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Flora of Baturité, Ceará: a Wet Island in the Brazilian Semiarid

Abstract

The biota of the humid mountain ranges of the Brazilian semiarid is still poorly understood. In order to fill this scientific gap, we carried out an extensive survey along altitudinal ranges (400m - 1,000 m) on both the windward and the leeward slopes of the Baturité Mountain Range, in the state of Ceará state. We registered 400 plant species and 92 families. The Myrtaceae (36 spp.), Fabaceae (25 spp.), Rubiaceae (20 spp.) and Bromeliaceae (15 spp.) families predominated on the windward slope; while Fabaceae (19 spp.), Myrtaceae (14 spp.) and Euphorbiaceae (11 spp.) were the most abundant on the leeward slope. As we expected, the species richness of trees, shrubs, subshrubs, epiphytes and terrestrial herbs was positively correlated with the altitude (R2 > 0.60). Above 800 m, we registered 273 species exclusive to the windward slope, 81 exclusive to the leeward slope, and 46 shared species. Therefore, management actions must consider the spatial heterogeneity, distribution and taxa richness.

Keywords:
Deciduous Tropical Forest; Evergreen Tropical Forest; Semideciduous Tropical Forest; Steppic Savanna

1. INTRODUCTION AND OBJECTIVES

The mountain ranges in the semiarid domain of Northeastern Brazil stand out from the surrounding flattened landscape expressing a climatic exception (Mantovani et al., 2017Mantovani W, Anjos L, Monteiro RF, Araujo FS. A conservação da biodiversidade no domínio caatinga. In: Mantovani W, Monteiro RF, Anjos L, Cariello MO, editors. Pesquisas em unidades de conservação no domínio da Caatinga subsídios à gestão. Fortaleza: Edições UFC; 2017. p. 81-122.; Moro et al., 2015Moro MF, Macedo MB, Moura-Fé MM, Castro AS, Costa RC. Vegetação, unidades fitoecológicas e diversidade paisagística do estado do Ceará. Rodriguésia 2015; 66(3): 717-743. 10.1590/2175-7860201566305
https://doi.org/10.1590/2175-78602015663...
). They occupy approximately 5% of the northeastern surface, being scattered throughout the states of Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas and Bahia (Souza & Oliveira, 2006Souza MJN, Oliveira VPV. Os enclaves úmidos e sub-úmidos do semi-árido do nordeste brasileiro. Revista Mercator 2006; 5(9): 85-102.). On windward slopes and at higher altitudes, the climate is cooler and wetter, whereas drier climates occur on leeward slopes and at lower altitudes (Nimer, 1989Nimer E. Climatologia do Brasil. 2nd ed. Rio de Janeiro: IBGE-SUPREN; 1989.). This climatic variation leads to a spatially heterogeneous flora along the altitudinal gradient, consequently increasing the local species richness (Ferraz & Rodal, 2006Ferraz EMN, Rodal MJN. Caracterização fisionômica-estrutural de um remanescente de floresta ombrófila montana de Pernambuco, Brasil. Acta Botanica Brasilica 2006; 20(4): 911-926. 10.1590/S0102-33062006000400015
https://doi.org/10.1590/S0102-3306200600...
; Homeier et al., 2010Homeier J, Breckle SW, Günter S, Rollenbeck RT, Leuschner C. Tree diversity, forest structure and productivity along altitudinal and topographical gradients in a species-rich Ecuadorian montane rain forest. Biotropica 2010; 42(2): 140-148. 10.1111/j.1744-7429.2009.00547.x
https://doi.org/10.1111/j.1744-7429.2009...
).

Evergreen and Semideciduous Forests have been commonly reported along this altitudinal gradient (Rodal & Sales, 2008Rodal MJN, Sales MF. Panorama of the montane forests of Pernambuco, Brazil. In: Thomas WW, editor. The Atlantic coastal forest of northeastern Brazil. New York: Botanical Garden Press; 2008. p. 541-559.; Silva & Figueiredo, 2013Silva MEA, Figueiredo MF. Flora fanerogâmica de um enclave úmido no Ceará: Serra da Meruoca. Enciclopédia Biosfera 2013; 9(17): 2811-2820.). Since this flora is a remnant of the Brazilian Atlantic Rain Forest (Barbosa et al., 2004Barbosa MRDV, Agra MDF, Sampaio EVSB, Cunha JD, Andrade LD. Diversidade florística na Mata do Pau-Ferro, Areia, Paraíba. Brasília: Ministério do Meio Ambiente; 2004 [cited 2019 Oct. 29]. p. 111-122. Available from: Available from: https://bit.ly/2WFrhMI
https://bit.ly/2WFrhMI...
), these mountain ranges have been considered a priority for the conservation of Brazilian biodiversity (Lopes et al., 2017Lopes SDF, Ramos MB, Almeida GRD. The role of mountains as refugia for biodiversity in Brazilian Caatinga: conservationist implications. Tropical Conservation Science 2017; 10: 1-12. 10.1177/1940082917702651
https://doi.org/10.1177/1940082917702651...
; MMA, 2000Ministério do Meio Ambiente - MMA. Avaliação e ações prioritárias para a conservação da biodiversidade da floresta atlântica e campos sulinos. Brasília: MMA/SBF; 2000.). Especially in the Brazilian Northeast, the mountainous areas are critically important for the preservation of regional ecosystems, because they represent natural refuges for biota (Silva et al., 2014Silva FKG, Lopes FS, Lopez LCS, Melo JIM, Trovão DMDBM. Patterns of species richness and conservation in the Caatinga along elevational gradients in a semiarid ecosystem. Journal of Arid Environments 2014; 110, 47-52. 10.1016/j.jaridenv.2014.05.011
https://doi.org/10.1016/j.jaridenv.2014....
).

Thirty-five percent (35%) of the mountain ranges are located within Ceará State (Sobrinho, 1971Vasconcelos Sobrinho J. As regiões naturais do Nordeste, o meio a civilização. Recife: Conselho do Desenvolvimento de Pernambuco; 1971.), but information on its biota composition is still incipient (Bétard et al., 2007Bétard F, Peulvast JP, Claudino-Sales V. Caracterização morfopedólogica de uma serra úmida no semi-árido do nordeste brasileiro: o caso do Maciço de Baturité-CE. Mercator-Revista de Geografia da UFC 2007; 6(12): 107-126.; Kamimura et al., 2017Kamimura VA, Moraes PLR, Ribeiro HL, Joly CA, Assis MA. Tree diversity and elevational gradient: the case of Lauraceae in the Atlantic Rainforest. Flora 2017; 234: 84-91. 10.1016/j.flora.2017.05.013
https://doi.org/10.1016/j.flora.2017.05....
; Lima & Mansano, 2011Lima JR, Mansano VDF. A família Leguminosae na Serra de Baturité, Ceará, uma área de Floresta Atlântica no semiárido brasileiro. Rodriguésia 2011; 62(3): 563-613. 10.1590/2175-7860201162309
https://doi.org/10.1590/2175-78602011623...
). Plant cover and soils of most local wetlands have suffered from intensive and chronic anthropogenic disturbances (Souza & Oliveira, 2006Souza MJN, Oliveira VPV. Os enclaves úmidos e sub-úmidos do semi-árido do nordeste brasileiro. Revista Mercator 2006; 5(9): 85-102.). Therefore, reports on floristic composition are crucial for implementing sustainable use actions, since the biodiversity degradation on the Baturité Mountain Range has been happening since the colonial period (Mantovani, 2006Mantovani W. Conservação de biodiversidade: importância das serras úmidas no Nordeste semi-árido brasileiro. In: Oliveira TS, Araújo FS, editors. Diversidade e conservação da biota na Serra de Baturité, Ceará. Fortaleza: Edições UFC-COELCE ; 2006. p. 3-15.; Oliveira et al., 2006Oliveira TS, Figueiredo MA, Nogueira RS, Sousa SC, Souza SSG, Romero RE. Histórico dos impactos antrópicos e aspectos geoambientais da serra de Baturité, Ceará. In: Oliveira TS, Araújo FS, editors. Diversidade e conservação da biota na Serra de Baturité, Ceará. Fortaleza: Edições UFC-COELCE ; 2006. p. 19-70.). Our study shows the need for registering, cataloguing and presenting the biodiversity of such peculiar environments. Thus, we aim to analyze the plant composition and species richness along the altitudinal gradient on the windward and leeward slopes of Baturité Mountain Range, in Ceará State. We also intend to spread information on the local flora and on the protected areas of Northeastern Brazil.

2. MATERIALS AND METHODS

2.1. Study site

The Baturité Mountain Range is a residual relief with an extension of 800 km2 and moderate altitudes (800 - 1,115 m). It is a Precambrian crystalline complex located in the northeast of Ceará State, 70 km from the coast (Bétard et al., 2007Bétard F, Peulvast JP, Claudino-Sales V. Caracterização morfopedólogica de uma serra úmida no semi-árido do nordeste brasileiro: o caso do Maciço de Baturité-CE. Mercator-Revista de Geografia da UFC 2007; 6(12): 107-126.; Souza & Oliveira, 2006Souza MJN, Oliveira VPV. Os enclaves úmidos e sub-úmidos do semi-árido do nordeste brasileiro. Revista Mercator 2006; 5(9): 85-102.). From the elevation of 600 m up, the mountain range is a strict protected area, the APA de Baturité. The first protected area was established by state law no. 20,956, on September 18, 1990, which was later altered by state law no. 27,290, on December 15, 2003. Currently, it encompasses 32,690 hectares.

