Introduction
The Atlantic Forest is considered a biodiversity hotspot due to the occurrence of a high number of endemic and threatened species (Myers et al. 2000). The natural distribution of the Atlantic Forest was originally continuous along the Brazilian coast and extended to some areas in Argentina and Paraguay (Fundação SOS Mata Atlântica 2013). Nowadays, it is one of the eight most critically endangered hotspots, as only 12,5% of the original forest remains (Fundação SOS Mata Atlântica 2014). Nevertheless, it is the richest Brazilian phytogeographic domain in plant diversity, with 8,728 endemic species of angiosperms (Flora do Brasil 2020, under construction).
Some studies have emphasized the division of the Atlantic Forest in northern and southern regions (Cracraft 1985, Colombi et al. 2010, Martins 2011) with a limit along the Doce river, in the north of Espírito Santo state. The floristic differences between these two regions were highlighted by Oliveira-Filho & Fontes (2000) and Oliveira-Filho et al. (2005), who observed several taxa with Amazonian affinities occuring in the northern region, while taxa with subtropical Andean affinities are mostly found in the southern region (Berry et al. 2004, Santos et al. 2007, Menini Neto et al. 2016). Other authors recognize three main centers of endemism in the Atlantic Forest: northern (Pernambuco and Alagoas states), southern (from Rio de Janeiro to Santa Catarina states), and central (southern Bahia and Espírito Santo states) (Murray-Smith et al. 2009, Thomas et al. 1998).
Recently, Saiter et al. (2016a) divided the central region of the Atlantic Forest in three subregions based on the composition of tree species, indicating the most important abiotic factors affecting that division. Contrary to previous knowledge, the rivers did not play an important role in this division, which lies about 100 km north of the Doce river and is best explained by climatic factors, such as moisture, elevation and temperature (Saiter et al. 2016a). One of the regions, referred to as Bahia Coastal Forests (BCF) by Saiter et al. (2016a), encompasses the northern extreme of Espírito Santo and most of the coast of the state of Bahia. Due to high biodiversity and the incidence of endemic and threatened species, this region has been designated as a hot-point within the Atlantic Forest hotspot (Martini et al. 2007). Many studies to this date on different groups of organisms support this claim, as the region is considered an area of endemism for frogs (Carnaval et al. 2009), birds (Silva et al. 2004), Myrtaceae (Murray-Smith et al. 2009) and trees in general (Thomas et al. 2003).
An estimate of endemic vascular plant species present in two protected areas in southern Bahia showed that the distribution of 59% of the species is restricted to Bahia and Espírito Santo states (Thomas et al. 1998). These results motivated the compilation of a preliminary list of endemic angiosperm species in the region comprising Espírito Santo to the north of the Doce river and the south of Bahia, including inland Seasonal Dry Forests (Thomas et al. 2003) (Figure 1a). This list consisted of 395 endemic species of angiosperms in the region (Thomas et al. 2003), among which several are known to occur only in dry forests, such as Chrysophyllum subspinosum Monach. (Sapotaceae) and Colicodendron bahianum Cornejo & Ilts (Capparaceae). That study reported eight endemic genera in this area: Arapatiella R.S.Cowan, Brodriguesia R.S.Cowan and Harleyodendron R.S.Cowan (Fabaceae), Atractantha McClure, Anomochloa Brogn., Alvimia Calderón ex Soderstr. & Londoño, and Sucrea Soderstr. (Poaceae), and Trigoniodendron E.F.Guim & Miguel (Trigoniaceae) (Thomas et al, 2003).

Figure 1 a) Study area in which Thomas et al. (2003) based their list of endemic species. b) Bahia Coastal Forest (BCF) according to Saiter et al. (2016a). c) Municipalities with at least 50% of its area within the BCF (light grey), and those added to the study area based on IBGE (1993) (dark grey). Coordinates: UTM. Datum WGS 84.
Checklists of endemic taxa are critical to overcome issues that hinder conservation strategies in highly biodiverse regions, such as: 1) basic taxonomic information (Linnean shortfall, or "which species are there?"), and 2) lack of geographic information (Wallacean shortfall, or "where do these species occur?") (Whittaker et al. 2005). Both shortfalls are part of reality in Bahia Coastal Forests, world-renowned as one of the highest diversity areas in tree species (Thomas et al. 1998, Martini et al. 2007, Murray-Smith et al. 2009).
We aimed to provide a list of endemic species of angiosperms in Bahia Coastal Forest following Saiter et al. (2016a) to verify whether the current list agrees with the preliminary checklist by Thomas et al. (2003). We expect this updated list to further highlight the importance of BCF as a critical area for biodiversity conservation in the Brazilian Atlantic forest.
Material and Methods
1. Study area
To delimit the study area we used the BCF sensuSaiter et al. (2016a) (Figure 1b) with a slight modification to include the municipalities whose territory was at least 50% inside that perimeter. Municipalities were used as a base for the retrieval of taxonomic information from online datasets (see data collection). To remove the arbitrary northern limit from Saiter et al. (2016a) (Figure 1b), the study area was expanded to include seven municipalities complying with the 50% territory criterion (Figure 1c). Municipal limits were defined based on data from the Brazilian Institute for Geography and Statistics (IBGE 1993) (Figure 1c).
The resulting area comprised 156 municipalities, which account for approximately 99,000 km2 (Figure 1c) at southern Bahia state, and a small fraction from northern Espírito Santo state and northeastern Minas Gerais state, contrasting with an area of about 119,000 km2 at southern Bahia which included more inland areas, and northen Espírito Santo with a southern limit at the Doce River (Thomas et al. 2003) (Figure 1a).
