Areas of endemism in the Espinhaço Range in Minas Gerais, Brazil

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Flora 206 (2011) 782– 791

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Areas of endemism in the Espinhac o Range in Minas Gerais, Brazil

Livia Echternachta,b,c,∗,1, Marcelo Trovóa,1, Caetano T. Oliveirab,2, José Rubens Pirania,1

a Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Rua do Matão 277, 05508-090 – São Paulo, SP, Brazilb Departamento de Botânica, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Av. Antônio Carlos 6627, 31270-901 – Belo Horizonte, MG, Brazilc UMR 7207 CNRS MNHN UPMC, Centre de Recherche en Paléobiodiversité et Paléoenvironnements, MNHN, 57 rue Cuvier, CC48, F-75005 – Paris, France

a r t i c l e i n f o

Article history:Received 15 September 2010Accepted 2 February 2011

Keywords:BiodiversityBiogeographyCerradoConservationEndemic floraPlant distribution

a b s t r a c t

The Espinhac o Range, a mountain chain located in the Brazilian states of Minas Gerais and Bahia, containsone of the richest floras in Brazil, with a high frequency of endemic species. Since 2005 it is designated asUNESCO biosphere reserve and is situated at the joint border of two global hotspots for biodiversity con-servation. Endemic species with congruent occurrence patterns were identified in order to establish areasof higher endemism within the Espinhac o Range. Taxonomic reviews were analyzed in order to identifyendemic taxa and a dataset was elaborated containing 1765 records from 178 endemic species of vascu-lar plants, representing 17 families and including the geographic coordinates for each record. Two mapswere produced showing species richness and collection effort in 15′ quadrats. The congruent occurrenceswere identified and a third map was provided, delimiting 10 candidate areas of endemism for a “par-simony analysis of endemicity” (PAE). One most parsimonious cladogram is then retrieved, evidencingthree major clades corresponding to the northern, central and southern portions of the Espinhac o, in addi-tion to four subclades included into the central clade. We finally identified six major areas of endemism.Furthermore, there is a strong correlation between species richness and collector effort, revealing whichareas are in need of further field inventories.

© 2011 Elsevier GmbH. All rights reserved.

Introduction

Areas of endemism are defined as regions that contain two ormore species that do not occur anywhere else, and which have con-gruent geographic occurrence (García-Barros et al., 2002; Platnick,1991). The identification of such areas provides important informa-tion for biogeography and for studies and practices in biodiversityconservation. Where endemic species are frequent, congruentoccurrence patterns can easily be recognized. These patterns, how-ever, must be delimited in order to propose areas of endemism,to test them and to establish their inter-relationships. Generally,Brazilian biomes exhibit numerous endemic species with congru-ent occurrence. Areas of endemism have already been proposed forthe Amazonian and Atlantic Forest domains considering both faunaand flora elements, but no studies were conducted until now forthe central Brazilian savannas, most commonly known as Cerrado(Fiaschi and Pirani, 2009).

∗ Corresponding author at: Departamento de Botânica, Instituto de Biociências,Universidade de São Paulo, Rua do Matão 277, 05508-090 – São Paulo, SP, Brazil.

E-mail addresses: [email protected] (L. Echternacht), [email protected](M. Trovó), [email protected] (C.T. Oliveira), [email protected] (J.R. Pirani).

1 Tel.: +55 11 3091 7545.2 Tel.: +55 31 3409 2678.

The Espinhac o Range in Minas Gerais (ERMG) is located betweenthe Cerrado and the Atlantic Forest domains, both consideredas biodiversity hotspots (Mittermeier et al., 2005; Myers et al.,2000). The ERMG is also recognized as a Biosphere Reserveby UNESCO (2005). Distributional patterns for some floweringplant families have already been identified in the Espinhac oRange of Minas Gerais (Hensold, 1988; Rapini et al., 2002;Verola, 2008; Versieux and Wendt, 2007). However, there areno available analyses that propose areas of endemism and testtheir relationships using a large set of vascular plant families,and looks for biogeographical patterns for the vegetation as awhole.

The limits of the Espinhac o Range are well established ina geomorphological sense, except for the problem of con-sidering or not the Iron Quadrangle (Quadrilátero Ferrífero)as a southern part of the ERMG. The Iron Quadrangle isindeed a mountain complex located between Ouro Brancoand Belo Horizonte municipalities, and is usually excludedfrom the ERMG by geologists. The geological origins of thetwo regions are different, as the Iron Quadrangle belongsto the Minas Super-group, while the ERMG belongs to theEspinhac o Super-group (Almeida-Abreu, 1995; Gontijo, 2008;Knauer, 2007; Renger et al., 1994). However, these areasexhibit great biological similarities, so that many biologistsconsider them as a single unit (Giulietti and Pirani, 1988;Giulietti et al., 1997; Hensold, 1988; Rapini et al., 2002;

0367-2530/$ – see front matter © 2011 Elsevier GmbH. All rights reserved.doi:10.1016/j.flora.2011.04.003


L. Echternacht et al. / Flora 206 (2011) 782– 791 783

Salino and Almeida, 2008; Vasconcelos, 2009). This biolog-ical similarity has never been tested in a biogeographicalanalysis.

In the taxonomic treatment of Paepalanthus subg. Xeractis (Eri-ocaulaceae), Hensold (1988) sub-divided the ERMG between OuroPreto and Diamantina into four regions mainly based on topog-raphy: (1) the Diamantina Plateau; (2) the north of the Serra doCipó – from Paraúna river up to the north of the Breu mountain;(3) the south of the Serra do Cipó – southward from Breu moun-tain; and (4) the southeast and southwest of the ERMG – mountainsbetween Belo Horizonte and Ouro Preto (Iron Quadrangle). Rapiniet al. (2002) proposed a similar subdivision, based on terrain topog-raphy and on political limits, but considering all parts of the ERMG:(1) North – from Espinosa to Itacambira towns; (2) DiamantinaPlateau – from Itamarandiba to Presidente Kubitschek, includingSerra do Cabral; (3) Serra do Cipó – from Congonhas do Norte toSanta Luzia; and (4) South – from Caeté to Ouro Preto. Despite thesecurrent hypotheses, areas of endemism have not yet been formallyproposed for the ERMG.