Both the altitude and the geographical position favor an orographic effect at the top and on the windward slope (north-eastward) of the mountain, whereby a humidity of > 1,000 mm × year-1 is carried by South Atlantic trade winds. On the leeward slope (westward), the precipitation is below 1,000 mm × year-1 (Mantovani, 2006Mantovani W. Conservação de biodiversidade: importância das serras úmidas no Nordeste semi-árido brasileiro. In: Oliveira TS, Araújo FS, editors. Diversidade e conservação da biota na Serra de Baturité, Ceará. Fortaleza: Edições UFC-COELCE ; 2006. p. 3-15.; Santos et al., 2012Santos FLA, Medeiros EM, Souza MJN. Contexto hidroclimático do enclave úmido do maciço de Baturité-Ceará: potencialidades e limitações ao uso da terra. Revista Geonorte 2012; 3(9): 1056-1065.; Souza & Oliveira, 2006Souza MJN, Oliveira VPV. Os enclaves úmidos e sub-úmidos do semi-árido do nordeste brasileiro. Revista Mercator 2006; 5(9): 85-102.). In order to register the local species composition and richness, we selected three well-conserved areas on each slope, at the following altitudinal ranges: 400 - 600 m a.s.l., 600 - 800 m a.s.l. and above 800 m a.s.l. We chose these areas in order to comprise the climatic and physiognomic variations along the altitudinal range on both slopes (Table 1).

Table 1
Location and characterization of the studied areas on the Baturité Mountain Range, Ceará State, Brazil.

The Forested Steppic Savanna (FSS; “Caatinga”) is composed of a thorny deciduous vegetation, predominant at lower altitudes. Along the altitudinal range, the FSS is gradually replaced by a forest vegetation along the altitudinal range. On the windward slope, there are: i) a Seasonal Semidecidous Submountain Forest (SSSF) 400 - 600 m; and ii) a Seasonal Evergreen Mountain Forest (SEMF) above 600 m. On the leeward slope, there are: i) a transition from the FSS to the SDSF (Seasonal Deciduous Submountain Forest) 400 - 600 m; ii) a Seasonal Deciduous Mountain Forest (SDMF) 600 - 800 m; and iii) a Seasonal Evergreen Mountain Forest (SEMF) above 800 m (see Table 1).

2.2. Data collection

In order to list the plants, we sampled 200 quadrants in each of the six areas, according to the procedures suggested by Araújo et al. (2006Araújo FS, Gomes VS, Silveira AP, Figueiredo MA, Oliveira RF, Bruno MMA et al. Efeito da variação topoclimática na fisionomia e estrutura da vegetação da serra de Baturité, Ceará. In: Oliveira TS, Araújo FS, editors. Diversidade e conservação da Biota na Serra de Baturité, Ceará. Fortaleza: Edições UFC-COELCE; 2006. p. 75-136.). In addition, we completed our list researching the samples of the EAC Herbarium of the Universidade Federal do Ceará. We also revised and updated the taxonomic identifications with the aid of specialists from the following herbaria: EAC, PEUFR, IPA and CEPEC. The taxonomic classification that we used follows the APG IV system (2016)Angiosperm Phylogeny Group - APG IV. An udpdate of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 2016; 181: 1-20. 10.1111/boj.12385
https://doi.org/10.1111/boj.12385...
. The names of botanical families, genera, species and authorships were confirmed in the International Plant Names Index (Royal Botanic Gardens, Kew et al., 2015Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; & Australian National Botanic Gardens. International Plant Names Index [Internet]. 2015 [cited 2019 Oct. 29]. Available from: Available from: http://www.ipni.org
http://www.ipni.org...
) and in the list of Brazilian Flora 2020 (JBRJ, 2016Jardim Botânico do Rio De Janeiro - JBRJ. Flora do Brasil 2020 em construção. Rio de Janeiro: Jardim Botânico do Rio de Janeiro; 2016 [cited 2019 Oct. 29]. Available from: Available from: http://floradobrasil.jbrj.gov.br
http://floradobrasil.jbrj.gov.br...
).

To categorize the vegetation physiognomy, we used Whittaker’s classification of plant growth forms (1975Whittaker RH. Communities and ecosystems. 2nd ed. New York: Macmillan; 1975.). The growth forms are adequate indicators of communities, since they can reflect global and local climatic conditions. We adjusted Whittaker’s (1975)Whittaker RH. Communities and ecosystems. 2nd ed. New York: Macmillan; 1975. system as following: i) trees: > 3 m-height woody plants; ii) shrubs: > 3 m-height woody plants with main branches developing at approximately 50 cm above ground; iii) subshrubs: < 2 m-height plants with a woody main stem and herbaceous secondary branching; iv) terrestrial herbs: land plants with herbaceous aerial stems; v) epiphytes: plants with herbaceous stems that use other plants as support; vi) vines: plants with prolonged stems that twine around a substrate; and vii) hemiparasites: photosynthetic plants that withdraw sap from their host plants.

We obtained precipitation data from the Fundação Cearense de Meteorologia e Recursos Hídricos (FUNCEME, 2017Fundação Cearense de Meteorologia e Recursos Hídricos - Funceme. Dados dos postos pluviométricos do Ceará. 2017 [cited 2019 Oct. 29]. Available from: Available from: https://bit.ly/2LzMrWm
https://bit.ly/2LzMrWm...
), at the following local stations, close to our studied areas: Baturité-n22, Pacoti-n105, Guaramiranga-54, Mulungu-n98 and Caridade-n31. Data showed a seasonal rainfall regime with precipitation concentrated from January to May, driven by the Intertropical Convergence Zone. However, stations located at distinct altitudes and different mountain slopes showed remarkable differences in the annual rainfall regime. Greater precipitation was found in the northeastern slope and at higher portions of the mountain range. The soil classification of the studied areas followed that of Oliveira et al. (2006Oliveira TS, Figueiredo MA, Nogueira RS, Sousa SC, Souza SSG, Romero RE. Histórico dos impactos antrópicos e aspectos geoambientais da serra de Baturité, Ceará. In: Oliveira TS, Araújo FS, editors. Diversidade e conservação da biota na Serra de Baturité, Ceará. Fortaleza: Edições UFC-COELCE ; 2006. p. 19-70.). Altitudinal ranges and coordinates were measured in the field with a GPS navigation device (Table 1).

2.3. Data analysis

We organized our list by family, species, vernacular name, growth form, altitudinal range, phytophysiognomy and collector number. Samples are deposited at EAC. We estimated the global richness, as well as the richness at family and species levels, and by phytophysiognomy. We analyzed the relationship between species richness, growth form (dependent variable), and altitude (independent or predictive variable) through a simple linear regression using the Bioestat 5.0 Software (Ayres et al., 2007Ayres M, Ayres M Jr, Ayres DL, Santos ADA. Bioestat 5.0: aplicações estatísticas nas áreas das ciências bio-médicas. Belém: Instituto Mamirauá; 2007.)

3. RESULTS AND DISCUSSION

We registered 400 morphospecies distributed within 92 families. A total of 23 taxa (13%) were identified only to the genus due to the lack of or inadequacy of reproductive organs. The families with greatest species richness were Myrtaceae (43 spp.), Fabaceae (38 spp.), Euphorbiaceae (21 spp.), Rubiaceae (20 spp.), Melastomataceae (14 spp.), and Bromeliaceae, Erythroxylaceae and Orchidaceae, with 10 species each (Table 2). The high richness of vascular plants confirms the documented pattern for mountainous areas of the Brazilian semiarid: they are more diverse than the surrounding Caatinga, since they contain a mix of Caatinga and Atlantic Forest species (Carnaval et al. 2009Carnaval AC, Hickerson MJ, Haddad CFB, Rodrigues MT, Moritz C. Stability predicts genetic diversity in the Brazilian Atlantic forest hotspot. Science 2009; 323(5915): 785-789. 10.1126/science.1166955
https://doi.org/10.1126/science.1166955...
; Leite et al., 2016Leite YLR, Costa LP, Loss AC, Rocha RG, Batalha-Filho H, Bastos AC et al. Neotropical forest expansion during the last glacial period challenges refuge hypothesis. PNAS 2016; 113(4): 1008-1013. 10.1073/pnas.1513062113
https://doi.org/10.1073/pnas.1513062113...
; Lopes et al., 2017Lopes SDF, Ramos MB, Almeida GRD. The role of mountains as refugia for biodiversity in Brazilian Caatinga: conservationist implications. Tropical Conservation Science 2017; 10: 1-12. 10.1177/1940082917702651
https://doi.org/10.1177/1940082917702651...
). In addition, the richness of 400 species of vascular plants, found on the Baturité Mountain Range, is higher than the species richness registered for other similar Brazilian forests, e.g., Pau-Ferro Ecological Reserve, with 309 species, in Paraíba State (Barbosa et al., 2004Barbosa MRDV, Agra MDF, Sampaio EVSB, Cunha JD, Andrade LD. Diversidade florística na Mata do Pau-Ferro, Areia, Paraíba. Brasília: Ministério do Meio Ambiente; 2004 [cited 2019 Oct. 29]. p. 111-122. Available from: Available from: https://bit.ly/2WFrhMI
https://bit.ly/2WFrhMI...
), Brejo Madre de Deus, with 293 species, in Pernambuco State (Nascimento et al., 2012Nascimento LM, Rodal MJN, Silva AG. Florística de uma floresta estacional no Planalto da Borborema, nordeste do Brasil. Rodriguésia 2012; 63(2): 429-440. 10.1590/S2175-78602012000200015
https://doi.org/10.1590/S2175-7860201200...
) and Meruoca Sierra, with 100 species, in Ceará State (Silva & Figueiredo, 2013Silva MEA, Figueiredo MF. Flora fanerogâmica de um enclave úmido no Ceará: Serra da Meruoca. Enciclopédia Biosfera 2013; 9(17): 2811-2820.), highlighting the importance of this montane forests for the conservation of tropical biodiversity.