2. Data collection
To compile the checklist of endemic species of angiosperms in BCF, we started by performing a data search in the Brazilian Plant List (Flora do Brasil 2020) on March 9, 2017, using the following filters simultaneously: "group: angiosperms", "endemism: only endemic to Brazil", "origin: native", "state: Bahia", and "phytogeographic domain: Atlantic Forest". We decided not to search only for species endemic to the Atlantic Forest in Bahia to avoid excluding species that could also occur in the north of Espírito Santo and in a small area in northeastern Minas Gerais (Figure 1c). The initial search resulted in 3,338 species, 59 subspecies and 154 varieties (3,551 taxa). These records were then checked in the SpeciesLink database (splink.cria.org.br). Each taxon map generated by the SpeciesLink website was visually checked, first to remove taxa that were clearly not endemic to BCF, i.e. those with several records outside the study area. Each of the 1,336 remaining taxa were then mapped with QGis 2.14 (Quantum GIS Development Team 2016) and we verified whether the points of occurrence were enclosed in the limits of the study area, which resulted in the further removal of 701 taxa.
From the 635 taxa left, the records with incongruent data were verified by experts or checked using recent taxonomic studies (Appendix 1). This step resulted in 30 additional non-endemic species discarded from the list. The taxa for which no scientific records were found in Google Scholar (scholar.google.com), Biodiversity Heritage Library (biodiversitylibrary.com), SpeciesLink (splink.cria.org.br), and the Brazilian Plant List (Flora do Brasil 2020) were complemented by a search for location records in the CEPEC herbarium. Once this verification was completed, 53 species from the list had to be removed because no records were available and a further five species were removed due to lack of precise geographic information.
Results
The final checklist consisted of 547 angiosperm taxa endemic to BCF (535 species, six subspecies, and six varieties) in 69 families (none endemic) and 230 genera (Table 1). Seven genera are endemic to BCF: Bahiella J.F.Morales (Apocynaceae), Santosia R.M.King & H.Rob. (Asteraceae), Harleyodendron (Fabaceae, Figure 2f), Cubitanthus Barringer (Linderniaceae), Anomochloa and Parianella Hollowell, F.M.Ferreira & R.P.Oliveira (Poaceae), and Andreadoxa Kallunki (Rutaceae). The genera with the highest number of endemic species in the BCF were Aechmea Ruiz & Pav. (Bromeliaceae) (33 spp.), Pavonia Cav. (Malvaceae) and Myrcia DC. (Myrtaceae), each with 16 species. The three families with the highest numbers of endemic taxa (species + infraspecific taxa) were Bromeliaceae (108), Fabaceae (41) and Myrtaceae (32).
Table 1 Checklist of angiosperms endemic to Bahia Coastal Forest sensu Saiter et al. (2016a).
Acanthaceae |
---|
Aphelandra ignea Nees ex Steud. |
Clistax bahiensis Profice & Leitman |
Harpochilus phaeocarpus Nees |
Herpetacanthus magnobracteolatus Indriunas & Kameyama |
Herpetacanthus strongyloides Indriunas & Kameyama |
Herpetacanthus tetrandrus (Nees & Mart.) Herter |
Justicia antirrhina Nees & Mart. |
Justicia physogaster Lindau |
Lepidagathis cuneiformis Kameyama |
Mendoncia bahiensis Profice |
Mendoncia blanchetiana Profice |
Pseuderanthemum albiflorum (Hook.) Radlk. |
Pseuderanthemum verbenaceum (Nees & Mart.) Radlk. |
Ruellia sessilifolia (Nees) Lindau |
Schaueria gonyostachya (Nees & Mart.) Nees |
Schaueria hirsuta Nees |
Schaueria marginata Nees |
Schaueria pyramidalis A.L.A.Côrtes |
Achariaceae |
Kuhlmanniodendron macrocarpum Groppo, Favaretto & Fiaschi |
Amaranthaceae |
Lecosia formicarum Pedersen |
Amaryllidaceae |
Griffinia arifolia Ravenna |
Griffinia parviflora Ker Gawl. |
Griffinia paubrasilica Ravenna |
Annonaceae |
Annona bahiensis (Maas & Westra) H.Rainer * |
Duguetia magnolioidea Maas * |
Duguetia restingae Maas * |
Duguetia reticulata Maas * |
Guatteria stenocarpa Lobão, Maas & Mello-Silva |
Hornschuchia cauliflora Maas & Setten * |
Hornschuchia leptandra D.M.Johnson * |
Hornschuchia obliqua Maas & Setten * |
Hornschuchia polyantha Maas * |
Hornschuchia santosii D.M.Johnson * |
Malmea obovata R.E.Fr. * |
Pseudoxandra bahiensis Maas * |
Unonopsis bahiensis Maas & Orava |
Xylopia involucrata M.C.Dias & Kinoshita * |
Apocynaceae |
Aspidosperma thomasii Marc.-Ferr. * |
Bahiella blanchetii (A.DC.) J.F.Morales |
