Malpighiaceae in the Raso da Catarina Ecoregion , Bahia , Brazil

This study presents a floristic survey of Malpighiaceae species from in the Raso da Catarina Ecoregion in the state of Bahia, Brazil. After extensive field and herbarium studies, we identified 18 genera and 32 species of Malpighiaceae in the study area: Aspicarpa harleyi W.R.Anderson, Banisteriopsis C.B.Rob. (3 spp.), Barnebya harleyi W.R.Anderson & B.Gates, Bronwenia ferruginea (Cav.) W.R.Anderson & C.C.Davis, Bunchosia pernambucana W.R.Anderson, Byrsonima Rich. ex Kunth (4 spp.), Carolus chasei (W.R.Anderson) W.R.Anderson, Diplopterys lutea (Griseb.) W.R.Anderson & C.C.Davis, Galphimia brasiliensis (L.) A.Juss., Heteropterys Kunth (5 spp.), Janusia anisandra (A.Juss.) Griseb., Mascagnia Bertero (2 spp.), Mcvaughia bahiana W.R.Anderson, Peixotoa hispidula A.Juss., Ptilochaeta Turcz. (2 spp.), Stigmaphyllon A.Juss. (3 spp.), Tetrapterys Cav. (2 spp.) and Thryallis longifolia Mart. Among the species recorded, three represent new records for the Caatinga biome, 25 are endemic to Brazil, and six are exclusive to the Caatinga. We present an identification key to all species, comments on morphology, taxonomy, distribution and phenology, besides illustrations to most species.


Introduction
Malpighiaceae Juss.comprises approximately 75 genera and 1,300 species distributed in tropical and subtropical regions worldwide (Anderson, W. 2013).Malpighiaceae species are recognized by the presence of hairs unicellular and medifixed, straight or rarely stellate in addition to "T", "Y" or "V"-shaped (malpighiaceous hairs), simple opposite leaves, sepals generally with two oil secreting glands (elaiophores) on the base of the abaxial surface, petals unguiculate, gynoecium 3-carpellate, commonly with 3 styles, 1 ovule per locule and generally schizocarpic fruits splitting into three winged mericarps to, occasionally, nuts or drupes (Anderson, W. 1979a(Anderson, W. , 1981)).Its greatest species diversity is found within the Neotropics (Anderson, W. 2013), and especially in Brazil with 60% are endemics comprising 44 genera and ca.570 species (Mamede et al. 2017).In northeastern Brazil, this family is represented by 31 genera and 241 species, of which 22 genera and 82 species occur in the Caatinga biome from the state of in Bahia (Mamede et al. 2017).
The first monograph on Malpighiaceae was "Monographie de la famille des Malpighiacées" written by Jussieu (1843), and the second, and last revision was published as part of the "Das Pflanzenreich" by Niedenzu (1928), considered a masterwork to family.

Results and Discussion
Malpighiaceae is represented by 18 genera and 32 species in the southern part (state of BA) of the RCE.The genera represented by the most species were Heteropterys Kunth (five spp.) and Byrsonima Rich.ex Kunth (four spp.).Three new occurrences for the Caatinga domain were recorded for species previously cited only for the phytogeographic dominions of Amazônia, Cerrado and/or Atlantic Forest: Byrsonima Malpighiaceae is positioned within the order Malpighiales along with 41 families (Xi et al. 2012).The monophyly of the family is strongly supported by molecular (Cameron et al. 2001, Davis et al. 2001) and morphological data (Davis & Anderson 2010), with the most conspicous synapomoprhy being its conserved floral morphology in Neotropical lineages due to their mutualism with oil-collecting bees (Anderson, W. 1979a).After major advances in the study of the phylogenetic relationships of the group, Malpighiaceae was divided into 16 informal clades, which does not correspond to any classification system proposed for the family so far (Davis & Anderson 2010).
Despite the existence of studies about Malpighiaceae from the Brazilian semiarid areas, regional studies on this family within the state of Bahia are scarce, especially those that include identification keys and morphological descriptions for the family as a whole (pers.obs.).Given the representation of the Malpighiaceae in the Brazilian flora, this study aimed to elaborate a taxonomic study of this family within the Raso da Catarina Ecoregion (RCE) in the state of Bahia.We present morphological descriptions, an identification key, and comments on distribution, ecology and taxonomy of all studied species.

