ff Serra Sul
Dioclea apurensis Kunth ® Li Vr, Vh N1, N2, N3, N4, N5, N8, S11A, S11C, S11D, ST
D. bicolor Benth. ® Li Vr, Ff N1, N3, S11A, ST, SB
D. megacarpa Rolfe Li Vr, Ff ST
Enterolobium schomburgkii (Benth.) Benth. Ar Vr, Ff S11B, S11C
Galactia jussiaeana Kunth ® Sb Vr, Ff N1, N2, S11A, S11B, S11C, S11D, ST, SB
Inga alba (Sw.) Willd. Ar Vr, Vh, Ff S11C
I. capitata Desv. Ar Ff N5, S11C
I. thibaudiana subsp. thibaudiana DC. Ar Ff S11C, S11D, SB
Machaerium acutifolium Vogel Ar Ff Serra Sul
Mimosa acutistipula var. ferrea Barneby ® Ar-Arb Vr, Ff N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11D, ST, SB
M. camporum Benth. Er Vr, Aa N1, N5
M. candollei R.Grether Er Vr, Aa N1, N5
M. guilandinae var. spruceana (Benth.) Barneby Li Vr, Vh N1, N4
M. pigra var. pigra L. Arb Vh, Aa N1
M. piresii Barneby ® Arb Vr, Vh S11A
M. pudica var. hispida Brenan+ Arb Aa N1, N5
M. setosa var. paludosa (Benth.) Barneby Arb Vr, Aa N1, N4, N5
M. skinneri var. carajarum Barneby ® Er Vh N1, N2, N3, N4, N5, N8, S11A, S11B, S11C, S11D, SB
M. somnians var. viscida (Willd.) Barneby ® Arb Vr, Ff N1, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST
M. xanthocentra var. mansii (Mart.) Barneby ® Sb Vr, Vh, Ff N1, N3, N4, N5, N7, N8, S11A, S11D, ST
Parkia platycephala Benth. ® Ar-Arb Vr, Ff N1, S11A, SB
Periandra coccinea (Schrad.) Benth. Li Vr, Ff N1, N3, N4, N5, N6, N7, N8, S11C, S11D, ST, SB
P. mediterranea (Vell.) Taub. ® Arb Vr, Vh N1, N2, N3, N4, N5, N6, N8, S11A, S11C, S11D, ST, SB
Senna macranthera (DC. ex Collad.) H.S.Irwin & Barneby Arb Vr, Aa S11B, S11D, ST
S. pendula (Willd.) H.S.Irwin & Barneby Arb Ff, Aa N5, S11B, ST
S. quinquangulata (Rich.) H.S.Irwin & Barneby Arb Ff, Aa N4
S. silvestris (Vell.) H.S.Irwin & Barneby Arb Aa N4, Serra Sul, ST
Stryphnodendron pulcherrimum (Willd.) Hochr. Ar Ff Serra Norte, S11C, ST
Stylosanthes capitata Vogel ® Sb Vr, Ff, Aa N1, S11D
S. hispida Rich. Sb Vr, Vh, Ff N1, N2, N4, N8, ST, SB
S. humilis Kunth Sb Vr, Ff, Aa N1, S11D
Tachigali vulgaris L.G.Silva & H.C.Lima Ar Vr, Ff N1, N5, N8, SB
Zornia latifolia Sm. Sb Vr, Ff N1, N4, N5
Gentianaceae - Guimarães et al. (2018)
Chelonanthus purpurascens (Aubl.) Struwe, S.Nilsson & V.A.Albert ® Sb Vr N1, N2, N3, S11A, S11B, S11C, S11D, ST
C. viridiflorus (Mart.) Gilg Sb Vh N1, N3, N4
Curtia tenuifolia (Aubl.) Knobl. Er Vh N1, N2
Schultesia benthamiana Klotzsch ex Griseb. ® Er Vr, Vh N1, N2, N3, N4, N5, N8, S11A, S11B, S11C, S11D, ST, SB
Gesneriaceae - Chautems et al. (2018)
Diastema sp. Er Ff S11C
Goyazia rupicola Taub. Er Vr, Ff N1, S11D
Mandirola rupestris (Gardner) Roalson & Boggan Er Vr S11C
Nautilocalyx kohlerioides (Leeuwenb.) Wiehler Li Ff ST
Phinaea albolineata (Hook.) Benth. ex Hemsl. Er Ff N3, S11A, S11B, S11C
Sinningia minima A.O.Araujo & Chautems ® Er Ff N1, N3, N4, N5, N8, S11A, S11B, S11C, S11D
Gnetaceae - Mota & Giulietti (2016a)
Gnetum nodiflorum Brongn. ® Li Ff N1, N2, N4, N8
Heliotropiaceae - Watanabe & Hiura (2018)
Euploca humistrata (Cham.) J.I.M.Melo & Semir Er Vh SB
E. lagoensis (Warm.) Diane & Hilger Er Vh N5, SB
Humiriaceae - Antunes & Holanda (2018)
Sacoglottis guianensis Benth. Ar Ff N4
S. mattogrossensis Malme ® Ar Ff S11D, SB
Hydrocharitaceae - Hall & Gil (2016)
Apalanthe granatensis (Humb. & Bonpl.) Planch. Er Vh N5, S11A
Ottelia cf. brasiliensis (Planch.) Walp. Er Vh N1
Hypericaceae - Marinho et al. (2017)
Vismia cayennensis (Jacq.) Pers. Ar Ff S11C, S11D,
V. gracilis Hieron. ® Ar Ff N1, N4
Vismia cf. schultesii N.Robson ® Arb Ff N1, N2, N3, S11C, S11D
Iridaceae - Damasceno & Gil (2016)
Cipura xanthomelas Maxim ex Klatt ® Er Vr S11A, S11B, S11C, S11D, ST
Lacistemataceae - Marinho & Amorim (2016b)
Lacistema aggregatum (P.J.Bergius) Rusby Arb Ff S11D
Lamiaceae - Harley (2016)
Aegiphila integrifolia (Jacq.) Moldenke ® Ar-Arb Ff N2, N3, N4, N5, N7, N8, ST
A. racemosa Vell. Li Ff N1, N5
Amasonia campestris (Aubl.) Moldenke ® Sb Ff N1, N2, N3, N4, N5
A. hirta Benth. Sb Ff N1, N3, N8
A. lasiocaulos Mart. & Schauer ex Schauer Sb Ff N1, N4, N5, ST, SB
Cantinoa americana (Aubl.) Harley & J.F.B.Pastore + Er Aa SB
C. mutabilis (Rich.) Harley & J.F.B.Pastore Er Aa ST, SB
Hyptis atrorubens Poit. Er Vr, Vh, Ff N1, N5, S11B, S11D, ST
H. brevipes Poit. Er Vh, Ff, Aa ST, SB
H. parkeri Benth. ® Er Vh N3, N4, N5, ST
H. recurvata Poit. ® Er Vr, Vh N1, SB
Leonotis nepetifolia (L.) R.Br. ++ Er Aa N5
Marsypianthes chamaedrys (Vahl) Kuntze Er Vr, Aa N1, N3, N5, N7, S11D, SB
Mesosphaerum pectinatum (L.) Kuntze + Er Aa N1, N3, N4, S11D
M. suaveolens (L.) Kuntze + Sb Aa N8, ST, SB
Vitex triflora Vahl ® Ar-Arb Vr, Ff N1, N4, N5, S11A, S11B, S11D, ST, SB
Lauraceae - Moraes (2018)
Cassytha filiformis L. Par Vr N1, N4, N5, S11C, S11D
Mezilaurus itauba (Meisn.) Taub. ex Mez Ar Vr, Ff N1, N3, N4, N5, S11A, S11D
Ocotea camphoromoea Rohwer Ar Ff N1, N2, S11A, S11D
Ocotea leucoxylon (Sw.) Laness. Ar Ff S11C
Ocotea puberula (Rich.) Nees Ar Ff S11D
Lentibulariaceae - Mota & Zappi (2018)
Genlisea filiformis A.St.-Hil. Er Vh N3, N7, S11D
Utricularia amethystina Salzm. ex A.St.-Hil. & Girard Er Vh N3, N7, S11D, SB
U. breviscapa C.Wright ex Griseb. Er Vh N6
U. calycifida Benj. ® Er Vh N1, N2, N5
U. costata P.Taylor Er Vh N1, N3, N5, S11B, ST, SB
U. gibba L. Er Vh N1, S11D, SB
U. neottioides A.St.-Hil. & Girard ® Er Vh N1, N3, N4, N6, S11A, S11C, S11D, ST
U. nigrescens Sylvén Er Vh N1, N3, S11D
U. physoceras P.Taylor ® Er Vh N1, N2, N3, N4, N5, N6, N7, N8, S11B, S11C, S11D, ST, SB
U. pusilla Vahl ® Er Vh N3, N4, N5, N6, S11D
U. subulata L. Er Vh N1, N2, N3, N4, N5, N7, N8, S11A, S11B, S11C, S11D, ST
U. trichophylla Spruce ex Oliv. Er Vh S11A, S11B
Linderniaceae - Scatigna & Mota (2016)
Lindernia brachyphylla Pennell ® Er Vh N1, N2, N3, N5, N6, N7, S11B, S11D, ST, SB
L. crustacea (L.) F.Muell. Er Vr N4, N5, N7, N8, S11D, ST, SB
L. diffusa (L.) Wettst. Er Vh N4
Loganiaceae - Zappi & Setubal (2016)
Spigelia flemmingiana Cham. & Schltdl. Er Aa N4, N8
Strychnos cogens Benth. Er Vr, Ff ST
Loranthaceae - Caires (2018)
Oryctanthus alveolatus (Kunth) Kuijt Par Vr, Ff N3, N6
O. florulentus (Rich.) Tiegh. Par Vr, Ff S11C
Passovia disjectifolia (Rizzini) Kuijt Par Ff S11A
P. pedunculata (Jacq.) Kuijt ® Par Vr N1, N3, N4, N6
P. pyrifolia (Kunth) Tiegh. Par Vr, Ff SB
Peristethium reticulatum (Rizzini) Caires Par Vr, Ff S11D, SB
Psittacanthus eucalyptifolius (Kunth) G.Don Par Ff N3, N4, N5, N6
Struthanthus marginatus (Desr.) G.Don Par Vr, Ff N4, S11A, S11B, S11D
S. polyrrhizus Mart. Par Vr S11D
Lythraceae - Cavalcanti et al. (2016)
Cuphea annulata Koehne ® Sb-Arb Vr, Ff N1, N3, N4, N5, N7, N8, S11A, S11B, S11C, S11D, ST
C. carajasensis Lourteig ® Sb Vr N1, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
C. carthagenensis (Jacq.) J.F.Macbr. + □ Sb Aa N1, N4, N5, S11D
Malpighiaceae - Amorim et al. (2018)
Banisteriopsis malifolia (Nees ex Mart.) B.Gates ® Li Vr N2, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST
B. muricata (Cav.) Cuatrec. ® Li Vr, Ff N1
B. stellaris (Griseb.) B.Gates ® Li Vr N1, N3, N4, N5, S11A, S11B, S11C, S11D, ST, SB
Byrsonima chrysophylla Kunth ® Ar Vr N1, N2, N4, N5, N8, S11C, S11D, SB
B. crispa A.Juss. Ar Vr, Ff S11D
B. stipulacea A.Juss. ® Ar Ff N1
Coleostachys genipifolia A.Juss. Arb Ff N1, N4
Diplopterys pubipetala (A.Juss.) W.R.Anderson & C.Davis ® Li Vr, Ff N1, N2, N3, N5, S11B, S11C, ST
Heteropterys nervosa A.Juss. ® Li Vr N1, S11A, S11B, S11C, S11D,
H. trigoniifolia A.Juss. Li Vr, Ff N4, ST
Niedenzuella acutifolia (Cav.) W.R.Anderson Li Vr, Ff S11B, S11D
Peixotoa reticulata Griseb. Arb Vr N7, ST
Stigmaphyllon paraense C.E.Anderson Li Vr, Ff N4, SB
Malvaceae - Fernandes-Júnior & Cruz (2018)
Eriotheca globosa (Aubl.) A.Robyns Ar Vr N1
Helicteres brevispira A.St.-Hil. Arb Vr N1
H. eitenii Leane Arb Vr N1
H. guazumifolia Kunth Arb Vr SB
Hibiscus furcellatus Lam. Arb Vh SB
Melochia arenosa Benth. Arb Vr, Vh N6, S11C, ST, SB
M. melissifolia Benth. Arb Vr, Vh N1, N2, N3, N4, N8
M. splendens A.St.-Hil. & Naudin. Arb Vh, Aa N1, N2, N3, N4, N5, N6, N7, S11A, S11B, S11D, ST
Pavonia malacophylla (Link & Otto) Garcke Arb Ff SB
Pseudobombax longiflorum (Mart.) A.Robyns Arb Vr S11D
P. marginatum (A.St.-Hil., Juss. & Cambess.) A.Robyns Arb Vr ST
Sida linifolia Juss. ex Cav.∆ □ Sb Vr ST
S. rhombifolia L. + □ Sb Vr, Aa N3, N4
S. tuberculata R.E.Fr. Sb Vr SB
S. urens L. Sb Vr, Ff N4
Urena lobata L. + Arb Aa N1
Waltheria indica L. + □ Arb Aa N1, N4, ST
Marantaceae - Saka (2017)
Maranta ruiziana Körn. Er Ff N4
Monotagma plurispicatum (Körn.) K.Schum. Er Ff N1, N4, N5, N7, S11A, S11B, S11D
Marcgraviaceae - Viana & Cruz (2017)
Norantea guianensis Aubl. ® Li Vr N1, N4, S11A, S11C, S11D
Souroubea guianensis Aubl. Li Ff N1
Mayacaceae - Mota e Koch (2016)
Mayaca fluviatilis Aubl. ® Er Vh N1, N3, N4, N5, S11A, S11B, S11D, SB
M. kunthii Seub. ® Er Vh N1, N3, N7, S11D, SB
M. longipes Mart. ex Seub. ® Er Vh N4, N5
Melastomataceae - Rocha et al. (2017)
Aciotis acuminifolia (Mart. ex DC.) Triana Er Vr, Vh N1, N2, N3, N4, S11A, S11B, S11D, SB
Acisanthera crassipes (Naudin) Wurdack ® Er Vh N1, N3, ST, SB
Bellucia egensis (DC.) Penneys, Michelangeli, Judd, & Almeda Ar Ff SB
Brasilianthus carajensis Almeda & Michelangeli ® Er Vr, Vh N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
Clidemia capitellata (Bonpl.) D.Don Arb Ff N1, N4
C. hirta (L.) D.Don Arb Vr, Ff S11A, S11D
C. microthyrsa R.O.Williams Arb Ff S11B, S11D
Desmoscelis villosa (Aubl.) Naudin Arb Vh N3
Henriettea ramiflora (Sw.) DC. Ar Vh, Ff N5, S11D
Leandra micropetala (Naudin) Cong. Arb Ff N1, N5, S11A, S11B, S11D
Macairea radula (Bonpl.) DC. Arb Vr, Vh S11D
Miconia affinis DC. Ar Vr, Ff S11B, S11C, S11D
M. alata (Aubl.) DC. Arb Vr, Ff N3, S11D
M. albicans (Sw.) Steud. Arb Vr, Vh N1, S11D
M. alternans Naudin Arb Vh N3, N4, S11C, S11D, ST, SB
M. bracteata (DC.) Triana Arb Ff N1
M. chrysophylla (Rich.) Urb. Ar Vr, Ff N2, N5, S11D
M. ciliata (Rich.) DC. Arb Vr N1, N4
M. cuspidata Mart. ex Naudin Ar Vr, Ff N1, N4, N5, S11A, S11D
M. elata (Sw.) DC. Ar Vr, Vh N1, N5
M. heliotropoides Triana Arb Ff N1, N7, S11C, S11D
M. holosericea (L.) DC. Ar Vr, Vh, Ff N3, N4, N5, N7, N8, S11A, S11B, S11C, S11D, ST
M. ibaguensis (Bonpl.) Triana Arb Vh S11D
M. lepidota DC. Ar Vh, Ff N1, N2, N4, S11D
M. manauara R.Goldenb., Caddah & Michelang. Ar Vr, Ff N1
M. minutiflora (Bonpl.) DC. Ar Ff N2, S11D
M. nervosa (Sm.) Triana Arb Ff S11D
M. prasina (Sw.) DC. Ar Vh N1, N5, S11D
M. splendens (Sw.) Griseb. Ar Ff Serra Norte
M. tomentosa (Rich.) D.Don ex DC. Ar Ff N1, N3, S11A, S11C, S11D
Miconia sp. Arb Ff N1, N2, N3, N4, N5, S11A, S11B, S11C, S11D, ST, SB
Mouriri cearensis subsp. carajasica Morley Ar Ff N1, N5, S11A, S11B, S11C, S11D
M. vernicosa Naudin Ar Ff N1, N2, N5, N6, S11D,
Nepsera aquatica (Aubl.) Naudin Arb Vh N1, N5
Pleroma carajasense K.Rocha, R.Goldenb. & F.S.Mey Arb Vr N1, N2, N4, N5, N8, S11A, S11C, S11D
