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Between Caatinga and Atlantic Forest: Floristic Composition in a Transition Area in Northeast Brazil

Abstract

The objective of this study was to carry out an inventory of the vascular plant species of Serra do Jatobá in the municipality of Milagres, Bahia. The survey included species of all habits and respective degrees of endemism, phytogeographic domains and threat categories, to contribute data about the flora of the Bahia semiarid region to enable management and preservation of species, to support the future creation of conservation units in Bahia inselbergs. We recorded 317 species, 208 genera and 71 families, with predominance of species belonging to the Bromeliaceae, Fabaceae, Euphorbiaceae and Asteraceae families. We identified 53 endemic species of the Caatinga, and 18 exclusive to the Atlantic Forest. Of these species, 24 are endemic to Bahia and five species are considered vulnerable. The floristic survey of Serra do Jatobá demonstrated high biodiversity, reinforcing the importance of ongoing research in the area for the implementation of a conservation unit.

Keywords:
Caatinga; Endangered species; Endemism; Flora; Semiarid

1. INTRODUCTION AND OBJECTIVES

Brazil is considered the country with one of the greatest biodiversities in the world, sheltering about 37 thousand species of vascular plants (Flora e Funga do Brasil, 2022Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. [ cited 2022 jul. 18]. Available from: http://floradobrasil.jbrj.gov.br.
http://floradobrasil.jbrj.gov.br...
). Floristic inventory efforts have increased in the last decade, resulting in the production of complete lists, making an important contribution to the conservation of the country’s biodiversity (Forzza et al., 2012Forzza RC, Baumgratz JFA, Bicudo CEM, Canhos DA, Carvalho Junior AA, Coelho MAN, et al. New Brazilian floristic list highlights conservation challenges. BioScience 2012; 62(1): 39-45. ; Stehmann & Sobral, 2017Stehmann JR, Sobral M. Biodiversidade no Brasil. In: Simões CM, Schenkel EP, Mentz LA, Petrovick PR, editors. Farmacognosia do Produto Natural ao Medicamento. Porto Alegre: Artmed; 2017.; Fernandes et al., 2020Fernandes MF, Cardoso D, Queiroz LP. An updated plant checklist of the Brazilian Caatinga seasonally dry forests and woodlands reveals high species richness and endemism. Journal of Arid Environments 2020; 174: 104079. ; Flora e Funga do Brasil, 2022Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. [ cited 2022 jul. 18]. Available from: http://floradobrasil.jbrj.gov.br.
http://floradobrasil.jbrj.gov.br...
). Nevertheless, the existence of these lists does not eliminate the need for punctual and long-term floristic studies, investigations focused on specific areas, and ecological analyses to characterize the vegetation, because the recording of a plant species in a determined locale adds data on the distribution and patterns of occurrence, thus shedding light on the biogeography of biomes (Kendal & Snelson, 2009Kendall PB, Snelson B. The role of floristic survey data and quantitative analysis in identification and description of ecological communities under threatened species legislation: A case study from north-eastern New South Wales. Ecological Management and Restoration 2009; 10: 16-26. ; Moro et al., 2011Moro MF, Araújo FS, Rodal M, Martins FR. Síntese dos estudos florísticos e fitossociológicos realizados no semiárido brasileiro. In: Eisenlohr PV, Felfili JM, Melo MMRF, Andrade LA, Meira Neto JAAM, editors. Fitossociologia no Brasil. Métodos e Estudo de Caso, Viçosa: UFV; 2011.). The resulting datasets are important for delineation and conservation of the plant communities with different phytophysiognomies or habits (Kendal & Snelson, 2009Kendall PB, Snelson B. The role of floristic survey data and quantitative analysis in identification and description of ecological communities under threatened species legislation: A case study from north-eastern New South Wales. Ecological Management and Restoration 2009; 10: 16-26. ; Moro et al., 2011Moro MF, Araújo FS, Rodal M, Martins FR. Síntese dos estudos florísticos e fitossociológicos realizados no semiárido brasileiro. In: Eisenlohr PV, Felfili JM, Melo MMRF, Andrade LA, Meira Neto JAAM, editors. Fitossociologia no Brasil. Métodos e Estudo de Caso, Viçosa: UFV; 2011.; Chaves et al., 2013Chaves ADCG, Santos RMS, Santos JO, Fernandes AA, Maracajá PB. A importância dos levantamentos florístico e fitossociológico para a conservação e preservação das florestas. Agropecuária Científica no Semiárido 2013; 9(2): 43-48. ).