The aims of this work are to identify congruent occurrencepatterns of endemic vascular plant species from the ERMG, andto propose areas of endemism. The areas outlined by Hensold(1988) and Rapini et al. (2002) were tested through evaluationof the occurrence of endemic species from several plant fami-lies, in order to evidence common biogeographic patterns for thewhole vegetation. We also intended to verify the effects of col-lector effort on the degree of endemism richness. The results areexpected to answer the following questions: (I) Can the ERMG,including the Iron Quadrangle, be considered as a single biogeo-graphical unit? (II) Is it possible to delimit areas of endemismwithin the ERMG? (III) Which are the hierarchical relationshipsbetween these areas? (IV) Are there strong relationships betweencollector effort and endemism richness? (V) Which are the mostpoorly collected regions in the ERMG? The reconstruction ofareas of endemism is the first step towards the development ofcladistic biogeography, which may test them as evolutionary geo-graphical units. At the same time, the identification of poorlycollected areas in the ERMG could promote future research effortsin order to provide a homogeneous knowledge of the flora in theregion.

Materials and methods

Studied area

This work adopts the sensu lato concept of ERMG (i.e., includingthe Iron Quadrangle) extending across nearly 460 km from north tosouth and 20–100 km from west to east. Elevations range between800 and 1500 m along the ERMG, with some isolated mountaintops reaching over 2000 m. “Campos Rupestres” (a landscape com-posed mainly by grasslands and rocky outcrops) is the vegetationphysiognomy covering most of the landscape. The lower altitu-dinal limits of the “Campos Rupestres” vegetation vary between800 m (Giulietti et al., 1997) and 1000 m (Martinelli, 2007). Thelower altitude of 900 m, as used here, delimits a continuous areathrough the Espinhac o Range and allows to outline the mainlymountainous complex (Rapini et al., 2008). A typical subtropicalseasonality defines the climate, with a dry season during winterand a rainy season during summer. Soils are acid and dystrophic,and derived mainly from quartzitic rocks; but in the southern por-tion the soils are predominantly ferruginous. Together with opengrasslands and rocky outcrops, semi-deciduous Atlantic forests andCerrados (woody savanna) compose most of the vegetation (Ribeiroand Walter, 2008).

Fig. 1. Map showing species occurrence points over the 15′ quadrats, of the endemicspecies in the Espinhac o Range in Minas Gerais, Brazil.


784 L. Echternacht et al. / Flora 206 (2011) 782– 791

Fig. 2. (a) Map showing 15′ quadrats for endemic species richness of the Espinhac o Range in Minas Gerais, Brazil. Letters (A-O) indicate the quadrats discussed in the results.(b). Map showing the 15′ quadrats for collection richness of endemic species of the Espinhac o Range. Letters (a)–(e) indicate quadrats discussed in the results.

Species occurrence dataset

Taxonomic reviews were analyzed in order to identify endemicspecies of vascular plants from the ERMG. The final dataset includes1765 records from 178 species, representing 17 families (Appendix1). Only recent taxonomic reviews with a complete list of ana-lyzed material were included, in order to ensure reliable dataon species identification and occurrence range (Baumgratz, 1997;Braz, 2005; Costa, 2005; Evangelista, 2001; Feres, 2001; Forzza,2001; Hensold, 1988; Kirkbride, 1976; Konno, 2005; Lovo, 2009;Maio, 1996; Marcondes-Fernandes Neto, 1988; Parra, 2000; Rolim,2004; Roque and Pirani, 2001; Roque, 1999; Sakuragui, 1998;Sales, 1993; Salimena, 2000; Sano, 1999; Scalon, 2007; Souza,1998; Sylvestre, 2001; Tissot-Squalli, 1996; Zanin, 2001; Zappi,

1994). All records were included in BRAHMS 6.03 software (Filer,2008) containing species information and geographical coordinateswith minute precision. Missing geographical coordinates were esti-mated from locality description using Google Earth for Mac OS X(2006). The full dataset is available on request.

Areas of endemism and parsimony analysis of endemicity

Records were plotted in a map (Fig. 1) using ArcView GIS 3.2(ESRI, 1999) and then translated into 15′ quadrats maps using DIVA-GIS 1.4 software (Hijmans et al., 2000). The quadrat size was definedin accordance with the minute precision of the geographical coor-dinates. Maps were obtained for species richness (Fig. 2a) and forcollection effort (Fig. 2b). Only quadrats with at least two endemic



Fig. 3. Map showing the areas of endemism of the Espinhac o Range in Minas Gerais, Brazil. The areas were numbered from 1 to 10. The Venn diagrams of sharing specieswere elaborated for the clades obtained by parsimony analysis of endemicity (PAE).

species were used to circumscribe areas of endemism. Pearson’scorrelation index between species richness and collection effortmaps was inferred.

First, the quadrats were tested by a “parsimony analysis ofendemicity” (PAE), but no resolution has been found in the resultingcladogram, probably because they were too small for our datasetand the biogeographic information was lost. Then, we groupedthe quadrats into 10 areas of endemism, so that they should keepthe biogeographic information, reflecting natural geographic units.The quadrats were delimited according to congruent occurrenceof exclusive species and topographical units, which were outlinedusing the 900 m limit for the ERMG. The delimitation included only

the area of occurrence of the species studied, so that several areasare not continuous along the landscape (e.g., area 1, 2, 4, 5 and 8 –locations see Fig. 3). In addition, some areas apparently continuous(e.g., 1 and 2, 6 and 7, 7 and 8) were separated in order to test theirbiogeographic connection. These areas of endemism were used ascandidate areas for the PAE.

