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Rodriguésia

On-line version ISSN 2175-7860

Rodriguésia vol.66 no.1 Rio de Janeiro Jan./Mar. 2015

https://doi.org/10.1590/2175-7860201566104 

Original Papers

Floristic surveys of Restinga Forests in southern Bahia, Brazil, reveal the effects of geography on community composition

Levantamento florístico de florestas de Restinga do Sul da Bahia, Brasil, revela o efeito da geografia na composição das comunidades

Moabe Ferreira Fernandes1  3 

Luciano Paganucci de Queiroz2 

1Universidade Estadual de Feira de Santana, Programa de Pós-Graduação em Botânica, 44036-900, Feira de Santana, BA, Brazil

2Universidade Estadual de Feira de Santana, Depto. Ciências Biológicas, 44036-900, Feira de Santana, BA, Brazil.

2175-7860-rod-66-01-0051-suppl01.pdf


ABSTRACT

The Restinga forests of southern Bahia state, Brazil, grow on sandy coastal Quaternary sediments. As their floras are relatively poorly known, the present study assessed their floristic compositions. We surveyed four sites at Maraú and Itacaré and identified 302 angiosperm species belonging to 184 genera of 75 families. The most species rich families were: Fabaceae (35 species), Myrtaceae (25), Rubiaceae (21), Sapotaceae (13), Bromeliaceae (12), Annonaceae (11), Erythroxylaceae (10), Melastomataceae (9), and Apocynaceae (8). Local floras include elements with distributions restricted to the Atlantic Forest domain, those disjunct between the Amazon and Atlantic Forest domains, and those also occurring in moist forests and dry vegetation of central Brazil. The hypothesis that the floristic compositions of restinga forests are influenced by neighboring wet forests was tested using cluster and principal component analyses of eleven restinga forests and nine Atlantic wet forest sites. The results supported five main groups, with most of them including both restinga forests and their adjacent wet forest sites, thus corroborating the hypothesis that wet forests in geographical proximity greatly influence the floristic compositions of restinga forests.

Key words: coastal vegetation; Atlantic Forest domain; flora; similarity

RESUMO

As florestas de Restinga do baixo-sul da Bahia, Brasil, encontram-se sobre sedimentos arenosos do Quaternário costeiro. Como sua flora é relativamente pouco conhecida, o presente estudo avaliou sua composição florística. Foram inventariadas quatro áreas nos municípios de Maraú e Itacaré e identificadas 302 espécies de angiospermas, distribuídas em 184 gêneros e 75 famílias. As famílias mais ricas em espécies foram: Fabaceae (35 espécies), Myrtaceae (25), Rubiaceae (21), Sapotaceae (13), Bromeliaceae (12), Annonaceae (11), Erythroxylaceae (10), Melastomataceae (9) e Apocynaceae (8). A flora local inclui elementos de distribuição restrita à Mata Atlântica, disjunta entre Amazônia e Mata Atlântica e florestas úmidas e a vegetação seca do Brasil central. A hipótese de que a composição florística das florestas de Restinga é influenciada pelas florestas pluviais geograficamente próximas foi testada usando análises de agrupamento e de componentes principais com onze áreas de florestas de Restinga e nove de florestas pluviais da Mata Altântica. Os resultados sustentaram cinco grupos principais, a maioria incluindo áreas de floresta de restinga e florestas pluviais adjacentes, corroborando a hipótese de que a proximidade geográfica aos estoques florísticos das florestas pluviais tem grande efeito na composição das florestas de Restinga.

Palavras-Chave: vegetação costeira; Domínio Mata Atlântica; flora; similaridade

Introduction

The word “Restinga” designates vegetation established on sandy Quaternary substrates subject to marine or fluvial-marine influences (Flexor et al. 1984; Sugiyama 1998). This definition encompasses a variety of plant communities that can vary greatly in terms of their physiognomic, floristic, and structural features (Assis et al. 2004). These vegetation complexes are organized along a sea-to-continent gradient, with positive correlations between species richness and size (Araújo 2000).

Restinga vegetation has great ecological value, serving as shelter for plant populations that are rare in other forest types in the Atlantic Forest domain (hereinafter Atlantic domain; Scarano 2009) and for endangered animals (Rocha et al. 2005) - and are used by traditional coastal communities for food, medicines, and raw materials for craft work (Menezes et al. 2009). These vegetation forms are also responsible for the maintenance of water resources (Dorneles & Weachter 2004; Scherer et al. 2005).

Despite their great social and ecological importance, Restingas are some of the least-known vegetation types in Brazil in terms of their diversity and conservation status (Rocha et al. 2005), and basic information about areas occupied by Restinga vegetation fragments is lacking at both regional and continental scales (Rocha et al. 2007).

Studies encompassing Restinga vegetation are still incipient and have been unequally distributed across Brazil’s geopolitical regions. Southern and southeastern Restinga sites have been more intensively studied (Lacerda et al. 1984; Araújo 2000; Martins et al. 2008; Lima et al. 2011), although Guedes et al. (2006) highlighted our lack of knowledge about the floristic and phytosociological structures of restinga vegetation in São Paulo State. Most of the works produced so far have been limited to floristic and vegetational aspects, with little emphasis on environmental patterns and processes. Bahia State has the longest coastline in the country (ca. 1200 km), but studies there have been scarce and concentrated mostly along its northern shore (e.g., Britto et al. 1993; Queiroz 2007; Menezes et al. 2009; Queiroz et al. 2012).

Restingas are highly diverse habitats, although they generally harbor fewer species than other forest types within the Atlantic domain (Almeida Jr. et al 2009; Lima et al. 2011). Restinga vegetation is normally established on geologically recent sediments originating from deposition by marine regressive movements during the Pleistocene and Holocene periods. The young ages of these habitats have not provided sufficient time for speciation to occur – a factor often cited as the principal explanation for their scarcity of endemic taxa (Scarano 2002; Castro et al. 2012). These communities are therefore composed mostly of elements from other vegetation types – with the Atlantic domain being considered the main source of their species (generally more than 50% of the total numbers of species) (Assis et al. 2004). However, there are also reports of contributions from Cerrado (Neotropical Savanna), Caatinga (Seasonally Dry Tropical Forests and Woodlands), and Amazonia floras (Cerqueira 2000; Sacramento et al. 2007; Castro et al. 2012).

Taking into consideration the young geological age and scarcity of endemism in Restinga vegetation, we hypothesize that the flora of Restinga forests is composed mostly of elements that have migrated from neighboring wet forest areas rather than arising from in situ speciation. This hypothesis can be tested by demonstrating that the floras of Restinga forests are more similar to those of geographically close rain forests than to other (more remote) Restinga forest sites, and we present here comparative analyses to that end. We also report the results of floristic studies of Restinga forests from southern Bahia that can help fill gaps in our knowledge of the floristic compositions of Restinga forests in an important center of diversity in the Atlantic domain.