Transitional environments are common between the Caatinga phytogeographic domain (CPD), inserted in the Brazilian semiarid region and present in all states of the Northeast as well as the northern part of the state of Minas Gerais (Cardoso & Queiroz, 2008Cardoso DBOS, Queiroz LP. Floristic composition of seasonally dry tropical forest fragments in central Bahia, northeastern Brazil. Journal of the Botanical Research Institute of Texas 2008; 1(2): 551-573.; Pereira et al., 2020Pereira TTC, Brasil RD, Oliveira AM, Poeiras LM, Almeida ICC. Propostas e desafios para definição de áreas prioritárias para conservação da biodiversidade no norte de Minas Gerais (Brasil). Revista Brasileira de Meio Ambiente 2020; 8(1): 53-69. ; Ganem, 2022Ganem RS. Caatinga: Estratégias de conservação. [cited 2022 jul. 12]. Available from: Available from: https://bit.ly/2WMONpj
https://bit.ly/2WMONpj...
). The Atlantic Forest phytogeographic domain (AFPD), which covers around 15% of Brazil’s territory and is distributed along the entire coastline from the northern state of Piauí to the southern state of Rio Grande do Sul, along with isolated areas in Goiás, Mato Grosso and Minas Gerais (SOS Atlantic Forest, 2018SOS Mata Atlântica. Florestas: A Mata Atlântica 2016. [cited 2022 jul. 6]. Available from: Available from: ttps://www.sosma.org.br/nossa-causa/a-mata-atlantica/ .
ttps://www.sosma.org.br/nossa-causa/a-ma...
). Despite advances in floristic knowledge of Bahia, both in the Caatinga (Cardoso & Queiroz, 2007Cardoso DBOS, Queiroz LP. Diversidade de Leguminosae nas caatingas de Tucano, Bahia: Implicações para a fitogeografia do Semi-Árido do Nordeste do Brasil. Rodriguésia 2007; 58(2): 379-391. ; Costa et al., 2015Costa GMD, Cardoso D, Queiroz LPD, Conceição AA. Variações locais na riqueza florística em duas ecorregiões de caatinga. Rodriguésia 2015; 66(3): 685-709, 2015. ; Vitório et al., 2019Vitório C, Marinho L, Costa G, Aona L. Flowering plants of Contendas do Sincorá National Forest (Caatinga, northeastern Brazil). Brazilian Journal of Botany 2019; 42(4): 717-725. ) and Atlantic Forest phytogeographic domain (Mori et al., 1981Mori SA, Boom BA, Prance GT. Distribution patterns and conservation of eastern Brazilian coastal forest tree species. Brittonia 1981; 33: 233-245. ; Carvalho-Sobrinho & Queiroz, 2005Sobrinho JGC, Queiroz LP. Composição florística de um fragmento de Mata Atlântica na Serra da Jibóia, Santa Terezinha, Bahia, Brasil. Sitientibus 2005; 5: 20-28.; Thomas et al., 2009Thomas WW, Jardim JJG, Fiaschi P, Neto EM, Amorim AA. Composição florística e estrutura do componente arbóreo de uma área transicional de Floresta Atlântica no sul da Bahia, Brasil. Brazilian Journal of Botany 2009; 32(1): 65-78. ; Alves et al., 2015Alves M, Oliveira RB, Teixeira SR, Guedes MLS, Roque N. Levantamento florístico de um remanescente de Mata Atlântica no litoral norte do Estado da Bahia, Brasil. Hoehnea 2015; 42(3): 581-595.; Moreira et al., 2020Moreira DM, Costa G, Souza JS, Aona LYS. Floristic survey in an Atlantic Forest remnant in the Recôncavo da Bahia, Bahia State, Brazil. Hoehnea 2020; 47: e572019. ), efforts are still lacking to produce systematic floristic lists for these transitional areas between CPD and AFPD.

A gap also exists in floristic knowledge of the Caatinga phytogeographic domain in Bahia, 54% of whose area is composed of this phytogeographic domain. Many Caatinga areas have undergone poor or no sampling (IBGE, 2004IBGE. Instituto Brasileiro de Geografia e Estatística. Mapa de biomas do Brasil: primeira aproximação. Rio de Janeiro: IBGE; 2004.; Moro et al., 2014Moro MF, Lughadha EN, Filer DL, Araújo FS, Martins FR. A catalogue of the vascular plants of the Caatinga Phytogeographical Domain: a synthesis of floristic and phytosociological surveys. Phytotaxa 2014: 160(1): 1-118. ). Only a few studies have investigated all plant habits of Caatinga areas in Bahia, such as Ramalho et al. (2009Ramalho CI, Andrade AP, Félix LP, Lacerda AV, Maracajá PB. Flora arbóreo-arbustiva em áreas de Caatinga no semi-árido baiano, Brasil. Revista Caatinga 2009; 22(3): 178-186. ), Fabricante et al. (2012Fabricante JR, Andrade LA, Dias-Terceiro RG. Divergências na composição e na estrutura do componente arbustivo-arbóreo entre duas áreas de caatinga na região do Submédio São Francisco (Petrolina, PE/ Juazeiro, BA). Biotemas 2012; 25(3): 97-109. ), Costa et al. (2015Costa GMD, Cardoso D, Queiroz LPD, Conceição AA. Variações locais na riqueza florística em duas ecorregiões de caatinga. Rodriguésia 2015; 66(3): 685-709, 2015. ) and Vitório et al. (2019Vitório C, Marinho L, Costa G, Aona L. Flowering plants of Contendas do Sincorá National Forest (Caatinga, northeastern Brazil). Brazilian Journal of Botany 2019; 42(4): 717-725. ). Knowledge about the Caatinga declines considerably in areas of transition to the Atlantic Forest, where studies focused on epiphytes have revealed high diversity in these areas, as found by Souza et al. (2021Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. ) and Carvalho et al. (2020Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. ) both performed at Serra do Jatobá, Milagres, Bahia.