P. stenocarpum Triana Arb Vh S11A, S11B, S11D
Pterolepis trichotoma (Rottb.) Cogn. Er Ff, Aa N3, S11A, S11B, S11D, ST, SB
Rhynchanthera hispida Naudin Arb Vr, Ff N2, ST
Tibouchina edmundoi Brade Arb Vr N1, N5, S11A, S11C, S11D, SB
Tibouchina sp. Ar Vr ST
Tococa guianensis Aubl. Arb Vr N3
Meliaceae - Fernandes-Júnior et al. (2017)
Guarea macrophylla Vahl Ar Ff N1
G. pubescens (Rich.) A.Juss. Ar Ff N4
G. silvatica C.DC. Ar Ff S11A, S11B
Trichilia micrantha Benth. ® Ar Vr, Ff N1
Menispermaceae Lima & Teixeira (2018)
Abuta grandifolia (Mart.) Sandwith ® Li Vr N1, S11D
Menyanthaceae - Giulietti (2016b)
Nymphoides humboldtiana (Kunth) Kuntze ® Er Vh N1, N4, N5, N7, N8, S11A, S11C, S11D
Metteniusaceae - Cruz e Viana (2016)
Emmotum nitens (Benth.) Miers Ar Ff N1, N4, N5, N7, N8, S11C, S11D, ST
Monimiaceae - Mota (2017b)
Mollinedia ovata Ruiz & Pav. Ar Ff N7, S11A, S11B
Moraceae - Romaniuc-Neto & Gaglioti (2018)
Ficus americana subsp. guianensis (Desv. ex Ham.) C.C.Berg Ar Ff N4, S11A, S11D
Myrtaceae - Trindade et al. (2018)
Calyptranthes bipennis O.Berg ® Ar Ff S11C
Campomanesia aromatica (Aubl.) Griseb. Ar Ff S11D
Eugenia anastomosans DC. ® Ar-Arb Ff N2, N5
E. cupulata Amshoff Ar Ff S11A
E. densiracemosa Mazine & Faria Ar-Arb Ff S11D
E. egensis DC. Ar Ff ST
E. flavescens DC. ® Ar Vr, Ff N1, N4, N8, S11D, ST
E. florida DC. Ar-Arb Ff N1
E. punicifolia (Kunth) DC. ® Arb Vr, Ff N1, N4, N5, S11D
Eugenia sp. Arb Ff S11C
Myrcia amazonica DC. Ar-Arb Vr, Ff N2, N5
M. atramentifera Barb.Rodr. Ar-Arb Ff N2, N5
M. bracteata (Rich.) DC. ® Ar-Arb Vr, Ff N3, N7, S11D, ST
M. clusiifolia (Kunth) DC. Ar Ff N4, N5
M. cuprea (O.Berg) Kiaersk. ® Ar-Arb Ff S11D, ST, SB
M. grandis McVaugh Ar Vr N1, N4
M. guianensis (Aubl.) DC. ® Ar Vr N1, N3, N4, S11A
M. inaequiloba (DC.) Lemée Ar Ff N1
Myrcia aff. maguirei (McVaugh) E.Lucas & C.W.Wilson Arb Ff N3
M. multiflora (Lam.) DC. ® Ar-Arb Vr, Ff N1, N2, N3, N4, N5, S11A, S11D
M. paivae O.Berg Ar Ff N5
M. splendens (Sw.) DC. Ar Vr, Ff N1, N2, N4, S11D, SB
M. subsessilis O.Berg Ar Ff N1, N2
M. sylvatica (G.Mey.) DC. ® Ar Vr, Ff N1, N4, S11D, ST
M. tomentosa (Aubl.) DC. Ar Vr, Ff N3, S11D, SB
Myrciaria floribunda (H.West ex Willd.) O.Berg Ar Ff N4, SB
Nyctaginaceae - Giulietti & Nogueira (2017)
Guapira venosa (Choisy) Lundell Ar Ff N1, N5
Neea floribunda Poepp. & Endl. ® Ar Ff N5, ST
N. macrophylla Poepp. & Endl. ® Ar-Arb Vr, Ff N1, N2, N5, S11A, S11B, S11D, ST
N. oppositifolia Ruiz & Pav. ® Ar-Arb Vr, Ff N1, N5, N8, S11A, S11B, S11C, S11D, ST, SB
Nympheaceae - Lima (2018)
Nymphaea rudgeana G.Mey. Er Vh S11B
Ochnaceae - Zappi (2018b)
Ouratea castaneifolia (DC.) Engl. ® Arb Vr, Ff N1, N4, S11A, S11C, SB
O. racemiformis Ule ® Ar-Arb Vh, Ff N1, N3, N4, N6, S11D, SB
Sauvagesia tenella Lam. ® Er Vr N1, N2, N4, N6, N8, S11A, S11D, SB
Olacaceae - Meirelles & Fernandes-Júnior (2017)
Heisteria ovata Benth. ® Ar-Arb Ff N1, N2, N3, N4, N5, S11A, S11B, S11C, S11D
Ximenia americana L. ® Ar-Arb Vr, Ff N1, S11A, S11D
Onagraceae - Lovo & Zappi (2018)
Ludwigia decurrens Walter Er Vh, Ff, Aa S11D
L. erecta (L.) H.Hara Arb Vh, Aa N4
L. hyssopifolia (G.Don.) Exell Er Vh N1
L. leptocarpa (Nutt.) H.Hara Er Vh, Aa N4
L. nervosa (Poir.) H.Hara ® Arb Vh S11A
L. octovalvis (Jacq.) P.H.Raven+ ® Er Vh, Aa N1, N2, S11D, ST
L. torulosa (Arn.) H.Hara Er Vh N5, SB
Opiliaceae - Ramalho & Zappi (2017)
Agonandra silvatica Ducke ® Ar-Arb Ff N1, N7, S11A
Orchidaceae - Koch et al. (2018)
Campylocentrum fasciola (Lindl.) Cogn. ® Er Ff N5
Catasetum discolor (Lindl.) Lindl. ® Er Vr N1, N2, N4, N8, S11A, S11D, ST, SB
Cranichis muscosa Sw. Er Vr Serra Sul
Cyrtopodium andersonii (Lamb. ex Andrews) R.Br. Er Vr, Ff N1, ST
Dichaea panamensis Lindl. Er Ff S11A, S11D
Encyclia randii (Barb.Rodr.) Porto & Brade ® Er Vr, Ff S11D
Epidendrum amblostomoides Hoehne Er Ff S11A
E. nocturnum Jacq. ® Er Vr, Ff S11A, S11C, S11D, SB
E. purpurascens Focke ® Er Vr, Ff N3, S11C, S11D, SB
Erycina pusilla (L.) N.H.Williams & M.W.Chase Er Vr S11D
Habenaria ludibundiciliata J.A.N.Bat. & Bianch. Er Vr N2, N7, ST
H. aff. nuda Lindl. ® Er Vr N8
Ionopsis utricularioides (Sw.) Lindl. Er Vr, Ff S11D
Laelia marginata (Lindl.) L.O.Williams Er Ff S11A
Macroclinium wullschlaegelianum (Focke) Dodson Er Vr N1
Mesadenella cuspidata (Lindl.) Garay Er Vr N3, S11D
Mormodes paraensis Salazar & J.B.F.Silva Er Ff
Notylia barkeri Lindl. Er Ff ST
N. lyrata S.Moore Er Ff S11D
Peristeria guttata Knowles & Westc. Er Ff N1
Polystachya concreta (Jacq.) Garay & H.R.Sweet Er Ff S11A, S11D
Prosthechea fragrans (Sw.) W.E.Higgins Er Vr, Ff S11D
Scaphyglottis prolifera (R.Br.) Cogn. Er Ff S11C, S11D
S. stellata Lodd. ex Lindl. Er Vr, Ff N1, N3
Sobralia liliastrum Lindl. ® Er Vr, Ff N1, N2, N4, N5, S11D
Trichocentrum sprucei (Lindl.) M.W.Chase & N.H.Willians ® Er Ff N4
Uleiochis longipedicellata A.Cardoso & IlK.-Borg. Er Vr, Ff N5
Orobanchaceae - Scatigna & Mota (2017)
Buchnera carajasensis Scatigna & N.Mota ® Er Vr, Vh N2, N3, N6, N7, S11A, S11B, S11C, S11D, ST, SB
Oxalidaceae - Hall et al. (2017)
Oxalis barrelieri L. Er Vr, Ff SB
O. mucronulata Norlind Er Vr, Ff N4
Passifloraceae - Koch & Ilkiu-Borges (2016)
Passiflora ceratocarpa F.Silveira Er Vr, Ff N7, SB
P. foetida L. - Harley 57373∆ □ Er Ff N7
P. glandulosa Cav. ® Er Vr S11D, ST
P. tholozanii Sacco ® Er Ff N1, N2, N3, S11D, ST
Passiflora sp. Er Ff N7
Phyllanthaceae - Secco & Silveira (2016)
Phyllanthus hyssopifolioides Kunth Er Vh N1, N3, N4, N5, N6, ST, SB
P. minutulus Müll.Arg. Er Vh N1, N3, N4, N5, S11B, S11C, S11D, ST, SB
P. orbiculatus Rich. Er Ff, Aa N1, SB
P. stipulatus (Raf.) G.L.Webster Er Vr N4
Phytollacaceae - Meirelles (2016)
Phytolacca thyrsiflora Fenzl ex J.A.Schmidt + Er Aa N4, N5, N8, SB
Picramniaceae - Pirani & Devecchi (2016) Ff
Picramnia ferrea Pirani & W.W.Thomas ® Ar-Arb Vr, Ff N1, N5, S11A, S11B, S11C, S11D, ST, SB
Piperaceae - Monteiro (2018)
Peperomia albopilosa D.Monteiro ® Er Vr, Ff N3, N4, N6, S11A, S11B, S11C, S11D
P. circinnata Link Er Ff N3, S11D
P. macrostachya (Vahl) A.Dietr. Er Ff, Aa N5, N6, S11B, S11C, S11D, SB
P. magnoliifolia (Jacq.) A.Dietr. Er Vr, Ff S11C, S11D
P. obtusifolia (L.) A.Dietr. ® Er Ff N4, S11C, S11D
P. pseudoserratirhachis D.Monteiro Er Ff N3, N4, N7, N8, S11D
P. uaupesensis Yunck. Er Vr, Ff N1, N4, S11A, S11D
Piper aduncum L. Arb Vr, Ff N5
P. aequale Vahl Arb Vr, Ff N1, N5, S11A
P. aleyreanum C.DC. Arb Vr,Ff, Aa N1, N5, S11A, S11B
P. arboreum var. arboreum Aubl. Arb Vr,Ff, Aa N1, S11B, S11D
P. arboreum var. hirtellum Yunck. Arb Vr, Ff, Aa S11D
P. carniconnectivum C.DC. Arb Vr,Vh, Aa N5
P. demeraranum (Miq.) C.DC. Arb Ff S11A, S11B
P. kegelianum (Miq.) C.DC. Arb Ff N1
P. krukoffii Yunck. Arb Vr, Ff S11A, S11B
P. nematanthera C.DC. Arb Ff, Aa S11D
P. obliquum Ruiz & Pav. Arb Ff N1, N4, S11A
P. pellitum C.DC. Arb Ff N5, S11B
Plantaginaceae - Scatigna & Mota (2017)
Bacopa monnierioides (Cham.) B.L.Rob. ® Er Vh N7, S11D, SB, ST
B. myriophylloides (Benth.) Wettst. ® Er Vh N1, SB
B. reflexa (Benth.) Edwall ® Er Vh N3, N4, N5, S11B, S11D, ST
Scoparia dulcis L. + Er Aa N4, S11D
Stemodia verticillata (Mill.) Hassl. Er Aa N1, N3
Poaceae - Viana et al. (2018)
Acroceras zizanioides (Kunth) Dandy Er Vh N5
Andropogon bicornis L. + □ Er Vr, Aa N1, N3, N4, N5, S11D
A. leucostachyus Kunth + □ Er Vr, Aa N3, ST, SB
A. virgatus Desv. ex Ham. Er Vr, Vh N3
Anthaenantia lanata (Kunth) Benth. ® Er Vr N2, S11D, SB
Axonopus aureus P.Beauv. Er Vr SB
A. capillaris (Lam.) Chase + □ Er Vr, Ff, Aa N1, N3, N4, N5, N6, N8, S11D
A. carajasensis M.N.Bastos Er Vr N1, S11D
A. compressus (Sw.) P.Beauv. ∆ □ Er Vr N3
A. longispicus (Döll) Kuhlm. Er Vr N1, N2, N3, N4, N5, N8, S11D, ST, SB
A. pressus (Nees ex Steud.) Parodi Er Vr N1
A. purpusii (Mez) Chase Er Vr N1, N4, N5, N6
A. rupestris Davidse ® Er Vr, Vh N1, N3, N6, N7, N8, ST, SB
Cenchrus polystachios (L.) Morrone ++ Er Aa N3, N4
Chloris barbata Sw. + Er Aa S11D
Coleataenia scabrida (Döll) Zuloaga Er Vh N5, ST, SB
C. stenodes (Griseb.) Soreng Er Vh S11D, ST, SB
Digitaria ciliaris (Retz.) Koeler + Er Aa S11D
D. insularis (L.) Fedde + □ Er Aa S11D, SB
D. violascens Link ++ Er Aa N4, S11D
Eleusine indica (L.) Gaertn. ++ Er Aa S11D
Eragrostis bahiensis Schrad. ex Schult. Er Vr, Vh N1, N4, N5
E. curvula (Schrad.) Nees ++ □ Er Aa N1
E. maypurensis (Kunth) Steud. ® Er Vr N1, N4, N5, N8, S11D
E. pilosa (L.) P.Beuv. ex Roem. & Schult. ++ Er Aa S11D
E. rufescens Schrad. ex Schult. Er Vr, Aa N1, N4, N6, S11D
E. tenella (L.) P.Beauv. ex Roem. & Schult. ++ Er Aa S11D
E. unioloides (Retz.) Nees ex Steud. ++ Er Aa N1, N2
Hildaea breviscrobs (Döll) C.Silva & R.P.Oliveira ® Er Ff S11D
H. pallens (Sw.) C.Silva & R.P.Oliveira Er Ff N7, N8
H. tenuis (J.Presl & C.Presl) C.Silva & R.P.Oliveira Er Ff N3, N5, S11B, ST
Hildaea sp. Er Vr, Ff N4, N5, S11A, S11B, ST
Homolepis aturensis (Kunth) Chase + Er Ff, Aa N3, N4
Ichnanthus calvescens (Nees ex Trin.) Döll ® Er Vr, Ff N4, N7, S11D, ST, SB
I. leptophyllus Döll Er Vr, Ff S11A, ST
Isachne polygonoides (Lam.) Döll Er Vh N1, N3, N5, N7, N8, S11B, S11D, SB
Lasiacis ligulata Hitchc. & Chase Er Ff N2
Luziola peruviana Juss. ex J.F.Gmel. ® Er Vh SB
Melinis minutiflora P.Beauv. ++ □ Er Aa N5, SB
Mesosetum annuum Swallen Er Vr N1, N3, N6, N7, S11A, S11B, S11D, SB
M. cayennense Steud. ® Er Vr N1, N3, N4, N5, S11C, S11D, ST, SB
M. filifolium F.T.Hubb. Er Vr N1, N3, N6, N7, S11A, S11D, ST,
Olyra latifolia L. ® Er Ff N4, S11D
Oryza glumipatula Steud. Er Vh N7, S11A, S11D
Otachyrium versicolor (Döll) Henrard Er Vh S11A
Panicum aquarum Zuloaga & Morrone Er Vh SB
P. millegrana Poir. ® Er Ff N5, S11D, ST
Paratheria prostrata Griseb. ® Er Vh S11D, SB
Parodiolyra micrantha (Kunth) Davidse & Zuloaga ® Er Ff N1, N2, N4
Paspalum axillare Swallen ® Er Vr, Ff N3, N4, N5, S11A, S11C
P. carajasense S.Denham Er Vr N1, N4, N5, N7, N8
Paspalum cangarum C.O. Moura, P.L. Viana, R.C. Oliveira Er Vr S11A, S11B, SB
P. cinerascens (Döll) A.G.Burm. & M.Bastos Er Vr, Ff N1, S11A, S11D, ST
P. conjugatum P.J.Bergius + Er Aa N1, N7, N8, ST, SB
P. densum Poir. Er Aa N2, N4, N5
P. expansum Döll Er Vr, Ff N7, SB
P. foliiforme S.Denham ® Er Vr N3, N4, N7, N8, SB
P. gardnerianum Nees Er Vr N1
P. lanciflorum Trin. Er Vr N1, N3, N4, S11A, S11C, S11D, SB
P. melanospermum Desv. ex Poir. Er Vr, Aa N1, N2, N4, N7
P. multicaule Poir. + Er Vr, Aa N1, N3, N4, N6, N7, SB
P. pallens Swallen Er Vh N7, S11D, ST, SB
P. paniculatum L. + □ Er Aa ST
P. parviflorum Rhode ex Flüggé Er Vr, Vh N1, N4, N5
P. reticulinerve Renvoize ® Er Vr N1, N3
P. spissum Swallen Er Vr N1, N3, N4, N5, N6, S11A, S11B, S11D
P. virgatum L. + Er Aa N1, S11D
Paspalum sp. Er Vr, Ff S11D, SB
Rhytachne gonzalezii Davidse ® Er Vh N1, N4, N7, N8, S11A, S11B, S11D, SB
Rugoloa pilosa (Sw.) Zuloaga + Er Vr, Ff, Aa N3, N4, N6, N7, N8, S11C, S11D, SB
R. polygonata (Schrad.) Zuloaga Er Vh N1, N4, N5, N8