Materials and methods

Study areas

The survey areas encompassed four sites in southern Bahia State, Brazil (Fig. 1), in the municipalities of Itacaré (Mata do Santo Amaro - MAS) and Maraú (Mata do Caubi - MC; Mata da Estrada Maraú-Itacaré – MMI; and Mata da Piracanga - MP; Tab. 1). The region is part of the Atlantic domain, with a mean annual precipitation between 1,200 and 2,400 mm, with the rainy season extending from March to July, without a dry period; the mean annual temperature is approximately 25 °C, with small oscillations between the minimum and maximum mean temperatures of 20 and 26 °C - an Af climate type according to the Köppen (1948) system (C.E.I. 1993).

Table 1 Restinga and wet forests in evaluated in the multivariate analyses, with their acronyms, locations, coordinates, and vegetation typesTabela 1 - Áreas de florestas de restinga e florestas pluviais usadas na análise multivariadas, com sigla, localização, coordenadas e tipos vegetacionais. 

Sigla Município Estado Latitude Longitude Formação Referência
ResSGA(CE) São Gonçalo do Amarante CE -3.606944444 -38.87027778 Restinga Castro et al. (2012)
ResMara(PB) Mataraca PB -6.486111111 -34.94166667 Restinga Oliveira-Filho & Carvalho (1993)
OmbSVF(PE) São Vicente Férrer PE -7.633333333 -35.5 Ombrófila Montana Ferraz & Rodal (2008)
ResCSA(PE) Cabo de Santo Agostinho PE -8.125 -35.01527778 Restinga Sacramento et al. (2007)
ResTam(PE) Tamandaré PE -8.788888889 -35.1125 Restinga Silva et al. (2008)
ResCon(BA) Conde BA -11.81277778 -37.64027778 Restinga Menezes et al. (2009)
OmbSTe(BA) Santa Terezinha BA -12.85 -39.96666667 Ombrófila Submontana Carvalho-Sobrinho & Queiroz (2005), Neves (2005)
ResMaIt(BA) Maraú-Itacaré BA -14.21694444 -38.99833333 Restinga Este estudo
OmbBP(BA) Barro Preto BA -14.76666667 -39.53333333 Ombrófila Montana Amorim et al.(2009)
OmbUna(BA) Una BA -15.16666667 -39.05 Ombrófila Tabuleiro Amorim et al.(2008)
OmbAra(BA) Arataca BA -15.16666667 -39.33333333 Ombrófila Montana Amorim et al.(2009)
OmbCam(BA) Camaca BA -15.38333333 -39.55 Ombrófila Montana Amorim et al.(2009)
OmbLin(ES) Linhares ES -19.2 -40.03333333 Ombrófila Tabuleiro Peixoto et al. (2008)
ResVit(ES) Vitória ES -20.25833333 -40.275 Restinga Pereira & Assis (2000)
ResSJB(RJ) São João da Barra RJ -21.73333333 -41.03333333 Restinga Assumpção & Nascimento (2000)
OmbCMa(RJ) Cachoeiras de Macaú RJ -22.48333333 -42.88333333 Ombrófila Submontana Kurtz & Araújo (2000)
ResMar(RJ) Maricá RJ -22.91805556 -42.81694444 Restinga Lemos et al. (2001)
ResBer(SP) Bertioga SP -23.75083333 -45.97 Restinga Martins et al. (2008)
ResBer2(SP) Bertioga SP -23.75083333 -45.97 Restinga Martins et al. (2008)
OmbSBa(SP) Sete Barras SP -24.23555556 -48.07833333 Ombrófila Submontana Ziparro et al. (2005)

The Restinga Forests in the survey region grow on well-preserved coastal ridges ranging from 5 to 8 m a.s.l. The substrate is sandy, and originated from ocean regressions following the Pleistocene transgression (Martin et al. 1980). Forest fragments are surrounded by open, seasonally flooded grasslands (Silva & Britez 2005) that grow in lowland areas between mountain ridges, with pools forming during the rainy period. These grasslands are characterized by a well-developed herbaceous layer with sparsely distributed woody plants (Silva & Britez 2005).

The four Restinga forest fragments studied here are classified as non-flooding Restinga forests, according Silva & Britez (2005), although all of them show micro-topographical variations with certain localities subject to periodic flooding. The forest canopy is approximately 15 m tall, although some emergent species may reach 22 m.

Floristic Surveys

Floristic surveys were carried out between March/2013 and February/2014 during nine field trips. The plant specimens were gathered and processed as described by Mori et al. (1989). Biological forms were assigned to each species following Whittaker (1975): trees (woody, ≥ 3 m tall), shrubs (woody, <3 m tall, usually multi-stemmed), herbs (self-standing, not woody), vines (climbing, woody or not woody), and epiphytes (growing on other plants). Trees and shrubs were collected even if they were not fertile, but other biological forms were sampled only when flowering and/or fruiting.

All collected materials were deposited in the State University of Feira de Santana herbarium (HUEFS). Identifications were made using taxonomic monographs and floras, by comparisons with identified specimens in the HUEFS and CEPEC (Herbário André Maurício Vieira de Carvalho, Centro de Pesquisas do Cacau) herbaria, and by consulting specialists of specific taxonomic groups. The species are arranged according the APG III (2009) system, with taxa names being updated according to the online databases of the Lista de Espécies da Flora do Brasil (<http://floradobrasil.jbrj.gov.br>) and the Missouri Botanical Gardens (<http://tropicos.org>). The occurrence of each species in specific Brazilian phytogeographical domains (Amazonia, Atlantic Forest, Caatinga, Cerrado, Pantanal, and Pampa) was determined by consulting the Lista de Espécies da Flora do Brasil website (<http://floradobrasil.jbrj.gov.br>). Species were considered endemic to the Atlantic domain following Stehmann et al. (2009).

Similarity and correlation analyses

In order to test the putative effects of geographical distances from wet forests on the floristic compositions of Restinga forests, we prepared a binary matrix (presence/absence) based on the floristic lists of eleven Restinga forest and nine wet forest sites (Tab. 1; Fig. 1). All wet forest sites were within the Atlantic domain and less than 200 km from the Restinga forests sites investigated.

Figure 1 Map of Brazil showing the locations of the areas used in the similarity analyses; circles indicate Restinga forests, and triangles wet forest areas. The inset shows the borders of the municipalities of Itacaré and Maraú (Bahia State), highlighting the distribution of the sandy Quaternary sediments and Restinga forest sites (circles) where the floristic surveys were conducted. Brazilian states with areas considered in the similarity and cluster analyses are indicated by darker grey shading: BA= Bahia; CE= Ceará; ES=Espírito Santo; PB=Paraíba; PE=Pernambuco; RJ= Rio de Janeiro; SP= São Paulo.Figura 1 – Mapa do Brasil mostrando a localização das áreas comparadas nas análises de similaridade; círculos indicam áreas de florestas de Restinga e triângulos áreas de florestas pluviais. Em detalhe, limites dos municípios de Itacaré e Maraú (estado da Bahia), destacando a extensão de sedimentos arenosos do Quaternário e os sítios de florestas de Restinga (círculos) onde foram realizadas amostragem florística. Estados com áreas usadas nas análises de similaridade e agrupamento estão em cinza mais escuro: BA= Bahia; CE= Ceará; ES=Espírito Santo; PB=Paraíba; PE=Pernambuco; RJ= Rio de Janeiro; SP= São Paulo. 