Serra do Jatobá is a transitional area between the Caatinga and Atlantic Forest biomes located in the municipality of Milagres, Bahia. A few studies of the area’s flora among those mentioned above have revealed high diversity. Carvalho et al. (2020Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. ) reported high diversity of epiphytes, while Souza et al. (2021Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. ) presented one of the most comprehensive investigations of Bromeliaceae in comparison with studies in the Caatinga and Atlantic Forest domains. Our objective was to investigate the floristic composition of the Serra do Jatobá region in Milagres, Bahia, covering species with all habits, with their respective degrees of endemism, phytogeographic domains and threat categories, to contribute data on the flora of the semiarid region of Bahia for management and preservation of species and future creation of conservation units in Bahia inselbergs.

2. MATERIAL AND METHODS

2.1. Study area

The study was conducted in a Caatinga area located in the Serra do Jatobá region in the municipality of Milagres, Bahia, Brazil, with central coordinates 12º55’24”S and 39º46’43”W. The vegetation consists of five phytophysiognomies: anthropized Caatinga with pasture; conserved Caatinga; slope vegetation, semideciduous seasonal forest; and an inselberg, typified as “whaleback”, with altitude of 392 m, length of 1,670 m and width of 1,417 m (Souza et al., 2021Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. ) (Figure 1). Its origin comes from outcrops of granitic rock arranged in groups (Ab´saber 1974Ab´Saber AN. O domínio morfoclimático semi-árido das Caatingas brasileiras. Série Geomorfologia 1974; 43:37.). Its lithology originates from the Lower Precambrian, consisting of gneisses, magmatite and granite intrusions (Santos and Salgado 2010Santos JM, Salgado AA. Gênese da superfície erosiva em ambiente semi-árido - Milagres/ BA: considerações preliminares. Revista de Geografia 2010; 1:250-262.). The local landscape contains inselbergs, which are small monolithic mountains or groups of them with geographic formation consisting mainly of granitic or gneissic rocks (Porembski et al. 1998Porembski S, Martinelli G, Ohlemuller R, Barthlott W. Diversity and ecology of saxicolous vegetation mats on inselbergs in the Brazilian Atlantic Rainforest. Biodiversity and Distribution 1998; 4:107-119.).

Figure 1
a) Map and location of the Jatobá community, Milagres, Bahia. Brazil. b) Front view of inselberg typified as “whaleback”. c) Vegetation on the top of the inselberg. d) Conserved Caatinga area. e) Pasture area in the upper part of the inselberg.

The area has human interference, because part is used for farming and livestock grazing by the local community, had been submitted to burning on various occasions in the past 50 years (Carvalho et al., 2020Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. ). The municipality has a dryness index of 58.1, average yearly rainfall of 480.5 mm and average annual temperature of 23.0 ºC (SEI, 1999SEI. Superintendência de estudos econômicos e sociais do Estado da Bahia. Balanço hídrico do Estado da Bahia. Salvador: SEI; 1999.). The climate is classified as BSh (hot semiarid climate) according to the scale of Köppen & Geiger (1928Köppen W, Geiger R. Klimate der Erde. Gotha: Verlag Justus Perthes; 1928.).

2.2. Floristic inventory

Specimens were was collected between July 2018 and February 2022. Collections were made by random walks in the area. Collections were periodic, covering the dry and rainy seasons. All vascular plants occurring in the area were collected. We collected plants belonging to all the habits. The specimens collected were herborized and vouchers were deposited in the collection of the Recôncavo da Bahia Herbarium (HURB). The plants were identified by consulting the specialized literature, comparison with previously identified material in online herbaria databases (SpeciesLink, 2022SPECIESLINK. Specieslink. CRIA Centro de Referência em Informação Ambiental. [cited 2022 jul. 6]. Available from: Available from: http://www.splink.org. br/index .
http://www.splink.org. br/index...
; Reflora, 2022Reflora. Herbário Virtual. [cited 2022 jul. 6]. Available from: Available from: http://reflora.jbrj.gov.br/reflora/herbarioVirtual .
http://reflora.jbrj.gov.br/reflora/herba...
) and/or consultation with specialists of each taxonomic group. All data associated with species, such as geographic distribution, habits, domains of occurrence, and origin were checked in the Flora and Funga from Brazil (http://floradobrasil.jbrj.gov.br).

The list of species indicates the scientific names, families and habits of the plants collected during this study along with the same information about specimens previously collected in the area (Carvalho et al., 2020Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. ; Souza et al., 2021Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. ). The classification followed the APG IV system (2016)APG (Angiosperm Phylogeny Group). An update 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. for angiosperms and PPG I (2016)PPG I (The Pteridophyte Phylogeny Group) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 2016; 54: 563-603. for ferns.

3. RESULTS

We recorded 317 species, distributed in 208 genera and 71 families, with predominance of Bromeliaceae (37 spp.), Fabaceae (36 spp.), Euphorbiaceae (22 spp.), Asteraceae (15 spp.), Malvaceae and Myrtaceae (11 spp.) (Table 1, Figure 2 and 3). The richest genera were Tillandsia L. (Bromeliaceae, 13 spp.), Aechmea Ruiz & Pav. (Bromeliaceae), Croton L. (Euphorbiaceae) and Eugenia L. (Myrtaceae) with five spp.