Sacciolepis myuros (Lam.) Chase Er Vh ST
Sorghum halepense (L.) Pers. ++ Er Aa S11D
Sporobolus indicus (L.) R.Br. ++ Er Aa S11D
S. multiramosus Longhi-Wagner & Boechat ® Er Vr N1, N2, N3, N4, N5, N6, N7, N8, S11B, S11D, ST, SB
Steinchisma laxum (Sw.) Zuloaga Er Vr, Vh, Aa N1, N4, N7, S11B, S11C, S11D, ST, SB
Steirachne barbata (Trin.) Renvoize Er Vr N1
Streptostachys asperifolia Desv. Er Vr, Ff N1, N3, N4, N5, N6, N7, S11D, ST,
Trachypogon spicatus (L.f.) Kuntze Er Vr N1, N3, N4, N5, N6, N8, S11D, SB
Trichanthecium cf. arctum (Swallen) Zuloaga & Morrone Er Vr, Vh N1, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, SB
T. cyanescens (Nees ex Trin.) Zuloaga & Morrone Er Vr, Vh N3, N4, N6, S11B, ST, SB
T. parvifolium (Lam.) Zuloaga & Morrone Er Vh N1, N4, N6, S11B, S11D, ST, SB
Tristachya chrysothrix Nees Er Vr S11D
Urochloa brizantha (Hochst. ex A.Rich.) R.D.Webster ++ □ Er Aa N4
U. decumbens (Stapf) R.D.Webster ++ □ Er Aa Serra Norte
U. maxima (Jacq.) R.D.Webster ++ Er Aa SB
Polygalaceae - Pastore & Silveira (2016)
Bredemeyera divaricata (DC.) J.F.B.Pastore Li Vr N1, S11B
B. floribunda Willd. ® ∆ □ Li Vr N4, N6, S11A, ST, SB
Caamembeca spectabilis var. spectabilis (DC.) J.F.B.Pastore ® Er Vr, Ff N1, N4, N5, N7, N8, S11A, S11B, S11C, S11D, SB
Polygala adenophora DC. ® Er Vh N1, N3, N4, N5, N6, N7, S11A, S11B, S11C, S11D, ST, SB
Securidaca diversifolia (L.) S.F.Blake Li Vr, Ff N1, N6, N7, S11A, S11B, S11C, S11D,
S. rivinifolia var. parvifolia A.W.Benn. Li Vr, Ff N5, N6, ST
S. rivinifolia var. rivinifolia A.St.-Hil. & Moq. ® Li Vr, Ff N3, ST, SB
Polygonaceae - Melo (2018)
Coccoloba coronata Jacq. Ar-Arb Ff S11D
C. parimensis Benth. Li Ff S11C
Polygonum acuminatum Kunth Er Vh N3
P. hydropiperoides Michx. Er Vh SB
Pontederiaceae - Sousa & Giulietti (2016)
Heteranthera oblongifolia Mart. ex Schult. & Schult.f. Er Vh S11D
H. reniformis Ruiz & Pav. Er Vh S11D
Portulacaceae - Giulietti & Coelho (2018)
Portulaca sedifolia N.E.Br. ® Er Vr, Vh N4, N6, N7, N8, S11D, ST, SB
Primulaceae - Freitas & Luma (2017)
Clavija lancifolia subsp. chermontiana (Standl.) B.Ståhl Arb Ff N4, S11A, S11C, S11D
C. macrophylla (Link ex Roem. & Schult.) Miq. Arb Ff N4
Cybianthus detergens Mart. ® Ar-Arb Ff N1, N4, S11C, S11D
C. penduliflorus Mart. Ar-Arb Vr, Ff N1, N3, N4, N7, S11D
Cybianthus sp. Arb Vr N1
Proteaceae - Hall (2016)
Roupala montana Aubl. ® Ar Vr, Ff N1, N3
Quiinaceae - Botelho & Rocha (2018)
Quiina pteridophylla (Radlk.) Pires Ar Ff N1
Rubiaceae - Zappi et al. (2017)
Alibertia edulis (Rich.) A.Rich. ex DC. Ar-Arb Vr, Ff N1, N3, N5, S11A, S11C, S11D, ST
Augusta longifolia (Spreng.) Rehder Arb Ff S11B, S11D
Borreria alata (Aubl.) DC. + ® Sb Aa N5, N7, S11B, S11D, ST, SB
B. carajasensis E.L.Cabral & L.M.Miguel ® Sb Vr N1, S11A, S11B, S11C, S11D, ST, SB
B. elaiosulcata E.L.Cabral & L.M.Miguel ® Sb Vr N1, N2, N3, N4, N5, N6, N8, S11A, S11B, S11C, S11D, ST, SB
B. heteranthera E.L.Cabral & Sobrado ® Er Vr N1, N4, N5, S11A, S11B, S11C, S11D, ST, SB
B. hispida Spruce ex K.Schum. ® Er Vr, Aa N1, N4, N7, S11A, S11B, S11C, S11D, ST
B. hyssopifolia (Roem. & Schult.) Bacigalupo & E.L.Cabral Er Vh Serra Norte, Serra Sul
B. ocymifolia (Willd. ex Roem. & Schult.) Bacigalupo & E.L.Cabral + ® Sb Aa N3, S11D
B. paraensis E.L.Cabral & Bacigalupo® Er Vr N1, N2, N3, N4, N5, N6, N7, N8, S11B, S11C, S11D, ST, SB
B. semiamplexicaulis E.L.Cabral® Er Vr, Vh N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11D, SB
B. spinosa Cham. & Schltdl. ex DC. Sb Vr S11A
B. verticillata (L.) G.Mey. + ® ∆ □ Sb Aa N1, N2, N3, N5, S11B, S11D, SB
Borreria sp. Sb Vr N5, ST
Carajasia cangae R.M.Salas, E.L.Cabral & Dessein ® Er Vr S11A, S11B, S11C, S11D
Chomelia ribesioides Benth. ex A.Gray ® Sb Vr, Vh N2, S11A, S11C, SB
Cordiera myrciifolia (K.Schum.) C.H.Perss. & Delprete ® Arb Vr, Ff N1, N2, N6, S11A, S11C, S11D, ST
Faramea capillipes Müll.Arg. Arb Ff N1, N2, N5, S11A, S11B, S11D, ST
F. multiflora A.Rich. Arb Ff S11B, S11C, ST, SB
Geophila cordifolia Miq. Er Ff N5
Hexasepalum teres (Walter) J.H.Kirkbr. Er Vr, Aa N3
Limnosipanea spruceana Hook.f. ® Er Vh SB
Margaritopsis inconspicua C.M.Taylor Arb Ff S11B
Mitracarpus carajasensis E.L.Cabral, Sobrado & E.B.Souza ® Er Vr, Ff N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
Palicourea deflexa (DC.) Borhidi Arb Ff N3, S11A, S11B, S11D, SB
P. guianensis Aubl. Ar-Arb Ff N2, S11A, S11B, S11C, S11D
P. marcgravii A.St.-Hil. ® Arb Ff S11D, ST
P. racemosa (Aubl.) Borhidi® Arb Ff S11A, S11D, SB
Perama carajensis J.H.Kirkbr. ® Er Vr, Ff N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
Psychotria appendiculata Müll.Arg. Arb Ff S11D, ST
P. carthagenensis Jacq. Arb Ff N2, N7, S11B, S11D
P. colorata (Willd. ex Schult.) Müll.Arg. Arb Ff N1, N5, S11A, S11C, S11D, ST, SB
P. hoffmannseggiana (Willd. ex Schult.) Müll.Arg. ® Sb Ff N1, N3, N4, N5, S11B, S11C, S11D, ST, SB
P. iodotricha Müll.Arg Sb Ff S11D
P. lupulina Benth. Sb Ff S11D, SB
P. prunifolia (Kunth) Steyerm. Sb Ff S11A, SB
P. trichosepala Müll.Arg. Sb Ff N1, ST, SB
P. variegata Steyerm. Er Ff N4
Remijia amazonica K.Schum. ® Arb Ff S11A, S11C, S11D
Richardia scabra L. Er Aa SB
Rudgea longiflora Benth. Arb Ff S11A, S11B, S11D
Sabicea grisea Cham. & Schltdl. ® Li Vr, Ff N2, ST
Schizocalyx cuspidatus (A.St.-Hil.) Kainul. & B.Bremer Ar Vr, Ff N1, S11A, S11B, S11C, S11D
Spermacoce exilis (L.O.Williams) C.D.Adams Er Ff, Aa N7, ST, SB
Tocoyena formosa (Cham. & Schltdl.) K.Schum. ® Ar-Arb Vr N1, SB
Uncaria guianensis (Aubl.) J.F.Gmel. Li Ff S11A, S11D
Rutaceae - Pirani & Devecchi (2018)
Dictyoloma vandellianum A.Juss. ® Arb Vr, Aa Serra Norte
Ertela trifolia (L.) Kuntze ® Sb Ff N8, ST, SB
Esenbeckia cowanii Kaastra Arb Vr, Ff N1
Pilocarpus carajaensis Skorupa® Arb Vr, Ff N4, N5, S11D
P. microphyllus Stapf ex Wardleworth ® Arb Ff N1, N2, N3, N6
Spiranthera parviflora Sandwith Arb Ff N1, N2, N3
Zanthoxylum apiculatum (Sandwith) P.G.Waterman Ar Ff Serra Norte
Z. monogynum A.St.-Hil. Ar Ff Serra Norte
Salicaceae - Marquete & Zappi (2018)
Casearia arborea (Rich.) Urb. Ar Vr, Ff N5, S11D
C. grandiflora Cambess. Ar Vr,Ff, Aa N2, N3, N7, S11D
C. javitensis Kunth ® Ar Ff N1, N2, S11B, S11D
C. pitumba Sleumer Ar Ff S11D
Ryania pyrifera (Rich.) Uittien & Sleumer Ar-Arb Ff N1, N5, N7
R. speciosa Vahl Ar-Arb Ff N1
Santalaceae - Caires (2017)
Dendrophthora warmingii (Eichler) Kuijt Par Ff ST
Phoradendron crassifolium (Pohl ex DC.) Eichler Par Ff N3
P. dipterum Eichler Par Ff N3
P. mucronatum (DC.) Krug & Urb. Par Ff N1
P. obtusissimum (Miq.) Eichler Par Ff Serra Norte
P. piperoides (Kunth) Trel. Par Ff Serra Sul
P. quadrangulare (Kunth) Griseb. ® Par Vr, Ff N1, N2, N3, N4, S11A, S11B, S11D, ST
P. tunaeforme (DC.) Eichler ® Par Vr, Ff S11A, S11B, S11C, S11D
Sapindaceae - Barbosa et al. (2018)
Allophylus latifolius Huber Ar Ff N2, S11B, S11D
A. racemosus Sw. Ar Vr N1, S11D
A. semidentatus (Miq.) Radlk. Ar Ff S11A, S11D, ST
Matayba guianensis Aubl. Ar Vr, Ff N5, S11A, S11B, S11C, S11D
M. inelegans Spruce ex Radlk. Ar Vr, Ff S11D
M. spruceana (Benth.) Radlk. Ar Ff N1
Serjania caracasana (Jacq.) Willd. Li Vr, Ff S11A, S11B, S11C, S11D, ST
S. lethalis A.St.-Hil. Li Vr, Ff S11B
Sapotaceae - Terra-Araújo & Zappi (2018)
Pouteria ramiflora (Mart.) Radlk. ® Ar Vr N1, N2, N3, N4, N6, S11A, S11B, S11D, ST
Simaroubaceae - Devecchi & Pirani (2016)
Simaba guianensis Aubl. Ar-Arb Vr, Ff N1, N4, N5
Simarouba amara Aubl. ® Ar Vr N1, N6, S11A, S11C, S11D
Siparunaceae - Hall & Meirelles (2017)
Siparuna ficoides S.S.Renner & Hausner Ar Ff SB
Smilacaceae - Andreata & Watanabe (2018)
Smilax irrorata Mart. ex Griseb. ® Li Vr, Ff N1, N2, N6, S11A, S11C, S11D, ST, SB
S. syphilitica Humb. & Bonpl. ex Willd. Li Ff N1, N3, S11A, S11B, S11C, S11D
Solanaceae - Giacomin & Gomes (2018)
Hawkesiophyton ulei (Dammer) Hunz. Arb Ff S11A, S11D
Physalis angulata L. ++ Er Aa N5, SB
Schwenckia americana L. Er Vr, Ff S11D, ST
Solanum acanthodes Hook.f. Arb Ff Serra Norte
S. americanum Mill. + □ Er Aa N1, N4, SB
S. campaniforme Roem. & Schult. ® Arb Vr, Ff N1, N2, N5, N6, N7, S11A, S11D, ST
S. coriaceum Dunal Li Ff N1
S. crinitum Lam. Ar-Arb Vr, Ff N4, N5, S11D
S. incarceratum Ruiz & Pav. Arb Vr, Ff SB
S. leucocarpon Dunal Ar-Arb Ff, Aa S11D
S. paniculatum L. Arb Vr, Aa ST
S. rugosum Dunal ® Ar-Arb Vr, Aa N4, S11D
S. schlechtendalianum Walp. Arb Vr, Ff, Aa N1, N3, N4, S11D
S. semotum M.Nee Ar-Arb Ff N1, N5, ST
S. sisymbriifolium Lam. Arb Vr, Aa S11D, SB
S. subinerme Jacq. ® Arb Vr, Ff N1, N2, ST
S. tegore Aubl. Ar Ff Serra Norte
S. torvum Sw. + Arb Vr, Aa S11D
S. uncinellum Lindl. Li Ff N1, N4, S11D, SB
S. velutinum Dunal ® Arb Vr, Ff N5, S11C, S11D, ST, SB
Styracaceae - Viana & Mota (2016)
Styrax griseus P.W.Fritsch Arb Vr N1, N3, N4, N7, S11A, S11B, S11C, S11D
S. pohlii A.DC. Ar-Arb Ff N1, N5, S11A, S11B, S11C, S11D
Thymelaceae - Mota & Giulietti (2016b)
Daphnopsis filipedunculata Nevling & Barringer ® Ar Ff N1, N2, N3, N4, N5, N6, N7
Trigoniaceae - Nunes & Gil (2017)
Trigonia nivea Cambess. Sb Ff N1, N3, N4, N5, ST
Turneraceae - Rocha (2018)
Turnera coerulea var. surinamensis (Urb.) Arbo & Fernández ® Arb Vr SB
T. glaziovii Urb. ® Arb Vr, Ff N2, N3, N4, N6, S11A, S11B, S11C, S11D, ST
T. laciniata Arbo Sb Vr S11D
T. melochioides var. arenaria Urb. Sb Vr N1, N3, N4, N8, ST
T. melochioides var. latifolia Urb. Sb Vr N5, N6, N7, S11D, ST
Urticaceae - Gaglioti et al. (2016)
Urera caracasana (Jacq.) Griseb. Arb Ff S11D, SB
Velloziaceae - Mello-Silva (2018)
Vellozia glauca Pohl ® Er Vr N1, N2, N3, N4, N5, N6, N7, N8, S11B, S11D, ST, SB
Verbenaceae - Carodos et al. (2018)
Lantana hirsuta M.Martens & Galeotti Arb Aa SB
L. paranaensis (Moldenke) R.W.Sanders. Arb Aa N1, N2, N5
Lippia grata Schauer ® Arb Vr, Ff N1, N4, N5, S11A, S11B, S11C, S11D, ST
Stachytarpheta cayannensis (Rich.) Vahl + ® Arb Aa S11D, ST
Vitaceae - Lombardi (2016)
Cissus apendiculata Lombardi ® Li Vr, Ff N1, N4, N7, S11A, S11B, S11C, ST,
C. erosa Rich. ® Li Vr, Ff N1, N4, N5, S11A, S11B, S11D, SB
C. sulcicaulis (Baker) Planch. - Fernandes 179 Li Ff N5
C. tinctoria Mart. in Spix & Mart. Li Vr, Ff N1, N3, N4, N5
C. verticillata (L.) Nicolson & C.E.Jarvis Li Aa N5
Vochysiaceae - Shimizu & Gonçalves (2017)
Callisthene microphylla Warm. ® Ar Ff N1, N3, N4, N5, N6, N8, S11B, S11C, S11D, ST, SB
Qualea multiflora Mart. Ar Ff S11D
Q. parviflora Mart. Ar Ff N4, S11A, S11B, S11D
Vochysia haenkeana Mart. ® Ar Ff N1, S11D
Xyridaceae - Mota & Wanderley (2016)
Xyris brachysepala Kral ® Er Vr, Vh N1, N2, N3, N4, N5, N6, N7, N8, S11A, S11B, S11C, S11D, ST, SB
X. lacerata Pohl ex Griseb. - Falcão 453 Er Vr, Vh SB
X. macrocephala Vahl ® Er Vh N3, N4, N6, S11A, S11C, ST
X. savanesis Miq. - Falcão 451 Er Vh SB
Species total by individual outcrop N1(392), N2(130), N3(224), N4(282), N5(267), N6(100), N7(114), N8(104), S11A(240), S11B(209), S11C(186), S11D(416), ST (228) e SB (230).