Our results show that the richness increases towards the top of the mountain, and that it is greater on the windward slope at the Evergreen Forest. We registered a total of 255 species (64%) and 69 families (75%) at higher altitudes, above 800 m in the Baturité Mountain Range - joined data from Arvoredo and Lagoa sites. On the windward slope, below 800 m-on Sinimbu and Taveiras sites together-, we registered 175 species (44%) and 62 families (67%); whereas, on the leeward slope, below 800 m-on Jardim and Salva-Vidas sites together-, we registered 127 species (32%) and 50 families (54%) (Table 2). Such differences reflect the combined effects of ocean winds, altitudinal variation and position of the slope. These results also corroborate the pattern documented by Lopes et al. (2008Lopes CGR, Ferraz EMN, Araújo EL. Physiognomic-structural characterization of dry-and humid-forest fragments (Atlantic Coastal Forest) in Pernambuco State, NE Brazil. Plant Ecology 2008; 198(1): 1-18. 10.1007/s11258-007-9380-z
https://doi.org/10.1007/s11258-007-9380-...
), Lima et al. (2011Lima JR, Sampaio EVSB, Rodal MJN, Araújo FS. Physiognomy and structure of a seasonal deciduous forest the Ibiapaba plateau, Ceará, Brazil. Rodriguésia 2011; 62(2): 379-389. 10.1590/2175-7860201162212
https://doi.org/10.1590/2175-78602011622...
), Kamimura et al., (2017Kamimura VA, Moraes PLR, Ribeiro HL, Joly CA, Assis MA. Tree diversity and elevational gradient: the case of Lauraceae in the Atlantic Rainforest. Flora 2017; 234: 84-91. 10.1016/j.flora.2017.05.013
https://doi.org/10.1016/j.flora.2017.05....
) and BFG (2015)Brazil Flora Group - BFG. Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 2015; 66: 1085-1113. 10.1590/2175-7860201566411
https://doi.org/10.1590/2175-78602015664...
, that indicates dry forests have significantly less species and families compared to humid-forests.

In the Seasonal Evergreen Forest located both on the windward and leeward slopes-above 600 m and 800 m, respectively (Table 1)-, the most diverse families were Myrtaceae (36 spp.), Fabaceae (25 spp.), Rubiaceae (20 spp.), Bromeliaceae (15 spp.), Melastomataceae (14 spp.), Euphorbiaceae (13 spp.) and Orchidaceae (10 spp.). On the leeward slope, where a Seasonal Deciduous Forest and a Forested Steppic Savanna predominate, Fabaceae (19 spp.), Myrtaceae (14 spp.) and Euphorbiaceae (11 spp.) were the most diverse families. In the Fabaceae family, the subfamilies presented distinct species richness on windward and leeward slopes. In the former, Mimosoideae (8 spp.) and Faboideae (7 spp.) predominated, whereas in the latter the most representative were Mimosoideae (13 spp.) and Caesalpinioideae (11 spp.).

In the Baturité Mountain Range leeward slope, where Deciduous Forests and Forested Savannas were the most representative (see Alcoforado-Filho et al., 2003Alcoforado-Filho FG, Sampaio EVSB, Rodal MJN. Florística e fitossociologia de um remanescente de vegetação caducifólia espinhosa arbórea em Caruaru, Pernambuco. Acta Botanica Brasilica 2003; 17(2): 287-303. 10.1590/S0102-33062003000200011
https://doi.org/10.1590/S0102-3306200300...
; Cestaro & Soares, 2004Cestaro LA, Soares JJ. Variações florística e estrutural e relações fitogeográficas de um fragmento de floresta decídua no Rio Grande do Norte, Brasil. Acta Botanica Brasilica 2004; 18(2): 203-218. 10.1590/S0102-33062004000200001
https://doi.org/10.1590/S0102-3306200400...
; Ferraz et al., 2004Ferraz EMN, Araújo EL, Silva SI. Floristic similarities between lowland and montane areas of Atlantic Coastal Forest in Northeastern Brazil. Plant Ecology 2004; 174(1): 59-70.), the subfamilies Mimosoideae and Caesalpinioideae, predominated. Even though Myrtaceae was among the most diverse families on the leeward slope, its richness was higher on the windward slope. Furthermore, most of its species (84%) occur above 600 m, indicating that the richness of Myrtaceae in the Brazilian semiarid is more associated with areas with higher water availability than with the surrounding Caatinga. The area of occurrence of Myrtaceae in the Evergreen Forest, both on the windward and on the leeward slopes, above 600 m and 800 m, respectively, confirms the pattern reported by Peixoto & Gentry (1990Peixoto AL, Gentry A. Diversidade e composição florística da mata de tabuleiro na Reserva Florestal de Linhares (Espírito Santo, Brasil). Revista Brasileira de Botânica 1990; 13(1): 19-25.), also observed in the Atlantic Domain as a whole. Furthermore, Myrtaceae, Fabaceae, Rubiaceae, Bromeliaceae, Melastomataceae, Euphorbiaceae and Orchidaceae are abundant in Tropical Rainforests as well, including lowland and highland Seasonal Evergreen Forests in the States of Pernambuco and Paraíba (Rodal & Nascimento 2002Rodal MJ, Nascimento LM. Levantamento florístico da floresta serrana da reserva biológica de Serra Negra, microrregião de Itaparica, Pernambuco, Brasil. Acta Botanica Brasilica 2002; 16(4): 481-500. 10.1590/S0102-33062002000400009
https://doi.org/10.1590/S0102-3306200200...
; Andrade & Rodal, 2004Andrade KVSA, Rodal MJN. Fisionomia e estrutura de um remanescente de floresta estacional semidecidual de terras baixas no nordeste do Brasil. Revista Brasileira de Botânica 2004; 27(3): 463-474. 10.1590/S0100-84042004000300007
https://doi.org/10.1590/S0100-8404200400...
; Barbosa et al., 2004Barbosa MRDV, Agra MDF, Sampaio EVSB, Cunha JD, Andrade LD. Diversidade florística na Mata do Pau-Ferro, Areia, Paraíba. Brasília: Ministério do Meio Ambiente; 2004 [cited 2019 Oct. 29]. p. 111-122. Available from: Available from: https://bit.ly/2WFrhMI
https://bit.ly/2WFrhMI...
; Ferraz & Rodal, 2006Ferraz EMN, Rodal MJN. Caracterização fisionômica-estrutural de um remanescente de floresta ombrófila montana de Pernambuco, Brasil. Acta Botanica Brasilica 2006; 20(4): 911-926. 10.1590/S0102-33062006000400015
https://doi.org/10.1590/S0102-3306200600...
; Nascimento et al., 2012Nascimento LM, Rodal MJN, Silva AG. Florística de uma floresta estacional no Planalto da Borborema, nordeste do Brasil. Rodriguésia 2012; 63(2): 429-440. 10.1590/S2175-78602012000200015
https://doi.org/10.1590/S2175-7860201200...
; Rodal & Sales, 2008Rodal MJN, Sales MF. Panorama of the montane forests of Pernambuco, Brazil. In: Thomas WW, editor. The Atlantic coastal forest of northeastern Brazil. New York: Botanical Garden Press; 2008. p. 541-559.). Species and family similarities to this type of forest are likely associated with water availability on the soil, from rainfall or dew.