Table 1
List of vascular species in Serra do Jatobá, Milagres, Bahia, Brazil with their respective habits, endemism, phytogeographic domains and conservation status (CS). H - Habit; Sub - subshrub; Vin - Vine; Hrb - Herb; Tree - Tree; Shrub - Shrub; SB - Substrate; T - Terrestrial; R - Rupicolous; E - Epiphyte; P - Parasite; HE - Hemiepiphyte; HP - Hemiparasite; * - Naturalized; LE - Local endemism; BA - Bahia; NE - Northeast; PD - Phytogeographic domain; AM - Amazon Forest; CA - Caatinga; CE - Cerrado; PN - Pantanal; AF - Atlantic Forest; PM - Pampa; LC - ‘Least Concern’; VU - ‘Vulnerable’; DD - ‘Deficient Data’; EN - ‘Endangered’.

Figure 2
Species occurring in Serra do Jatobá - Bahia. (a) Acianthera ochreata (Orchidaceae). (b) Alcantarea nahoumii (Bromeliaceae). (c) Arrojadoa penicillata (Cactaceae). (d) Aspidosperma pyrifolium (Apocynaceae). (e) Cnidoscolus urens (Euphorbiaceae). (f) Dichorisandra saxatilis (Commelinaceae). (g) Eugenia punicifolia (Myrtaceae). (h) Euphorbia heterodoxa (Euphorbiaceae). (i) Gomphrena celosioides (Amaranthaceae). (j) Karawata multiflora (Bromeliaceae). (k) Marsdenia caatingae (Apocynaceae) (l) Oxalis frutescens (Oxalidaceae).

Figure 3
Species occurring in Serra do Jatobá - Bahia. (a) Pilosocereus pentaedrophorus (Cactaceae). (b) Poecilanthe ulei (Fabaceae). (c) Psittacanthus cordatus (Loranthaceae). (d) Sida galheirensis (Malvaceae). (e) Stizophyllum perforatum (Bignoniaceae). (f) Stylosanthes viscosa (Fabaceae). (g) Syagrus vagans (Arecaceae). (h) Tillandsia chapeuensis (Bromeliaceae) (i) Trichilia hirta (Meliaceae). (j) Vellozia candida (Velloziaceae) (k) Waltheria indica (Malvaceae).

Regarding habits of the sampled species, herbs accounted for 34.81%, shrubs for 28.16%, subshrubs for 13.29%, vines for 12.65% and trees for 10.44%. Therefore, about 60% of the plants in the area belong to the non-woody stratum and 40% to the woody stratum. Bromeliaceae was predominant among the herbs (37 spp.); for shrubs, the main family was Euphorbiaceae (13 spp.), followed by Fabaceae (9 spp.); among trees, the family Fabaceae stood out (11 spp.); while for vines, Apocynaceae was the most common family (5 spp.); and for subshrubs, Fabaceae was the main family (4 spp.).

Seven species were considered naturalized/exotic (Table 1), while 40 were found to be endemic to Northeast Brazil, of which 24 were exclusive to Bahia (Table 1). Twenty-seven species are distributed in all Brazilian phytogeographic domains, 53 species were endemic to the Caatinga phytogeographic domain, and 18 species were endemic to the Atlantic Forest (Table 1).

We found in the studied area seven species exclusive to ombrophilous forests: Aechmea bicolor L.B.Sm., Aechmea disjuncta (L.B.Sm.) Leme & J.A.Siqueira, Adenocalymma coriaceum A.DC., Billbergia macrocalyx Hook., Lomariopsis marginata (Schrad.) Kuhn and Psidium cauliflorum Landrum & Sobral.

On the general list, 21 species were exclusively related to dry vegetation (Caatinga stricto sensu, deciduous and semideciduous forests): Amorimia rigida (A.Juss.) W.R.Anderson, Amorimia velutina W.R.Anderson, Bromelia laciniosa Mart. ex Schult. & Schult.f., Colicodendron yco Mart., Croton grewioides Baill., Cryptanthus bahianus L.B.Sm., Eugenia rosea DC., Eugenia zigzag K.Cout. & Sobral, Hippeastrum stylosum Herb., Hohenbergia lanata E.Pereira & Moutinho, Ipomoea franciscana Choisy, Ipomoea pintoi O’Donell, Luetzelburgia purpurea D.B.O.S.Cardoso et al., Neoglaziovia variegata (Arruda) Mez, Orthophytum maracasense L.B.Sm., Pouteria citriodora Alves-Araújo, Psidium schenckianum Kiaersk., Pusillanthus pubescens (Rizzini) Caires, Ruellia asperula (Mart. ex Nees) Lindau, Ruellia bahiensis (Nees) Morong and Spondias bahiensis P. Carvalho et al.