The new collecting efforts and study of the resulting specimens by specialists helped to detect families that were not yet listed by Viana et al. (2016) at the onset of the FCC Project, such as Apodanthaceae, Bixaceae, Cannabaceae, Droseraceae, Meliaceae, Monimiaceae, Quiinaceae and Siparunaceae. On the other hand, despite having been listed by Viana et al. (2016), Haloragaceae and Heliconiaceae were found to occur only in the forest, thus these have not been monographed. After verifying herbarium material and checking the complete Carajás database, it was also found that some plant families listed in the introductory paper (Viana et al. 2016), such as Peraceae and Ranunculaceae, are not found in the canga. Additionally, Elaeocarpaceae and Rhizophoraceae monographs, despite not having been listed in the introductory paper, were published despite the fact that no species recorded in the canga were represented in these families. Therefore, the Peraceae, Ranunculaceae, Elaeocarpaceae and Rhizophoraceae were not included in the present list.

APG IV (APG 2016) was the base for the classification used in the seed plant list, except for Passifloraceae (segregated from Turneraceae), Ochnaceae (published separately from Quiinaceae) and Boraginaceae (represented by Cordiaceae and Heliotropiaceae). Species that appeared as indeterminate in the FCC treatments and were subsequently published as new were updated in the present list, and recent nomenclatural updates were also provided for relevant names.

