Asteraceae in the northern Espinhaço Range , Brazil : richness , endemism and conservation

Floristic inventories focussing on the Espinhaço Range have revealed an extensive diversity for Asteraceae and emphasize the high degree of endemism of its species. is study aims to explore the species-rich Asteraceae through a oristic survey, and by identifying endemic records for the municipality of Morro do Chapéu, Bahia. Samples were collected during six eld trips in di erent phytophysiognomies and approximately 1,400 exsiccatae from the main herbaria collections were examined. Asteraceae in Morro do Chapéu is represented by 18 tribes, 72 genera and 119 species. Eupatorieae and Vernonieae are the most species rich tribes and currently comprise 30 and 28 species, respectively, representing 50 % of the local Asteraceae ora. Baccharis represents the richest genus with seven species, followed by Lepidaploa and Mikania with six species each. Species belonging to Eupatorieae, namely Acritopappus santosii, Acritopappus sp. nov., Lapidia apicifolia, Stylotrichium edmundoi, Scherya bahiensis, Trichogonia tombadorensis and, additionally, a new species of Vernonieae, Stilpnopappus sp. nov., are endemic to the municipality. We provide a checklist and identi cation key for the species, and report the occurrence of endemism and their importance for the biological conservation.


Introduction
Asteraceae is considered the most species-rich angiosperm family with approximately 24,700 species distributed among 1,600-1,700 genera (Funk et al. 2009).It exhibits a wide range of evolutionary strategies and di erent life forms (herbs, shrubs, subshrubs, trees and lianas), and has a cosmopolitan distribution, although most of its species occur in grassland vegetation in temperate and semi-arid climates of tropical and subtropical zones (Bremer 1994;Funk et al. 2009).
e family is recognized by a set of morphological characteristics that includes owers arranged in a head, introrse anthers connate into a tube, secondary presentation of pollen and a 2-carpellate inferior ovary with a basal and erect ovule which develops in a cypsela usually possessing a pappus (Roque & Bautista 2008;Funk et al. 2009).It is recognized as a monophyletic group, and molecular studies ( Twenty-eight tribes of Asteraceae have been recorded in Brazil, represented by 287 genera and 2,086 species; 66 genera and 1,315 species are endemic and occur in all phytogeographic areas of the country (BFG 2015).e greatest diversity of Asteraceae is in areas of cerrado (savanna), campos rupestres Asteraceae of Morro do Chapéu, Bahia, Brazil: richness, endemism and conservation (rocky elds), campos sulinos (southern grasslands), caatinga and restinga (sand dunes) (Hind & Miranda 2008).Floristic surveys carried out in the Espinhaço Range have shown high species richness for Asteraceae, but particularly for Eupatorieae and Vernonieae, the most speciose tribes, followed by Helianteae and Astereae (Harley & Simmons 1986;Giulietti et al. 1987;Grandi et al. 1988;Pirani et al. 1994;Hind 1995;2003 Roque et al. 2016).ese studies have con rmed that many species are common, yet have distributions restricted to the Espinhaço Range, whereas other species are micro-endemic to only one region, either in the state of Bahia or Minas Gerais (Giulietti et al. 1987; Giulietti & Pirani 1988;Harley 1995).
Approximately 125 genera and 500 species of Asteraceae have been documented in the campos rupestres of Espinhaço Range, of which 10 genera and about 75 species are endemic to Chapada Diamantina in Bahia (Roque et al. 2016).us, the focus of the present study was to perform a oristic inventory of Asteraceae and identify endemic records for the municipality of Morro do Chapéu in Chapada Diamantina, Bahia, Brazil.
Several vegetation types associated with a series of elevational strata comprise the Espinhaço Range, which is considered the center of diversity for many groups of angiosperms and hosts about 15 % of the vascular ora of Brazil (Rapini et al. 2008;Silveira et al. 2015).Some 30 % of the plant species of the Espinhaço Range are endemic to the campos rupestres (Giulietti et al. 1987), a phytophysiognomy composed of herbs and shrubs growing in sandy and acidic soils between large rocky outcrops (Giulietti & Pirani 1988;Harley 1995;Pirani et al. 2003).
Chapada Diamantina belongs to the northern region of the Espinhaço Range, and can be divided into di erent types of natural vegetation, including caatinga, forest, cerrado and campos rupestres, all of which are extremely biologically diverse (Harley 1995).
The municipality of Morro do Chapéu is located in Chapada Diamantina within the semi-arid region of Bahia, and is characterized by tabular reliefs ranging between 480 m and 1,293 m in elevation (Rocha & Costa 1995).e landscape re ects the climatic particularities within the so-called "Polígono das Secas" including mountainous formations with pronounced scarps, deep valleys and high plateaus (Barbosa 1995).Due to the spatial and temporal variation inherent in the pluviometric regime of the region, including its intensity, di erent vegetation formations can develop on the same type of soil.Furthermore, the region is characterized by structurally compartmentalized reliefs, which generate di erent mesoclimates in adjacent areas (Silva 1995).
Morro do Chapéu is classified as an area of high priority for preservation in the Chapada Diamantina due to the fact that it possesses a unique vegetation typology of the caatinga (Maury 2002).According to Queiroz et