A presence/absence matrix for each species in each one of thesecandidate areas (Appendix 1) was elaborated using MacClade 4.06software (Maddison and Maddison, 2003). A hypothetical area withno species was used to root the cladogram. The PAE was per-formed with an exact search using PAUP* 4.10 software (Swofford,2000). The former candidate areas were then tested. The result-



Fig. 4. Relationships between the candidate areas of endemism in theEspinhac o Range in Minas Gerais using parsimony analysis of endemicity (PAE).CMD = Conceic ão do Mato Dentro; DP = Diamantina plateau; NMC = Northern Moun-tains Complex; SCa = Serra do Cabral, SCi = Serra do Cipó, SMC = Southern MountainsComplex.

ing clades were considered as the final macro-regional areas ofendemism. Thereafter, Venn diagrams were prepared, showing theshared species between the areas of endemism.

We should emphasize that our goal was not to establishareas of endemism as historical entities (according to Harold andMoor, 1994, p. 262). We rather propose distributional units, asprimary biogeographical homologies (de Pinna, 1991; Morrone,2001) that may be tested through cladistic biogeography as sec-ondary homologies, when phylogenetic hypotheses for the ERMG’sendemic taxa will be available.

Results

Endemic taxa are found throughout the whole range of theERMG, as seen in Fig. 2a. The quadrats with high numbers ofendemic species are located in the Diamantina Plateau and in theSerra do Cipó. Within the Diamantina Plateau, quadrats F and Cshow 50 and 49 endemic species respectively, followed by quadratsG (29), I (22), E (21), D (18) and H (17). Within the Serra do Cipó,quadrat N had 55 endemic species, followed by quadrats L (24), K(18), J (11) and M (11). Secondary centers of endemism are pre-dicted in the region of Grão-Mogol (quadrat A, with 14 endemicspecies), in the Serra do Cabral (quadrat B, with 10 species), and inthe region of Ouro Preto (quadrats O and P, with 10 and 13 speciesrespectively).

Fig. 2b shows that, except for five quadrats, the ERMG is coveredby relatively homogeneous collection efforts from 2 to 75 recordsper quadrat. The highest record number is observed for the quadratE (451), in the Serra do Cipó region, followed by quadrat D (81). Inthe Diamantina Plateau the most collected regions are representedby quadrats B (258) and C (187). In the northern mountain complex,quadrat A is the most collected and contains 83 records. Speciesrichness (Fig. 2a) and collection effort maps (Fig. 2b) exhibit rela-tively congruent distribution patterns, and this is confirmed by ahigh Pearson’s correlation index (0.87).

Ten candidate areas of endemism are then outlined (Fig. 3).Some of them roughly correspond to the biogeographic units sug-gested by Hensold (1988) and Rapini et al. (2002). The PAE of theseareas resulted in a single most parsimonious tree (Fig. 4) with 242steps and consistency index of 0.74. According to this cladogram,the candidate areas of endemism are regrouped into six biogeo-graphic macro-areas defined below.

Areas 1 and 2 – Northern Mountains Complex (NMC): Theseareas comprise the northern portion of the ERMG. Latitudinal lim-its are given by the municipalities of Monte Azul (S 15◦10′) andItacambira (S 17◦10′). Longitudinal limits are given by the munic-ipalities of Grão-Mogol (W 42◦52′) and Montes Claros (W 43◦50′).Area 3 – Serra do Cabral (SCa): Here, the latitudinal limits aregiven by the municipalities of Francisco Dumont (S 17◦18′) andCorinto (S 18◦11′). Longitudinal limits are given by the Das VelhasRiver (W 44◦30′) and the BR 135 road (W 44◦10′).Areas 4 and 5 – Diamantina Plateau (DP): This region includesthe major continuous area within the ERMG. Latitudinal limitsare given by the municipality of Diamantina (S 17◦50′) and bythe Paraúna River (S 18◦37′). Longitudinal limits are given bythe municipalities of Monjolos (W 44◦04′) and Itamarandiba (W42◦41′).Areas 6 and 7 – Serra do Cipó (SCi): Latitudinal limits are given bythe Paraúna River and the municipality of Jaboticatubas (S 19◦33′).Longitudinal limits are given by the Cipó River (W 43◦59′) and thevalley between the Pico do Breu and the Campo River (W 43◦38′).Area 8 – Conceic ão do Mato Dentro (CMD): It comprises the east-ern slopes of Serra do Cipó and adjacent areas. Latitudinal limitsare given by the municipalities of Conceic ão do Mato Dentro (S19◦00′) and Itabira (S 19◦40). Longitudinal limits are given by thevalley between the Pico do Breu and the Campo River (43◦38′) andthe municipality of Itabira (W 43◦16′).Areas 9 and 10 – Southern Mountains Complex (SMC): It involvesthe region known as the “Iron Quadrangle”. Longitudinal limits aregiven by the municipalities of Belo Horizonte (S 19◦44′) and OuroBranco (S 20◦31′). Latitudinal limits are given by the municipalitiesof Catas Altas (W 43◦23′) and Moeda (44◦01′).

The PAE provided three main clades clustered in a polytomy:the SMC, the NMC and the Central sector. Within the Central sector,CMD was retrieved as sister to the clade [SCa (SCi, DP)]. Accordingto the Venn diagrams depicted in Fig. 3, the NMC and SMC have nospecies in common. Most of the species are restricted to the cen-tral portion of the ERMG, which shares more species with the SMCthan with the NMC. Comparing SCi, DP, and CMD, only four speciesoccur in all of these three areas, most of the species are exclusivefrom the former two areas, and most of the shared species occurin the SCi and DP. Comparing SCi, DP, and SCa, the same patternarises, with seven species shared by the three areas and no speciessimultaneously occurring in the SCi and SCa.