We excluded from the dataset any taxa identified only to the genus or family levels, or species with inaccurate identifications (affine [aff.] or confer [cf.]). Infraspecific ranks were not considered, and subspecies and different varieties were therefore regarded as the same species. The consolidated floristic list of the 20 areas comprised 2,431 species (Supplementary material).

The sites were compared by cluster analysis using Jaccard’s coefficient (Mueller-Dombois & Ellenberg 1974) and the Unweighted Pair Group Method using Arithmetic averages (UPGMA) with bootstrap support calculated from 1,000 replications using Past software (Hammer et al. 2001). Principal Component Analysis (PCA) was also performed using the same dataset and software cited above.

The Mantel Test using XLSTAT software was employed to assess correlations between the geographic and floristic distances of all sites (Addinsoft 2007). Floristic distances were estimated using the Jaccard coefficient, calculated with Past software (Hammer et al. 2001). Geographical distances were assessed by considering a central coordinate for each site, and the distances between areas were then measured using the Google Earth ruler tool. The significance of the Mantel test was calculated using the Monte Carlo test, with 10,000 permutations.

Results

The floristics of restinga forests in southeastern Bahia

A total of 302 angiosperm species belonging to 184 genera and 75 families were surveyed (Tab. 2), with 295 taxa being identified to the species level. The richest families in terms of their numbers of species were: Fabaceae (35 species), Myrtaceae (25), Rubiaceae (21), Sapotaceae (13), Bromeliaceae (12), Annonaceae (11), Erythroxylaceae (10), Melastomataceae (9), and Apocynaceae (8). Together, these families comprised 47.7% of the total number of species sampled. The richest genera were Myrcia (12 species), Erythroxylum (10), Aechmea (8), Miconia (8), and Psychotria (7).

Table 2 List of the Angiosperm species collected at Maraú-Itacaré: MC= Mata do Caubi; MMI= Mata estrada Maraú- -Itacaré; MP= Mata Piracanga; MAS= Mata do Santo Amaro. Brazilian phytogeographic domains: AM= Amazonian, CA= Caatinga, CE= Cerrado, MA= Atlantic Forest, PAM=Pampa, PAN= Pantanal. Collectors: EM= Eloína Neri de Matos; MF= Moabe Ferreira Fernandes. *= Species endemic to southeastern Bahia and northern Espírito Santo.Tabela 2 - Lista das especies de Angiospermas amostradas nas florestas de Restinga da regiao de Marau-Itacare: MC= Mata do Caubi; MMI= Mata estrada Marau-Itacare; MP= Mata Piracanga; MAS= Mata do Santo Amaro. Dominios fitogeograficos: AM=Amazonia, CA=Caatinga, CE=Cerrado, MA=Mata Atlantica, PAM=Pampa, PAN=Pantanal. Coletores: EM= Eloina Neri de Matos; MF= Moabe Ferreira Fernandes. *=Endemicas do sul da Bahia e norte do Espirito Santo. 