Bahiella infundibuliflora J.F.Morales |
Lacmellea bahiensis J.F.Morales |
Marsdenia carvalhoi Morillo & Carnevali |
Matelea riparia Morillo |
Matelea santosii Morillo & Fontella |
Oxypetalum laciniatum Rapini & Farinaccio |
Rauvolfia atlantica Emygdio |
Araceae |
Anthurium bromelicola subsp. bahiense Mayo et al. * |
Anthurium illepidum Schott |
Anthurium molle E.G.Gonç. & J.G.Jardim |
Anthurium teimosoanum E.G.Gonç. & J.G.Jardim |
Asterostigma riedelianum (Schott) Kuntze |
Dracontioides salvianii E.G.Gonç. |
Philodendron aemulum Schott |
Zomicarpa steigeriana Maxim. ex Schott |
Araliaceae |
Schefflera aurata Fiaschi |
Arecaceae |
Bactris soeiroana Noblick ex A.J.Hend. |
Geonoma pohliana subsp. rubescens (Wendland ex Drude) Henderson |
Geonoma pohliana subsp. unaensis Henderson |
Syagrus × camposportoana (Bondar) Glassman |
Syagrus itapebiensis (Noblick & Lorenzi) Noblick & Meerow |
Syagrus santosii K. Soares & C.A.Guim. |
Aristolochiaceae |
Aristolochia brunneomaculata I.Abreu & Giul. |
Aristolochia longispathulata F.González * |
Asparagaceae |
Hagenbachia brasiliensis Nees & Mart. |
Asteraceae |
Acmella paniculata (Wall. ex DC.) R.K.Jansen |
Austroeupatorium morii R.M.King & H.Rob. |
Barrosoa atlantica R.M.King & H.Rob. * |
Diacranthera hebeclinia H.Rob. |
Litothamnus ellipticus R.M.King & H.Rob. * |
Mikania amorimii Borges & Forzza |
Mikania kubitzkii R.M.King & H.Rob. * |
Mikania mattos-silvae R.M.King & H.Rob. * |
Mikania santosii R.M.King & H.Rob. * |
Piptocarpha riedelii (Sch.Bip.) Baker |
Santosia talmonii R.M.King & H.Rob. * |
Vernonanthura vinhae (H.Rob.) H.Rob. * |
Begoniaceae |
Begonia delicata Gregório & J.A.S.Costa |
Begonia elianeae Gregório & J.A.S.Costa |
Begonia epibaterium Mart. ex A.DC. |
Begonia goldingiana L.Kollmann & A.P.Fontana |
Begonia mattos-silvae L.B.Sm. ex S.F.Sm. & Wassh. * |
Begonia pinheironis L.B.Sm. ex S.F.Sm. & Wassh. * |
Begonia russelliana L.B.Sm. ex S.F.Sm. & Wassh. * |
Begonia saxifraga A.DC. |
Begonia subacida Irmsch. |
Begonia sylvatica Meisn. ex A.DC. |
Bignoniaceae |
Handroanthus parviflorus Espírito-Santo & M.M. Silva-Castro |
Bromeliaceae |
Aechmea amicorum B.R.Silva & H.Luther |
Aechmea amorimii Leme * |
Aechmea ampla L.B.Sm. |
Aechmea andersoniana Leme & H.Luther |
Aechmea andersonii H.Luther & Leme |
Aechmea carvalhoi E.Pereira & Leme |
Aechmea correia-araujoi E.Pereira & Moutinho |
Aechmea curranii (L.B.Sm.) L.B.Sm. & M.A.Spencer |
Aechmea digitata L.B.Sm. & R.W.Read |
Aechmea discordiae Leme |
Aechmea disjuncta (L.B.Sm.) Leme & J.A.Siqueira |
Aechmea echinata (Leme) Leme * |
Aechmea farinosa (Regel) L.B.Sm. |
Aechmea glandulosa Leme |
Aechmea gregaria Leme & L.Kollmann |
Aechmea guaratingensis Leme & L.Kollmann |
Aechmea heterosepala Leme |
Aechmea incompta Leme & H.Luther |
Aechmea ituberaensis Leme & L.Kollmann |
Aechmea laevigata Leme |
Aechmea lanata (L.B.Sm.) L.B.Sm. & M.A.Spencer |
Aechmea limae Leme |
Aechmea lymanii W.Weber |
Aechmea miniata Beer ex Baker |
Aechmea mira Leme & H.Luther |
Aechmea mollis L.B.Sm. |
Aechmea pendulispica Leme & L.Kollmann |
Aechmea ramusculosa Leme |
Aechmea tentaculifera Leme, Amorim & J.A. Siqueira |
Aechmea turbinocalyx Mez |
Aechmea viridipetala A.F.Costa & Amorim |
Aechmea viridostigma Leme & H.Luther |
Aechmea weberi (E.Pereira & Leme) Leme |
Alcantarea cerosa Leme, A.P.Fontana & O.A.B.Ribeiro |
Alcantarea pataxoana Versieux |
Araeococcus montanus Leme |
Araeococcus nigropurpureus Leme & J.A.Siqueira |
Araeococcus sessiliflorus Leme & J.A.Siqueira |
Billbergia fosteriana L.B.Sm. |
Billbergia macrocalyx Hook. |
Canistrum fosterianum L.B.Sm. |
Canistrum guzmanioides Leme |
Canistrum lanigerum H.Luther & Leme |
Canistrum sandrae Leme |
Canistrum seidelianum W.Weber |
Cryptanthus colnagoi Rauh & Leme |
Cryptanthus coriaceus Leme |
Cryptanthus ilhanus Leme |
Cryptanthus lyman-smithii Leme |
Cryptanthus pseudopetiolatus Philcox * |
Cryptanthus ruthiae Philcox |
Cryptanthus ubairensis I.Ramírez |
Cryptanthus vexatus Leme |
Cryptanthus viridovinosus Leme |
Cryptanthus walkerianus Leme & L.Kollmann |
Hohenbergia barbarespina Leme & Fraga |
Hohenbergia brachycephala L.B.Sm. |
Hohenbergia burle-marxii Leme & W.Till |
Hohenbergia capitata Schult. & Schult.f. |
Hohenbergia castellanosii L.B.Sm. & Read |
Hohenbergia correia-araujoi E.Pereira & Moutinho |
Hohenbergia flava Leme & C.C.Paula |
Hohenbergia hatschbachii Leme * |
Hohenbergia itamarajuensis Leme & Baracho |
Hohenbergia littoralis L.B.Sm. |