Discussion

The ERMG as an area of endemism

According to the results, the ERMG sensu lato cannot be con-sidered as one area of endemism as a whole, because none of theanalyzed species occurs throughout the entire ERMG. In fact, nospecies are shared between the NMC and the SMC. Actually, thegeological, climatic and biotic conditions are very different betweenthe northern, central and southern areas of the ERMG. The southernsector is more humid, with lower average temperature and higheraverage elevations, and is more strongly influenced by the Atlanticbiome than the northern sector (for a detailed description of theclimatic and biotic influences over the Espinhac o Range, see Alvesand Kolbek, 2010). These conditions gradually change northward,so that many species are shared between the northern and thecentral sectors, or between the central and the southern sectors,supporting the inclusion of the Iron Quadrangle (southern sector)as a biogeographical unit within the ERMG.



Areas of endemism within the ERMG

We succeeded in recognizing several areas of endemism withinthe ERMG, sharing many exclusive species with congruent andhierarchically organized distributions. There are several differencesbetween the three major clades of endemism areas. The NMC isseparated from the central and SMC sectors by lowlands (less than900 m) with contrasting climate and soil conditions, which can rep-resent a barrier for many endemic mountain plants. Lowlands havelong been considered by many authors as putative barriers for dis-persal of many mountain species, including plant species of theEspinhac o Range (e.g., Harley, 1988). Moreover, the NMC is theregion with the lowest number of endemic species, which is likelyrelated to its smaller area and lower average altitude. The SMCalso shows many differences from the central and NMC sectors,as a more complex geological formation with higher ferruginouscontent, and a greater influence from the Atlantic biome, involvingrepresentative species from the coastal mountains (e.g., Calibrachoaspp. – Solanaceae; Cecropia hololeuca Miq. – Urticaceae; Chusqueapinifolia (Nees) Nees – Poaceae; Gaultheria spp. – Ericaceae), whilethe remaining parts of the ERMG exhibit more obvious floristicconnections to the Cerrado and Caatinga biomes. This latitudinalpattern of the influence of the biomes over the ERMG was alsorecovered for Bromeliaceae (Versieux and Wendt, 2007). Thereforeit is obvious that the northern, central and southern ERMG hostdifferent and contrasted environments and floristic compositions.

The CMD area includes four exclusive endemic species withcongruent occurrence, as well as different climate and flora compo-sition compared to its neighbor areas. This may be related to greaterinfluences from the Atlantic biome. It was therefore considered as aseparate area of endemism. Because of the geographical proximityof this area to the SCi, their clustering was expected. However, theCMD likely emerges at the base of the clade, due to the small num-ber of endemic species found in comparison with the remainingareas (see Venn diagram in Fig. 3), which makes the relationshipbetween this region and the rest of ERMG weaker than betweenthe other areas themselves.

The SCa is located between the NMC and the DP; however itappears as sister to the clade formed by DP and SCi, linked there-fore to the central and not to the northern sector. The existence ofan endemism unit formed by the DP and SCi is corroborated, withmany shared endemic species. Actually it can be considered as thelargest continuous area of the Espinhac o ridge, showing a relativeclimatic, geological and biological homogeneity. However, theseboth macro-regions form distinct biogeographical units, with manyunique species. It may be presumed, therefore, that the ParaúnaRiver valley may represent a barrier to a larger geographical occur-rence of species. In addition, the SCi sector has higher moisturedue to a greater influence of the Atlantic forest, while the DP unitis dryer due to a greater influence from the surrounding Cerradobiome. These factors are likely to restrict a wider distribution oflocally endemic taxa. Despite of the common influence from theAtlantic biome observed in the SCi and SMC, a stronger relationshipbetween these two areas has not been found.

The use in the analysis of only recently reviewed taxa, frommonographs containing a complete list of examined specimens,would ensure a high reliability for the results. However, the choiceof the taxa sampled does not exclude the lack of bias in the providedresults. The selected family with the greatest number of endemicsis Eriocaulaceae, which is more diverse in the central portion ofthe range, giving it a biogeographical cohesion. Nevertheless, con-trasted patterns are observed for other taxonomic groups (Rapiniet al., 2008). The pteridophytes, for example, have a richer florain the Iron Quadrangle and the diversity is progressively smallernorthward (Salino and Almeida, 2008). In addition, other very rep-resentative angiosperm families from the ERMG were not included

at all, such as Xyridaceae and Velloziaceae, while Fabaceae andMelastomataceae, with many endemic species in the ERMG, arerepresented by just a few taxa. This is due to the lack of recent tax-onomic reviews with complete lists of analyzed material. We thusprioritized the reliability of the results rather than a more completedataset that could not faithfully represent the endemic flora. Mean-while, as the knowledge of taxonomy and distribution of speciesin the ERMG is increasing, the detection of consistent patterns ofendemism will become more robust.

Species richness and records

Some regions in the ERMG concentrate far more endemicspecies than others, as discussed above. This observation mayhave two causes: (1) there is actually a higher concentration ofendemic species in those regions, thus requiring hypotheses aboutthe historical, biotic and/or abiotic factors that might have pro-moted speciation and might be responsible for the maintenanceof endemic species, or (2) the highest concentration of endemicspecies in such areas is nothing but an artifact of uneven samplingeffort along the Espinhac o Range.