Study area
The Raso da Catarina Ecoregion (RCE) comprises an area of 30.800 km 2 , being one of the eight Ecoregions recognized for the Caatinga biome in Brazil.In the North-south direction it is narrow and elongated.In the North, West and East it is limited to the southern hinterland depression.The northeastern portion borderswith the Borborema Plateau in the state of Pernambuco, and the southern part borders with hinterlands in the state of the Bahia.The Ecoregion is a sedimentary basin with a very flat relief, showing canyons formed by sandstone outcrops only in the western part.Altitudes above sea level vary from 400 to 600 m in the southern part (state of Bahia) and from 350 to 700 m in the northern part (Jatobá basin, state of Pernambuco).In the southern part (state of Bahia) most of the soils are composed of sand (deep, excessively drained, acid and very low fertility) and oxisol (deep, well drained, acid and low fertility) whereas in the northern part (Pernambuco) sandy soils prevail.Water availability is scarce in the Figure 1.Location of conservation units of the Raso da Catarina Ecoregion, state of Bahia, Brazil (Varjão et al. 2013, modified) coccolobifolia Kunth, Peixotoa hispidula A.Juss. and Tetrapterys paludosa A.Juss.Among the catalogued species 25 are endemic from Brazil.Aspicarpa harleyi W.R. Anderson
Distribution, ecology and phenology: Aspicarpa harleyi is endemic to Brazil and is restricted to the state of Bahia where it is associated with the phytogeographic dominions of Caatinga and Cerrado (Mamede et al. 2017).In the Ecorregion Raso da Catarina, the species was recorded only in the ESRC and on sandy soils in preserved areas.It was collected with flowers and fruits only in March.
Conservation: Due to the degree of endemism, the species was assessed and categorized as "Vulnerable (VU)" according to Biodiversitas (2005), "Threatened with Extinction" (MMA 2008) and "Near Threatened (NT)" by CNCFlora in 2017.
Taxonomy: In the study area it can be recognized by leaves with abaxial surface densely sericeous, silver bright, flowers with pink petals, white in age, and samara with a muricate nut.
Distribution, ecology and phenology: Banisteriopsis stellaris is endemic to Brazil, occurring in the North, Northeast, Midwest and Southeast regions in the phytogeographic dominions of Amazon rainforest, Caatinga and Cerrado (Mamede et al. 2017).In the Ecorregion Raso da Catarina, the species can be found in open, anthropized and preserved areas with sandy or sandy-loam soils.It was collected with flowers during almost the entire year and with fruits in March, June, July August and December.
Taxonomy: It is abundant in the study area and can be differentiated from its congeners by show up entirely glabrous or glabrescent leaves.
Distribution, ecology and phenology: Barnebya harleyi occurs only in the Northeast Region of Brazil in the states of Alagoas, Bahia, Ceará, Paraíba, Pernambuco, Piauí and Sergipe in the Caatinga environments (Mamede et al. 2017).The species in found in preserved areas of the RCE in sandy soils.It was collected with flowers in March, May, July and August and with fruits in May, June, September and November.