Eleven species were exclusively related to rocky outcrops (inselbergs, campos de altitude and campos rupestres): Alcantarea nahoumii (Leme) J.R.Grant, Dichorisandra saxatilis Aona & M.C.E.Amaral, Dyckia maracasensis Ule, Encholirium spectabile Mart. ex Schult. & Schult.f., Euphorbia heterodoxa Müll.Arg., Euphorbia phosphorea Mart., Mandevilla bahiensis (Woodson) M.F.Sales & Kin.-Gouv., Mandevilla sancta (Stadelm.) Woodson, Marsdenia caatingae Morillo, Melocactus ernestii Vaupel, Orthophytum saxicola Ule, Tillandsia milagrensis Leme and Tillandsia chapeuensis Rauh.

Of the species identified, 29 are classified as threatened according to Flora e Funga do Brasil (2022)Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. [ cited 2022 jul. 18]. Available from: http://floradobrasil.jbrj.gov.br.
http://floradobrasil.jbrj.gov.br...
and/or are on the list of threatened endemic species of Bahia (SEMA 2017SEMA. Secretaria do Meio Ambiente. Lista Oficial das Espécies Endêmicas da Flora Ameaçadas de Extinção do Estado da Bahia 2017 [cited 2022 jul. 6]. Available from: Available from: http://www.meioambiente.ba.gov.br/2017/08/11254/Governo-publica-a-Lista-das-Especies-da-Flora-Ameacadas-de-Extincao-do-Estado-da-Bahia.html .
http://www.meioambiente.ba.gov.br/2017/0...
). Of these, 20 are considered to be of ‘Least Concern’ (LC), three species (Aechmea disjuncta (L.B.Sm.) Leme & J.A.Siqueira, Dichorisandra saxatilis and Tillandsia milagrensis) are classified as ‘Endangered’ (EN) and Amorimia velutina W.R.Anderson is indicated as having ‘Deficient Data’ (DD). Finally, the species Alcantarea nahoumii (Leme) J.R.Grant (Figure 1b), Aosa gilgiana (Urb.) Weigend, Alternanthera multicaulis (Mart.) Kuntze, Eugenia rosea DC. and Orthophytum rubrum L.B.Sm are considered ‘Vulnerable’ (VU) (Table 1).

4. DISCUSSION

Floristic inventories conducted in the Caatinga and Atlantic Rain Forest domains have revealed high richness and endemism of species, even those with high anthropization degree due to human activities over the years (Amorim et al., 2005Amorim AM, Jardim JG, Clifton BC, Fiaschi P, Thomas WW, Carvalho AMV. The vascular plants of a forest fragment in southern Bahia, Brazil, . Sida Contributions to Botany 2005; 21: 1726-1752.; Cardoso & Queiroz, 2007Cardoso DBOS, Queiroz LP. Diversidade de Leguminosae nas caatingas de Tucano, Bahia: Implicações para a fitogeografia do Semi-Árido do Nordeste do Brasil. Rodriguésia 2007; 58(2): 379-391. ; Amorim et al., 2009Amorim, A.M., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O., Thomas, W.W. Angiospermas em remanescentes de floresta montana no sul da Bahia, Brasil. Biota Neotropica 2009; 9(3): 313-348.; Thomas et al. 2009Thomas WW, Jardim JJG, Fiaschi P, Neto EM, Amorim AA. Composição florística e estrutura do componente arbóreo de uma área transicional de Floresta Atlântica no sul da Bahia, Brasil. Brazilian Journal of Botany 2009; 32(1): 65-78. ; Gomes & Alves, 2010Gomes P, Alves M. Floristic diversity of two crystalline rocky outcrops in the Brazilian northeast semi-arid region. Brazilian Journal of Botany 2010; 33(4): 661-676. ; Costa et al., 2015Costa GMD, Cardoso D, Queiroz LPD, Conceição AA. Variações locais na riqueza florística em duas ecorregiões de caatinga. Rodriguésia 2015; 66(3): 685-709, 2015. ; Queiroz et al., 2015Queiroz RT, Moro MF, Loiola MIB. Evaluating the relative importance of woody versus non-woody plants for alpha-diversity in a semiarid ecosystem in Brazil. Plant Ecology and Evolution 2015; 148(3): 361-376.; Oliveira et al. 2016Oliveira EVS, Gomes LA, Déda RM, Melo LMS, Silva ACC, Farias MCV & Prata, APN. Floristic survey of the Mata do Junco Wildlife Refuge, Capela, Sergipe State, Brazil. Hoehnea 2016; 43(4): 645-667.; Vitório et al., 2019Vitório C, Marinho L, Costa G, Aona L. Flowering plants of Contendas do Sincorá National Forest (Caatinga, northeastern Brazil). Brazilian Journal of Botany 2019; 42(4): 717-725. ). An updated list of Caatinga plants indicates the presence of 3,347 species, 962 genera and 153 families, with 15% endemism (Fernandes et al., 2020Fernandes MF, Cardoso D, Queiroz LP. An updated plant checklist of the Brazilian Caatinga seasonally dry forests and woodlands reveals high species richness and endemism. Journal of Arid Environments 2020; 174: 104079. ). The floristic composition of Serra do Jatobá is composed of 316 species, a relatively high number in comparison to other studies of Caatinga areas, which have varied from 211 to 450 species (Gomes & Alves, 2010Gomes P, Alves M. Floristic diversity of two crystalline rocky outcrops in the Brazilian northeast semi-arid region. Brazilian Journal of Botany 2010; 33(4): 661-676. ; Costa et al., 2015Costa GMD, Cardoso D, Queiroz LPD, Conceição AA. Variações locais na riqueza florística em duas ecorregiões de caatinga. Rodriguésia 2015; 66(3): 685-709, 2015. ; Queiroz et al., 2015Queiroz RT, Moro MF, Loiola MIB. Evaluating the relative importance of woody versus non-woody plants for alpha-diversity in a semiarid ecosystem in Brazil. Plant Ecology and Evolution 2015; 148(3): 361-376.; Vitório et al., 2019Vitório C, Marinho L, Costa G, Aona L. Flowering plants of Contendas do Sincorá National Forest (Caatinga, northeastern Brazil). Brazilian Journal of Botany 2019; 42(4): 717-725. ). It is worth mentioning that the size of the area and the way of sampling directly influence the quantification of the species in the cited studies.