The high correlation between Fig. 2a and b favors the secondhypothesis: the sites that have been more extensively collectedare the richest in species. When investigating the research historyin the ERMG, it is well known that over the past decades sam-pling effort was concentrated in the DP and SCi areas. Selectinga few small regions to be initially studied has resulted in efficientplans for conservation of their biodiversity, through improvementof the creation of conservation units, such as the National Park ofthe Serra do Cipó. However, it brought bias on the observation ofpatterns when studying the diversity throughout the entire ERMG.Currently, the SCi and the DP are much better studied when com-pared to the other ERMG regions, into which the sampling effortmust increase, in order to homogenize the biodiversity knowledgealong the Espinhac o Range and to allow more reliable analyses ofits unique flora. In spite of it, the sampling effort throughout theDP and the SCi is not homogeneous but heavily concentrated inhighways vicinities (Madeira et al., 2008). Some regions inside theDP and the SCi are surprisingly poor collected and increasing thecollector effort in these regions has provided new discoveries ofendemic taxa populations (Echternacht et al., 2010).

The correlation between Fig. 2a and b, however, is not direct. Theless sampled quadrats may have a contrasted richness of endemicspecies, favoring the first hypothesis. A cyclical reasoning can bedeveloped relating collection and species richness. Regions withhigher diversity usually attract more researchers, whose samplingeffort becomes unequal in relation to poorer regions. For example,in the Asclepiadoideae (Apocynaceae), the areas with the great-est number of records are located in the SMC, which is preciselythe region of greatest species richness (Rapini et al., 2002). Nev-ertheless, the SCi and DP are still the richest regions in endemicspecies. This suggests that these both areas may be more favorableto endemism than other areas in the ERMG. Another example thatreinforces the first hypothesis is the region of Grão-Mogol. Despitethe study of the flora at an advanced stage (Pirani et al., 2003), theregion exhibits a lower level of endemism in comparison with theSCi and the DP. The hotspot concept itself relates to regions in theglobe with a greater endemic biota (Myers et al., 2000; Mittermeieret al., 2005). We consider also the possibility of detecting evenpatterns in a much more local scale within the ERMG.

The richness of endemic species is likely higher than reportedin under-sampled areas, and differences on the variations betweenthe quadrats of Fig. 2a and b could be not so great. We pro-pose nevertheless that there are some areas more favorable to thedevelopment of endemic species than others. The causes for thesedifferences may be historical (meaning biogeographical processes



taking place in ancient times, like major past climatic changes,vicariance and dispersal events), or ecological (abiotic and bioticpresent interactions, such as greater complexity of micro-habitatsdue to higher altitudinal ranges, climate and soil).

The ERMG hosts a mosaic of phytophysiognomies (Alves andKolbek, 2010) that favors biological diversity (Rapini et al., 2008)and the development of micro-endemism. The “Campos Rupestres”are constituted by several habitats with different substrates, floris-tic compositions, presence of rocky outcrops or sandy sediments,dry or periodically humid soils, affecting the biotic communitiesin small ranges (Conceic ão and Pirani, 2005, 2007). Some stud-ies with an ecological approach performed in Espinhac o sites atthe Chapada Diamantina (Bahia state) demonstrated that micro-endemic species are highly frequent, and that floristic similarity isnot directly correlated with distance, indicating that other factorsmay influence distributional patterns (Conceic ão et al., 2007). Nev-ertheless, in a larger scale study, the Eupatorieae tribe (Asteraceae),which also exhibits a rich micro-endemic flora at high altitudinalfields, showed that floristic similarity among the ERMG’s locali-ties is influenced by geographical distance (Almeida et al., 2004).However there is no study that explicitly correlates habitat com-plexity with species richness or with micro-endemism richness inthe region.

Implications for biodiversity conservation

The ERMG deserves special attention for biodiversity conser-vation. The delimitation of several areas of endemism within theEspinhac o Range suggests that small areas are specially involvedcontributing to its entire biodiversity. The biological richness of aregion can contribute to the diversity of its neighboring regions.Thus, to maximize the conservation of plant diversity in the ERMG,several areas must be selected along its entire extension.

The few conservation units already created in the ERMG arenot enough to protect the diversity of the entire mountain range,and more parks and reserves must be created (Silva et al., 2008;Versieux and Wendt, 2007). In addition, the existing conserva-tion units lack the ideal governmental support, as most of themdo not have their territories yet expropriated, and/or lack admin-istrative and physical infrastructure. The entire ERMG has greateconomic importance to the mining industry, particularly the SMC,which provides most of the iron ore produced in Brazil. However,

the conservation efforts in these regions of mining interest arestill insufficient (Jacobi and do Carmo, 2008). Most ERMG endemicspecies are endangered (Mendonc a and Lins, 2000), which furtherhighlights the need for major efforts to conserve this region. Finally,the conservation units that already exist in the ERMG must beeffectively established and new protected areas must be created,so that all the Espinhac o Range and its biodiversity can becomesignificantly protected.

This work represents an advance to knowledge about areas ofendemism within the ERMG. We believe that further more detailedstudies are needed to achieve a better understanding of vegeta-tion history and the current distribution of endemic species alongthe Espinhac o Range. Increasing the dataset with a greater num-ber of species and more accurately geo-referentiated records mustrefine the result scale, enabling the detection of local patterns anddefining more precisely the areas of endemism and priority areasfor conservation. Inclusion of faunistic data can also provide fur-ther corroboration and resolution for the areas of endemism withinthe ERMG. The selected method reinforces the importance of pub-lishing complete lists of examined material in taxonomic reviews,or making them available to scientists and authorities, in orderto improve knowledge about the representativeness of taxon col-lections. In addition, a greater collection effort must be appliedin those less collected regions, in order to produce a homoge-neous sampling effort of the flora, so that results can improve theirreliability.

Acknowledgements

We would like to thank to the researchers that have worked andstill work in the ERMG, especially the taxonomists who drew up thetaxonomic reviews from which this work was prepared; to PedroFiaschi for improvements on an earlier version of the manuscript; toMarco Túlio Ferreira for the English revision. Financial support wasprovided by Fundac ão de Amparo à Pesquisa do Estado de São Paulo(FAPESP), by Coordenac ão de Aperfeic oamento de Pessoal de NívelSuperior (CAPES) and by Conselho Nacional de DesenvolvimentoCientífico e Tecnológico (CNPq).