Bronwenia ferruginea
Distribution, ecology and phenology: Endemic to Brazil, Bronwenia ferruginea is distributed in the Northeast and Southeast regions in environments of the Caatinga and Atlantic Forest (Mamede et al. 2017).In the RCE, the species was found along a trail in open areas on sandy soils.It was collected only one time in October with flowers and fruits.
Taxonomy: It is recognized by the presence of flowers with androecium consisting of stamens monomorphic, gynoecium with three parallel styles with apex truncate and samara bearing a pair of winglets on the nut.
Specimens examined: BRAZIL, BAHIA: Glória, Serra do Retiro, depois da Estação XIII, 09º20'05.Distribution, ecology and phenology: Bunchosia pernambucana is endemic to Brazil with a distribution restricted to the Northeast Region in the phytogeographic dominions of Caatinga and Atlantic Forest (Mamede et al. 2017).In the RCE, it was found in open and closed preserved areas on sandy soils.It was collected with flowers and fruits in March, April, May, June, July and December and in June, July and November only with fruits.
Conservation: The species was categorized as "Endangered" (EN) in 2013 when collected only in a conservation unit (CNCFlora 2017, Amorim et al. 2013).
Economic potential: The species has pharmacological activity and has medicinal potential, where as the fruits provide food for the fauna (Silva Júnior 2015).
Distribution, ecology and phenology: Byrsonima gardneriana occurs only in Brazil, and is widely distributed in the North Northeast and Southeast regions, associated with the phytogeographic dominions of Amazon rainforest, Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).The species is found in sandy soils in preserved or anthropized areas in the study area.It was collected with flowers and fruits in August and November, and only with flowers in May and August.
Economic potential: In the RCE, Byrsonima gardneriana has nutritional potential, with its fruits being used by the local population, and according to Guilhon-Simplicio & Pereira (2011), it has pharmacological activity.
Taxonomy: It is distinguished from its congeners by a predominantly shrubby habit and flowers with predominantly white petals.
Distribution, ecology and phenology: The species is not endemic in the Brazil..In Brazil, it is widely distributed in the North, Northeast, Midwest to the Southeast regions and is found in the phytogeographical dominions of Amazon rainforest, Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the study area, Byrsonima sericea is found in open environments along trails on sandy soils.It was collected with flowers in January, February, March, June and December and with fruits in January, March and May.
Economic potential: Byrsonima sericea has nutritional potential, the root and bark are used to dye fabrics, and it features ethnopharmacological and pharmacological activity (Guilhon-Simplicio & Pereira 2011).
Taxonomy: It is an easily recognized plant in the RCE by being generally a tree with the abaxial surface of the leaves sericeousferruginous and the petals completely yellow.The species exhibits morphological variation in the presence of elaiophores at the base of the sepals, and there may be individuals who do not have this character.
Distribution, ecology and phenology: The species is found in South America from Colombia and Venezuela to Brazil.In Brazil, Byrsonima verbascifolia occurs in the North, Northeast, Midwest, Southeast and South regions in the dominions of Amazon rainforest, Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the RCE the species is found in open areas along trails or in well-preserved areas with sandy soils.It was collected only with flowers in January and with flowers and fruits in February.
Taxonomy: The main characteristics that distinguish it from its congeners are leaves with the abaxial surface densely tomentose and flowers with yellow-orange petals.
Distribution, ecology and phenology: Carolus chasei is endemic to Brazil and occurs only in the Northeast region in the state of Bahia, in the phytogeographic dominion of Caatinga (Anderson, W. 1993).The species is associated with sandy soils in preserved areas in RCE.It was collected with flowers in February, March and June and with fruits in April, June, July, August and September, with a flowering peak recorded in March in RCE.
Distribution, ecology and phenology: The species occurs in Argentina, Brazil, Bolivia, Paraguay and Peru (Almeida 2013).It is widely distributed in the North, Northeast, Midwest, Southeast and South regions of Brazil in the dominions of Amazon rainforest, Caatinga and Cerrado (Mamede et al. 2017).Diplopterys lutea is associated with sandy soils and sandstone outcrops in preserved or open areas.It was collected with flowers in August and with fruits in May and November.
Taxonomy: The species can be recognized by umbelliform corymbose inflorescences, flowers with petals yellow and posterior petal yellow with red-orange at base, the gynoecium with the anterior style pubescent proximally to the middle, samaras with hairs irritating at the touch and bearing 1-6 lateral winglets on the nut.In the study area, the leaves are persistent or deciduous in periods of flowering and fruiting.
Distribution, ecology and phenology: Galphimia brasiliensis is endemic to Brazil and is restricted to the Northeast Region in the states of Bahia, Paraíba, Pernambuco, Piauí and Sergipe in the phytogeographic dominions of Caatinga, Cerrado and Atlantic Forest (Anderson, C. 2007, Mamede et al. 2017).The species can be found in sandy soils in preserved or anthropized areas such as the borders of trails.It was collected in the study area with flowers and fruits in March, August and September.
Taxonomy: It can be easily recognized in the RCE by subshrub habit, absence of elaiophores at the base of the sepals and petals yellow, often marked with 1 reddish line abaxially.
Specimens examined: BRAZIL, BAHIA: Paulo Afonso, Estação Ecológica Raso da Catarina, Trilha secundária da estrada em direção à base velha do ICMBio, 09º39'39.8"S,38º34'14.7"W,589 m, 10.VI.2014, fr., J.V. Santos 359 (HUNEB).Distribution, ecology and phenology: Heteropterys caducibracteata is endemic to Brazil and is distributed in the Northeast Region in the states of Bahia and Piauí, where it occurs only in the Caatinga (Mamede et al. 2017).In the study area, the species is found on sandy rocky soils in preserved areas.It was collected once when it was bearing fruit in March.
Distribution, ecology and phenology: A species endemic to Brazil, Heteropterys catingarum is distributed in the Northeast region in the phytogeographic dominions of Caatinga (Amorim 2003).In the RCE, the species is associated with sandy soils in anthropized areas of roadsides.It was collected only once when it was with flowers in October.
Distribution, ecology and phenology: According to Mamede et al. ( 2017), the species is endemic to Brazil, being distributed in the Northeast and Midwest regions in the Caatinga and the Cerrado.In the study area, the species is found on sandy soils in preserved areas of the RCE.It was collected in April with flowers and in November with flowers and fruit.