Family/Species Habit Phytogeographic domains Voucher MC MMI MP MSA
Acanthaceae
Aphelandra nitida Nees & Mart. Herb MA MF513 x x x x
Ruellia affinis (Schrad.) Lindau* Shrub MA MF130 x     x
Achariaceae
Carpotroche brasiliensis (Raddi) A Gray Tree CE, AM, MA MF464       x
Anacardiaceae              
Tapirira guianensis Aubl. Tree CA, CE, AM, MA MF90 x x x x
Thyrsodium spruceanum Benth. Tree AM, MA MF460   x x x
Annonaceae
Annona acutiflora Mart. Tree MA MF540   x x  
Annona bahiensis (Maas & Westra) H.Rainer Tree MA MF541 x x x x
Annona salzmannii A.DC. Tree MA MF211 x x x x
Duguetia restingae Maas* Tree MA EM3535       x
Guatteria australis A.St.-Hil. Tree MA MF98 x x x x
Guatteria oligocarpa Mart. Tree MA MF152 x x x x
Guatteria tomentosa Rusby Tree AM, MA MF299 x     x
Unonopsis riedeliana R.E.Fr. Tree MA MF317     x x
Xylopia ochrantha Mart. Tree MA MF420     x x
Xylopia sericea A.St.-Hil. Tree CE, AM, MA MF138   x    
Xylopia sp. Tree - MF355     x  
Apocynaceae
Aspidosperma discolor A.DC. Tree CA, CE, AM, MA MF270 x x   x
Couma rigida Mull.Arg.* Tree CA, MA MF283 x   x  
Ditassa crassifolia Decne. Vine CE, MA MF227 x x x  
Himatanthus bracteatus (A.DC.) Woodson Tree AM, MA MF248   x x x
Mandevilla scabra (Hoffmanns. ex Roem. & Schult.) K.Schum. Vine CA, CE, AM, MA MF79 x x    
Rauvolfia grandiflora Mart. Tree MA MF122     x x
Tabernaemontana flavicans Willd. ex Roem. & Schult. Tree CE, AM, MA MF364   x   x
Tabernaemontana salzmannii A.DC. Tree CE, MA MF112       x
Aquifoliaceae
Ilex floribunda Reissek ex Maxim. Tree MA MF430 x x x x
Araceae
Anthurium gladiifolium Schott Herb MA MF345       x
Heteropsis oblongifolia Kunth Herb CE, AM, MA MF91 x x x x
Rhodospatha latifolia Poepp. Herb CA, CE, AM, MA EM3543       x
Araliaceae
Schefflera selloi (Marchal) Frodin & Fiaschi* Tree MA MF261 x x x  
Arecaceae
Attalea funifera Mart. ex Spreng. Tree MA MF542 x x x x
Bactris hirta Mart. Shrub AM, MA MF408   x x  
Bactris horridispatha Noblick ex A.J.Hend.* Shrub MA MF343 x     x
Euterpe edulis Mart. Tree CE, MA MF543 x x x x
Geonoma pauciflora Mart. Shrub MA EM3536     x  
Geonoma pohliana Mart. Shrub MA MF292   x    
Asteraceae
Piptocarpha riedelii (Sch.Bip.) Baker* Vine MA MF479 x x    
Vernonanthura divaricata (Spreng.) H.Rob. Shrub CE, MA MF417 x     x
Vernonanthura vinhae (H.Rob.) H.Rob.* Shrub MA MF213 x x x  
Bignoniaceae
Anemopaegma chamberlaynii (Sims) Bureau & K.Schum. Vine CA, CE, MA MF140   x   x
Handroanthus umbellatus (Sond.) Mattos Tree CA, CE, MA MF390   x   x
Jacaranda obovata Cham. Tree CA, CE, MA MF500 x x x x
Lundia cordata (Vell.) DC. Vine CA, MA MF95 x     x
Tabebuia elliptica (DC.) Sandwith Tree CA, CE, MA MF77 x x x x
Tabebuia stenocalyx Sprague & Stapf Tree MA MF74 x x x x
Boraginaceae
Cordia glabrifolia M.Stapf* Tree MA EM3554       x
Bromeliaceae
Aechmea amorimii Leme* Epiphyte MA MF357       x
Aechmea andersonii H.Luther & Leme* Epiphyte MA EM3534       x
Aechmea aquilega (Salisb.) Griseb. Herb CA, CE, AM, MA EM3540 x x    
Aechmea bicolor L.B.Sm.* Epiphyte MA MF296       x
Aechmea blanchetiana (Baker) L.B.Sm.* Herb MA MF134 x x x  
Aechmea bromeliifolia (Rudge) Baker Herb CA, CE, AM, MA MF133 x x   x
Aechmea marauensis Leme Herb MA MF110     x x
Aechmea mertensii (G.Mey.) Schult. & Schult.f. Epiphyte AM, MA MF450       x
Guzmania lingulata (L.) Mez Epiphyte AM, MA EM3537 x x x x
Vriesea duvaliana E.Morren* Epiphyte MA EM3544     x  
Vriesea ensiformis (Vell.) Beer Epiphyte MA EM3552     x  
Vriesea procera (Mart. ex Schult. & Schult.f.) Wittm. Herb CA, CE, MA MF224 x x x  
Burseraceae
Protium bahianum Daly* Tree MA MF478   x x x
Protium heptaphyllum (Aubl.) Marchand Tree CA, CE, AM, MA MF359 x x x x
Tetragastris occhionii (Rizzini) Daly* Tree MA MF318     x x
Callophylaceae
Kielmeyera itacarensis Saddi * Tree MA MF537 x      
Cannabaceae
Trema micrantha (L.) Blume Tree AM, CA, CE, MA, PAN, PAM MF358       x
Caricaceae
Jacaratia heptaphylla (Vell.) A.DC. Tree MA MF83 x   x x
Caryocaraceae
Anthodiscus amazonicus Gleason & A.C.Sm. Tree AM, MA MF342   x   x
Celastraceae
Maytenus distichophylla Mart. ex Reissek Tree MA MF256 x x x x
Chrysobalanaceae
Couepia belemii Prance* Tree MA MF129 x x x x
Couepia ovalifolia (Schott) Benth. ex Hook.f. Tree CE, MA MF223   x x  
Licania hoehnei Pilg. Tree CE, MA MF531 x     x
Licania hypoleuca Benth. Tree AM, MA MF454 x x x x
Licania lamentanda Prance* Tree MA EM3553       x
Licania littoralis Warm. Tree MA EM3559       x
Licania sp. Tree - MF356   x    
Parinari alvimii Prance* Tree MA MF240     x  
Clusiaceae
Clusia nemorosa G.Mey. Tree CA, CE, AM, MA MF415 x x x x
Clusia sellowiana Schltdl. Tree MA MF491     x x
Garcinia gardneriana (Planch. & Triana) Zappi Tree CA, CE, AM, MA MF305 x x x x
Garcinia macrophylla Mart. Tree AM, MA MF333 x      
Symphonia globulifera L.f. Tree AM, MA MF281 x x x x
Combretaceae
Buchenavia hoehneana N.F.Mattos Tree MA MF389   x   x
Conocarpus erectus L. Tree AM, MA MF243       x
Terminalia glabrescens Mart. Tree CA, CE, AM, MA MF353 x x x x
Comelinaceae
Dichorisandra procera Mart. ex Schult & Schult.f. Herb MA MF286 x x x x
Cucurbitaceae
Cayaponia petiolulata Cogn.* Vine MA MF 103       x
Gurania lobata (L.) Pruski Vine AM, MA MF398 x   x x
Cyclanthaceae
Evodianthus funifer (Poit.) Lindm. Herb AM, MA MF105 x x x x
Dichapetalaceae
Tapura sp. Tree - MF336     x  
Dilleniaceae
Davilla flexuosa A.St.-Hil. Vine MA MF477 x x x x
Doliocarpus validus Kubitzki* Vine MA MF360   x   x
Elaeocarpaceae
Sloanea guianensis (Aubl.) Benth. Tree CE, AM, MA MF285 x x    
Ericaceae
Agarista revoluta (Spreng.) J.D. Hook. ex Nied. Shrub MA MF380 x x   x
Erythroxylaceae
Erythroxylum compressum Peyr.* Shrub MA MF216       x
Erythroxylum cuspidifolium Mart. Tree MA EM3562       x
Erythroxylum martii Peyr.* Shrub MA MF73 x x    
Erythroxylum mattos-silvae Plowman* Tree MA MF344 x x x  
Erythroxylum mikanii Peyr. Shrub MA EM3545     x  
Erythroxylum nobile O.E.Schulz Tree MA EM3567   x    
Erythroxylum passerinum Mart. Tree MA EM3539 x x    
Erythroxylum splendidum Plowman* Tree MA MF78 x     x
Erythroxylum squamatum Sw. Tree CA, CE, AM, MA MF87 x      