In this study, the predominant habit types were shrubs, with approximately 38.05%, followed by herbs with 33.58% and trees with 12.31%. The proportion of non-woody species (undershrubs, herbs and vines) was greater than the woody species (shrubs and trees), a result also reported by Souza & Rodal (2010Souza JAN, Rodal MJN. Levantamento florístico em trecho de vegetação ripária de caatinga no Rio Pajeú, Floresta/Pernambuco-Brasil. Revista Caatinga 2010; 23(4): 54-62.) and Queiroz et al. (2015Queiroz RT, Moro MF, Loiola MIB. Evaluating the relative importance of woody versus non-woody plants for alpha-diversity in a semiarid ecosystem in Brazil. Plant Ecology and Evolution 2015; 148(3): 361-376.). Queiroz et al. (2015)Queiroz RT, Moro MF, Loiola MIB. Evaluating the relative importance of woody versus non-woody plants for alpha-diversity in a semiarid ecosystem in Brazil. Plant Ecology and Evolution 2015; 148(3): 361-376. pointed to the fact that the majority of the floristic studies of Caatinga areas have been focused on woody plants, leaving a gap in information about non-woody plants, which have been found to be predominant with high diversity. Thus, it is important to conduct further floristic studies of plants with all habits to obtain more details about the vegetation studied here.

Although the study area is a transition between caatinga and Atlantic Forest, the main groups found follow the caatinga pattern. The high richness found for the family Fabaceae reinforces its status as the main Caatinga family (Table 1). We found 36 species of Fabaceae distributed in 25 genera, with predominance of tree and shrub habits (Table 1). The great diversity of the Fabaceae family in the Caatinga can be explained by the morphological adaptations of its species, to enable them to withstand the periodic water shortages, high temperatures and low humidity (Cardoso & Queiroz, 2010Cardoso DBOS, Queiroz LP. Caatinga no contexto de uma metacomunidade: evidências da biogeografia, padrões filogenéticos e abundância de espécies em Leguminosas. In: Carvalho CJB, Almeida EAB, editors. Biogeografia da América do Sul. Padrões e processos. São Paulo: Roca; 2010.).

Euphorbiaceae has been reported to be among the most representative families in various studies of Caatinga areas (Table 1), and it also stood out in this study, with 22 species, mainly in the Croton L. (five spp.) (Table 1). Croton species are common elements of the secondary vegetation, resulting from a natural process of regeneration of vegetation. They are important floristic elements of arid and semiarid regions such as the Caatinga domain (Carneiro-Torres et al., 2011Carneiro-Torres DS, Cordeiro I, Giulietti AM, Berry PE, Riina R. Three new species of Croton (Euphorbiaceae ss) from the Brazilian Caatinga. Brittonia 2011; 63(1): 122-132. ).

We also found Myrtaceae to have high richness, with 11 species (Table 1). It is a family with wide distribution in various Brazilian domain, with great economic, ecological and medicinal importance (Morais et al., 2014Morais LMF, Conceição GM, Nascimento JM. Família Myrtaceae: Análise morfológica e distribuição geográfica de uma coleção botânica. Agrarian Academy 2014; 1(1): 317-346.). The genus Eugenia L. (five spp.) was predominant and the species Eugenia rosea DC., Eugenia zigzag K.Cout. & Sobral and Algrizea macrochlamys (DC.) Proença & NicLugh stood out for not only being endemic to Caatinga areas, but also endemic to Bahia. The presence of E. zigzag in Serra do Jatobá is a new record of a species only recently described, with only two records in the municipality of Livramento de Nossa Senhora (BA) (Sobral et al., 2018Sobral M, Faria JEQ, Coutinho K. Five new Brazilian species of Eugenia (Myrtaceae). Phytotaxa 2018; 347(1): 59-70. ).

Verbenaceae (eight spp.) was mainly represented by Lippia L. (three spp.) (Table 1). According to Gomes et al. (2011Gomes SVF, Nogueira PCL, Moraes VRS. Aspectos químicos e biológicos do gênero Lippia enfatizando Lippia gracilis Schauer. Eclética Química 2011; 36(1): 64-77, 2011. ), Lippia is a genus with large floristic richness in Caatinga areas. Fifteen species were found for the Asteraceae family, with the standouts being two species endemic to the Northeast region of Brazil (Lepidaploa chalybaea (Mart. ex DC.) H.Rob. and Moquiniastrum oligocephalum (Gardner) G.Sancho) (Table 1).