Appendix A. Presence/absence matrix of the endemicspecies in each one of the 10 candidate areas of endemismin the Espinhac o Range in Minas Gerais used in theparsimony analysis of endemicity

Species 1 2 3 4 5 6 7 8 9 10

Actinocephalus actinocephaloides (Silveira) Costa 0 0 0 0 1 0 0 0 0 0Actinocephalus aggregatus Costa 0 0 0 0 0 1 1 0 0 0Actinocephalus arenicola (Silveira) Costa 0 0 0 1 0 0 0 1 0 0Actinocephalus barbiger (Silveira) Costa 0 0 0 0 0 0 1 0 0 0Actinocephalus brachypus (Bong.) Sano 0 0 1 1 0 0 0 0 0 0Actinocephalus cabralensis (Silveira) Sano 0 0 1 0 0 0 0 0 0 0Actinocephalus callophyllus (Silveira) Sano 0 0 0 0 1 0 0 0 0 0Actinocephalus ciliatus (Silveira) Sano 0 0 0 1 1 0 0 0 0 0Actinocephalus cipoensis (Silveira) Sano 0 0 0 0 0 0 1 0 0 0Actinocephalus compactus (Gardn.) Sano 0 0 0 1 0 0 0 0 0 0Actinocephalus coutoensis (Silveira) Sano 0 0 0 0 1 0 0 0 0 0Actinocephalus deflexus Costa 0 0 0 0 0 1 0 0 0 0Actinocephalus diffusus (Silveira) Sano 0 0 0 0 0 0 0 1 0 0Actinocephalus fimbriatus (Silveira) Sano 0 0 0 0 1 0 0 0 0 0Actinocephalus geniculatus (Bong.) Costa 0 0 0 1 1 1 1 0 0 1Actinocephalus glabrescens (Silveira) Sano 0 0 0 0 1 0 0 0 0 0Actinocephalus glareosus (Bong.) Costa 0 0 0 1 1 1 1 0 0 0Actinocephalus graminifolius Costa 0 0 0 0 0 0 0 1 0 0Actinocephalus heteropus (Silveira) Costa 0 0 0 0 1 0 0 0 0 0Actinocephalus heterotrichus (Silveira) Sano 0 0 0 0 1 0 1 0 0 0Actinocephalus incanus (Bong.) Costa 0 0 0 1 1 0 0 0 0 0Actinocephalus nodifer (Silveira) Sano 0 0 0 0 1 0 0 0 0 0Actinocephalus perbrancchiatus (Silveira) Costa 0 0 0 1 0 0 0 0 0 0Actinocephalus rhizomatosus (Silveira) Costa 0 0 0 1 0 0 0 0 0 0