Erythroxylum tenue Plowman* Shrub MA MF320 x x    
Euphorbiaceae
Actinostemon concolor (Spreng.) Mull. Arg. Tree CA, CE, AM, MA MF372     x x
Alchornea glandulosa subsp. iricurana (Casar.) Secco Tree MA MF298 x      
Astraea sp. Tree - MF225   x x  
Croton macrobothrys Baill. Tree MA MF447   x x  
Croton sellowii Baill. Shrub CA, MA MF271 x x x x
Microstachys corniculata (Vahl) Griseb. Herb CA, CE, AM, MA MF350 x      
Microstachys heterodoxa (Mull.Arg.) Esser* Shrub CA, MA MF379   x    
Fabaceae
Abarema filamentosa (Benth.) Pittier Tree MA MF88 x x x x
Abarema jupunba (Willd.) Britton & Killip Tree AM, MA MF93     x x
Abarema turbinata (Benth.) Barneby & J.W. Grimes* Tree MA MF104 x x x  
Albizia pedicellaris (DC.) L.Rico Tree CE, AM, MA MF113       x
Andira marauensis N.F.Mattos* Tree MA MF76   x x  
Andira nitida Mart. ex Benth. Tree MA MF323 x x x x
Arapatiella psilophylla (Harms) R.S.Cowan* Tree MA MF128       x
Chamaecrista ensiformis (Vell.) H.S.Irwin & Barneby Tree CA, CE, AM, MA MF157       x
Clitoria falcata Lam. Vine CE, AM, MA MF337   x   x
Copaifera lucens Dwyer Tree MA MF384       x
Dalbergia nigra (Vell.) Allemao ex Benth. Tree MA MF403       x
Diplotropis incexis Rizzini & A.Mattos Tree MA MF228 x x x  
Harleyodendron unifoliolatum R.S.Cowan Tree MA MF468       x
Hymenolobium alagoanum Ducke Tree MA MF141 x x    
Inga capitata Desv. Tree AM, MA MF99 x x x x
Inga edulis Mart. Tree CA, CE, AM, MA MF348   x   x
Inga pleiogyna T.D.Penn.* Tree MA MF151 x x   x
Inga tenuis (Vell.) Mart. Tree MA MF392 x   x x
Inga thibaudiana DC. Tree CA, CE, AM, MA MF86 x     x
Leptolobium bijugum (Spreng.) Vogel Tree MA MF275 x      
Machaerium salzmannii Benth. Shrub MA MF102       x
Macrolobium latifolium Vogel* Tree MA MF101 x x   x
Mimosa ceratonia L. Vine CE, MA MF207       x
Moldenhawera blanchetiana Tul.* Tree MA MF162       x
Ormosia fastigiata Tul. Tree CE, MA MF255 x x x x
Parkia bahiae H.C.Hopkins* Tree MA MF81 x x x x
Parkia pendula (Willd.) Benth. ex Walp. Tree AM, MA MF206     x x
Piptadenia adiantoides (Spreng.) J.F.Macbr. Vine CA, CE, MA MF125       x
Pterocarpus rohrii Vahl Tree CE, AM, MA MF385   x x  
Senna pinheiroi H.S.Irwin & Barneby Shrub CA, MA MF144 x     x
Swartzia apetala Raddi Tree CA, CE, MA MF80 x   x x
Swartzia polita (R.S.Cowan) Torke* Tree MA MF302 x x x x
Swartzia simplex (Sw.) Spreng. Tree MA MF346 x x    
Tachigali densiflora (Benth.) L.G.Silva & H.C.Lima Tree MA MF282 x     x
Zollernia glabra (Spreng.) Yakovlev Tree MA MF273 x x x  
Gentianaceae
Chelonanthus purpurascens (Aubl.) Struwe et al. Herb CA, CE, AM, MA MF118       x
Heliconiaceae
Heliconia psittacorum L.f. Herb AM, CA, CE, MA, PAN EM3566       x
Humiriaceae
Humiria balsamifera (Aubl.) J.St.-Hil. Tree CA, CE, AM, MA MF136 x x x x
Sacoglottis mattogrossensis Malme Tree CE, AM, MA MF485 x      
Vantanea bahiaensis Cuatrec.* Tree MA MF301   x    
Hypericaceae              
Vismia guianensis (Aubl.) Choisy Tree CA, CE, AM, MA MF100 x x x x
Vismia macrophylla Kunth Tree AM, MA MF149       x
Icacinaceae
Emmotum affine Miers Tree MA MF316 x x x x
Lacistemataceae
Lacistema robustum Schnizl. Tree MA MF117 x     x
Lamiaceae
Aegiphila integrifolia (Jacq.) Moldenke Vine CA, CE, AM, MA MF470   x    
Aegiphila macrantha Ducke Vine AM, MA MF525       x
Vitex polygama Cham. Tree CA, CE, AM, MA MF242 x x x  
Lauraceae
Endlicheria glomerata Mez Tree CA, CE, MA MF544 x      
Nectandra membranacea (Sw.) Griseb. Tree CA, CE, AM, MA MF352 x x    
Ocotea canaliculata ( Rich.) Mez Tree AM, MA MF276 x      
Ocotea notata (Nees & Mart.) Mez Tree MA MF326 x x x x
Ocotea nutans (Nees) Mez Tree CE, MA MF295 x   x  
Persea caesia Meisn. Tree MA MF309   x    
Lecythidaceae
Eschweilera ovata (Cambess.) Mart. ex Miers Tree CA, CE, AM, MA MF165 x x x x
Lecythis lurida (Miers) S.A.Mori Tree AM, MA MF373     x  
Lecythis pisonis Cambess. Tree AM, MA MF394       x
Malpighiaceae              
Byrsonima cacaophila W.R.Anderson* Tree MA MF289   x   x
Byrsonima sericea DC. Tree CA, CE, MA MF250 x x x x
Heteropterys imperata Amorim* Vine MA MF347 x   x x
Stigmaphyllon blanchetii C.E.Anderson Vine CA, CE, MA MF147   x   x
Malvaceae
Apeiba tibourbou Aubl. Tree CA, CE, AM, MA MF121       x
Eriotheca macrophylla (K.Schum.) A.Robyns Tree MA MF510 x x x x
Melastomataceae
Henriettea succosa (Aubl.) DC. Tree AM, MA MF123 x x   x
Miconia albicans (Sw.) Triana Shrub CA, CE, AM, MA MF509   x    
Miconia amoena Triana Tree MA MF215     x x
Miconia ciliata (Rich.) DC. Tree CA, CE, AM, MA MF114 x      
Miconia cinnamomifolia (DC.) Naudin Tree MA MF461 x x   x
Miconia compressa Naudin Shrub MA MF108   x x x
Miconia holosericea (L.) DC. Tree CE, AM, MA MF418 x x    
Miconia prasina (Sw.) DC. Tree CA, CE, AM, MA MF458 x     x
Miconia tomentosa (Rich.) D.Don Tree CE, AM, MA MF284       x
Meliaceae
Guarea guidonia (L.) Sleumer Tree CA, CE, AM, MA MF322 x x x x
Trichilia lepidota Mart. Tree MA MF232 x x x x
Trichilia ramalhoi Rizzini Tree MA MF145     x x
Moraceae
Brosimum rubescens Taub. Tree AM, MA EM3560 x x x x
Ficus bahiensis C.C.Berg & Carauta Tree CA, CE, MA MF257 x x x  
Ficus gomelleira Kunth Tree CA, CE, AM, MA MF204 x x x x
Ficus hirsuta Schott Tree CA, MA MF97   x   x
Sorocea racemosa Gaudich. Tree MA MF452     x x
Myristicaceae
Virola officinalis Warb. Tree MA MF367       x
Myrtaceae              
Calycolpus legrandii Mattos Tree MA MF205       x
Calyptranthes restingae Sobral Tree MA MF274   x x  
Campomanesia dichotoma (O.Berg) Mattos Tree CA, CE, MA MF307 x   x x
Eugenia astringens Cambess. Tree MA MF545   x x x
Eugenia candolleana DC. Tree CA, CE, MA MF547     x  
Eugenia hirta O.Berg Tree MA MF221   x    
Eugenia itacarensis Mattos* Tree MA MF436   x    
Eugenia longifolia DC.* Tree MA EM3568 x x   x
Eugenia sp. Tree - MF131 x      
Marlierea glabra Cambess. Tree MA EM3550 x x x  
Marlierea obscura O.Berg Tree MA MF539 x x    
Marlierea obversa D.Legrand Tree MA MF506 x     x
Myrcia bergiana O.Berg Tree CA, MA MF107 x x x x
Myrcia grazielae NicLugh.* Tree MA MF546 x     x
Myrcia guianensis (Aubl.) DC. Tree CA, CE, AM, MA MF212 x x x x