Aspidosperma pyrifolium Mart., Bauhinia cheilantha (Bong.) Steud., Cordia glazioviana (Taub.) Gottschling & J. S. Mill., Croton blanchetianus Baill., and Mimosa caesalpiniifolia Benth. occurred below 600 m on the windward slope. These taxa are commonly found in the Caatinga. Some species are strictly distributed in wetter areas, above 800 m on the windward slope and above 600 m on the leeward slope, such as: Albizia polycephala (Benth.) Killip ex Record, Apeiba tibourbou Aubl., Byrsonima crispa A. Juss., Cassia ferruginea (Schrad.) Schrad. ex DC. var. ferruginea, Cupania racemosa (Vell.) Radlk., Guazuma ulmifolia Lam., Inga marginata Willd., Myrciaria ferrugínea O.Berg, Ouratea polygyna Engl., Podocarpus sellowii Klotzsch ex. Endl., Pouteria macrophylla (Lam.) Eyma, Vismia guianensis (Aubl.) Choisy and Zanthoxylum rhoifolium Lam. We note that Podocarpus sellowii was found at one collecting site only (Arvoredo). At lower altitudes of the windward slope, some exclusive species occurred, such as: Alchornea glandulosa subsp. iricurana (Casar.) Secco, Attalea speciosa Mart. ex Spreng., Casearia grandiflora Cambess., Chrysophyllum gonocarpus (Mart. & Eichler ex Miq.) Engl., Coccoloba parimensis Benth., Coussarea contracta (Walp.) Müll.Arg. var. contracta, Oreopanax capitatus (Jacq.) Decne. & Planch., Parkia pendula (Willd.) Benth. ex Walp., Protium warmingianum Marchand., Pseudobombax marginatum (A.St.-Hil.) A. Robyns and Zizyphus undulata Reissek.

Concerning growth forms, the studied flora was composed of 214 trees (54%), 82 shrubs, 49 terrestrial herbs, 23 epiphytic herbs, 18 vines, 10 subshrubs and 4 hemiparasites. There was a positive correlation between altitude and richness of trees, shrubs, subshrubs, epiphytic herbs and terrestrial herbs (R2 > 0.60 and p < 0.05). However, for vines and hemiparasites, the correlation was not statistically significant (Figure 1). The richness of tree species is one of the most striking characteristics of the Tropical Forest typologies, a pattern documented in other Brazilian Atlantic forests (BFG, 2015Brazil Flora Group - BFG. Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 2015; 66: 1085-1113. 10.1590/2175-7860201566411
https://doi.org/10.1590/2175-78602015664...
; Nascimento et al., 2012Nascimento LM, Rodal MJN, Silva AG. Florística de uma floresta estacional no Planalto da Borborema, nordeste do Brasil. Rodriguésia 2012; 63(2): 429-440. 10.1590/S2175-78602012000200015
https://doi.org/10.1590/S2175-7860201200...
).

Figure 1
Linear regression of altitude and number of plants of each growth form [(a), (b), (c), (d), (e), (f), (g)] and dispersion diagrams (h) of plants.

shr: shrub; subshr: subshrub; vi: vine; th: terrestrial herb; eh: epiphytic herb; hm: hemiparasite; W: windward; L: leeward; T: top.


The species richness and composition found in our study showed that, on the windward slope above 600 m, and on the leeward slope above 800 m, the flora is more similar. In contrast, below these altitudinal levels, on both slopes, the flora differed. The positive correlation between diversity and altitude is likely a response to greater water availability in higher elevations of the altitudinal gradient. The mountains of Northeastern Brazil are relatively low, with some altitudes of approximately 1,000 m a.s.l. However, they have a more favorable water balance, milder temperatures, and lower rates of evapotranspiration and evening condensation than the lower area of the countryside depression (Mantovani et al., 2017Mantovani W, Anjos L, Monteiro RF, Araujo FS. A conservação da biodiversidade no domínio caatinga. In: Mantovani W, Monteiro RF, Anjos L, Cariello MO, editors. Pesquisas em unidades de conservação no domínio da Caatinga subsídios à gestão. Fortaleza: Edições UFC; 2017. p. 81-122.; Moro et al., 2015Moro MF, Macedo MB, Moura-Fé MM, Castro AS, Costa RC. Vegetação, unidades fitoecológicas e diversidade paisagística do estado do Ceará. Rodriguésia 2015; 66(3): 717-743. 10.1590/2175-7860201566305
https://doi.org/10.1590/2175-78602015663...
; Souza & Oliveira, 2006Souza MJN, Oliveira VPV. Os enclaves úmidos e sub-úmidos do semi-árido do nordeste brasileiro. Revista Mercator 2006; 5(9): 85-102.). This may explain why the plant species richness in the semiarid domain of Northeastern Brazil increases with higher altitudes.

It is also worth mentioning that the greater richness of epiphytic herbs (18 out of 23 species) at altitudes above 800 m reflects a higher humidity. According to Gentry (1988Gentry AH. Changes in plant community diversity and floristic composition on environmental and geographical gradients. Annals of the Missouri Botanical Garden 1988; 75(1): 1-34. 10.2307/2399464
https://doi.org/10.2307/2399464...
), there is a positive correlation between diversity and precipitation in the Neotropical Region. In addition, our results are in accordance with those found in other studies on the Mountain Forests of Pernambuco State (Ferraz et al., 1998Ferraz EMN, Rodal MJN, Sampaio EVSB, Pereira RDCA. Composição florística em trechos de vegetação de caatinga e brejo de altitude na região do Vale do Pajeú, Pernambuco. Brazilian Journal of Botany 1998; 21(1): 7-15. 10.1590/S0100-84041998000100002
https://doi.org/10.1590/S0100-8404199800...
; Ferraz et al., 2003Ferraz EMN, Rodal MJN, Sampaio EVSB. Physiognomy and structure of vegetation along an altitudinal gradient in the semi-arid region of northeastern Brazil. Phytocoenologia 2003; 33(1): 71-92. 10.1127/0340-269X/2003/0033-0071
https://doi.org/10.1127/0340-269X/2003/0...
; Rodal & Nascimento, 2002Rodal MJ, Nascimento LM. Levantamento florístico da floresta serrana da reserva biológica de Serra Negra, microrregião de Itaparica, Pernambuco, Brasil. Acta Botanica Brasilica 2002; 16(4): 481-500. 10.1590/S0102-33062002000400009
https://doi.org/10.1590/S0102-3306200200...
). According to Ferraz et al. (2004)Ferraz EMN, Araújo EL, Silva SI. Floristic similarities between lowland and montane areas of Atlantic Coastal Forest in Northeastern Brazil. Plant Ecology 2004; 174(1): 59-70. and Lopes et al (2008Lopes CGR, Ferraz EMN, Araújo EL. Physiognomic-structural characterization of dry-and humid-forest fragments (Atlantic Coastal Forest) in Pernambuco State, NE Brazil. Plant Ecology 2008; 198(1): 1-18. 10.1007/s11258-007-9380-z
https://doi.org/10.1007/s11258-007-9380-...
), under similar climatic and edaphic regimes, the Evergreen and the Semidecidous Montane Forests can be found closely associated in Northeastern Brazil, but demonstrate different floristic and structural compositions. Thus, protection efforts must contemplate both humid and dry forest areas.

4. CONCLUSION

In conclusion, 273 species (68%) were found exclusively at the top of the mountain and on the windward slope; 81 species (20%) were exclusively found on the leeward slope; and 46 species (12%) were found on both slopes, with a total of 400 species on the Baturité Mountain Range, in Ceará State. Our results highlight that the management actions, the restoration of degraded areas and the establishment of integral conservation on the Baturité Mountain Range must consider the spatial heterogeneity described in our work; that is, the differential plant distribution and richness both along the altitudinal gradient and between slope positions. Adequate conservation efforts should consider the total richness and the local heterogeneity.

ACKOWLEGMENTS

We are grateful to the taxonomists who gently identified the botanical material; to the land owners for allowing the data collection (Arvoredo, Lagoa, Sinimbu, Labirinto dos Taveiras, Jardim and Salva-Vidas sites); and to the Companhia de Eletricidade do Estado do Ceará (COELCE) for funding this study.

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  • FINANCIAL SUPPORT

    Companhia de Eletricidade do Estado do Ceará (COELCE).

Appendix A

List of families and species registered at distinct altitudinal levels on the Baturité Mountain Range, Ceará, Brazil.