Floristic survey
Six field expeditions were carried out between November of 2014 and March of 2016, to survey species of Asteraceae.Fieldwork was schedule taking into account the main seasonal and pluviometric variation in the region.The main herbaria storing floral material from Chapada Diamantina were visited, and included ALCB, CEPEC, HUEFS, HRB, RB, R, SP and SPF (acronyms according to Thiers 2017), and approximately 1,400 exsiccatae were analyzed.
e principal collection sites for species of Asteraceae, and a set of photographs showing the di erent landscapes of the municipality of Morro do Chapéu, are provided in Figure 1. e map provided here was generated with ArcMap (version 10).Terminology for forest phytophysiognomies is based on the classi cation system proposed by IBGE (2012), whereas that for campos rupestres followed Giulietti et al. (1987) and that of caatinga vegetation followed Rizzini (1997).
e collection strategy involved random walking around the main access road and secondary smaller trails.Collected material was geo-referenced and processed according to Peixoto & Maia (2003).All material was deposited in the ALCB herbarium with duplicates being sent to HUEFS.Species identi cation was aided by speci c literature, protologues, comparisons with dried material and virtual collections.Asteraceae of Morro do Chapéu, Bahia, Brazil: richness, endemism and conservation      Asteraceae of Morro do Chapéu, Bahia, Brazil: richness, endemism and conservation

Diversity and phytophysiognomies: specifics regarding Asteraceae
Studies on the ora of Morro do Chapéu reveal that the vegetation found in the northeastern region of the municipality is predominantly composed of arborealshrubby caatinga.Meanwhile, extensive areas of campos rupestres can be found in the central region, whereas semi-deciduous forests are located in the southwest and southeast.Associated with these phytophysiognomies, gallery forests and ecotonal vegetation exhibit complex areas of transition depending on topography and elevation (Silva 1995;França & Melo 2013).
e distribution of Dasyphlyllum diamantinense, included in a basal tribe of Asteraceae, is restricted to the region of Chapada Diamantina (Saavedra et al. 2014), where it can be found in semi-deciduous forest throughout the study area.
e other species of this genus that occur in the region (D.brasiliense and D. sprengelianum) can be found in carrasco and caatinga areas.
Considered an endemic of Bahia, Trixis pruskii occurs predominantly in gallery forest in sandy soils or between rocks.e other species of this genus that occur in the region (T.antimenorrhoea, T. calycina and T. vauthieri) are found in other phytophysiognomies, such as border parts of semi-deciduous seasonal forest, or ecotones between campo rupestre and caatinga vegetation.
Richterago is an endemic genus of Brazil, and although more than half of its species are restricted to Espinhaço Range (Roque & Pirani 2014), only one species, Richterago discoidea, was found to occur in the study area.Moquiniastrum shows two of the six recognized species in the State of Bahia., M. blanchetianum and M. oligocephalum are found in transitional areas between campo rupestre and caatinga at elevations above 1000 m on sandy soils, but without rocky outcrops.
The genera Stilpnopappus and Vernonanthura, represented by four species each, and the genus Eremanthus, represented by two, were the predominant groups of Vernonieae in Morro do Chapéu, where they occurred in transitional areas.
The absence of species of Lychnophora, and the low number of species of Lessingianthus, reveal an unusual pattern compared to other areas in Espinhaço Range, since both genera have their greatest diversity in cerrado and campo rupestre phytophysiognomies (Giulietti et al.According to Fiaschi & Pirani (2009), Lychnophora and Eremanthus are key elements for understanding the origin of the ora endemic to the cerrado.erefore, the lack, or low representation, of, these genera may be associated with the di culty in delimiting areas of cerrado sensu stricto in Morro do Chapéu.e species Acritopappus confertus, Aspilia martii, Calea candolleana, Eremanthus capitatus, Lepidaploa aurea and Lepidaploa lilacina are widely distributed throughout di erent phytophysiognomies of Morro do Chapéu, such as campos rupestres, transition zones between campo rupestre and the caatinga, and the outermost parts of semi-deciduous seasonal forests.
On the other hand, Agrianthus empetrifolius, Cyrtocymura harleyi and Stilpnopappus tomentosus are found predominantly in campo rupestre vegetation where they are restricted to rocky outcrops.Scherya bahiensis, Stilpnopappus sp.nov.and Stylotrichium edmundoi are considered micro-endemics of a speci c area of campos rupestres called the "Tabuleiro do Tigre" (Fig. 1D).Apart from these species, Acritopappus santosii, Lapidia apicifolia and Trichogonia tombadorensis are also identi ed as endemic to Morro do Chapéu.According to Conceição et al. (2005), this microendemism is particularly in uential in campos rupestres where a signi cant part of the diversity is composed of less frequently occurring species.
Among the patterns of geographic distribution proposed for the flora of campos rupestres by Giulietti & Pirani (1988), one specific pattern can be highlighted, that comprising species whose occurrence is restricted to Espinhaço Range.This pattern can be divided into two subgroups.The first group encompasses species whose occurrence is restricted to campos rupestres, but at the same time are widely distributed throughout the Espinhaço Range.The second group encompasses species that are endemic to one specific locality or to only a few regions of the Espinhaço Range located in the state of Minas Gerais, or in the Chapada Diamantina in the state of Bahia.Thus, the endemic species of Morro do Chapéu can be subsumed into the second subgroup.As pointed out by Conceição & Pirani (2005), the contiguity between different habitat types produces a high degree of heterogeneity in floristic structure and composition in certain restricted areas, reflecting variation in substrates.This phenomenon, therefore, contributes to the high number of endemic species of campo rupestre.
According to Zappi (2008), the implementation of an environmental protection area and the development of management plans have to be regarded as extremely relevant in the case of Morro do Chapéu, where the unique enclaves of the caatinga and campo rupestre are endangered.In this sense, further studies on the taxonomy, ecology and biogeography of the representative and endemic groups found in the campos rupestres of Espinhaço Range is considered essential.ese studies can provide crucial information regarding evolutionary patterns that have determined the origin of rare species, and thus can provide insight into conservation.Furthermore, they can provide important criteria for the development of strategies for biodiversity conservation (