Hohenbergia pabstii L.B.Sm. & Read |
Hohenbergia reconcavensis Leme & Fraga |
Hohenbergia sandrae Leme |
Lymania alvimii (L.B.Sm. & R.W.Read) R.W.Read * |
Lymania azurea Leme |
Lymania brachycaulis (E.Morren ex Baker) L.F.Sousa |
Lymania corallina (Brong. ex Beer) R.W.Read |
Lymania globosa Leme |
Lymania languida Leme |
Lymania marantoides (L.B.Sm.) R.W.Read |
Lymania spiculata Leme & Forzza * |
Neoregelia azevedoi Leme |
Neoregelia crispata Leme * |
Neoregelia longisepala E.Pereira & I.A.Penna * |
Neoregelia rothinessa Leme, H.Luther & W.Till |
Neoregelia silvomontana Leme & J.A.Siqueira |
Neoregelia viridolineata Leme |
Neoregelia wilsoniana M.B.Foster |
Orthophytum buranhense Leme & A.P.Fontana |
Orthophytum guaratingense Leme & L.Kollmann |
Orthophytum rubrum L.B.Sm. |
Portea alatisepala Philcox * |
Portea filifera L.B.Sm. |
Portea grandiflora Philcox * |
Portea kermesina K.Koch |
Portea nana Leme & H.Luther |
Quesnelia alborosea A.F.Costa & T.Fontoura |
Quesnelia clavata Amorim & Leme |
Quesnelia dubia Leme |
Quesnelia koltesii Amorim & Leme |
Ronnbergia brasiliensis E.Pereira & I.A.Penna * |
Ronnbergia carvalhoi Martinelli & Leme * |
Ronnbergia neoregelioides Leme |
Ronnbergia silvana Leme |
Vriesea dictyographa Leme |
Vriesea graciliscapa W.Weber |
Vriesea longisepala A.F.Costa |
Vriesea minuta Leme |
Vriesea minutiflora Leme |
Vriesea roberto-seidelii W.Weber |
Vriesea ruschii L.B.Sm. subsp. ruschii |
Vriesea sandrae Leme |
Vriesea silvana Leme |
Burseraceae |
Protium icicariba var. talmonii Daly * |
Trattinnickia lorenziana Daly & M.F.F.Melo |
Cactaceae |
Rhipsalis hileiabaiana (N.P.Taylor & Barthlott) N.Korotkova & Barthlott * |
Calophyllaceae |
Kielmeyera itacarensis Saddi |
Kielmeyera marauensis Saddi |
Capparaceae |
Colicodendron martianum Cornejo |
Caryocaraceae |
Caryocar edule Casar. |
Chrysobalanaceae |
Couepia bondarii Prance * |
Couepia coarctata Prance * |
Couepia impressa subsp. cabraliae Prance * |
Couepia longipetiolata Prance * |
Licania bahiensis Prance |
Licania lamentanda Prance * |
Licania littoralis Warm. var. littoralis |
Licania santosii Prance * |
Licania turbinata Benth. |
Parinari alvimii Prance * |
Clusiaceae |
Tovomita iaspidis L.Marinho & Amorim |
Tovomita megantha L.Marinho & Amorim |
Commelinaceae |
Dichorisandra jardimii Aona & M.C.E.Amaral |
Dichorisandra leucophthalmos Hook. |
Dichorisandra leucosepala Aona & M.C.E.Amaral |
Dichorisandra marantoides Aona & Faden |
Dichorisandra ordinatiflora Aona & Faden |
Dichorisandra radicalis Nees & Mart. |
Dichorisandra subtilis Aona & M.C.E.Amaral |
Connaraceae |
Connarus blanchetii var. laurifolius (Baker) Forero |
Connarus cuneifolius Baker |
Connarus portosegurensis Forero |
Rourea bahiensis Forero * |
Rourea carvalhoi Forero et al. * |
Rourea discolor Baker |
Rourea macrocalyx Carbonó et al. * |
Rourea tenuis G.Schellenb. |
Cucurbitaceae |
Cayaponia nitida Gomes-Klein & Pirani |
Fevillea bahiensis G.Rob. & Wunderlin |
Gurania wawrei Cogn. |
Cyclanthaceae |
Asplundia maximiliani Harling |
Cyperaceae |
Becquerelia discolor Kunth |
Hypolytrum bahiense M.Alves & W.W.Thomas * |
Hypolytrum jardimii M.Alves & W.W.Thomas * |
Hypolytrum lucennoi M.Alves & W.W.Thomas |
Dichapetalaceae |
Stephanopodium magnifolium Prance * |
Tapura martiniae Amorim & D.Lisboa |
Tapura zei-limae Amorim & Fiaschi |
Dilleniaceae |
Davilla bahiana Aymard |
Davilla macrocarpa Eichler |
Dioscoreaceae |
Dioscorea macrothyrsa Uline |
Ebenaceae |
Diospyros amabi B.Walln. |
Diospyros riedelii (Hiern) B.Walln. |
Diospyros scottmorii B.Walln. |
Eriocaulaceae |
Actinocephalus ochrocephalus (Körn.) Sano |
Erythroxylaceae |
Erythroxylum compressum Peyr. |
Erythroxylum leal-costae Plowman |
Erythroxylum martii Peyr. |
Erythroxylum mattos-silvae Plowman * |
Erythroxylum membranaceum Plowman * |
Erythroxylum santosii Plowman * |
Erythroxylum splendidum Plowman * |
Euphorbiaceae |
Actinostemon lasiocarpus (Müll.Arg.) Baill. |
Algernonia bahiensis (Emmerich) G.L.Webster * |
Bernardia gambosa Müll.Arg. |
Bernardia micrantha Pax & K.Hoffm. |
Croton sapiifolius Müll.Arg. |
Croton thomasii Riina & P.E.Berry |
Dalechampia armbrusteri G.L.Webster |
Dalechampia viridissima G.L.Webster * |
Gymnanthes gaudichaudii Müll.Arg. |
Ophthalmoblapton pedunculare Müll.Arg. |
Fabaceae |
Andira carvalhoi R.T.Penn. & H.C.Lima * |
Andira marauensis N.F.Mattos * |
Arapatiella psilophylla (Harms) R.S.Cowan * |
Canavalia cassidea G.P.Lewis * |