We found 36 epiphyte species in Serra do Jatobá (Table 1), with predominance of species of the families Bromeliaceae (30 spp.) and Orchidaceae (four spp.), a result also observed by Carvalho et al. (2020Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. ). The epiphyte species have an important ecological role because they influence the nutrient cycling and provide shelter and food for many animals (Freitas et al., 2016Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, et al. A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 2016; 58: 65-79, 2016. ; Souza et al., 2021Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. ). The high number of species found belonging to the Bromeliaceae (37 spp.) can be related to the presence of a phorophyte (licuri palm) and the fact the study area is located in an ecotone with varying altitudes and different phytophysiognomies, as also reported by Carvalho et al. (2020)Carvalho AJA, Souza EH, Costa GM, Aona LYS, Soares ACF. Vascular epiphytes on licuri palms (Syagrus coronata (Mart.) Becc.) in a toposequence: Caatinga conservation indicator species. Brazilian Journal of Botany 2020; 43(4): 1061-1075. and Souza et al. (2021)Souza EH, Carvalho AJA, Gama EVS, Hora Neto AR, Aona LYS. Macambiras e espécies simpatricas da Serra do Jatobá, Milagres, Bahia. Revista Macambira 2021; 5(1): 1-12. . Besides this, the richness of Bromeliaceae can be associated with the fact it is one of the main plant families that compose dry environments like inselbergs (Porembski, 2007Porembski S. Tropical inselbergs: habitat types, adaptive strategies and diversity patterns. Brazilian Journal of Botany 2007; 30(4): 579-586. ; Verçoza & Bastos, 2013Verçoza FC, Bastos M. Bromeliaceae e Cactaceae dos afloramentos rochosos do Costão de Itacoatiara, Parque Estadual da Serra da Tiririca, Niterói, Rio de Janeiro, Brasil. Natureza on line 2013; 11(1): 7-11.), as well as the sampling effort. In a study focused on the flora of inselbergs in the Milagres region, França et al. (1997França F, Melo ED, Santos CD. Flora de inselbergs da região de Milagres, Bahia, Brasil: I. Caracterização da vegetação e lista de espécies de dois inselbergs. Sitientibus 1997; 17: 163-184.) also observed that the Bromeliaceae was among the most representative families. The family contributed to the high number of herbaceous species, since all those listed have this habit (Table 1).

In this study, we observed 11 species endemic to Caatinga vegetation, such as Croton grewioides Baill., Colicodendron yco Mart., Eugenia rosea DC. and Neoglaziovia variegata (Arruda) Mez, and 53 species endemic to the Caatinga phytogeographic domain, such as Arrojadoa penicillata (Gürke) Britton & Rose (Figure 1c), Encyclia jenischiana (Rchb.f.) Porto & Brade, Mimosa arenosa (Willd.) Poir. (Table 1), and Syagrus vagans (Bondar) A.D.Hawkes (Figure 2g).

We found 24 species endemic to Bahia, such as Algrizea macrochlamys (DC.) Proença & NicLugh., Dichorisandra saxatilis Aona & M.C.E.Amaral (Figure 1f), Mandevilla bahiensis (Woodson) M.F.Sales & Kin.-Gouv. and Orthophytum saxicola Ule (Table 1).

In relation to the phytogeographic domains of the species found, 18 species are found in the AFPD and 30 species in both the CPD and AFPD (Table 1). The fact that the Serra do Jatobá is a boundary area with an Atlantic Forest fragment can explain this result, because although it is a Caatinga area, the region also shares many Atlantic Forest species.

We also identified the exotic species Melia azedarach L., of the Meliaceae family. Melia azedarach is an exotic/naturalized species occurring throughout Brazil, native to Asia and northern Australia (Batcher, 2000Batcher MS. Element stewardship abstract for Melia azedarach. The Nature Conservancy; 2000.), and is considered to be one of the worst invasive exotic species in many regions of the world (Sampaio & Schmidt, 2013Sampaio AB, Schmidt IB. Espécies exóticas invasoras em unidades de conservação federais do Brasil. Biodiversidade Brasileira 2013; 3(2): 32-49.). Exotic species are defined as those that do not occur naturally in a geographic region, instead relying on human transport (Moro et al., 2012Moro MF, Souza VC, Oliveira-Filho ATD, Queiroz LPD, Fraga CND, Rodal MJN, Martins FR. Alienígenas na sala: o que fazer com espécies exóticas em trabalhos de taxonomia, florística e fitossociologia? Acta Botanica Brasilica 2012; 26(4): 991-999. ). They can alternative ecosystems and cause loss of biodiversity (MMA, 2006MMA. Ministério do meio ambiente. Espécies Exóticas Invasoras: Situação Brasileira. Brasília: Ministério do Meio Ambiente, Secretaria de Biodiversidade e Florestas; 2006.).