WINDWARD ALTITUDE TOP LEEWARD FAMILY/SPECIES/AUTHOR VN GF 400-600 600-800 > 800B > 800S 600-800 400-600 C TAV SIN ARV LAG JAR SAL 1. Acanthaceae Dicliptera ciliaris Juss. subshr x V. Gomes, 912.2 Justicia aequilabris (Nees) Lindau subshr x V. Gomes, 562.2 Justicia sp. shr x V. Gomes, 2109-8 Ruellia bahiensis (Nees) Morong subshr x V. Gomes, 398 2. Alstroemeriaceae Bomarea edulis (Tussac) Herb. th x V. Gomes, 1271 3. Amaranthaceae Alternanthera brasiliana L. th x V. Gomes, 2109-1 Cyathula achyranthoides (Kunth) Moq. th x V. Gomes, 737 Iresine diffusa Humb. & Bonpl. ex Willd. Cabeça-branca th x V. Gomes, 744 4. Amaryllidaceae Hippeastrum stylosum Herb. Açucena th x x F. S. Araújo, 1612 5. Anacardiaceae Astronium fraxinifolium Schott Gonçalo-alves tre x M.A.Figueiredo, 18463 Myracrodruon urundeuva Allemão Aroeira tre x L.W.Lima-Verde, 3526 Thyrsodium spruceanum Benth. Cajazeira-brava tre x x x x V. Gomes, 1113 6. Annonaceae Cymbopetalum brasiliense (Vell.) Benth. ex Baill. Gitó-da-mata shr x x x x V. Gomes, 766 Duguetia riedeliana R.E.Fr. Ata-brava tre x V. Gomes, 936 Guatteria pogonopus Mart. Sabonete shr x x V. Gomes, 1274 Xylopia frutescens Aubl. Imbiriba tre x A. Silveira, 470 Xylopia sericea A.St.-Hil. Imbiriba tre x x A. Silveira, 144 7. Apocynaceae Aspidosperma multiflorum A.DC. Piquiá tre x x V. Gomes, 61-3 Aspidosperma pyrifolium Mart. Pereiro tre x V. Gomes, 2109-4 Aspidosperma ulei Markgr. Piquiá tre x V. Gomes, 5-32 Blepharodon bicolor Decne. vi x A. Silveira, 744 Condylocarpon isthmicum (Vell.) A.DC. vi x x V. Gomes, 1029 Macoubea sp. vi x A. Silveira, 948 8. Aquifoliaceae Ilex sapotifolia Reissek Pinho-branco, Pereira tre x x x x V. Gomes, 1011-03 9. Araceae Anthurium scandens (Aubl.) Engl. eh x x V. Gomes, 1129 Anthurium sinuatum Benth. ex Schott th x V. Gomes 1205-9 Monstera adansonii var. klotzchiana (Schott) Madison th x V. Gomes, 2209-1 Monstera praetermissa E.G.Gonç. & Temponi eh x V. Gomes, 780 Philodendron pedatum (Hook) Kunth th x V. Gomes, 902-1 Philodendron ornatum Schott th x V. Gomes, 902-2 Anthurium pentaphyllum (Aubl.) G.Don th x V. Gomes, 1026 10. Araliaceae Oreopanax capitatus (Jacq.) Decne. & Planch. Piroá tre x V. Gomes, 2704-2 Schefflera morototoni (Aubl.) Maguire et al. var. morototoni Gargaúba tre x x x V. Gomes, 1003 11. Arecaceae Attalea speciosa Mart. ex Spreng. Babaçu tre x Lima, J. R., 1127 Geonoma pohliana Mart. Palmeirinha-da-serra, Guaricana tre x x x V. Gomes, 663 Syagrus comosa (Mart.) Mart. Coco-babão, Catolé tre x x V. Gomes, 4-278 12. Asteraceae Cyrtocymura scorpioides (Lam.) H.Rob. Assa-peixe subshr x L.W. Lima-Verde, 3479-8 Gymnanthemum amygdalinum (Delile) Sch.Bip. ex Walp. Boldo shr x A. Silveira, 388 Trichogoniopsis adenantha (DC.) R.M.King & H. Rob. th x A. Silveira, 969 Vernonanthura brasiliana (L.) H.Rob. Catirina shr x x A. Silveira, 459 Wedelia alagoensis Baker Camará-de-flecha subshr x V. Gomes, 6-2 13. Balanophoraceae Langsdorffia hypogaea Mart. th x V. Gomes, 707-1 14. Begoniaceae Begonia reniformis Dryand. Begônia th x A. Silveira, 300 15. Bignoniaceae Handroanthus impetiginosus Mattos Pau-d’arco-roxo tre x x x A. Silveira, 863 Handroanthus serratifolius (A.H.Gentry) S.Grose Pau-d’arco-amarelo tre x x x V. Gomes, 597 Jacaranda brasiliana (Lam.) Pers. Caroba tre x x x x A. Silveira, 219 Lundia cordata (Vell.) DC. Cipó-de-cesta vi x A. Silveira, 295 Lundia sp. vi x V. Gomes, 894 16. Bixaceae Cochlospermum vitifolium (Willd.) Spreng. Pacotê tre x x x L. W. Lima-Verde, 3515 17. Boraginaceae Cordia alliodora (Ruiz & Pav.) Cham. tre x V. Gomes, 4-338 Cordia anabaptista Cham. Freijó tre x V. Gomes, 495 Cordia glazioviana (Taub.) Gottschling & J.S.Mill. Pau-branco-louro tre x L. W. Lima-Verde, 3492 Cordia rufescens A.DC. tre x A. Silveira, 355 Cordia taguahyensis Vell. tre x x A. Silveira, 817 Cordia toqueve Aubl. tre x V. Gomes, 1235 Cordia trichotoma (Vell.) Arráb. ex Steud. Freijó tre x x X x A. Silveira, 1605 18. Bromeliaceae Aechmea aquilega (Salisb.) Griseb. Croatá eh x x x x x V. Gomes, 546 Aechmea bromeliifolia (Rudge) Baker Croatá eh x x V. Gomes, 712 Guzmania lingulata (L.) Mez Croatá eh x A. Silveira, 1029 Guzmania monostachia (L.) Rusby ex Mez Croatá eh x x x V. Gomes, 307 Racinaea spiculosa (Griseb,) M.A.Spencer & L.B.Sm. Croatá eh x V. Gomes, 0607-15 Tillandsia juncea (Ruiz & Pav.) Poiret. Croatá eh x V. Gomes, 899 Tillandsia recurvata (L.) L. Croatá eh x x x V. Gomes, 2109-9 Tillandsia stricta Sol. var. stricta Croatá eh x x x V. Gomes, 2209-11 Vriesea oleosa Leme Croatá eh x V. Gomes, 726 Vriesea rodigasiana E. Morren Croatá eh x V. Gomes, 376 19. Burseraceae Commiphora leptophloeos (Mart.) J. B. Gillett Imburana tre x x A. Silveira, 947 Protium heptaphyllum (Aubl.) Marchand subsp. heptaphyllum Almécega tre x x x V. Gomes, 1120 Protium warmingianum Marchand. Almécega tre x V. Gomes, 887 20. Cactaceae Cereus jamacaru DC. subsp. jamacaru Cardeiro, Mandacaru shr x x V. Gomes, 4-399 Epiphyllum phyllanthus (L.) Haw. eh x V. Gomes, 625 Hylocereus setaceus (Salm – Dyck) R. Bauer vi x Lima-Verde, 3472 Pereskia aculeata Mill. vi x Lima-Verde, 3596 Pilosocereus catingicola subsp. salvadorensis (Werderm.) Zappi Cardeiro shr x V. Gomes, 5-277 Rhipsalis baccifera (J.M.Muell.) Stearn. subsp. baccifera Chororongo eh x x A.Silveira, 406 21. Capparaceae Cynophalla flexuosa (L.) J.Presl. Feijão-bravo tre x x V. Gomes, 1160 22. Caricaceae Jacaratia spinosa (Aubl.) A.DC. Jacaratiá tre x x A. Castro, 30996 23. Celastraceae Maytenus distichophylla Mart. ex Reissek Folha-dura tre x V. Gomes, 442 Maytenus erythroxyla Reissek Jerimum tre x x x A. Silveira, 851 Maytenus gonoclada Mart. Folha-dura tre x V. Gomes, 912 Maytenus impressa Reissek tre x x x V. Gomes, 2009 Maytenus obtusifolia Mart. Jerimum tre x x x x x V. Gomes, 2704-5 Maytenus schumanniana Loes. Jerimum tre x V. Gomes, 1-273 Maytenus sp. tre x x 24. Chrysobalanaceae Hirtella racemosa var. hexandra (Willd. ex Roem. & Schult.) Prance shr x x V. Gomes, 779 Licania sp. tre x V. Gomes, 924 25. Clusiaceae Clusia dardanoi G.Mariz & Maguire Gitó-da-mata tre x x V. Gomes, 1144 Clusia nemorosa G.Mey. Orelha-de-burro tre x x x x A. Silveira, 815 Garcinia gardneriana (Planch. & Triana) Zappi Bacupari tre x x x x V. Gomes, 1014 26. Combretaceae Buchenavia tetraphylla (Aubl.) R.A.Howard. Amarelão tre x x x x A. Silveira, 182 27. Commelinaceae Aneilema brasiliense C.B.Clarke th x V. Gomes, 1305-9 Commelina benghalensis L. th x x V. Gomes, 665 Dichorisandra hexandra (Aubl.) Kuntze ex Hand.