The determination of species habits followed the categories: trees (including palms with stem), shrubs, subshrubs, herbs, lianas and parasites. Some species were attributed more than one habit, following publications of the FCC. When the information was incomplete or lacking, available databases were consulted, such as Flora do Brasil (2020, under construction) and WCSP (2018).

Species distribution in the study area was discriminated between canga blocks of the Serra Norte (N1, N2, N3, N4, N5, N6, N7, N8), Serra Sul (S11A, S11B, S11C, S11D), Serra do Tarzan and Serra da Bocaina (Fig. 1). Species with known distribution restricted to the canga of the FLONA of Carajás and PNCF were indicated as endemic. Species that occurred in canga outside the study area (e.g., Serra de Campos, Serra do Cristalino, Serra Arqueada), even if considered endemic of the canga of Carajás were not indicated as endemic within this study.

The species were assigned to one or more of the following vegetation categories proposed for the canga of Carajás by Mota et al. (2015): a. open rupestrial vegetation, represented by herbaceous to shrubby plants found on the more or less bare iron-rich, rocky soil; b. hydromorphic vegetation, including lakes, marshes and seasonal or perennial streams; and c. forest formations including deciduous to semi-deciduous forest groves and low forest that grow over the iron outcrops.

Floristic comparisons between different canga sites in Brazil were carried out by compiling a list for the Quadrilátero Ferrífero, in Minas Gerais (Carmo & Jacobi 2016, Messias & Carmo 2015 and Viana & Lombardi 2007), and for the Corumbá region, Mato Grosso do Sul (Takahasi 2015). Not fully determined species (cf. or aff.) were excluded from the comparisons. Nomenclatural and taxonomic adjustments between the lists were performed when needed using the Flora do Brasil 2020 (under construction) database.

The indication of invasive exotic or native species follows the Brazilian law (Instrução Normativa 4 (2011) and 11 (2014)), with modifications suggested Giulietti et al. (2018). Data from the FCC monographs was considered for this purpose. Species with DNA samples banked at ITV were also indicated.

Results

The list of seed plants of the canga of Serra dos Carajás comprises 116 families, 419 genera and 856 species (Tab. 1). Eight species that were not monographed in the FCC Project are included here: Philodendron blanchetianum (Araceae), Ipomoea procumbens,I. reticulata,I. setifera (Convolvulaceae), Passiflora foetida (Passifloraceae), Cissus sulcicaulis (Vitaceae), Xyris lacerata and X. savanensis (Xyridaceae), with the first record of C. sulcicaulis for the state of Pará. Amongst these species, 258 count with extracted total DNA sample (Tab. 1).