Canavalia dolichothyrsa G.P.Lewis * |
Chamaecrista amabilis H.S.Irwin & Barneby * |
Chamaecrista amorimii Barneby * |
Chamaecrista onusta H.S.Irwin & Barneby * |
Chamaecrista salvatoris (H.S.Irwin & Barneby) H.S.Irwin & Barneby |
Copaifera majorina Dwyer |
Dahlstedtia bahiana (A.M.G.Azevedo) M.J.Silva & A.M.G.Azevedo |
Harleyodendron unifoliolatum R.S.Cowan * |
Inga aptera (Vinha) T.D.Penn. * |
Inga conchifolia L.P.Queiroz |
Inga pedunculata (Vinha) T.D.Penn. * |
Inga pleiogyna T.D.Penn. |
Machaerium aureum Filardi & H.C.Lima |
Moldenhawera blanchetiana var. multijuga L.P.Queiroz et al. |
Moldenhawera intermedia G.P.Lewis & L.P.Queiroz |
Moldenhawera luschnathiana Yakovlev |
Moldenhawera nutans L.P.Queiroz et al. |
Muellera longiunguiculata (MJ.Silva & AMG.Azevedo) MJ.Silva & AMG.Azevedo |
Ormosia lewisii D.B.O.S.Cardoso, C.H.Stirt. & Torke * |
Ormosia limae D.B.O.S.Cardoso & L.P.Queiroz |
Ormosia timboensis D.B.O.S.Cardoso, Meireles & H.C.Lima |
Parapiptadenia ilheusana G.P.Lewis * |
Phanera carvalhoi (Vaz) Vaz |
Piptadenia killipii var. cacaophila G.P.Lewis * |
Piptadenia ramosissima Benth. |
Piptadenia santosii Barneby ex G.P.Lewis * |
Schnella lilacina (Wunderlin & Eilers) Wunderlin |
Senegalia amorimii M.J.F.Barros & M.P.Morim |
Senegalia olivensana (G.P.Lewis) Seigler & Ebinger |
Senegalia piptadenioides (G.P.Lewis) Seigler & Ebinger |
Swartzia alternifoliolata Mansano |
Swartzia arenophila R.B.Pinto, Torke & Mansano |
Swartzia curranii R.S.Cowan |
Swartzia pinheiroana R.S.Cowan * |
Swartzia riedelii R.S.Cowan |
Swartzia thomasii R.B.Pinto, Torke & Mansano |
Zollernia magnifica A.M.Carvalho & Barneby * |
Gentianaceae |
Macrocarpaea atlantica J.R.Grant & V.Trunz |
Macrocarpaea dolichophylla J.R.Grant & V.Trunz |
Macrocarpaea orbiculata J.R.Grant & V.Trunz |
Icacinaceae |
Pleurisanthes brasiliensis (Val.) Tiegh. |
Iridaceae |
Neomarica brachypus (Baker) Sprague |
Neomarica floscella A.Gil & M.C.E.Amaral |
Neomarica portosecurensis (Ravenna) Chukr * |
Neomarica unca (Ravenna) A.Gil |
Lauraceae |
Nectandra micranthera Rohwer |
Ocotea montana (Meisn.) Mez |
Ocotea ramosissima L.C.S.Assis e Mello-Silva |
Ocotea sperata P.L.R.Moraes et van der Werff |
Ocotea thinicola van der Werff et P.L.R.Moraes |
Lecythidaceae |
Eschweilera complanata S.A.Mori |
Eschweilera mattos-silvae S.A.Mori * |
Eschweilera sphaerocarpa M.Ribeiro & S.A.Mori |
Linderniaceae |
Cubitanthus alatus (Cham. & Schltdl.) Barringer |
Loganiaceae |
Spigelia genuflexa Popovkin & Struwe |
Strychnos alvimiana Krukoff & Barneby * |
Strychnos setosa Krukoff & Barneby |
Loranthaceae |
Psittacanthus excrenulatus Rizzini |
Psittacanthus salvadorensis Kuijt |
Struthanthus longiflorus Rizzini |
Malpighiaceae |
Bunchosia itacarensis W.R.Anderson * |
Heteropterys sanctorum W.R.Anderson * |
Peixotoa sericea C.E.Anderson * |
Stigmaphyllon hispidum C.E.Anderson |
Stigmaphyllon macropodum A.Juss. |
Malvaceae |
Byttneria cristobaliana Dorr |
Pavonia cauliflora (Nees) Fryxell ex G.L.Esteves |
Pavonia ciliata G.L.Esteves & Krapov. * |
Pavonia crispa Krapov. * |
Pavonia gerleniae Gonçalez & M.C.Duarte |
Pavonia goetheoides (Hassl.) Fryxell ex G.L.Esteves |
Pavonia latibracteolata Krapov. * |
Pavonia longifolia A.St.-Hil. |
Pavonia macrobracteolata Gonçalez & M.C.Duarte |
Pavonia ovaliphylla G.L.Esteves & Krapov. * |
Pavonia paucidentata Fryxell |
Pavonia pilifera Krapov. |
Pavonia rubriphylla G.L.Esteves |
Pavonia sancti Krapov. |
Pavonia spectabilis Krapov. * |
Pavonia spiciformis Krapov. * |
Pavonia stipularis Krapov. * |
Marantaceae |
Goeppertia fasciata (Linden ex K.Koch) Borchs. & S.Suárez |
Goeppertia rufibarba (Fenzl) Borchs. & S.Suárez |
Ischnosiphon bahiensis L.Andersson * |
Monotagma grallatum Hagberg & R. Erikss. * |
Stromanthe bahiensis Yosh.-Arns, Mayo & J.M.A. Braga |
Marcgraviaceae |
Schwartzia geniculatiflora Gir.-Cañas & Fiaschi |
Melastomataceae |
Bertolonia alternifolia Baumgratz, Amorim & A.B.Jardim |
Bertolonia bullata Baumgratz, Amorim & A.B.Jardim |
Huberia carvalhoi Baumgratz |
Huberia sessilifolia R. Godenberg & Michelangelo |
Meriania inflata Michelangeli & R.Goldenb. |
Miconia lurida Cogn. |
Ossaea loligomorpha R.Goldenb. & Reginato |
Ossaea sulbahiensis D'El Rei Souza |
Physeterostemon aonae Amorim, Michelangeli & R.Goldenb. |
Physeterostemon fiaschii R.Goldenb. & Amorim |
Physeterostemon jardimii R.Goldenb. & Amorim |
Physeterostemon thomasii Amorim, Michelangeli & R.Goldenb. |