-Mazz. th x V. Gomes, 1270 28. Costaceae Costus spiralis (Jacq.) Roscoe th x x x x V. Gomes, 666 29. Cyperaceae Becquerelia cymosa Brongn. th x V. Gomes, 798 Cyperus cf. ligularis L. th x V. Gomes, 719 Rhynchospora cephalotes (L.) Vahl th x V. Gomes, 336 Scleria latifolia Sw. th x A. Silveira, 78 30. Dilleniaceae Doliocarpus dentatus (Aubl.) Standl. subsp. dentatus vi x V. Gomes, 776 31. Elaeocarpaceae Sloanea garckeana K.Schum. shr x V. Gomes, 883 32. Erythroxylaceae Erythroxylum affine A.St.-Hil. tre x x A. Silveira, 809 Erythroxylum citrifolium A. St.-Hil. tre x x x A. Silveira, 418 Erythroxylum macrochaetum Miq. Café-bravo shr x V. Gomes, 1002-1 Erythroxylum mucronatum Benth. tre x x x x V. Gomes, 112 Erythroxylum pulchrum A.St.-Hil. tre x V. Gomes, 1093 Erythroxylum simonis Plowman shr x V. Gomes, 1095 Erythroxylum squamatum Sw. shr x x A. Silveira, 111 Erythroxylum subrotundum A.St.-Hil. shr x V. Gomes, 209-6 Erythroxylum tenue Plowman shr x A. Silveira, 317 Erythroxylum sp.1 shr x A. Silveira, 923 33. Euphorbiaceae Acalypha sp. shr x V. Gomes, 739 Acalypha villosa Jacq. shr x A. Silveira, 427 Actinostemon concolor (Spreng.) Müll. Arg. tre x A. Silveira, 807 Actinostemon klotzschii (Didr.) Pax shr x A. Silveira, 580 Actinostemon verticillatus (Klotzsch) Baill. shr x A. Silveira, 549 Alchornea glandulosa subsp. iricurana (Casar.) Secco Murici-de-jacu, Sabiá-timbú tre x A. Silveira, 891 Aparisthmium cordatum (A.Juss.) Baill. Piroá tre x V. Gomes, 774 Bernardia tamanduana (Baill.) Müll. Arg. shr x V. Gomes, 738 Croton argyroglossus Baill. Marmeleiro tre x x V. Gomes, 698 Croton blanchetianus Baill. Marmeleiro-preto tre x x V. Gomes 1102-4 Croton floribundus Spreng. Marmeleiro-da-serra tre x x A. Silveira, 739 Croton sp. 1 Marmeleiro-da-serra tre x x x V. Gomes, 1138 Croton sp. 2 Marmeleiro shr x V. Gomes, 1-3 Hieronyma oblonga (Tul.) Müll Arg. Sabiá-da-mata tre x x A. Silveira, 442 Jatropha mollissima (Pohl) Baill. Pinhão-bravo shr x x A. Silveira, 611 Manihot carthaginensis subsp. glaziovii (Müll Arg.) Allem. tre x x x x x A. Silveira, 873 Sapium obovatum Klotzsch ex Müll. Arg. Burra-leitera tre x x A. Silveira, 854 Sebastiania commersoniana (Baill.) L.B.Sm. & Downs shr x V. Gomes, 0404-1 Sebastiania jacobinensis (Müll.Arg.) Müll.Arg. Pau-de-leite tre x A. Silveira, 849 Sebastiania macrocarpa Müll.Arg. shr x x x V. Gomes, 2009-5 Tragia volubilis L. vi x A. Silveira, 742 34. Fabaceae 34.1 Caesalpinioideae Bauhinia cf. cheilantha (Bong.) Steud. Mororó, Capa-bode shr x V. Gomes, 402 Bauhinia sp. Capa-bode shr x x V. Gomes, 5-217 Cassia ferruginea (Schrad.) Schrad. ex DC. var. ferruginea tre x M.A.Figueiredo, 15951 Cassia grandis L. f. tre x M. A. Figueiredo, 11711 Chamaecrista duckeana (P. Bezerra & Afr. Fern.) H.S. Irwin & Barneby Canafístula-brava th x V. Gomes, 548 Chamaecrista zygophylloides var. colligans (H.S.Irwin & Barneby) H.S.Irwin & Barneby shr x x J.R.Lima, 862 Chamaecrista sp. subshr x A. Silveira, 943 Copaifera langsdorffii Desf. Pau-d’óleo tre x x x x A. Silveira, 884 Hymenaea eriogyne Benth. Jatobá tre x x V. Gomes, 6-54 Libidibia ferrea (Mart. ex Tul.) L.P.Queiroz var. ferrea Jucá, Pau-ferro tre x A. Silveira, 951 Libidibia ferrea var. leiostachya (Benth.) L.P.Queiroz Pau-ferro tre x V. Gomes, 1162 Poincianella bracteosa (Tul.) L.P.Queiroz Catingueira tre x x V. Gomes, 5-411 Senna quinquangulata (Rich.) H.S.Irwin & Barneby Besouro tre x A. Silveira, 362 Senna splendida (Vogel) H.S.Irwin & Barneby São-João shr x M.A.Figueiredo, 8920 34.2 Faboideae Andira cf. nitida Mart. ex Benth. tre x V. Gomes, 850 Desmodium procumbens (Mill.) Hitchc. th x M.A.Figueiredo, 17662 Dioclea grandiflora Mart. ex Benth. Mucunã vi x x V. Gomes, 0903-2 Dioclea virgata (Rich.) Amshoff Mucunã vi x L.W. Lima-Verde, 110 Lonchocarpus sericeus (Poir.) Kunth ex DC. Ingá-brava tre x x M.A.Figueiredo, 15938 Machaerium hirtum (Vell.) Stellfeld Chifre-de-bode tre x x A. Silveira, 1001 Myroxylon peruiferum L.f. Bálsamo tre x V. Gomes, 4-778 Ormosia sp. tre x x x x V. Gomes, 1-221 Platymiscium floribundum Vogel tre x x V. Gomes, 907-1 34.3 Mimosoideae Abarema jupunba (Willd.) Britton & Killip var. jupunba tre x x x x x J.R.Lima, 385 Albizia polycephala (Benth.) Killip ex Record Camuzé tre x x x x x A. Silveira, 348 Anadenanthera colubrina var. cebil (Griseb.) Altschul Calumbi tre x x L.W.Lima-Verde, 3570 Chloroleucon dumosum (Benth.) G.P. Lewis Arapiraca tre x x A. Silveira, 867 Inga bollandii Sprague & Sandwith Ingaí tre x x x A. Silveira, 357 Inga ingoides (Rich.) Willd. Ingá tre x x x x V. Gomes, 329 Inga laurina (Sw.) Willd. Ingá tre x V. Gomes, 4-751 Inga marginata Willd. Ingá tre x M.R.Oliveira, 20976 Mimosa arenosa (Willd.) Poir. var. arenosa tre x x L.W.Lima-Verde, 3621 Mimosa caesalpiniifolia Benth. Sabiá tre x x x V. Gomes, 4 Parkia pendula (Willd.) Benth. ex Walp. Visgueiro tre x A. Silveira, 379 Piptadenia stipulacea (Benth.) Ducke Saia-velha tre x V. Gomes, 436 Senegalia polyphylla (DC.) Britton & Rose Espinheiro, Espinheiro-preto tre x x V. Gomes, 4-773 Senegalia riparia (Kunth) Britton & Rose ex Britton & Killip Unha-de-gato shr x V. Gomes, 5-689 Stryphnodendron guianense (Aubl.) Benth. subsp. guianense Favinha tre x x x A. Silveira, 403 35. Gentianaceae Chelonanthus purpurascens Aubl. shr x A. Silveira, 160 Voyria flavescens Griseb. th x V. Gomes 1127-2 36. Heliconiaceae Heliconia spathocircinata Aristeg. Cana-de-macaco th x L.W.Lima-Verde, 316 Heliconia pendula Wawra Heliconia th x V. Gomes 523 Heliconia psittacorum L. f. Pacavira th x x x x x V. Gomes, 338 37. Humiriaceae Sacoglottis sp. Folha-dura tre x V. Gomes,1205-3 38. Hypericaceae Vismia guianensis (Aubl.) Choisy Lacre-vermelho tre x x x V. Gomes, 761 39. Hypoxidaceae Hypoxis decumbens L. th x V. Gomes, 1102-8 40. Iridaceae Cipura paludosa Aubl. th x V. Gomes, 1102-7 41. Lamiaceae Aegiphila integrifolia (Jacq.) Moldenke Orelha-de-onça tre x V. Gomes, 1102-3 Hyptis pectinata (L.) Poit. Canela-de-juriti th x V. Gomes, 743 Vitex cf. capitata Vahl Guabiraba tre x A. Silveira , 557 Vitex flavescens Kunth Chapéu-de-sol tre x V. Gomes, 1305-1 Vitex cf. panshiniana Moldenke Gargaúba tre x V. Gomes, 277 Vitex triflora Vahl Guabiraba tre x x A. Silveira , 637 Vitex sp. Guabiraba tre x A. Silveira, 886 42. Lauraceae Cinnamomum triplinerve (Ruiz & Pav.) Kosterm. Louro-eucalipto tre x x A. Silveira, 771 Endlicheria sp. tre x V. Gomes, 1-331 Nectandra cuspidata Nees Louro tre x x V. Gomes, 1252 Ocotea daphnifolia (Meisn.) Mez Louro tre x A. Silveira, 339 Ocotea glauca (Ness & Mart.) Mez tre x V. Gomes, 913 Ocotea glomerata (Nees) Mez Louro tre x x x A. Silveira, 149 Ocotea longifolia Kunth Louro tre x x x V. Gomes, 83-2 Ocotea puberula (Rich.) Nees Jenipapo-bravo tre x V. Gomes, 1404-8 Ocotea sp. Louro tre x x V. Gomes, 501 43. Lythraceae Lafoensia pacari A.St.