Heteropterys trichanthera
Distribution, ecology and phenology: Heteropterys trichanthera is endemic to Brazil and distributed in the Northeast and Southeast regions where it occurs only in the Caatinga phytogeographic dominion (Mamede et al. 2017).In the study area, the species is associated with sandy soils in preserved or roadside environments with dense vegetation.It was collected in RCE with flowers and fruits in March, May and June.
Distribution, ecology and phenology: Endemic to Brazil, the species occurs in the Northeast, Southeast and Midwest regions in the phytogeographic dominions of Atlantic Forest and Caatinga (Pessoa et al. 2014, Mamede et al. 2017).Heteropterys trigoniifolia is found on sandy soils in preserved areas, or along trails, with dense vegetation.In the study area it was collected with flowers in June, fruits in September and flowers and fruits in July.
Distribution, ecology and phenology: Janusia anisandra is endemic to Brazil and occurs in the Northeast and Southeast regions in the dominions of Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the RCE, the species can be found in preserved areas and in dense vegetation along trails.It was collected only with flowers in May, August and November and with flowers and fruits in June and December.
Distribution, ecology and phenology: The species occurs in Bolivia and Brazil where it is distributed in the North and Southeast regions (Anderson & Davis 2005).It is treated here as a new occurrence for the Caatinga.In the study area, Mascagnia cordifolia is associated with sandy soils of open trails.It was collected only one time with fruits in December.
Taxonomy: It can be recognized by its chartaceous to coriaceous leaves, and samaras with orbicular wing of 2.3-2.6 cm in diameter, sparsely sericeous.
Distribution, ecology and phenology: Mascagnia sepium occurs in Argentina, Brazil, and Paraguay (Niedenzu 1928).In Brazil the species is widely distributed in the North, Northeast, Midwest, Southeast and South regions in the dominions of Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the RCE, the species can be found in anthropized environments on sandy soils.It was collected with flowers and fruits only in December.
Taxonomy: It is differentiated from its congeners by chartaceous leaves and samaras with orbicular wings well developed, ca.6 mm in diameter, densely sericeous.
Distribution, ecology and phenology: Mcvaughia bahiana is endemic to Brazil and has a geographic distribution restricted to the state of Bahia and Piauí, occurring only in the Caatinga biome (Mamede et al. 2017, Amorim pers. comm.).In the RCE, the species can be found on sandy soils in environments with preserved vegetation.It was collected with flowers and fruits only in August.
Taxonomy: Mcvaughia bahiana possesses characters unique among the species cataloged at RCE, and can be recognized by having flowers not entirely expanded lateral-anterior petals, that are cucullate and nested one inside the other, androecium comprising of seven fertile stamens and three staminodes, and nuts slightly twisted with apiculate laterally base.A.Juss., Ann. Sci. Nat., Bot. II, 13: 279. 1840. Iconography: Anderson, C. (1982: 86), Almeida (2013: 131).Figure 5L Scandent shrubs to 2 m tall or lianas.Leaves chartaceous to coriaceous, opposite; large stipules 5-15.5 × 4-8.5 mm, persistent or Distribution, ecology and phenology: According to Mamede et al. (2017), the species is endemic to Brazil, being distributed in the Northeast and Southeast regions in the Atlantic Forest, thus it is treated here as a new occurrence for the Caatinga in the semiarid region of Brazil.Peixotoa hispidula can be found in open areas along trails with remnant vegetation.It was collected in January and December with flowers and June with flowers and fruits.
Distribution, ecology and phenology: The species is endemic to Brazil, and is distributed in the Northeast, Midwest and Southeast regions, occurring in the phytogeographic dominions of Caatinga and Cerrado (Mamede et al. 2017).Ptilochaeta bahiensis is associated with sandy soils and preserved vegetation in two conservation units (EPASB and ESRC).In the study area it was collected only in November with flowers.
Distribution, ecology and phenology: Endemic to Brazil, Stigmaphyllon blanchetii is widely distributed in the Northeast and Southeast regions in the phytogeographic dominions of Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the study area, Stigmaphyllon blanchetii can be found on sandy soils in open areas of trails.It was collected with flowers only in February.
Distribution, ecology and phenology: Tetrapterys paludosa is endemic to Brazil, is distributed in the Northeast and Southeast regions and occurs only in the phytogeographic dominions of Cerrado and Atlantic Forest (Mamede et al. 2017).This species is treated here as a new occurrence for the Caatinga.In the RCE, it can be found on sandy soils in preserved vegetation in the EPASB.It was collected in November and December only with flowers.
Distribution, ecology and phenology: Tetrapterys ramiflora is endemic to Brazil and is distributed in the North, Northeast, Midwest and Southeast regions in the Caatinga, Cerrado and Atlantic Forest (Mamede et al. 2017).In the RCE this species can be found in open vegetation along trails on sandy soils.It was collected only in August bearing flowers.

Figure 2 .
Figure 2. Units of Conservation of the Raso da Catarina Ecoregion, part Bahia, Brazil. A. Environmental Protection Area Serra Branca; B. Biological Station of Canudos; C-D.Ecological Station Raso da Catarina; E. State Park of Canudos; F. RPPN Farm Flor de Lis.Photos by J.V. Santos.