Pleiochiton amorimii Reginato & R.Goldenb. |
Tibouchina bahiensis Wurdack * |
Tibouchina bradeana Renner |
Tibouchina paulo-alvinii Guimarães da Vinha * |
Tibouchina stipulacea Vinha * |
Tibouchina taperoensis Wurdack * |
Tibouchina tomentulosa Wurdack |
Meliaceae |
Guarea anomala T.D.Penn |
Trichilia florbranca T.D.Penn. * |
Trichilia magnifoliola T.D.Penn. * |
Menispermaceae |
Curarea crassa Barneby * |
Moraceae |
Dorstenia setosa Moric. |
Myrtaceae |
Calyptranthes blanchetiana O.Berg |
Eugenia barrana Sobral |
Eugenia beruttii (Mattos) Mattos |
Eugenia fissurata Mattos |
Eugenia itacarensis Mattos * |
Eugenia longifolia DC. |
Eugenia serraegrandis Sobral |
Eugenia sessilifolia DC. |
Eugenia unana Sobral |
Marlierea lealcostae G.M.Barroso & Peixoto |
Marlierea verticillaris O.Berg |
Myrcia abrantea (O.Berg) E.Lucas & Sobral |
Myrcia cataphyllata M.F.Santos |
Myrcia felisbertii (DC.) O.Berg |
Myrcia gigantea (O.Berg) Nied. |
Myrcia grazielae NicLugh. |
Myrcia marianae Staggemeier & E.Lucas |
Myrcia monoclada Sobral |
Myrcia pendula Sobral |
Myrcia pseudomarlierea Sobral |
Myrcia raminfinita L.Marinho & E.Lucas |
Myrcia spathulifolia Proença |
Myrcia stigmatosa O.Berg |
Myrcia teimosa Sobral |
Myrcia tetraphylla Sobral |
Myrcia thomasii B.S.Amorim & A.R.Lourenço |
Myrcia truncata Sobral |
Plinia callosa Sobral * |
Plinia longiacuminata Sobral |
Plinia muricata Sobral * |
Plinia rara Sobral * |
Plinia spiciflora (Nees & Mart.) Sobral |
Nyctaginaceae |
Neea alumnorum M.Pignal, Soares Filho & Romaniuc |
Neea duckei (Huber) Furlan |
Ochnaceae |
Ouratea bahiensis Sastre * |
Ouratea gigantophylla (Erhard) Engl. |
Ouratea longipes Sastre * |
Ouratea papulosa Sastre |
Ouratea platicaulis Sastre * |
Orchidaceae |
Anathallis velvetina Luer & Toscano |
Aspasia silvana F.Barros * |
Bifrenaria silvana V.P.Castro * |
Brassavola reginae Pabst |
Bulbophyllum teimosense E.C.Smidt & Borba |
Cattleya alaorii (Brieger & Bicalho) Van den Berg |
Cattleya grandis (Lindl. & Paxton) A.A.Chadwick |
Cattleya kerrii Brieger & Bicalho * |
Cirrhaea silvana V.P.Castro & Campacci * |
Coryanthes bahiensis Marçal & Chiron |
Coryanthes bueraremensis Campacci & Bohnke |
Elleanthus hymenophorus (Rchb.f.) Rchb.f. |
Encyclia fimbriata C.A.Bastos, Van den Berg & Meneguzzo |
Epidendrum garciae Pabst |
Gomesa silvana (V.P.Castro & Campacci) M.W.Chase & N.H.Williams |
Gongora meneziana V.P.Castro & G.Gerlach |
Koellensteinia abaetana L.P.Queiroz |
Leptotes bohnkiana Campacci |
Masdevallia sururuana Campacci |
Pabstiella dasilvae Chiron & Xim.Bols. |
Promenaea silvana F.Barros & Cath. * |
Prosthechea bohnkiana V.P.Castro & G.F.Carr |
Prosthechea bueraremensis (Campacci) Campacci |
Stanhopea bueraremensis Campacci & Marçal |
Stenia bohnkiana V.P.Castro & G.F.Carr |
Oxalidaceae |
Oxalis alata var. hirta Lourteig * |
Oxalis bela-vitoriae Lourteig * |
Oxalis kuhlmannii var. adpressipila Lourteig |
Passifloraceae |
Passiflora igrapiunensis T.S.Nunes & L.P.Queiroz |
Phyllanthaceae |
Discocarpus pedicellatus Fiaschi & Cordeiro |
Phyllanthus carvalhoi G.L.Webster |
Picramniaceae |
Picramnia coccinea W.W.Thomas * |
Piperaceae |
Peperomia epipremnifolia D.Monteiro & Leitman |
Peperomia riparia Yunck. |
Peperomia serpentarioides Miq. |
Peperomia sulbahiensis D.Monteiro & M.Coelho |
Piper bahianum Yunck. |
Piper robustipedunculum Yunck. |
Piper vellosoi Yunck. |
Poaceae |
Alvimia auriculata Soderstr. & Londoño * |
Alvimia gracilis Soderstr. & Londoño * |
Anomochloa marantoidea Brongn. * |
Arberella bahiensis Soderstr. & Zuloaga * |
Atractantha cardinalis Judz. * |
Atractantha radiata McClure * |
Chusquea clemirae A.C.Mota, R.P.Oliveira & L.G.Clark |
Diandrolyra pygmaea Soderstr. & Zuloaga ex R.P.Oliveira & L.G.Clark |
Digitaria doellii Mez * |
Eremetis robusta Hollowell, F.M.Ferreira & R.P.Oliveira |
Eremocaulon aureofimbriatum Soderstr. & Londoño * |
Ichnanthus longhi-wagnerae A.C.Mota & R.P.Oliveira |
Merostachys annulifera Send. |
Merostachys argentea Send. |
Merostachys lanata Send. |
Merostachys magnispicula Send. |
Merostachys medullosa Send. |
Merostachys ramosissima Send. |
Olyra bahiensis R.P.Oliveira & Longhi-Wagner |
Olyra filiformis Trin. |
Olyra latispicula Soderstr. & Zuloaga * |
Parianella carvalhoi (R.P.Oliveira & Longhi-Wagner) F.M.Ferreira & R.P. Oliveira |
Parianella lanceolata (Trin.) F.M.Ferreira & R.P.Oliveira |
Paspalum restingense Renvoize * |
Paspalum strigosum Döll |