Appendix A (Continued )Species 1 2 3 4 5 6 7 8 9 10

Actinocephalus rigidus (Bong.) Sano 1 1 0 1 0 0 0 0 0 0Actinocephalus robustus (Silveira) Sano 0 0 0 0 0 1 1 1 0 0Actinocephalus scytophyllus (Ruhland) Costa 0 0 0 0 0 1 0 0 0 0Actinocephalus stereophyllus (Ruhland) Sano 0 0 1 1 0 0 0 0 0 0Actinocephalus trichopeplus (Silveira) Costa 0 0 0 0 0 0 1 0 0 0Actinocephalus variabilis (Silveira) Costa 0 0 1 1 1 1 1 0 0 0Actinocephalus velutinus (Silveira) Costa 0 0 1 0 0 0 0 0 0 0Andropogon campestris Trin. 0 0 0 0 0 0 1 0 0 0Aspidosperma dispermum Muell. Arg. 1 0 1 1 1 0 1 0 0 0Asplenium schwackei Christ 0 0 0 0 0 0 0 0 0 1Byrsonima macrophylla (Pers.) W. R. Anderson 0 0 0 1 0 0 1 0 0 1Ceradenia warmingii (C. Chr.) Labiack 0 0 0 0 0 0 0 0 0 1Comanthera aciphylla (Bong.) L.R.Parra and Giul. 0 0 0 0 0 0 1 1 0 0Comanthera brasiliana (Giul.) L.R.Parra and Giul. 0 1 0 1 1 1 1 0 0 0Comanthera dealbata (Silveira) L.R.Parra and Giul. 0 0 0 0 1 0 0 0 0 0Comanthera elegans (Bong.) L.R.Parra and Giul. 1 0 1 1 0 0 0 0 0 0Comanthera magnifica (Giul.) L.R.Parra and Giul. 0 0 1 1 1 0 0 1 0 0Comanthera nitida (Bong.) L.R.Parra and Giul. 0 0 0 0 1 0 0 0 0 0Comanthera suberosa (Giul.) L.R.Parra and Giul. 0 0 1 1 0 0 1 1 0 0Declieuxia cacuminis Müller 1 0 0 0 1 0 0 0 0 0Declieuxia diamantinae Kirkbride 0 0 0 1 0 0 0 0 0 0Declieuxia diantheroides Standley 0 0 0 1 0 0 1 0 0 0Declieuxia gracilis Kirkbride 0 0 0 0 0 0 1 0 0 0Declieuxia hatschbachii Kirkbride 0 0 0 1 0 0 0 0 0 0Declieuxia irwinii Kirkbride 0 0 0 0 0 0 1 0 0 0Declieuxia juniperina Saint-Hilaire 0 0 0 1 1 0 0 0 0 0Declieuxia leiophylla Müller 0 0 0 0 0 0 1 0 0 0Declieuxia muscosa Saint-Hilaire 0 0 0 1 0 0 0 0 0 0Declieuxia passerina Martius and Zuccarini 0 1 0 1 0 0 1 0 0 0Declieuxia spergulifolia Martius and Zuccarini 0 0 0 1 0 0 0 0 0 1Ditassa aequicymosa E. Fourn. 0 0 0 1 0 0 1 0 1 1Ditassa cipoensis (Fontella) Rapini 0 0 0 0 0 1 1 0 0 0Ditassa cordeiroana Fontella 1 1 0 0 0 0 0 0 0 0Ditassa fasciculata E. Fourn. 0 1 1 1 1 0 1 0 0 0Ditassa itambensis Rapini 0 0 0 0 1 0 0 0 0 0Ditassa longicaulis (E.Fourn.) Rapini 0 0 0 1 0 0 0 0 0 0Ditassa longisepala (Hua) Fontella and E. A. Schwarz 0 0 0 0 0 0 0 0 1 1Encholirium biflorum (Mez) Forzza 0 0 0 1 0 0 1 0 0 0Encholirium heloisae (L.B.Sm.) Forzza and Wanderley 0 0 0 0 0 0 1 0 0 0Encholirium irwinii L.B.Sm. 1 0 0 0 0 0 0 0 0 0Encholirium magalhaesii L.B.Sm. 0 0 0 1 1 0 0 0 0 0Encholirium pedicellatum (Mez) Rauh 0 0 0 1 0 0 0 0 0 0Encholirium reflexum Forzza and Wanderley 1 0 0 1 0 0 0 0 0 0Encholirium structor (L.B.Sm.) Rauh 0 0 0 1 0 0 0 0 0 0Encholirium vogelii Rauh 0 0 0 0 0 0 0 1 0 0Hindsia irwinii Steyermark 1 0 0 0 0 0 0 0 0 0Huberia piranii Baumgratz 0 0 0 1 0 0 0 0 0 0Lippia bradei Moldenke 1 1 0 0 0 0 0 0 0 0Lippia elliptica Schauer 0 0 1 0 0 0 0 0 0 0Lippia florida Cham. 0 0 0 0 0 0 1 0 1 1Lippia pseudothea (A. St.-Hil.) Schauer 0 0 1 1 1 0 0 1 0 0Lippia rhodocnemis Mart. and Schauer 0 0 0 0 1 0 0 0 0 0Luxemburgia angustifolia Planch 0 0 0 1 0 0 1 0 0 0Luxemburgia bracteata Dwyer 0 0 0 0 1 0 0 0 0 1Luxemburgia ciliatibracteata Sastre 0 0 0 0 0 0 1 0 1 0Luxemburgia ciliosa (Mart.) Gardner 0 0 0 1 1 0 1 0 0 0Luxemburgia corymbosa A.St.-Hil., 0 0 0 0 0 0 0 0 0 1Luxemburgia damazioana Beauverd 0 0 0 0 0 0 1 1 1 0Luxemburgia flexuosa Sastre 0 0 0 1 1 0 0 0 0 0Luxemburgia nobilis Eichler 0 0 0 0 0 0 0 0 0 1Luxemburgia polyandra A.St.-Hil. 0 0 0 1 1 1 1 0 0 0Luxemburgia schwackeana Taub. 0 0 0 1 0 0 1 1 1 1Luxemburgia speciosa A.St.-Hil. 0 0 0 1 1 0 0 0 0 0Mandevilla densiflora (Pohl ex Malme) M. F. Sales 1 1 1 1 1 0 1 0 0 0Mandevilla rubra MGF ex M. F. Sales 0 0 0 1 0 0 0 0 0 0Mandevilla semirii M. F. Sales 1 1 0 0 0 0 0 0 0 0Micropolypodium perpusillum (Maxon) A. R. Sm. 0 0 0 0 0 0 0 0 0 1Minara bifurcata (Rapini) T.U.P.Konno and Rapini 0 0 0 1 0 0 0 0 0 0Minara diamantensis (Fontella) T.U.P.Konno and Rapini 0 0 0 1 0 0 0 0 0 0Minara grazielae (Fontella and Marquete) T.U.P.Konno and Rapini 0 0 0 1 0 0 0 0 0 0Minara hemipogonoides (E.Fourn.) T.U.P.Konno and Rapini 0 0 0 0 0 0 1 0 0 0Minara magisteriana (Rapini) T.U.P.Konno and Rapini 0 0 0 0 0 0 1 0 0 0Minara parva (Silveira) T.U.P.Konno and Rapini 0 0 0 1 0 1 1 0 0 0Minara polygaloides (Silveira) T.U.P.Konno and Rapini 0 0 0 0 0 1 1 0 0 0Minara refractifolia (K.Schum.) T.U.P.Konno and Rapini 0 0 0 1 1 0 0 0 0 0Minara semirii (Fontella) T.U.P.Konno and Rapini 0 0 0 0 0 0 1 0 0 0Ossaea cinnamomifolia (Nandin) Triana 0 0 0 0 0 0 1 0 1 1