Mariana Guerra Staudt, Maria Alves and Nádia Roque
In this sense, it is important to emphasize that Morro do Chapéu possesses unique oristic diversity, and a signi cant number of species of Asteraceae possess restricted distributions and high degrees of endemism.We stress that this oristic diversity is constantly at high risk of becoming endangered due to population reductions and loss of habitat, particularly considering the intense activities involving sand-extraction and growing exploitation for agriculture.

Figure 1 .
Figure 1.Map of the municipality of Morro do Chapéu with the main collection sites of the Asteraceae species and a set of pictures taken from the region's di erent landscapes: A -Gruta dos Brejões; B -Cachoeira do Ferro do Doido; C -Trilha da Guariba; D -Tabuleiro do Tigre; E -Parque Estadual de Morro do Chapéu (Cidade das Pedras); F -Trilha da Veredinha (A-D.Cardoso; B, C, D, F-M.Staudt; E-L.Barres).

In
Morro do Chapéu, Asteraceae is represented by 18 tribes, 72 genera and 119 species (Tab.1).Corroborating previous oristic surveys in Espinhaço Range (Giulietti et al. 1987; Hind 1995; 2003; Zappi et al. 2003; Moura & Roque 2014; Campos et al. 2016; Roque et al. 2016), Eupatorieae and Vernonieae were the most diverse tribes with 30 and 28 species respectively, and together comprise 50 % of the local diversity of Asteraceae.Heliantheae was the third most diverse tribe with 19 species, representing about a third of Heliantheae species registered in the state of Bahia by Alves & Roque (2016).Baccharis was the richest genus with seven species, followed by Lepidaploa and Mikania, with six species each; results similar to those observed in other areas of Chapada Diamantina (Harley 1995; Zappi et al. 2003; Moura & Roque 2014; Roque et al. 2016).Other speciose genera included Acritopappus with ve species; Stilpnopappus, Trixis, and Vernonanthura, with four species each; and Dasyphyllum and Trichogonia with three each.e remaining genera were represented by only one or two species each, and represented 60 % of the species of Asteraceae in the area.Seven genera found at Morro do Chapéu are monotypic (Albertinia, Bahianthus, Conocliniopsis, Lapidia, Prolobus, Scherya and Synedrellopsis), while 23 species are endemic to the state of Bahia, of which 19 belong to the tribes Eupatorieae and Vernonieae (Tab.1).Morro do Chapéu is the type locality for Acritopappus prunifolius, Cyrtocymura harleyi, Lepidaploa tombadorensis and Stilpnopappus semirianus.Moreover, Acritopappus santosii, Acritopappus sp.nov., Lapidia apicifolia, Stylotrichium edmundoi, Scherya bahiensis and Trichogonia tombadorensis, all belonging to the tribe Eupatorieae, and a new species, Stilpnopappus sp.nov. of to the tribe Vernonieae, are recorded as endemic to the municipality (Tab.1).Stilpnopappus suffruticosus is here recorded as a new record for the state of Bahia, and is classified as endangered in the Red Book (Martinelli & Moraes 2013), even as Paralychnophora harleyi, Stilpnopappus semirianus, Stylotrichium corymbosum and Stylotrichium edmundoi.