The richest plant family in the canga of the Serra dos Carajás was Poaceae (86 spp.), followed by Fabaceae (65), Rubiaceae (46), Cyperaceae (45), Melastomataceae (41), Asteraceae (34), Orchidaceae (27), Myrtaceae (26), Apocynaceae (21), Convolvulaceae and Solanaceae (20 spp. each), representing around 48% of the sampled species (Tab. 2; Fig. 2). Out of the total of 419 genera, 269 are represented by a single species, 64 by two, 38 by three, that together add up to nearly 60% of the generic richness of the FCC. The most representative genera, Miconia (20), Paspalum (19), Solanum (16), Myrcia (14), Cyperus (13), Rhynchospora,Piper,Borreria and Ipomoea (12 spp. each), Mimosa and Utricularia (11 spp. each), represent approximately 16% of the richness of the FCC (Fig. 3).

Table 2 List of the ten richest seed plant families in the canga of Carajás, Corumbá, Quadrilátero Ferrífero 

Serra dos Carajás Corumbá Quadrilátero Ferrífero
1 Poaceae Poaceae Asteraceae
2 Fabaceae Fabaceae Poaceae
3 Rubiaceae Cyperaceae Orchidaceae
4 Cyperaceae Malvaceae Fabaceae
5 Melastomataceae Euphorbiaceae Melastomataceae
6 Asteraceae Convolvulaceae Myrtaceae
7 Orchidaceae Apocynaceae Rubiaceae
8 Myrtaceae Malpighiaceae Cyperaceae
9 Apocynaceae Portulacaceae Apocynaceae
10 Convolvulaceae/Solanaceae Cactaceae Solanaceae

Figure 2 Top 10 families in the canga of the Serra dos Carajás 

Figure 3 Most representative genera of the canga of the Serra dos Carajás 

Regarding the large angiosperm groups, three aquatic species belong to order Nymphaeales. The Magnoliids are represented by 28 spp. (3.2%), including a high number of Piperaceae (19 spp.), that represent more than half of the species of this group. The Monocots comprise 239 spp. (28%), spread throughout the orders with the exception of Acorales and Petrosaviales. The Eudicots are the taxonomic group with higher species richness, including 585 species (68.3%) distributed in almost all groups from Rosids to Asterids. The only Gymnosperm recorded in the FCC was Gnetum nodiflorum.

When considering plant habit, the herbs are the predominant group, with 347 spp. (40.5%), while subshrubs were 62 spp. (7.2%), shrubs 153 spp. (17.9%). Among the species, 45 (5.1%) had variable habit, fitting more than one category. The trees, including five species of palm (Arecaceae) were represented by 119 spp. (14%). The lianas were 109 spp. (12.7%). Parasites were 22, representing 2.6% of the habit total.

In terms of habitat, out of 856 species recorded, 497 (247 exclusive) were recorded in forest formations, while 418 (128 exclusive) occur in open rupestrial vegetation and 174 (99 exclusive) occur in hydromorphic sites. It is worth recording that 319 species are found in more than one habitat. If considering rupestrial and hydromorphic vegetation as open areas, these concentrate 534 species of seed plants. Forest formations share a much more expressive number of species (223 spp.) with rupestrial vegetation when compared with hydromorphic vegetation (22 spp. in common with forest formations). Only 12 species were shared by all three habitat. Finally, there are 119 species that are associated to anthropized environments, of which 64 were recorded solely in those sites (Tab. 1).

Serra Norte was the location with higher species count, with 659 species of seed plants, followed by Serra Sul (545), Serra da Bocaina (230) and Serra do Tarzan (228). Together, the canga of PNCF (Serra da Bocaina and Serra do Tarzan) are home for 351 species. Looking in more detail at the first two locations, the blocks with highest richness were S11D (416 spp.) at the Serra Sul, followed by N1 (392), N4 (283) and N5 (267), at Serra Norte (Tab. 1). These blocks also figure among the better sampled areas: N1 (1737 specimens), S11D (1411), N4 (747) and N5 (648), and are amongst the largest in area.

Based on the FCC data and subsequent publications (Nunes et al. 2015), 24 species and one subspecies were marked as endemic from the canga of the FLONA of Carajás and the PNCF. These are Philodendron carajasense (Araceae), Cavalcantia glomerata, Lepidaploa paraensis, Parapiqueria cavalcantei (Asteraceae), Ipomoea cavalcantei (Convolvulaceae), Bulbostylis cangae, Eleocharis pedrovianae, Hypolytrum paraense (Cyperaceae), Erythroxylum carajasense, Erythroxylum nelson-rosae (Erythroxylaceae), Sinningia minima (Gesneriaceae), Cuphea carajasensis (Lythraceae), Mouriri cearensis subsp. carajasica (Melastomataceae), Uleiorchis longipedunculata (Orchidaceae), Picramnia ferrea (Picramniaceae), Peperomia albopilosa, Peperomia pseudoserratirhachis (Piperaceae), Axonopus carajasensis, Paspalum cangarum,P. carajasense (Poaceae), Borreria elaiosulcata,B. heteranthera,Carajasia cangae (Rubiaceae), Daphnopsis filipedunculata (Thymelaeaceae) and Xyris brachysepala (Xyridaceae) (Tab. 1).

Amongst potential invasive species, the record of 53 native, problematic species alongside 17 exotic invasives (Tab. 1) was made, and the outcrops with highest number of these species were S11D (36 spp.), N1 (23 spp.), N4 e N5 (22 spp.), Serra da Bocaina (19 spp.) and N3 (14 spp.).

A small number of shared species between FCC canga and other areas was found, with 96 shared species between FCC and the QF, totalling 11% of the species. Only 25, or 3% of species, were found to occur both in the FCC and Corumbá. The canga of Corumbá has also shown very little correspondence with the ones at QF, with 18 shared species (Tab. 1; Fig. 4).

Figure 4 Venn diagram showing the floristic relationships between canga sites in the Serra dos Carajás in Pará state, of the Quadrilátero Ferrífero in Minas Gerais and in Corumbá, Mato Grosso do Sul. 

Discussion

Taxonomic richness of the flora of the canga of Carajás

During the first comprehensive floristic study of the canga of the Serra de Carajás, Silva et al. (1991) recorded 231 species in 57 angiosperm families. The initial estimate from the introductory chapter of the FCC was of around 600 species in 111 families of angiosperms and one of gymnosperms (Viana et al. 2016). However, the accurate determination of specimens that were still unnamed or incorrectly named, together with a considerable contribution from recent collections (2015-2017) have collaborated to the increase of species recorded for FCC, with 856 species in 115 angiosperm and one gymnosperm families. The collections carried out as part of the FCC project have also provided data to understand the species distribution in the area. A pioneering list for the Serra Norte was presented by Secco & Mesquita (1983), including 85 species of seed plants. The present work increases the number of species ocurrences many fold, with 659 species recorded for the canga of this location.

When looking at family composition, the ten richest plant families of the FCC represent 48.1% of the flora, while the remaining half of the species is distributed in the 106 remaining families. When comparing with overall figures provided by BFG (2015), we find a similar pattern, where the tem largest families contribute towards around 60% of the flora while the remainder 233 families represent the other 40% of Brazil´s flowering plants.

The 856 species recorded in the FCC represent 2.6% of the Brazilian flora, 7% of the flora of the Brazilian Amazon domain and 13.1% of the seed plants that are recorded for the state of Pará (Flora 2020, under construction). The 419 genera from the FCC respectively 14%, 21.7% and 28.6% of the genera recorded for Brazil, the Brazilian Amazon and Pará (Flora 2020 under construction). Taking into account the small parcel of the Brazilian territory occupied by the canga of Carajás (~ 0.001%), this area can be highlighted as a site of extreme importance for its plant biodiversity.

There is a trend involving large number of genera represented by few species that may result from the high number of micro-habitat found in the canga of Carajás (sensuMota et al. 2015). This situation is reflected by the following five groups: genera that are typical from forest formations such as Tapirira (Anacardiaceae), Protium (Burseraceae), Hirtella (Chrysobalanaceae), Hypolytrum (Cyperaceae), Bellucia (Melastomataceae), Olyra (Poaceae) amongst others; genera exclusive from hydromorphic environments, such as Sagitaria (Alismataceae), Cabomba (Cabombaceae), Apalanthe and Ottelia (Hydrocharitaceae), Nymphoides (Menyanthaceae), Nymphaea (Nymphaeaceae), Limnosipanea (Rubiaceae), Helantium (Alismataceae), Lindernia (Linderniaceae), Mayaca (Mayacaceae), Bacopa (Plantaginaceae), Oryza (Poaceae), Genlisea (Lentibulariaceae); genera associated to open, not hydromporphic environments such as Cavalcantia and Lepidaploa (Asteraceae), Anemopaegma (Bignoniaceae), Cereus (Cactaceae), Lagenocarpus (Cyperaceae), Euploca (Heliotropiaceae), Cipura (Iridaceae), Cuphea (Lythraceae), Habenaria and Sobralia (Orchidaceae); rupicolous genera from forest interior such as Diastema, Goyazia and Phinea (Gesneriaceae), Epidendrum (Orchidaceae); open rocky area genera such as Parapiqueria (Asteraceae), Ananas and Dyckia (Bromeliaceae), Brasilianthus (Melastomataceae) and Carajasia (Rubiaceae), all these genera represented by three or less species within the study area. This situation contrasts strongly with what is found in the eastern Brazilian campo rupestre, where some genera contribute with many sympatric species (Zappi et al. 2017).

Amongst the better represented genera of seed-plants in the study area, forest groups such as Miconia (20 species), Solanum (16), Myrcia (14) and Piper (12) (Cardoso et al. 2017) appear in the forest formations that occur on the iron-rich canga, namely small groves and transitional forest between open areas and the adjacent ombrophilous forests (Tab. 1). The other genera with high number of species in the area are either herbaceous, such as Paspalum (19 spp.), Cyperus (13 spp.), Rhynchospora (12 spp.), Utricularia (11 spp.), or herbaceous to shrubby, such as Borreria (12 spp.) (Tab. 1).

Plant habits

Herbaceous habit (347 species, or 40.5% of the total number of seed plants) predominates over other plant habit types, following the pattern found in other Brazilian biomes with exception of the Amazon, such as Cerrado, Pantanal, Caatinga e Pampa (BFG 2015). The habit proportion found in the FCC is one tree (including those with variable habit between tree and shrub and the palms) for each four species of herb, subshrub, shrub or liana. This proportion is rather different from that recorded for the Brazilian Amazon Rainforest, where the proportion is of one tree for each two different plant habits (BFG 2015).

The diversity of the herbaceous layer is not exclusive to the FCC, being a strong feature of the open vegetation of eastern Brazil, such as the quartzitic outcrops of the Cadeia do Espinhaço, however, there the floristic composition is markedly dominated by subshrubs or shrubs with subterranean structures (xylopodia) adapted to strongly seasonal climates as well as to natural fires (Neves & Conceição 2010). The canga of Carajás substrate differs from the lateritic soil with deep water-table found in cerrado vegetation (Eiten 1972), and also from the shallow, stony soil of quartzitic campo rupestre, as there are evidences that the canga is more permeable, porous and capable to store water for longer periods of time (Carmo & Jacobi 2016).