Piresia palmula M.L.S.Carvalho & R.P.Oliveira |
Raddia distichophylla (Schrad. ex Nees) Chase * |
Raddia stolonifera R.P.Oliveira & Longhi-Wagner |
Polygalaceae |
Caamembeca martinelli (Marques & E.F.Guim.) J.F.B.Pastore * |
Caamembeca martinelli var. carnosa (Marques & E.F.Guim.) J.F.B.Pastore |
Securidaca revoluta (A.W.Benn.) Marques |
Primulaceae |
Cybianthus nemoralis (Mez) G.Agostini |
Rubiaceae |
Chomelia bahiae J.H.Kirkbr. |
Denscantia andrei (E.L.Cabral & Bacigalupo) E.L.Cabral & Bacigalupo * |
Denscantia macrobracteata (E.L.Cabral & Bacigalupo) E.L.Cabral & Bacigalupo * |
Faramea bicolor J.G.Jardim & Zappi |
Faramea nocturna J.G.Jardim & Zappi |
Ixora bahiensis Benth. |
Ixora cabraliensis Di Maio et Peixoto |
Malanea harleyi J.H.Kirkbr. * |
Mitracarpus anthospermoides K.Schum. |
Posoqueria bahiensis Macias & Kin.-Gouv. |
Psychotria martiusii Müll.Arg. |
Psychotria megalocalyx Müll.Arg. |
Psychotria salzmanniana Müll.Arg. |
Psychotria silvicola Müll.Arg. |
Psychotria strigosa Müll.Arg. |
Psychotria wawrana Müll.Arg. |
Rudgea hileiabaiana Zappi & Bruniera |
Rudgea ilheotica Müll.Arg. |
Rudgea involucrata Müll.Arg. |
Rudgea malpighiacea Standl. |
Rudgea mouririoides Standl. |
Salzmannia arborea J.G. Jardim |
Sphinctanthus insignis Steyerm. |
Rutaceae |
Andreadoxa flava Kallunki * |
Conchocarpus concinnus Kallunki * |
Conchocarpus dasyanthus Kallunki * |
Conchocarpus fissicalyx Pirani * |
Conchocarpus gaudichaudianus subsp. bahiensis Kallunki * |
Conchocarpus hirsutus Pirani * |
Conchocarpus inopinatus Pirani * |
Conchocarpus longipes Kallunki * |
Conchocarpus mastigophorus Kallunki * |
Conchocarpus modestus Kallunki * |
Conchocarpus punctatus Kallunki * |
Conchocarpus racemosus (Nees & Mart.) Kallunki & Pirani |
Conchocarpus santosii Pirani & Kallunki * |
Ertela bahiensis (Engl.) Kuntze |
Galipea revoluta Pirani |
Neoraputia calliantha Kallunki |
Neoraputia micrantha Kallunki |
Zanthoxylum nemorale Mart. |
Zanthoxylum retusum (Albuq.) P.G.Waterman |
Sapindaceae |
Cardiospermum integerrimum Radlk. * |
Paullinia livescens Radlk. |
Paullinia unifoliolata Perdiz & Ferrucci |
Serjania morii Acev.-Rodr. |
Serjania scopulifera Radlk. |
Sapotaceae |
Chromolucuma apiculata Alves-Araújo & M.Alves |
Pouteria atlantica Alves-Araújo & M.Alves |
Pouteria glauca T.D.Penn. |
Pouteria synsepala Popovkin & A.D.Faria |
Pouteria trifida Alves-Araújo & M.Alves |
Pradosia longipedicellata Alves-Araújo & M.Alves |
Verbenaceae |
Citharexylum obtusifolium Kuhlm. |
Stachytarpheta hirsutissima Link |
Vochysiaceae |
Vochysia talmonii M.C.Vianna et al. |
*Taxon also in Thomas et al. (2003).

Figure 2 Some species found in the checklist of endemic angiosperms of BCF: a) Kuhlmanniodendron macrocarpum (Achariaceae). b) Quesnelia koltesii (Bromeliaceae). c) Dichorisandra leucophtalmos (Commelinaceae). d) Tapura zei-limae (Dichapetalaceae). e) Arapatiella psilophylla (Fabaceae). f) Harleyodendron unifoliolatum (Fabaceae). g) Pavonia goetheoides (Malvaceae). h) Pleiochiton amorimii (Melastomataceae). i) Anomochloa marantoidea (Poaceae). Photos: a, b, d, g, h) A.Amorim; c) L.Aona; e, f) D.Cardoso; i) J.Jardim.
Discussion
The families with the highest numbers of endemic taxa in BCF are also among the ten richest angiosperm families in Brazil and in the Atlantic Forest (BFG 2015). Thomas et al. (2003) reported Fabaceae (53 spp.) as the richest family of a total of 65 families in the previous list. Compared with the BFG (2015), it is remarkable that Orchidaceae, one of the three richest families in Brazil and in the Atlantic Forest, was not among the richest families in this study, with only 25 taxa. This may be due to the lack of geographic records for 14 species in this family, which had to be discarded.
Among the 20 richest angiosperm genera in Brazil (BFG 2015), Aechmea and Myrcia are also among the genera with the highest number of endemic species in the BCF. In a preliminary list of the endemic angiosperm species from southern Bahia and northern Espírito Santo, Thomas et al. (2003) cited Conchocarpus J.C.Mikan (Rutaceae) (19 spp.) as the richest genus, followed by Pavonia Cav. (Malvaceae) (13 spp.), Couepia Aubl., and Erythroxylum P.Browne (seven spp. each). The occurrence of only 12 endemic species of Conchocarpus in the present checklist probably indicates that several endemic species of this genus are exclusively found at more inland seasonally dry forests (Kallunki & Pirani 1998).