Myrcia micropetala (Mart.) Nied. * Tree MA MF550     x x
Myrcia ramuliflora (O.Berg) N.Silveira* Tree MA MF234 x x    
Myrcia salzmannii O.Berg Tree MA MF538   x   x
Myrcia silvatica (G.Mey.) DC. Tree CE, AM, MA MF115 x     x
Myrcia sp. Tree - EM3541       x
Myrcia sp.2 Tree - MF425 x      
Myrcia spectabilis DC. Tree MA MF312 x x x x
Myrcia splendens (Sw.) DC. Tree AM, CA, CE, MA, PAN MF132 x x x x
Myrcia tomentosa (Aubl.) DC. Tree CA, CE, AM, MA MF168     x x
Psidium bahianum Landrum & Funch* Tree MA MF388       x
Nyctaginaceae
Guapira areolata (Heimerl) Lundell Tree CE, MA MF246 x x   x
Guapira opposita (Vell.) Reitz Tree CA, CE, AM, MA MF226 x x x x
Guapira pernambucensis (Casar.) Lundell Tree MA MF492   x x  
Guapira venosa (Choisy) Lundell Tree AM, MA MF401     x x
Neea floribunda Poepp. & Endl. Tree AM, MA MF167     x x
Neea macrophylla Poepp. & Endl. Tree CE, AM, MA MF288 x x x x
Ochnaceae
Ouratea bahiensis Sastre* Shrub MA MF89 x     x
Ouratea castaneifolia (DC.) Engl. Tree CA, CE, AM, MA MF368       x
Ouratea gigantophylla (Erhard) Engl.* Tree MA MF374 x x x x
Ouratea longipes Sastre* Shrub MA MF303     x  
Ouratea platicaulis Sastre* Tree MA MF497 x x x x
Ouratea rotundifolia (Gardner) Engl.* Shrub MA MF249   x    
Olacaceae
Cathedra bahiensis Sleumer* Tree MA MF222     x  
Heisteria perianthomega (Vell.) Sleumer Tree CA, CE, AM, MA MF210   x x x
Orchidaceae
Epistephium lucidum Cogn. Herb CA, CE, AM, MA MF376       x
Koellensteinia florida (Rchb.f.) Garay Herb MA MF109   x    
Passifloraceae
Passiflora sidifolia M.Roem. Vine MA MF96   x    
Passiflora silvestris Vell. Vine CE, MA MF328 x      
Passiflora watsoniana Mast. Vine MA MF457     x  
Peraceae
Pera glabrata (Schott) Poepp. ex Baill. Tree CA, CE, AM, MA MF251 x x x x
Pogonophora schomburgkiana Miers ex Benth. Tree CA, CE, AM, MA MF370 x x x x
Phyllantaceae
Richeria grandis Vahl Tree CA, CE, AM, MA MF164 x x x x
Piperaceae
Piper aduncum L. Tree AM, CA, CE, MA, PAN, PAM MF396       x
Piper amplum Kunth Shrub CE, MA MF399       x
Piper ilheusense Yunck. Shrub MA MF75 x x x  
Poaceae
Parodiolyra ramosissima (Trin.) Soderstr. & Zuloaga* Herb MA MF382   x    
Polygalaceae
Bredemeyera disperma (Vell.) J.F.B.Pastore Vine MA MF92 x   x  
Polygonaceae
Coccoloba laevis Casar. Tree CA, MA MF244 x     x
Coccoloba marginata Benth. Tree CE, AM, MA MF209 x x    
Coccoloba oblonga Lindau* Shrub MA MF218 x   x  
Coccoloba parimensis Benth. Shrub CA, CE, AM, MA MF279       x
Coccoloba rosea Meisn. Tree MA MF119 x x x x
Primulaceae
Cybianthus amplus (Mez) G.Agostini Tree AM, MA MF438   x x  
Cybianthus densiflorus Miq. Tree AM, MA MF245 x x x x
Rubiaceae
Amaioua intermedia Mart. ex Schult. & Schult.f. Tree CE, MA MF208 x x x  
Chiococca alba (L.) Hitchc. Shrub AM, CA, CE, MA, PAN MF427 x x x x
Coussarea graciliflora (Mart.) Mull.Arg. Shrub MA MF407   x    
Faramea coerulea (Nees & Mart.) DC. Shrub MA MF280   x    
Faramea nocturna J.G.Jardim & Zappi* Tree MA MF349   x x x
Margaritopsis chaenotricha (DC.) C.M.Taylor Shrub MA MF217 x      
Melanopsidium nigrum Colla Tree CE, MA MF453 x x    
Mitracarpus salzmannianus DC. Herb CA, CE, AM, MA MF71     x x
Pagamea guianensis Aubl. Shrub CA, CE, AM, MA MF377 x x x x
Pagamea harleyi Steyerm.* Tree MA MF262   x x  
Palicourea guianensis Aubl. Tree CE, AM, MA MF146 x x x x
Psychotria bahiensis DC. Shrub CA, CE, MA MF70 x x x x
Psychotria cupularis (Mull.Arg.) Standl. Tree AM, MA MF362   x    
Psychotria jambosioides Schltdl.* Shrub MA MF137 x x x x
Psychotria mapourioides DC. Tree AM, MA MF426 x x x x
Psychotria schlechtendaliana (Mull.Arg.) Mull.Arg. Shrub CA, MA MF124 x      
Psychotria stachyoides Benth. Shrub CA, CE, MA MF300 x      
Psychotria vellosiana Benth. Tree CA, CE, MA MF383   x    
Randia calycina Cham. Tree CE, AM, MA MF321     x x
Rudgea interrupta Benth. Shrub MA MF465 x x    
Salzmannia nitida DC. Shrub MA MF214 x x x  
Rutaceae
Rauia nodosa (Engl.) Kallunki Tree MA MF455       x
Salicaceae
Casearia arborea (Rich.) Urb. Tree CE, AM, MA MF435 x x x x
Casearia commersoniana Cambess. Tree CA, CE, AM, MA MF85 x x x x
Sapindaceae
Cupania oblongifolia Mart. Tree CE, AM, MA MF325       x
Cupania racemosa (Vell.) Radlk. Tree CA, CE, AM, MA MF126 x x    
Matayba discolor (Spreng.) Radlk. Tree MA MF84     x  
Paullinia racemosa Wawra Vine MA MF230 x x x x
Serjania salzmanniana Schltdl. Vine CE, AM, MA MF94 x x x x
Talisia macrophylla (Mart.) Radlk. Tree AM, MA MF429       x
Sapotaceae
Chrysophyllum flexuosum Mart. Tree MA MF519 x   x  
Diploon cuspidatum (Hoehne) Cronquist Tree AM, MA MF369 x x    
Ecclinusa guianensis Eyma Tree AM, MA MF351 x x x x
Manilkara salzmannii (A.DC.) H.J.Lam Tree MA MF315 x x x x
Micropholis crassipedicellata (Mart. & Eichler) Pierre Tree MA MF366       x
Micropholis emarginata T.D.Penn.* Tree CA, CE, MA MF445     x  
Micropholis gardneriana (A.DC.) Pierre Tree CA, CE, AM, MA MF294 x x    
Pouteria butyrocarpa (Kuhlm.) T.D.Penn.* Tree MA MF330       x
Pouteria coelomatica Rizzini Tree MA MF329 x x   x
Pouteria cuspidata (A.DC.) Baehni Tree CE, AM, MA MF304   x    
Pouteria guianensis Aubl. Tree AM, MA MF331   x x  
Pouteria macahensis T.D.Penn. Tree MA MF 414 x      
Pradosia lactescens (Vell.) Radlk. Tree MA MF332   x x x
Schoepfiaceae
Schoepfia brasiliensis A.DC. Tree CE, AM, MA MF265   x    
Siparunaceae
Siparuna cymosa Tolm. Tree AM, MA MF291       x
Smilacaceae
Smilax campestris Griseb. Vine CA, CE, MA, PAM MF441 x x x  
Solanaceae
Cestrum salzmannii Dunal Shrub MA MF405 x   x x
Solanum asperum Rich. Shrub CA, CE, AM, MA MF404     x x
Solanum rupincola Sendtn. Vine MA MF375   x x  
Stemonuraceae
Discophora guianensis Miers Tree AM, MA MF127   x   x
Thymaelaeceae
Daphnopsis racemosa Griseb. Tree CE, MA MF269   x x x
Triuridaceae
Peltophyllum luteum Gardner Herb MA MF72 x     x
Urticaceae
Cecropia pachystachya Trecul Tree AM, CA, CE, MA, PAN MF397 x x x  
Verbenaceae
Lippia macrophylla Cham.* Shrub CE, MA MF339 x x x  
Violaceae
Paypayrola blanchetiana Tul. Tree MA MF386 x      
Rinorea bahiensis (Moric.) Kuntze Tree MA MF148 x   x  