Taking into account the Amazonian context, the Carajás region is an area with high richness of herbaceous species (347 spp.). If we consider that 476 herbaceous species are referred for all the Amazonian campinarana areas in Brazil (FBO 2020, under construction), the FCC are very relevant in their contribution to herbaceous species richness for this domain, especially when considering the small area covered by this vegetation (approx. 120 km2). On the other hand, this information also suggests that the campinarana areas of the region may be under-sampled, calling for the need to develop similar taxonomic projects in the diverse vegetation types of the Amazon Rainforest.

Well defined rain seasonality may represent another particularity of the canga of Carajás. The region has a well defined rainy summer followed by a dry winter, with – of the total precipitation concentrated in the three rainiest months, typically between January and March (Viana et al. 2016). During the dry season, the open landscape is marked by the presence of deciduous or semi-deciduous shrubs and treelets, such as Cochlospermum orinocense (Bixaceae), Callisthene microphylla (Vochysiaceae), Mimosa acutistipula var. ferrea (Fabaceae), Erythroxylum spp. (Erythroxylaceae), Croton spp. (Euphorbiaceae), a smaller number of evergreen shrubs and lianas, such as Clusia spp. (Clusiaceae) and Norantea guianensis (Marcgraviaceae), and an empoverished herbaceous layer represented by perennial herbs such as Bromeliaceae (e.g.,Dyckia duckei, Pitcarnia burchellii), Cyperaceae (e.g.,Bulbostylis cangae), Orchidaceae (e.g., Catasetum discolor and Sobralia liliastrum), Poaceae (e.g.,Axonopus longispicus,Paspalum cinerascens), Velloziaceae (Vellozia glauca), among others. The rainy period is marked by the proliferation of annual herbaceous plants such as Cavalcantia glomerata, C. percymosa, Monogereion carajensis, Praxelis asperulacea (Asteraceae), Bulbostylis conifera and Rhynchospora acanthoma (Cyperaceae), Eriocaulon carajense (Eriocaulaceae), Utricularia physoceras (Lentibulariaceae), Lindernia brachyphylla (Linderniaceae), Brasilianthus carajensis (Melastomataceae), Paspalum cangarum, P. carajasense, Sporobolus multiramosus, Trichanthecium aff. arctum (Poaceae), Perama carajasensis (Rubiaceae) and Xyris brachysepala (Xyridaceae) that, together with the shrubs, treelets and lianas that are in full leaf and often in blossom during this period, transform what was a greyish place into a colourful landscape (Zappi 2017).

The flora and environments of the canga of Carajás

At the Serra dos Carajás, the canga is characterized by a mosaic formed by different phytophysionomies associated to the iron-rich substrate (Cleef & Silva 1994; Mota et al. 2015). However, in order to organize the species into vegetation types, we considered only the tree major groups proposed by Mota et al. (2015), without considering the subgroups proposed therein.

The importance of forest formations within canga is obviated by the fact that, from the 492 forest species recorded, 247 are exclusive to this environment. However, if adding up the two open formations (open rupestrial + hydromorphic vegetation), a total of 534 species, of which 271 are exclusive, a balance between open and forest vegetation can be observed.

While the hydromorphic vegetation contributes with the smaller number of recorded species (174 spp.), it is also the habitat with largest proportion of exclusive species (c. 57%) in relation to the forest (50%) and rupestrial (31%) habitats.

The role played by each type of habitat in the floristic richness fo the canga of Carajás is evident, and the deciduous to semi-deciduous low forests found as groves or transitional areas support a peculiar flora, that presents endemic taxa such as Hypolytrum paraense (Cyperaceae), Sinningia minima (Gesneriaceae), Mouriri cearensis subsp. carajasica (Melastomataceae), Peperomia pseudoserratirhachis (Piperaceae) and Daphnopsis filipedunculata (Thymelaeaceae) (Tab. 1).

Endemic taxa

The canga of Carajás is an area of ancient mountains in the Amazon of pre-Cretacean origin, when supergenic events in what is now southern Pará originated expressive lateritic crust in the region (Ab’Saber 1986; Viana et al. 2016). Isolated by a rainforest matrix, the blocks of canga of Carajás resemble vegetation islands over a peculiar substrate where geographic isolation events may have promoted speciation processes and generated endemic patterns. A list of 24 species and a subspecies are endemic to the canga of the FLONA of Carajás and PNCF, representing 2.9% of the flora of the study area (Tab. 1).

The two monospecific genera endemic to the study area, Parapiqueria (Asteraceae) and Carajasia (Rubiaceae), when added to two other genera, Monogereion (Asteraceae) and Brasilianthus (Melastomataceae), that are restricted to the canga of Carajás, however ocurring over a wider set of outcrops that neighbour the FCC study area (Cruz et al. 2016; Rocha et al. 2017) show how important the region is in terms of endemism. Other relevant point is that the 24 putative endemic species make up nearly 5% of the 443 species considered endemic for Pará state (FBO 2020, under construction), highlighting the importance of the contribution of the canga of the FLONA of Carajás and the PNCF towards the number of endemic seed plants at state level.

The 120 monographs published by the FCC project encompass 40 species that were marked not fully identified, of which around 50% were flagged by the authors as possible new to science (Cruz et al. 2015; Koch & Ilkiu-Borges 2016; Nunes et al. 2016; Reis et al. 2017; Watanabe et al. 2017; Zappi et al. 2017; Chautems et al. 2018; Coelho 2018; Costa et al. 2018; Fernandes et al. 2018; Mattos et al. 2018; Rocha et al. 2017; Koch et al. 2018; Viana et al. 2018). Of these, a number has been found only on the canga of Carajás, being thus potentially endemic. Efforts to solve these taxonomic problems are urged, together with widening the scope for fieldwork in and around the areas in order to ascertain the true level of endemism found in the region.

Potentially invasive species

Species considered as potential invasives (exotic and problem native species) add up to approximately 12% of the species analysed. The blocks of canga with larger number of these species coincide with the location of open mines, such as S11D, N4 e N5, or with areas that were transformed recently into conservation units, but that were formerly under strong anthropic pressure, such as the Serra da Bocaina, that was included in the PNCF in 2017. Areas that have a long history of occupation and others used as support for mining activities, such as N1 and N3 (STCP 2016), also feature among these.

The outcrops that have operating mines have suffered an increase of the number of invasives. As an example of this, we can cite some new occurrences for the FCC for the S11D mine at the Serra Sul, such as Ipomoea procumbens, first recorded in 2017 (Zappi 3510), Bidens pilosa, in 2012 (Arruda 1222), Digitaria ciliaris with two records, one in 2009 and other in 2015, both near the Geosol camp at the S11D site (Viana 4376 and Afonso 120), as well as Eragrostis pilosa, collected in 2015 (Afonso 118) and E. tenella, in 2009 and 2015 (Viana 4371 and Afonso 115). Furthermore, there are recent records associated probably to the increase of the area of the N4 mine, such as Digitaria violascens, with the first record in 2012 at N4WS (Viana 5315). All the above mentioned species were not collected before 2000 in the study area. An identification guide for invasive plants was prepared specifically for block S11D, comprising also has management proposals for control and erradication of exotic species (Giulietti et al. 2018).

The presence of many invasive species in the Serra da Bocaina, including Ageratum conyzoides,Emilia sonchifolia, Porophyllum ruderale, Cyperus agreggatus, Cantinoa americana, Phytolacca thyrsiflora and Paspalum conjugatum, may be associated to the long history of landscape change and degradation linked to intensive and disordinated land use, with forest felling, timber harvesting and frequent criminal fires. Before its inception as part of the PNCF in June 2017, the Serra da Bocaina did not have status as a protected area. It is desirable to provide adequate management to the exotic and native problem species in this conservation unit to avoid their dissemination in the canga area that is now under full legal protection.

Local diversity in Carajás

The area covered by this study is of approximately 120 km2, with four major subdivisions: Serra Sul (~ 46 km2), Serra Norte (~ 35 km2), Serra da Bocaina (~ 20 km2) and Serra do Tarzan (~ 8 km2), at altitudes between 500 and 800 m a.s.l. Within the FLONA of Carajás, the Serra Norte and Serra Sul encompass, respectively, 77% and 64% of the species of seed plants found in the study area. The area covered by the Serra Sul is larger than the Serra Norte (~ 46 km2 vs. 35 km2), however the comparison here shows that the Serra Sul harbours less species. Two factors may be used to explain this difference, namely that the sampling effort dedicated to the Serra Norte is comparably much larger (4,900 specimens vs. 2,806 in the Serra Sul) and that the discontinuity between the blocks of Serra Norte may contribute towards the higher richness documented there. The Serra Sul has been divided in four sectors (S11A, S11B, S11C and S11D) that are, however, interconnected.

A separate analysis of the blocks within the major outcrops highlights higher richness in S11D, N1, N4 and N5, and these also correspond to areas historically better sampled. The Serra Norte was the first accessible point of the region back at the end of the 1960´s to be investigated by botanist Paulo Bezerra Cavalcante at the onset of the Projeto Ferro Carajás (Viana et al. 2016), while block N1, one of the largest canga blocks, was repeatedly visited over 45 years of exploration in the region. It has the highest documented plant richness of all blocks, with 392 species.

Meanwhile, blocks N4 and N5 were studied from the implementation of the Projeto Ferro Carajás as part of the prospecting for the first operating iron mines in the area (STCP 2016). Due to the economic importance of this enterprise, many environmental impact reports were prepared for these plateaus and, together with the effort employed by the FCC projects culminated with the high number of species recorded in these blocks. However, a large part of the original area of canga of these blocks has been supressed through mining activities, and a comparison using Google Earth historical images shows that about 20% and 9% of the original area of N4 and N5, respectively, is left. A similar landscape is under construction in the Serra Sul, with the implementation of the S11D iron mine, where approximately 12% of the canga has already been supressed. The periodic evaluation of the composition of the flora and the populations of endemic species found within these plateaus is paramount to ensure that measures are in place for local flora conservation.

The Serra da Bocaina and Serra do Tarzan encompass the lowest number of plant species found in the study area, with respectively 27% e 26%. These are also the least sampled outcrops and have much smaller area than the Serra Norte and Serra Sul. They are the only areas of canga included in the PNCF and, together, comprise 351 species of seed plants, or nearly 41% of the species number recorded for the FCC. Despite being an important step towards plant conservation in the region of Carajás, the creation of this fully protected area has not ensured that several endemic canga species from the FLONA of Carajás, such as Parapiqueria cavalcantei (Asteraceae), Ipomoea cavalcantei (Convolvulaceae), Axoponus carajasensis and Paspalum carajasense (Poaceae), Carajasia cangae (Rubiaceae) and Daphnopsis filipedunculata (Thymelaeaceae).

Canga of Carajás, a unique vegetation

The 13 species shared by the three areas of canga compared here (Carajás, QF and Corumbá) are widely distributed plants: Cyperus aggregatus,C. laxus,C. sesquiflorus (Cyperaceae), Passiflora foetida (Passifloraceae) and Axonopus compressus (Poaceae) are found in all Brazilian biomes; Blepharodon pictum (Apocynaceae), Commelina erecta (Commelinaceae), Evolvulus filipes (Convolvulaceae), Bulbostylis conifera (Cyperaceae), Sida linifolia (Malvaceae), Bredemeyera floribunda (Polygalaceae) and Borreria verticillata (Rubiaceae) are in all biomes excepting the Pampa (BFG 2015); and Evolvulus lithospermoides (Convolvulaceae) is cited for the Amazon Rainforest and the Cerrado in open areas including grasslands, campo rupestre and Amazonian savanna (BFG 2015). Amongst the above cited species, C. aggregatus and B. verticillata are also considered potentially invasive (Tab. 1).