Among the endemic genera from southern Bahia and northern Espírito Santo reported by Thomas et al. (2003), such as Arapatiella and Brodriguesia (Fabaceae), Atractantha, Alvimia and Sucrea (Poaceae), and Trigoniodendron (Trigoniaceae), several are not endemic to BCF. For example, Brodriguesia, Atractantha and Trigoniodendron have been recorded outside our study area, respectively, in Sergipe, Amazonas, and near the Doce River. Similarly, Physeterostemon R.Goldenb. & Amorim (Melastomataceae), a recently described endemic genus from the Atlantic Forest of Bahia state (Amorim et al. 2014, Goldenberg et al. 2016, Goldenberg & Amorim 2006), was not considered endemic to BCF in this checklist, as among its species P. gomesii Amorim & R.Goldenb. occurs in the municipality of Boa Nova (Amorim et al. 2014), which lies slightly outside our study area.
In comparison with the preliminary list of Thomas et al. (2003), which included 395 endemic angiosperm taxa from an area that covered a larger part of northern Espírito Santo state, as far as the Doce River, and the southern Bahia Atlantic Forest, including more inland areas, and almost reaching the municipality of Salvador (i.e., an area ca. 20% larger than ours) (Figure 1a), the list presented here surprisingly shares only 143 taxa (Table 1). If we had used the Doce River as our area southern limit, it would have included at least 45 additional species restricted to the Linhares region, such as Cryptanthus beuckeri E.Morren (Bromeliaceae), Rourea luizalbertoi Forero, L.A.Vidal & Carbonó (Connaraceae), Simira eliezeriana Peixoto (Rubiaceae) and the endemic genus Riodocea Delprete (Thomas et al. 2003, Rolim et al. 2016). Despite being geographically close and having several species in common (Saiter et al. 2016b), the vegetation of Linhares and the Doce River floodplain are ecologically different from the BCF (Rolim et al. 2005, 2006). In floristic terms, the Linhares region may be more similar to northern Rio de Janeiro, southern Espírito Santo (Silva & Nascimento 2001, Saiter et al. 2016b) and eastern Minas Gerais states (see Krenák-Waitaká Forests sensuSaiter et al. 2016a).
Another difference between the two checklists is due to the time gap of about 15 years between studies. A total of 174 species in our checklist were described after 2003, probably due to increased efforts in collecting and studying plants in the Atlantic Forest in Bahia. For instance, about 41.5% of all new angiosperms in the Brazilian flora described between 1990 and 2006 are endemic to the Atlantic Forest (Sobral & Stehmann 2009). Moreover, 23 species of angiosperms endemic to BCF were described between 2015 and 2017 (e.g., Araújo et al. 2015, Santos et al. 2015, Abreu & Giulietti 2016, Amorim et al. 2016, Côrtes et al. 2016, Ferreira et al. 2016, Marinho & Lucas 2016, Popovkin et al. 2016, Ribeiro et al. 2016, Terra-Araujo et al. 2016, Daly & Melo 2017, Gonçalez et al. 2017).
Compared to the results of a search in the Brazilian Plant List (Flora do Brasil 2020, under construction) using the following filters: "group: angiosperms", "endemism: only endemic to Brazil", "occurrence: only occurs in", "origin: native", "state: Bahia", and "phytogeographic domain: Atlantic Forest", the number of species in our checklist (547 spp.) was much smaller than the 795 species identified by the search engine. Among the reasons for this large difference (248 spp.) it is worth mentioning that a) more than 50 taxa from our list had to be removed due to lack of precise geospatial data, such as the municipality name, which precluded us from citing them as endemic to BCF; b) among the taxa exclusive to the Brazilian Plant List (Flora do Brasil 2020, under construction) that are not restricted to BCF, several occur in more inland seasonally dry forests in the Atlantic Forest, such as Chrysophyllum subspinosum Monach. (Sapotaceae) and Colicodendron bahianum Cornejo & Iltis (Capparaceae); and c) at least 29 species exclusive to the Brazilian Plant List search occur close to, but outside the study area, such as Canistrum camacaense Martinelli & Leme (Bromeliaceae), Inga grazielae (Vinha) T.D.Penn. (Fabaceae), and Bertolonia carmoi Baumgratz (Melastomataceae), all from the municipality of Boa Nova, and Aechmea bicolor L.B.Sm. (Bromeliaceae), Heteropterys jardimii Amorim (Malpighiaceae), and Passiflora timboensis T.S.Nunes & L.P.Queiroz (Passifloraceae), from the municipality of Santa Teresinha.
There are additional reasons to anticipate that the difference in species numbers between the Brazilian Plant List and our list could be even greater, as the list presented here includes 84 taxa that also occur in northern Espírito Santo and in a small area in northeastern Minas Gerais (Figure 1c). Moreover, the Brazilian Plant List (Flora do Brasil 2020, under construction) is an ongoing project on which new taxa can be continuously added and identified as endemic, what could bring the total number of endemic angiosperm species using the same filters even larger. Alternatively, if the native distribution of several taxa is shown to extend beyond the Atlantic Forest domain in Bahia, the difference between the lists could decrease, as fewer species from the Brazilian Plant List would appear as endemic while applying the same search filters.
Developing knowledge on patterns of endemism for plant species is extremely important and must be taken into account in conservation strategies. This information is relevant for the establishment of new protected areas (Kerr 1997), restoration policies (Chazdon 2008) and in directing additional studies (Francisco-Ortega et al. 2010). Considering the key role of endemic species in the design of conservation policies (Myers et al. 2000, Van Der Werff & Consiglio 2004, Moraes et al. 2005, Lamoreux et al. 2006, Martinelli et al. 2008, Essl et al. 2009, Nowak & Nobis 2010), we expect this checklist could guide future studies and conservation strategies, as well as emphasize the importance of the BCF region as a key area for plant conservation.