Among biological forms, trees were the most abundant (213 species / 70.5%), followed by shrubs (39 / 12.9%), vines (24 / 7.9%), herbs (19 / 6.3%), and epiphytes (7 / 2.3%).

Phytogeographical domain assignments resulted in 144 species (47.7%) identified as being endemic to the Atlantic domain, 34 species (11.2%) disjunct between the Atlantic and Amazonia domains, 72 species (23.8%) continuously distributed throughout the Atlantic and Amazonian domains, and 45 (14.9%) throughout the Atlantic, Caatinga, and Cerrado domains. Among the species endemic to the Atlantic domain, 58 (19.2%) were restricted to the area between southern Bahia and northern Espírito Santo.

Five species encountered in the study area are probably endemic to Restinga vegetation: Abarema turbinata (Benth.) Barneby & J.W.Grimes, Leptolobium bijugum (Spreng.) Vogel, Parkia bahiae H.C.Hopkins (Fabaceae), Pagamea harleyi Steyerm. (Rubiaceae), and Schefflera selloi (Marchal) Frodin & Fiaschi (Araliaceae).

Similarity and correlation analyses

Similarity analysis indicated the formation of five groups (A – E) with low similarity values, but with high support values, except group A (Fig. 2). Four groups include Restinga and wet forests, with: Group A comprising the sites in Rio de Janeiro and southern Espírito Santo states; group B comprising the sites located in Bahia State north of Todos os Santos Bay to Paraíba State; group C comprising the only site examined in Ceará State; group D comprising sites in southern Bahia and northern Espírito Santo states; and group E comprising the sites in São Paulo State.

Figure 2 Results of the UPGMA cluster analysis that considered areas of Restinga and wet forests of the Atlantic domain (above), showing the formation of five major groups according to their Jaccard similarity indices. The numbers associated with the groups are bootstrap support values (1,000 replications). Principal component analysis (PCA) using the same 20 areas is indicated below, highlighting the four major groups (A, B, D and E) in the first two axes (34.4% of total variance). The only area of group C (São Gonçalo do Amarante, Ceará) is included within the polygons formed by groups A and B.Figura 2 – Resultado da análise de agrupamento (UPGMA) entre áreas de florestas de Restinga e florestas pluviais do domínio Mata Atlântica (acima) mostrando a formação de cinco grupos principais de acordo com índice de similaridade de Jaccard. Números nos grupos são valores de suporte de bootstrap (1.000 replicações). Análise de componentes principais (PCA) entre as mesmas 20 áreas é mostrada abaixo ressaltando os quatro grupos maiores (A, B, D e E) nos dois primeiros eixos (34,4% da variância total). A única área do grupo C (São Gonçalo do Amarante, Ceará) está incluída nos polígonos formados pelos grupos A e B. 

PCA axis 1 and 2 (accounting for 34.4% of the observed variance) supported the separation of groups D and E from a group formed by the A, B and C sites. The Mantel test indicated a significant negative correlation between geographic distance and floristic similarities (r = -0.452; p < 0.0001).

Discussion

The floras of the southern Bahia Restinga forests

The species richness of the study areas were found to be similar to those of other Restinga forest surveys (Martins et al. 2008; Castro et al. 2012), but approximately three times lower than neighboring wet forest formations of the Atlantic domain (Amorim et al. 2008; Amorim et al. 2009). The low diversities of Restinga forests have been highlighted by Guedes et al. (2006) and Lima et al. (2011), and seem to be determined by factors related to their nutrient-poor and well-drained sandy soils (Almeida Jr. et al. 2009). Edaphic conditions are therefore presumed to act as environmental filters that limit the establishment of lineages not adapted to the unique conditions of Restinga habitats.

The most species rich families encountered in our surveys (Fabaceae, Myrtaceae, Sapotaceae, and Rubiaceae) were also among the most important families reported in other Restinga forest sites (Assumpção & Nascimento 2000; Assis et al. 2004; Martins et al. 2008; Silva et al. 2008). Myrtaceae taxa are typically found on soils with low fertility (Berry 1915) and are important components of Restinga vegetation along the entire coast of Brazil. Erythroxylaceae and Melastomataceae, on the other hand, generally contribute very few species to the Restinga forest floras, although they are well-represented in wet forests of the Atlantic domain (Stehmann et al. 2009); Erythroxylaceae also shows high richness in the Cerrado domain (Ratter et al. 2003). Bromeliaceae was among the five richest families in the Restinga forests studied. This family is highly diverse in eastern Brazil (Smith & Downs 1979) and is the most diverse family of epiphytes in the Atlantic domain (Borgo & Silva 2003; Giongo & Weachter 2004) – but has relatively little importance in sites along the northern coast of Brazil (Castro et al. 2012). Thus, the high richness of families such Melastomataceae, Erythroxylaceae, and Bromeliaceae is probably idiosyncratic in southern Bahia Restinga forests.

Myrcia and Psychotria were among the most species rich genera in the study sites, and have likewise been reported as being highly diverse genera in Restinga forests (Martins et al. 2008) and other forest types in the Atlantic domain (Amorim et al. 2009; Stehmann et al. 2009). Aechmea, Erythroxylum, and Miconia were recorded here for the first time as being among the most diverse genera of Restinga forests.

Epiphytes are poorly represented in southern Bahia Restinga forests, in agreement with reports from coastal forests in the neighboring Espírito Santo State (Pereira et al. 1998).

The presence of species disjunct between Atlantic and Amazonian wet forests reinforces the hypothesis of past connections between these two forest blocks. It has been hypothesized that connections were possible during the wetter periods of the Quaternary due to the expansion of those wet forests and/or the emergence of ecological corridors (Bigarella & Andrade-Lima 1982). About 8% of the species found in forests in southern Bahia have this distribution pattern (Mori et al. 1981; Amorim et al. 2008), but less than 4% of Restinga species in Rio de Janeiro State (Araújo 2000). These findings indicate that proximity to southern Bahia wet forests (“Hileia Baiana”) accounts for higher proportions of species from Restinga forests in South Bahia showing Atlantic-Amazonia disjunction patterns than those in other areas (Pereira & Araújo 2000).

Some species encountered in the Restinga forests studied here occur from Atlantic domain forests through to the Caatinga and Cerrado domains, occasionally reaching the Amazon region. Oliveira-Filho & Ratter (1995) concluded that these distribution patterns could have been established through two distinct processes: by interchanges of species between deciduous and semi-deciduous forests (requiring the presence of high to medium fertility soils); or the expansion of species from gallery forests (that could have acted as ecological corridors for wet forests trees across central Brazil).