The FCC and QF shared 96 species, the majority of which are widely distributed in Brazil, are comprised by nearly a third (28 species) considered potentially invasive (exotic or problem natives) (Tab. 1): Asclepias curassavica (Apocynaceae), Ageratum conyzoides,Bidens pilosa, Emilia sonchifolia, Porophyllum ruderale (Asteraceae), Commelina benghalensis (Commelinaceae), Ipomoea procumbens (Convolvulaceae), Cyperus aggregatus,C. surinamensis (Cyperaceae), Chamaecrista nictitans, C. rotundifolia,Desmodium incanum (Fabaceae), Cuphea carthagenensis (Lythraceae), Sida rhombifolia, Waltheria indica (Malvaceae), Ludwigia octovalvis (Onagraceae), Andropogon bicornis,A. leucostachyus,Axonopus capillaris,Digitaria insularis,Eragrostis curvula,Erechtites hieracifolius,Melinis minutiflora,P. paniculatum,Urochloa brizantha,U. decumbens (Poaceae), Borreria verticillata (Rubiaceae), Solanum americanum (Solanaceae), Stachytarpheta cayannensis (Verbenaceae).

Between the families with higher number of species, only Poaceae, Fabaceae, Cyoeraceae and Apocynaceae appear among the top ten for all areas. The list of top families in Carajás and QF is similar, however the ranking of the families is different in the two areas, with Carajás three top families being Poaceae, Fabaceae and Rubiaceae, while QF has Asteraceae, Poaceae and Orchidaceae, as its top families (Tab. 2).

Some families recorded in the FCC are not present in the canga of the QF, such as Gnetaceae, Burseraceae, Caryocaraceae, Chrysobalanaceae, Combretaceae, Costaceae, Marantaceae, Marcgraviaceae, Simaroubaceae, Trigoniaceae. Gnetaceae is a pantropical family with its centre of diversity in the Amazon (Price 1996). Widely distributed in the Amazon, Gnetum nodiflorum,the only gymnosperm species found in the canga until the present moment was found in forest groves over canga in Carajás. Some angiosperm families not recorded in the QF have their diversity centre in the Amazon biome, such as Chrysobalanaceae, Burseraceae, Costaceae, Combretaceae, Marantaceae (Daly et al. 2012; Stace 2010; Costa et al. 2011).

Many of the genera with higher species diversity in FCC were not present or had low representativeness at the QF (Viana & Lombardi 2007; Carmo & Jacobi 2012; Messias & Carmo 2015): Philodendron and Anthurium (Araceae), Justicia and Ruellia (Acanthaceae), Rhynchospora (Cyperaceae), Peperomia and Piper (Piperaceae). From these, only Rhynchospora occupies open types of vegetation, while the others had most of their species associated to forest formations within the FCC.

Likewise, amongst the angiosperm families that are well represented in the QF, many were not recorded at the FCC: Amaryllidaceae, Apiaceae, Aquifoliaceae, Araliaceae, Brassicaceae, Campanulaceae, Clethraceae, Cunoniaceae, Ericaceae, Haloragaceae, Hypoxidaceae, Juncaceae, Pentaphyllaceae, Peraceae, Rhamnaceae, Rosaceae, Scrophulariaceae, Symplocaceae, Theaceae, Violaceae and Winteraceae (Viana & Lombardi 2007; Carmo & Jacobi 2012; Messias & Carmo 2015). Many of these families are associated to temperate climates, such as Aquifoliaceae, Campanulaceae, Clethraceae, Cunoniaceae, Ericaceae, Rosaceae, Symplocaceae, Theaceae, Violaceae and Winteraceae (Safford 1999), and this may explain their absence in the Carajás region, where monthly temperatures vary between 25,1ºC and 26.3ºC, with the absolute minimum between 15.6ºC and 18.3ºC recorded between July and October, and the maximum of 34.3ºC e 38.1ºC spread through all the remaining months (Viana et al. 2016).

Another important difference between the QF and FCC floras is the high floristic representativity of some of the families in the QF that is not obsered at the FCC. For instance, the Eriocaulaceae (QF 5% × 1.2% FCC), Xyridaceae (1.4% × 0.5%) and Velloziaceae (1% × 0.1%) show this pattern, as well as several genera that are absent or poorly represented at the FCC, such as Baccharis (QF 18 spp. × 0 FCC), Lippia (10 × 1), Microlicia (9 × 0), Ditassa (9 × 0) and Stachytarpheta (8 × 1) (Carmo & Jacobi 2012). In this case is possible to notice the strong influence of the Cerrado and campo rupestre from the Espinhaço Ragne in the floristic composition of the canga of the QF. The Eriocaulaceae, Xyridaceae and Velloziaceae appear among the families with more species in the quartzitic campo rupestre of the Espinhaço Range, while the abovementioned genera have an important rôle in the composition of several floras studied in that region (Giulietti et al. 1987; Giulietti & Pirani 1988) and may thus influence the QF canga composition.

Recent studies by Zappi et al. (2017) point to strong influence of the Atlantic Rainforest biome in the compositon of the QF canga. The data generated by FCC make it possible to envisage the influence of the Amazon Rainforest over the floritic composition of the canga of Carajás. To enable us to extend and deepen a discussion within this context it is necessary to perform quantitative analyses and to obtain more representative sample of other neotropical canga areas, involving different biogeographical context. Other important areas, such as the Rio Santo Antônio basin (transition between Cerrado and Mata Atlântica) and the Vale do Peixe Bravo (Caatinga), in Minas Gerais state, as well as the canga of Caetité, within the Bahian Caatinga, still lack suitable authoritative floristic lists (Carmo & Kamino 2015). For the moment, only the canga of Carajás (Amazon Rainforest) and the Quadrilátero Ferrífero (transition between Atlantic Rainforest and Cerrado) are relatively well sampled, but it is necessary to intensify collections and listing exercise for the rest of the areas of canga of Brazil.

Conclusion

The collecting effort invested in getting to know the areas of canga of the FLONA of Carajás and the PNCF and the involvement of 131 plant taxonomistas in the study of these specimens have built the most detailed floristic study so far for an area in the Brazilian Amazon. The results of the total publication of the FCC show an impressive species richness that is significantly higher than what was referred earlier, surpassing even the estimates made at the start of the project. Supposing this is a pattern that may encompass other plant formations in the Amazon, it is necessary to apply similar effort to other sites within this biome, especially focussing on conservation units, coupled with the involvement of taxonomists, to start to reveal the real biodiversity of the Amazon flora.

The floristic composition of the canga of Carajás has been revealed as different from the canga of the Quadrilátero Ferrífero, in Minas Gerais, and of Corumbá, in Mato Grosso do Sul. Despite the similarity between the iron-rich substrate where this vegetation occurs, the phytogeographic context appears to have a determining role in the floristic identity of each site.

The richness and singularity of the local flora, including its endemic species, frame the area of Carajás as an important region for the conservation of the Amazon flora. Associated to the threat that land transformation for mining poses towards this vegetation, as well as the man made fires and competition with exotic species through land use in the general region, the situation calls for rigorous planning towards conservation of the flora of the canga of Carajás. We hope that the detailed knowledge regarding taxonomy and distribution of the canga species can be of use to provide a basis for the management of this area, contributing to the informed dialogue between the productive sector and the organs that provide the environmental licencing in order to safeguard the country´s nature.

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Acknowledgements

We would like the Museu Paraense Emílio Goeldi (MPEG) and Instituto Tecnológico Vale (ITV) for the infrastructure and support that were fundamental to develop this project. The financial support was provided by the project that represented the base for the Term of Agreement between MPEG/ITVDS/FADESP (01205.000250/2014-10) and the one approved by CNPq (processo 455505/2014-4). The International Capacitation Programme (MPEG/MCTI) for the PCI and CAPES for the post-doc grant provided to the first author. To CNPq for the DTI, PCI and Productivity grant for several students and researchers at MPEG. To MCTI/CNPq/VALE-ITV for the Human Resources Formation in the Mining Sector Project, including DTI, PDI, MBEV grants for students and researchers from other institutions. To the PCI grantees from MPEG, Climbiê Hall, Júlia Meirelles, Ana Kelly Koch, Clebiana de Sá Nunes, Aluísio Fernandes Júnior, and the DTI grantees from ITV Mayara Pastore, Matheus Nogueira, Liziane Vilela Vasconcelos, Marcos Enoque, Aline Joseph Ramalho and Ana Paula Cruz. Without this large team of grant holders it would have been impossible to achieve such an expressive number of plant collections and to conclude the treatments for the FCC. To the short-term PCI grants from MPEG, Raymond Harley, Roberto Salas, Julio Lombardi, Fernanda Antunes Carvalho, José Floriano Pastore, Regina Célia Oliveira, André Amorin and Renato Goldenberg. To the Post-graduate Programme in Biological Sciences - Tropical Botany - UFRA/MPEG and students that developed their projects in line of the FCC. To Dr Marlúcia Bonifácio Martins (MPEG), Dr Ana Vilacy Galúcio (MPEG), Dr Anna Luiza Ilkiu-Borges and Dr Vera Lúcia Imperatriz Fonseca (ITV), their dedication to consolidate and oversee the Term of Agreement between MPEG and ITV. To Dr. José Siqueira for enabling the necessary conditions to the development of the technical cooperation between MPEG and ITV. To the herbarium curators of all institutions involved, especially to BHCB, IAN, INPA, RB, Alexandre Salino, Helena Joseane Raiol, Mike Hopkins and Rafaela Forzza for their support during the whole project. To the MG herbarium technical staff, Ione Bemerguy, Antônio Elielson Rocha, Maria de Fátima Almeida, Júlio Melo, Mário Rosa and in special to Raimundo Procópio Bahia (Doca, in memoriam) for the efficiency in processing the samples collected and distributing loans to the collaborating researchers. To the ICMBio, especially Frederico Drumond Martins, for the collecting permits and field support provided. To Alexandre Castilho and Leonardo for the logistic support in the study area granted for students and researchers from ITV. To the 131 taxonomists from Brazil and abroad that were committed with preparation of monographs, especially to Dr. André dos Santos Bragança Gil, Dr. Ricardo Secco, Dr. João Ubiratan Moreira Santos from MPEG, who supervised several students, contributing with important plant treatments for the FCC. The first and last authors thank Leonardo Vianna da Costa e Silva, for presenting them to the Serra dos Carajás, and to Deco Mota Viana, for all the happiness. To the Rodriguésia bulletin and to the Jardim Botânico do Rio de Janeiro for the successful partnership that accomplished the publication of the FCC. We would like to give a special vote of thanks to chief editor Dr. Vidal Mansano who, representing the Editorial Commitee of this bulletin, accepted our request to publish and followed up the process throughout the project, and to all editors and reviewers that took part in this venture, entrusting the specific editors of the flora with this important task. The quality of the publication would not have been the same if we did not count with total commitment from Simone Bittencourt.

Editor de área: Dr. Vidal Mansano

Received: March 21, 2018; Accepted: May 18, 2018

4 Author for correspondence: pedroviana@museu-goeldi.br

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