Appendix A (Continued )Species 1 2 3 4 5 6 7 8 9 10

Ossaea cogniauxii Del Rey Souza 0 0 0 0 0 0 0 0 0 1Ossaea coriacea (Nandin) Triana 0 0 0 0 0 0 0 0 1 1Paepalanthus anamariae Hensold 0 0 0 0 0 1 0 0 0 0Paepalanthus argenteus (Bong.) Koern. 0 0 0 1 1 1 1 0 0 0Paepalanthus ascendens Silveira 0 0 0 1 0 0 0 0 0 0Paepalanthus ater Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus augustus Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus aureus Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus bromelioides Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus calvulus (Ruhland) Hensold 0 0 0 0 0 1 0 0 0 0Paepalanthus chlorocephalus Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus chrysolepis Silveira 0 0 0 1 0 0 0 0 0 0Paepalanthus comans Silveira 0 0 0 1 0 0 0 0 0 0Paepalanthus conduplicatus Silveira 0 0 0 0 0 0 0 0 1 1Paepalanthus crinitus Tissot-Squalli 0 0 0 1 0 0 0 0 0 0Paepalanthus dianthoides Mart. ex Koern. 0 0 0 0 0 0 0 0 0 1Paepalanthus digitiformis Hensold 0 0 0 0 0 1 0 0 0 0Paepalanthus elatus (Bong.) Koern. 0 0 0 0 0 1 0 0 0 0Paepalanthus garimpensis Silveira 0 0 0 0 0 0 0 0 0 1Paepalanthus globulifer Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus homomallus (Bong.) Mart. ex Koern. 0 0 0 0 0 1 0 0 0 0Paepalanthus hydra Ruhland 0 0 0 0 0 0 0 0 1 1Paepalanthus langsdorffii (Bong.) Koern. 0 0 0 0 0 0 0 0 0 1Paepalanthus latifolius Koern. 0 0 0 1 0 0 0 0 0 0Paepalanthus latipes Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus lepidus Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus moedensis Silveira 0 0 0 0 0 0 0 0 1 0Paepalanthus mollis Kunth 0 0 0 0 1 0 0 0 0 1Paepalanthus nigrescens Silveira 0 0 0 1 0 0 1 1 0 0Paepalanthus parviflorus (Hensold) Hensold 0 0 0 0 0 0 1 1 0 0Paepalanthus plumosus (Bong.) Koern. 0 0 0 0 0 1 0 0 0 0Paepalanthus revolutus Hensold 0 0 0 0 0 1 0 0 0 0Paepalanthus scicaefolius Silveira 0 0 0 0 0 0 1 0 0 0Paepalanthus senaeanus Ruhland 0 0 0 0 0 0 1 1 0 0Paepalanthus spixianus Mart. 0 0 0 0 0 0 0 0 1 1Paepalanthus superbus Ruhland 0 0 0 0 0 1 0 0 0 0Paepalanthus trichopetalum Koern 0 0 0 1 1 0 0 0 0 0Paepalanthus uncinatus Gardner 0 0 0 0 0 0 0 1 0 0Paepalanthus vellozioides Koern 0 0 0 0 0 0 1 0 1 1Paepalanthus villosulus Koern 0 0 0 1 0 0 0 0 0 0Paepalanthus xiphophyllus Ruhl. 0 0 0 0 0 0 0 0 0 1Philodendron biribiriense Sakuragui and Mayo 0 0 0 1 0 0 0 0 0 0Philodendron cipoense Sakuragui and Mayo 0 0 0 1 0 0 1 0 0 0Philodendron rhizomatosum Sakuragui and Mayo 0 0 0 0 0 0 1 0 0 0Pilosocereus coerulescens (Lem.) Ritter 0 1 1 1 1 0 1 1 0 0Pilosocereus fulvilanatus (Buin. and Brederoo) Ritter 1 0 0 0 0 0 0 0 0 0Pilosocereus rosae P.J. Braun 0 0 1 0 0 0 0 0 0 0Pseudotrimezia barretoi R.C. Foster 0 0 0 1 0 0 0 0 0 0Pseudotrimezia brevistaminea Chukr 0 0 0 0 0 0 1 0 0 0Pseudotrimezia cipoana Ravenna 1 1 0 1 0 0 1 0 0 0Pseudotrimezia concava Ravenna 1 0 0 0 0 0 0 0 0 0Pseudotrimezia diamantinensis Ravenna 0 0 0 1 1 0 0 0 0 0Pseudotrimezia elegans Ravenna 0 0 0 0 1 0 0 0 0 0Pseudotrimezia fulva Ravenna 0 0 0 1 0 0 0 0 0 0Pseudotrimezia gracilis Chukr 0 0 0 0 0 0 1 0 0 0Pseudotrimezia laevis Ravenna 0 0 1 1 1 0 0 0 0 0Pseudotrimezia monticola (Klatt) Ravenna 0 0 0 0 1 0 0 0 0 0Pseudotrimezia pauli Chukr 0 0 0 1 1 0 0 0 0 0Pseudotrimezia planifolia Ravenna 0 0 0 1 0 0 0 0 0 0Pseudotrimezia pumila Ravenna 0 0 0 1 0 0 0 0 0 0Pseudotrimezia recurvata Ravenna 0 0 0 1 1 0 0 0 0 0Pseudotrimezia sublateralis Ravenna 0 0 1 1 0 0 0 0 0 0Pseudotrimezia synandra Ravenna 0 0 0 1 0 0 0 0 0 0Pseudotrimezia tenuissima 1 0 0 1 1 0 0 0 0 0Richterago angustifolia 0 0 1 1 0 0 1 0 0 0Richterago arenaria (Baker) Roque 0 0 1 1 1 0 1 0 0 0Richterago caulescens Roque 0 0 0 1 1 0 1 0 0 0Richterago conduplicata Roque 0 0 0 0 0 0 1 0 0 0Richterago elegans Roque 0 0 0 0 0 0 1 0 0 0Richterago hatschbachii (Zardini) Roque 0 0 0 1 0 1 0 0 0 0Richterago lanata Roque 0 0 0 0 0 0 1 0 0 0Richterago polyphylla (Baker) Roque 0 0 0 0 0 1 1 0 0 0Richterago stenophylla (Cabrera) Roque 0 0 0 1 1 0 0 0 0 0Staurogyne elegans (Ness) Kuntze 0 0 0 0 0 0 1 0 0 0Staurogyne hirsuta (Ness) Kuntze 0 0 0 0 0 0 1 0 0 0Staurogyne vauthieriana (Ness) Kuntze 0 0 0 0 0 0 1 0 1 1Stryphnodendron gracile Heringer and Rizzini 0 0 0 0 0 0 0 0 0 1



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Areas of endemism in the Espinhaço Range in Minas Gerais, Brazil

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