-Hil. tre x V. Gomes, 2009-13 44. Malpighiaceae Bunchosia acuminata Dobson tre x x x V. Gomes, 1303-5 Byrsonima crispa A.Juss. Murici tre x A. Silveira, 206 Byrsonima sericea DC. Murici tre x x x x x V. Gomes, 1112 Byrsonima stipulacea A.Juss. Murici tre x M. A. Figueiredo, 17050 Heteropterys tricanthera A.Juss. vi x V. Gomes, 1102-3 Tetrapterys mucronata Cav. Canela-brava vi x A. Silveira 436 45. Malvaceae Apeiba tibourbou Aubl. Jangada, Pau-de-jangada tre x x x x A. Silveira, 861 Callianthe bezerrae (Monteiro) Donnel subshr x M.A. Figueiredo, 349 Ceiba glaziovii (Kuntze) K.Schum. Barriguda tre x x V. Gomes, 508-3 Helicteres baruensis Jacq. Maria-preta shr x V. Gomes, 1102-5 Helicteres velutina K.Schum. Maria-preta shr x x V. Gomes, 795 Helicteres sp. shr x V. Gomes, 1102-6 Pseudobombax marginatum (A.St.-Hil.) A. Robyns Imbiratanha tre x x V. Gomes, 1103-6 Sida urens L. th x x M. A. Figueiredo, 15249 46. Marantaceae Calathea cylindrica (Roscoe) K.Schum. Bananinha th x V. Gomes, 2604-4 Calathea sp. Bananinha-de-salão th x x A. Silveira, 972 Ischnosiphon puberulus Loes. Taquari th x x x V. Gomes, 1119 Maranta leuconeura E. Morren Baratinha th x x A. Silveira, 971 47. Marcgraviaceae Norantea guianensis Aubl. shr x V. Gomes, 903 48. Melastomataceae Aciotis sp. th x V. Gomes, 778 Clidemia debilis Crueg. Lava-mato shr x V. Gomes, 2604-3 Clidemia dentata D.Don Lacre-branco shr x A. Silveira, 161 Clidemia hirta (L.) D.Don shr x V. Gomes, 2704-2 Miconia affinis DC. shr x x x x A. Silveira, 903 Miconia alata (Aubl.) DC. Canela-de-veado tre x x x A. Silveira, 404 Miconia aff. caudigera DC. Lacre-branco shr x A. Silveira, 353 Miconia holosericea (L.) DC. tre x x V. Gomes, 767 Miconia hypoleuca (Benth.) Triana shr x x V. Gomes, 1109 Miconia minutiflora (Bonpl.) DC. Lacre-branco shr x x A. Silveira, 10 Miconia nervosa (Sm.) Triana Língua-de-vaca shr x Silveira, 626 Miconia prasina (Sw.) DC. Lacre-branco tre x x x x A. Silveira, 380 Miconia sp. shr x V. Gomes, 2704-7 Tibouchina heteromalla (D.Don) Cogn. shr x A. Silveira, 894 49. Meliaceae Cedrela odorata L. Cedro tre x A. Silveira, 635 Trichilia emarginata (Turcz.) C.DC. tre x x V. Gomes, 2009-18 Trichilia ramalhoi Rizzini tre x V. Gomes, 1068 50. Menispermaceae Cissampelos andromorpha DC. vi x A. Silveira, 906 51. Moraceae Brosimum gaudichaudii Trécul Inharé tre x x x A. Silveira, 419 Ficus guianensis Desv. Gameleira tre x x x x x x V. Gomes, 797 52. Myrsinaceae Myrsine guianensis (Aubl.) Kuntze Cajueiro-bravo tre x V. Gomes, 700 Myrsine umbellata Mart. Coração-de-nego; Mium-de-sangue tre x x x x A. Silveira, 304 53. Myrtaceae Campomanesia aromatica (Aubl.) Griseb. Guabiraba tre x x x x Silveira, 902 Campomanesia ilhoensis Mattos tre x V. Gomes, 1002-9 Campomanesia sp1. Guabiraba tre x x x V. Gomes, 609 Campomanesia sp2. tre x Eugenia acutata Miq. tre x V. Gomes, 1140 Eugenia aurata O.Berg. Café-bravo tre x A. Silveira, 871 Eugenia cf. cachoeirensis tre x x x A. Silveira, 976 Eugenia cf. egensis DC. Folha-miúda tre x A. Silveira, 612 Eugenia cf. schottiana O. Berg tre x x A. Silveira, 924 Eugenia cf. uniflora L. shr x V. Gomes, 598 Eugenia flavescens DC. Folha-miúda tre x A. Silveira, 586 Eugenia florida DC. Café-bravo tre x x x V. Gomes, 438 Eugenia ligustrina (Sw.) Willd. Folha-miúda tre x x x x V. Gomes, 1079 Eugenia cf. paraensis O. Berg tre x A. Silveira, 607 Eugenia piresii Mattos tre x x V. Gomes, 790 Eugenia punicifolia (Kunth) DC. Folha-miúda tre x x V. Gomes, 49 Eugenia sp.1 tre x x x V. Gomes, 771-1 Eugenia sp. 2 Café-bravo tre x x A. Silveira, 949 Eugenia sp. 3 shr x V. Gomes, 775 Marlieria sp1 tre x V. Gomes, 1176 Marlieria sp2 tre x V. Gomes, 1-207 Myrcia sp. 1 shr x x V. Gomes, 2009-23 Myrcia sp. 2 Cabacinha tre x x x V. Gomes, 1152 Myrcia sp. 3 tre x V. Gomes, 2-561 Myrcia sp. 4 tre x V. Gomes, 2-230 Myrcia sp. 5 tre x V. Gomes, 2-798 Myrcia sp. 6 tre x x A. Silveira, 999 Myrcia alagoensis O. Berg tre x x x x V. Gomes, 1041 Myrcia multiflora (Lam.) DC. shr x x V. Gomes, 935 Myrcia pubiflora DC. tre x V. Gomes, 875 Myrcia rostrata DC. tre x x x x V. Gomes, 1147 Myrcia splendens (Sw.) DC. Folha-miúda tre x x x V. Gomes, 1197 Myrcia sylvatica (G.Mey.) DC. Folha-miúda-preta tre x x x A. Silveira, 458 Myrcia tomentosa (Aubl.) DC. Goiabinha tre x x A. Silveira, 387 Myrciaria ferruginea O.Berg tre x A. Silveira, 808 Myrciaria sp1 tre x x x A. Silveira, 338 Myrciaria sp2 tre x V. Gomes, 1205 Myrciaria sp3 tre x A. Silveira, 923 Myrciaria tenella (DC) O. Berg Sangue de boi tre x V. Gomes, 1216 Psidium guianense Sw. tre x V. Gomes, 1228 Psidium sartorianum (O.Berg.) Nied. tre x V. Gomes, 0206-15 Siphoneugenia sp. shr x A. Silveira 779 Syzygium jambos (L.) Alston tre x V. Gomes, 695 54. Nyctaginaceae Guapira sp. João-mole tre x V. Gomes, 1-462 Neea obovata Spruce ex. Heimerl João-mole tre x x x x x x V. Gomes, 1153 55. Ochnaceae Ouratea hexasperma (A.St.-Hil.) Baill. Cajuzinho tre x x x A. Silveira, 399 Ouratea polygyna Engl. Cajuzinho tre x V. Gomes, 726-1 56. Olacaceae Heisteria blanchetiana (Engl.) Sleumer tre x V. Gomes, 20-6 Heisteria perianthomega (Vell.) Sleumer Mium-de-sangue-branco tre x x A. Silveira, 973 Schoephia obliquifolia Turcz. tre x V. Gomes, 764 Ximenia americana L. tre x V. Gomes, 4-449 57. Oleaceae Chionanthus sp. tre x V. Gomes, 462 58. Opiliaceae Agonandra brasiliensis Miers. ex Benth. & Hook.f. Juá-mirim tre x V. Gomes, 72 59. Orchidaceae Alatiglossum barbatum (Lindl.) Baptista eh x V. Gomes, 523 Catasetum macrocarpum Rich. ex Kunth eh x x V. Gomes, 1009-1 Epidendrum armeniacum Lindl. eh x V. Gomes, 0607-3 Epidendrum nocturnum Jacq. eh x V. Gomes, 299 Gongora quinquenervis Ruiz & Pav. eh x x V. Gomes, 3031 Notylia lyrata S. Moore eh x x V. Gomes, 2209-16 Polystachia concreta (Jacq.) Garay & Sweet eh x x x A. Silveira, 897 Preschottia stachyodes (Sw.) Lindl. th x x x x V. Gomes, 716 Specklinia trifida (Lindl.) F.Barros eh x x V. Gomes, 621 Trichocentrum fuscum Lind. eh x V. Gomes, 716 60. Oxalidaceae Oxalis alstonii Lourteig. th x A. Silveira, 804 61. Passifloraceae Mitostemma brevifilis Gontsch. Maracujá-suspiro vi x A. Silveira, 741 62. Peraceae Pera glabrata (Schott) Poepp. ex Baill. Casquim tre x x x V. Gomes, 991 63. Phyllanthaceae Phyllanthus acutifolius Poir. ex Spreng. subshr x A. Silveira, 855 64. Phytolaccaceae Hilleria latifolia (Lam.) H.Walter th x A. Silveira, 829 65. Picramiaceae Picramnia gardneri Planch. tre x V. Gomes, 488 Picramnia glazioviana Engl. tre x V. Gomes, 922 66. Piperaceae Piper aduncum