Of the species on the floristic list of Serra do Jatobá that were evaluated regarding the threat categories, the majority were classified as having ‘Least Concern’ (LC), i.e., having low risk of extinction (Table 1). Alcantarea nahoumii (Figure 1b), Aosa gilgiana and Eugenia rosea are species that occur exclusively in Bahia (Table 1) and were classified as ‘vulnerable’, with risk of extinction (CNCFlora, 2022CNCFlora. Alcantarea nahoumii in Lista Vermelha da flora brasileira versão 2012.2. Centro Nacional de Conservação da Flora. [cited 2022 jul. 22]. Available from: Available from: http://cncflora.jbrj.gov.br/portal/pt-br/profile/Alcantarea nahoumii .
http://cncflora.jbrj.gov.br/portal/pt-br...
). According to Bastos et al. (2018Bastos MJSM, Bastos LP, Souza EH, Costa GM, Rocha MAC, Souza FVD, Costa MA PC. Spatial distribution and associated flora of Alcantarea nahoumii, a vulnerable endemic species to rocky outcrops of the Serra da Jibóia, Bahia, Brazil. Rodriguésia 2018; 69(2): 503-514. ), A. nahuomii is a species that establishes communities for colonization and secondary succession of species of the Bromeliaceae family in rocky outcrops. The species is restricted to a few municipalities in Bahia (Milagres, Castro Alves, Elísio Medrado, Santa Teresinha and Brejões) and is vulnerable due to the burning, extractive activity and absence of protection (Versieux, 2018Versieux LM. Contradicting Yourself In Taxonomy: New Evidence Allows the Recognition of an Overlooked Alcantarea Species Hypoth-esis from Bahia, Brazil. Journal of the Bromeliad Society 2018; 68(1): 11-21. ).

We found Orthophytum rubrum, this species was described in 1955, the last specimens were only collected in 1979, since then the species has not been found in the field. It was believed to have disappeared, due mainly to the significant loss of habitat. However, in 2017 the species was again recorded (HURB 18251), in the municipality of Brejões (Bahia), with a population composed of fewer than 30 individuals, in an area with the presence of livestock grazing. In 2021, a new expedition was carried out in the same place and no O. rubrum individuals were found. During that expedition, a thorough sweep was performed to collect seeds for replenishment of degraded areas, but without success. In Serra do Jatobá, we only observed this species on the top of the inselberg near a small forest fragment, with a small population (fewer than 30 individuals). Unfortunately, this population is also vulnerable due to grazing by cattle and goats.

When comparing the species found in this study with the updated checklist of Brazilian Caatinga Plants of Fernandes et al. (2020Fernandes MF, Cardoso D, Queiroz LP. An updated plant checklist of the Brazilian Caatinga seasonally dry forests and woodlands reveals high species richness and endemism. Journal of Arid Environments 2020; 174: 104079. ), we verified that 28 species found in Serra do Jatobá are not on the list. This indicates that despite the existence of consolidated species lists, it is essential to conduct new floristic inventories in Caatinga areas. Furthermore, the recording of a recently described species distributed in the state of Bahia (Eugenia zigzag) and the presence of endemic Bahia species (Alcantarea nahoumii, Aosa gilgiana and Eugenia rosea) that are threatened with extinction reinforce the importance of floristic inventories of Caatinga areas in Bahia.

Less than 2% of the Caatinga vegetation is protected in conservation units (Teixeira et al., 2021Teixeira LP, Lughadha EM, Silva, MVC, Moro MF, How much of the Caatinga is legally protected? An analysis of temporal and geographical coverage of protected areas in the Brazilian semiarid region. Acta Botanica Brasilica 2021; 35(3): 473-485.). This scenario is even more discouraging, since no inselberg area is protected within the limits of the caatinga (Barbosa-Silva et al., 2022Barbosa-Silva RG, Andrino CO, Azevedo L., Lucresia L, Lovo J, Hiura AL., Viana PL., Giannini T C, Zappi DC. A wide range of South American inselberg floras reveal cohesive biome patterns. Frontiers in Plant Science 2022; 1-15. ). The results of this study indicate that the Serra do Jatobá in Milagres, Bahia, is an area with high diversity and endemism, showing the need for ongoing research in this region for the future establishment of a conservation unit, mainly because it includes areas of inselberg.

ACKNOWLEDGEMENTS

The authors are grateful to the following taxonomists who helped in the species identification: Asteraceae (G.V.Fonseca), Bignoniaceae (R.S.Ribeiro), Boraginaceae (M.R.Sampaio), Fabaceae (R.T.Queiroz), Euphorbiaceae (M.Martins), Poaceae (C.Silva), Rubiaceae (A.N.Oliveira), Velloziacaee (C.A.F.Júnior), M.L.Guedes, D.Zappi and J.Jardim. We would also like to thank Antônio Ramos da Hora (Neto) and the Associação dos Produtores de Ouricuri do Jatobá (APOJ) for all their help during the fieldwork. This paper is part of ACMC MSc dissertation, and this study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior-Brasil (CAPES)-Finance Code 001 (CAPES - 88882.424415/2019-01).

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Edited by

Associate editor: Marcos Gervásio Pereira http://orcid.org/0000-0002-1402-3612

Publication Dates

  • Publication in this collection
    14 July 2023
  • Date of issue
    2023

History

  • Received
    19 Sept 2022
  • Accepted
    13 June 2023
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