Southern Bahia and northern Espírito Santo State show high levels of endemism and diversity (Thomas et al. 1998; Murray-Smith et al. 2008) – which has been attributed to the environmental stability of a putative ecological refuge during the climatic fluctuations of the Quaternary period (Carnaval & Moritz 2008). The few endemic species observed in Restinga vegetation (Lima et al. 2011) probably reflect the more recent origin of its geological substrate (Scarano 2002).

Are Restinga forests a subset of Atlantic wet forests?

Multivariate analyzes demonstrated (with high bootstrap support) that Restinga forest sites did not group together, yielding instead mixed groups of Restinga forests and neighboring Atlantic wet forest sites. The Mantel test results reinforced the influence of geography on the similarities between areas, demonstrating that geographically closer areas tend to be more similar – even though they did not necessarily share similar ecological conditions (as with Restinga and wet forests).

The low similarity values observed are the result of only small numbers of species being shared between sites, and reflect the fact that different areas had floristic particularities that are probably the result of the environmental heterogeneity that characterizes Brazilian Restinga sites (Araújo & Henriques 1984; Magnago et al. 2011).

The fact that Restinga forests: (1) have few endemic species; (2) are of recent geological origin; and (3) most species (~94%) also occur in wet forests in the Atlantic domain (Stehmann 2009), reinforce the hypothesis that Restingas are marginal habitats and that their floras are subsets of the floras of adjacent wet forest areas (Scarano 2009). These wet forests thus serve as sources of species able to overcome environmental filters imposed by unique restinga soil conditions (Assis et al. 2011). The proximity of wet forests that serve as sources of propagules should have strong influences on community assemblages and diversity patterns in Restinga forests (Assumpção & Nascimento 2000; Almeida Jr et al. 2009; Santos et al. 2012). This floristic continuity, together with evidence that tropical forest trees can disperse pollen over long distances (up to 20 km; see Ward et al. 2005 for a review), suggests that gene flow can occur between Restinga forest plants and populations in adjacent wet forests – and would account for low speciation rates and low endemism.

The Restinga forest sites of Bahia State clustered into two groups: 1) sites north of Todos os Santos Bay in group B and other sites from northeastern Brazil; and 2) sites in southern Bahia in group D and sites from northern Espírito Santo State. This finding reinforces previous reports that the Restinga vegetation of Bahia State forms two blocks – with sites located in the southern portion of that state being more similar to Restinga sites in Espírito Santo and Rio de Janeiro than to sites located in northern Bahia (Araújo 2000). The differentiation of these floristic blocks could reflect distinct climate types (a moist tropical climate with no dry season in the southern region, and a seasonal tropical climate with a dry winter in the northern region) (Peel et al. 2007; Alvares et al. 2013). Seasonality thus represents another factor that can influence the floristic composition of Restinga forests located north of Todos os Santo Bay, and this is corroborated by the presence of species that are typical of seasonal environments in group B sites, such as Commiphora leptophloeos (Mart.) J.B.Gillett (Burseraceae), Jatropha mollissima (Pohl) Baill. (Euphorbiaceae), and Syagrus coronata (Mart.) Becc (Arecaceae).

Substructuring according to elevation was noted in group D, with one group being formed by lowland areas (“tabuleiros”) and Restinga, and another group formed by montane and submontane forests. Despite the environmental heterogeneity attributable to elevation, this group is well-supported (95% bootstrap). Even though geographically distant, the inclusion of the Linhares area into this group was expected because of an apparent floristic gradient between the forests of northern Espírito Santo and those of southern Bahia State (Oliveira-Filho & Fontes 2000).

The areas located in the states of Rio de Janeiro and São Paulo formed two distinct groups by cluster analysis (groups A and E respectively). These two groups are located on the southeastern coast of Brazil (Villwock et al. 2005) an area that is characterized by the presence of the Serra do Mar mountain range. Coastal geology therefore also seems to influence the floristic composition of Restinga vegetation (Pereira & Araújo 2000). Despite their geological affinities, groups A and E are subjected to different climatic regimes (Peel et al. 2007; Alvares et al. 2013), which justifies their separation.

The isolated position of the Restinga forest located in Ceará State reinforces the importance of contiguous vegetation areas as propagule sources in the constitution of Restinga forest floras. The Restinga forest in this area is close to Caatinga (dryland) vegetation and shares many species that predominantly occur in the latter, such as Cereus jamacaru DC. (Cactaceae), Crotonblanchetianus Baill. (Euphorbiaceae), Margaritopsis carrascoana (Delprete & E.B.Souza) C.M.Taylor & E.B.Souza (Rubiaceae), and Sideroxylon obtusifolium (Humb. ex Roem. & Schult.) T.D. Penn. (Sapotaceae).

Supplementary material

Supplementary material is available at (<http://dx.doi.org/10.6084/m9.figshare.1246794>).

Acknowledgements

We are greatful to the following specialists for helping with plant identification: Adriana Q. Lobão (Annonaceae), Alessandro Rapini (Apocynaceae), Ana L. A. Côrtes (Acanthaceae), Anderson F. Machado (Cannabaceae, Moraceae and Urticaceae), André M. Amorim (Malpighiaceae and Violaceae), Cláudia E. Carneiro (Sapotaceae), Daniela S. Carneiro-Torres (Euphorbiaceae), Denis N. de Carvalho (Orchidaceae), Efigênia Melo (Dilleniaceae, Lacistemataceae and Polygonaceae), Eudes B. Mattos (Ericaceae), Fábio S. do Espírito-Santo (Bignoniaceae), Fabrício M. Ferreira (Poaceae), Fernanda O. Silva (Ochnaceae), Flávio França e Raymond M. Harley (Lamiaceae), Francisco S. Souza (Nyctaginaceae), Gabriela B. Siqueira (Gentianaceae), Herlon A. Santos (Carycaceae), Jefferson G. Carvalho-Sobrinho (Malvaceae), Jomar G.Jardim (Achariaceae and Rubiaceae), José F. B. Pastore (Polygalaceae), Juliana G. Freitas (Cyclanthaceae and Melastomataceae), Karoline C. de Santana (Myrtaceae), Larry Noblick (Arecaceae), Lucas Marinho (Clusiaceae and Hypericaceae), Maria B. B.Alves (Asteraceae), Matheus G. C. Nogueira (Bromeliaceae), Ricardo de O. Perdiz (Burseraceae and Sapindaceae), Tânia R. S. Silva (Verbenaceae), Teonildes S. Nunes (Passifloraceae), Thiago A. Pontes (Araceae), Thiago F. de Araújo (Erythroxylaceae). We are also greatful to André M. Amorim, Jomar G. Jardim, Marcelo Moro and two anonymous reviewers for their helpful considerations on the manuscript. This study is part of the M.Sc. thesis of MFF, developed at PPGBot-UEFS, with a fellowship from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES). The research was supported by the project “Padrões de diversidade de leguminosas nos biomas brasileiros: ligando taxonomia e moléculas para o entendimento da evolução da biota do Brasil” (Sisbiota CNPq processo 563084/2010-3 / Fapesb PES 0053/2011).

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Received: August 29, 2014; Accepted: November 23, 2014

3Author for correspondence: moabeffernandes@gmail.com

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