L. shr x A. Silveira, 463 Piper arboreum Aubl. shr x x A. Silveira, 390 Piper ovatum Vahl. subshr x V. Gomes, 0707-3 Peperomia circinnata Link th x V. Gomes, 0707-2 67. Plumbaginaceae Plumbago scandens L. subshr x V. Gomes, 735 68. Poaceae Merostachys sp. Taquara th x V. Gomes, 1124 Olyra latifolia L th x V. Gomes, 15-3 Parodiolyra micrantha (Kunth) Davidse & Zuloaga th x V. Gomes, 545 69. Podocarpaceae Podocarpus sellowii Klotzsch ex. Endl. tre x A. Silveira, 239 70. Polygalaceae Acanthocladus albicans A.W.Benn. Ameixa shr x V. Gomes, 793 Polygala paniculata L. Vique th x x x A. Silveira, 447 71. Polygonaceae Coccoloba parimensis Benth. tre x A. Silveira, 729 Coccoloba sp.1. tre x V. Gomes, 1197 Coccoloba sp. 2 shr x A. Silveira, 494 Ruprechtia laxiflora Meisn. Tubibeira tre x A. Silveira, 924 72. Portulacaceae Talinum paniculatum (Jacq.) Gaertn. th x A. Silveira, 953 73. Proteaceae Roupala sp. Carne-de-vaca; Rabugem tre x x V. Gomes, 729 74. Rhamnaceae Colubrina glandulosa Perkins Sabiaguaba shr x x V. Gomes, 1205-1 Ziziphus undulata Reissek Juá-mirim tre x V. Gomes, 6-26 75. Rosaceae r Prunus myrtifolia (L.) Urb. Pau-de-soinho tre x x x x V. Gomes, 2209-3 76. Rubiaceae Alseis floribunda Schott Guabiraba tre x x x x A. Silveira, 850 Amaioua intermedia Mart. ex Schult. & Schult. f Casquim tre x x x x V. Gomes, 1087 Chiococca alba (L.) Hitchc. shr x M.A.Figueiredo, 16670 Coussarea contracta (Walp.) Müll.Arg. var. contracta Folha-dura tre x V. Gomes, 6-42 Coutarea hexandra (Jacq.) K.Schum. Quina-quina tre x A. Silveira, 909 Faramea hyacinthina Mart. Folha-dura tre x x x x V. Gomes, 1003-5 Faramea sp. 1 Violeta, Folha-dura tre x x V. Gomes, 916 Faramea sp. 2 Folha-dura tre x x x A. Silveira, 774 Gonzalagunia dicocca Cham. & Schltdl. Canela-de-juriti th x V. Gomes, 1048 Guettarda angelica Mart. ex Müll.Arg. Espinho-branco shr x x x x A. Silveira, 721 Hamelia patens Jacq. th x A. Silveira, 827 Palicourea guianensis Aubl. Sombrião, Erva-de-rato-grande shr x x x V. Gomes 1103 Palicourea marcgravii A.St.-Hil. shr x V. Gomes, 1205-7 Psychotria bracteocardia (DC.) Müll.Arg. Sorriso-de-viúva th x x x V. Gomes, 0902-1 Psychotria capitata Ruiz & Pav. shr x x x x A. Silveira, 759 Psychotria carthagenensis Jacq. shr x x x x A. Silveira, 767 Psychotria colorata (Willd. ex Schult.) Müll. Arg. Erva-de-rato-falsa shr x A. Silveira, 734 Psychotria deflexa DC. shr x A. Silveira, 733 Psychotria hoffmannseggiana (Willd. ex Schult.) Müll. Arg shr x x A. Silveira, 742 Randia armata (Sw.) DC. Veludo-preto tre x x x x V. Gomes, 1102-4 77. Rutaceae Esenbeckia grandiflora Mart. tre x x x x A. Silveira, 811 Pilocarpus spicatus A.St.-Hil. tre x A. Silveira, 845 Rauia sp. tre x V. Gomes, 772 Zanthoxylum petiolare A.St.-Hil & Tul. Limãozinho tre x x x V. Gomes, 466 Zanthoxylum rhoifolium Lam. Limãozinho tre x A. Silveira, 908 78. Salicaceae Banara guianensis Aubl. Farinha-seca tre x x x x V. Gomes, 1125 Casearia commersoniana Cambess. shr x V. Gomes, 0607-1 Casearia grandiflora Cambess. shr x V. Gomes, 1114 Casearia sylvestris Sw. shr x x x x A. Silveira, 826 Casearia sp. 1 shr x V. Gomes, 2-753 Casearia sp. 2 shr x x V. Gomes, 6-263 Prockia crucis P.Browne ex L. shr x A. Silveira, 726 Xylosma ciliatifolia (Clos) Eichler Espinho-de-judeu shr x x V. Gomes, 0306-1 79. Santalaceae Phoradendron crassifolium (Pohl ex DC.) Eichler hm x x V. Gomes, 727 Phoradendron mucronatum (DC.) Krug & Urb. hm x V.Gomes, 2009-14 Phoradendron sp. 1 hm x x V. Gomes, 1130 Phoradendron sp. 2 hm x A. Silveira, 975 80. Sapindaceae Allophylus edulis (A.St.-Hil. et al.) Hieron. ex Niederl. shr x A. Silveira, 822 Aparisthmium cordatum (A.Juss.) Baill. Piroá tre x V. Gomes, 774 Cupania impressinervia Acev.-Rodr. tre x Araújo, F. S. 1604 Cupania racemosa (Vell.) Radlk. Cajueiro-bravo tre x V. Gomes, 736 Cupania longifolia Benth. Cajueiro-bravo tre x x V. Gomes, 0902-1 Paullinia uloptera Radlk. vi x V. Gomes, 441 Serjania hebecarpa Benth. vi x V. Gomes, 751 81. Sapotaceae Chrysophyllum flexuosum Mart. tre x V. Gomes, 580 Chrysophyllum gonocarpum (Mart. & Eichler ex Miq.) Engl. Jitó tre x V. Gomes, 1204-3 Chrysophyllum sp. Folha-dura tre x A. Silveira, 982 Manilkara rufula (Miq.) H.J.Lam Maçaranduba tre x x x x x A. Silveira, 537 Micropholis aff. guyanensis (A.DC.) Pierre tre x x V. Gomes, 504 Pouteria bangii (Rusby) T.D.Penn. Engasga-vaca tre x V. Gomes, 0206-3 Pouteria macrophylla (Lam.) Eyma tre x V. Gomes, 1002-1 Pouteria peduncularis (Mart. & Eichler ex Miq.) Baehni tre x A. Silveira, 108 Pouteria venosa (Mart.) Baehni subsp. venosa tre x V. Gomes, 1002-15 82. Schoepfiaceae Schoepfia brasiliensis A.DC. tre x x V. Gomes, 764 83. Simaroubaceae Simarouba amara Aubl. Paraíba tre x x x x A. Silveira, 1112 84. Siparunaceae Siparuna guianensis Aubl. Sabonete shr x V. Gomes, 1116 85. Smilacaceae Smilax sp. Japecanga vi x x V. Gomes, 1028 86. Solanaceae x Acnistus arborescens (L.) Schltdl. shr x V. Gomes, 1002-52 Brunfelsia uniflora (Pohl) D.Don shr x x A. Silveira, 749 Cestrum axillare Vell. Dominguinho shr x Araújo, F.S. 1608 Cestrum schlechtendalli G.Don. Dominguinho shr x V. Gomes, 1003 Solanum caavurana Vell. shr x x A. Silveira, 772 Solanum campaniforme Roem. & Schult. Caninana shr x A. Silveira, 426 Solanum paniculatum L. shr x A. Silveira, 491 Solanum rhytidoandrum Sendtn. Boldo, Jurubeba-preta shr x x V. Gomes, 1102-5 87. Sterculiaceae Basiloxylon brasiliensis (All.) K. Schum. Piroá tre x x V. Gomes, 1103-1 Guazuma ulmifolia Lam. Mutamba-brava tre x V. Gomes, 2-289 88. Symplocaceae Symplocos nitens (Pohl) Benth. tre x V. Gomes, 1143 89. Thymelaeaceae Daphnopsis racemosa Griseb. Embira-branca shr x x x x V. Gomes, 1106 90. Urticaceae Cecropia palmata Willd. Embaúba, Torém tre x x x A. Silveira, 11 Urera baccifera (L.) Gaudich. ex Wedd. Urtiga tre x V. Gomes, 1203-5 91. Verbenaceae Lantana camara L. Camará shr x A. Silveira, 732 Lantana radula Sw. Camará shr x V. Gomes, 2109-5 92. Zingiberaceae Renealmia chrysotricha Petersen Colônia-brava th x V. Gomes, 455 VN: vernacular name; GF: growth form; ter: tree; shr: shrub; subshr: subshrub; vi: vine; th: terrestrial herb; eh: epiphytic herb; hm: hemiparasite; C: main collector’s name and number; x: present species. Surveyed sites: TAV: Taveiras; SIN: Sinimbu; ARV: Arvoredo; LAG: Lagoa; JAR: Jardim; SAL: Salva-Vidas.

Edited by

Associate editor: Rodrigo Studart Corrêa 0000-0002-9422-2629

Publication Dates

  • Publication in this collection
    17 July 2020
  • Date of issue
    2020

History

  • Received
    28 June 2018
  • Accepted
    07 Nov 2018
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