Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

Pesquisas Agrárias e Ambientais

DOI: https://doi.org/10.31413/nativa.v9i4.12732 ISSN: 2318-7670

Atlantic Forest meets the Cerrado: floristic, structure and species distribution

of an ecotonal tree community

Gabriel Pavan SABINO1*, Vitor de Andrade KAMIMURA1,2, Gabriel Mendes MARCUSSO1,

Reinaldo MONTEIRO1

1Departamento de Botânica, Universidade Estadual Paulista "Júlio de Mesquita Filho", Rio Claro, SP, Brasil.

2 Departamento de Botânica, Universidade Federal de Minas Gerais, Belo Horizonte, MG, Brasil.

*E-mail: gpsabino@gmail.com

(ORCID: 0000-0003-1284-8781; 0000-0002-3276-5812; 0000-0002-7520-2876; 0000-0003-1019-6003)

Recebido em 08/07/2021; Aceito em 01/11/2021; Publicado em 11/11/2021.

ABSTRACT: We evaluated the floristic and structural composition of a tree community in an ecotone between

Cerrado (cerradão) and Atlantic Forest (seasonal semideciduous forest) domains located in Porto Ferreira State

Park (PFSP), southeastern Brazil. We compared the floristic relationships of this ecotone with those of previous

surveys carried out on the same vegetation types and checked the species distribution among the Brazilian

biomes. We sampled all living trees with PBH>10 cm in 64 10x10 m plots (0.64 ha), totaling 1,755 individuals

belonging to 101 species and 37 families. The richest families were Myrtaceae (13 spp.) and Fabaceae (11 spp.),

and Siparuna guianensis was the most abundant species (188 individuals). We reported two threatened species. A

great number of species are widely distributed, occurring in different Brazilian biomes. Floristic similarity values

were low among the selected studies, but our sampled community clustered with communities of cerradão and

ecotone areas of previous surveys. Our results corroborate that ecotonal areas have great tree diversity and the

predominance of widely distributed species. This fact, combined with the vegetation thickening verified

through historical photographs, reinforces that the study area belongs to an under-changing ecotone.

Keywords: ecotone; cerradão; seasonal semideciduous forest; phytogeography; floristic similarities.

Mata Atlântica encontrando o Cerrado: florística, estrutura e distribuição de

espécies de uma comunidade arbórea ecotonal

RESUMO: No presente estudo, avaliamos a composição florística e estrutura de uma comunidade arbórea de

um ecótono entre o Cerrado (cerradão) e a Floresta Atlântica (floresta estacional semidecidual) localizado no

Parque Estadual de Porto Ferreira (PEPF), sudeste do Brasil. Verificamos também as relações florísticas com

outros levantamentos realizados nestas fitofisionomias, e analisamos a distribuição das espécies nos domínios

fitogeográficos brasileiros. Para isso amostramos todos os indivíduos vivos com PAP >10cm em 64 parcelas

de 10x10m (0,64 ha), totalizando 1755 indivíduos pertencentes a 101 espécies e 37 famílias. As famílias mais

ricas em espécies foram Myrtaceae (13 espécies) e Fabaceae (11) sendo Siparuna guianensis a espécie mais

abundante (188 indivíduos). Registramos duas espécies ameaçadas de extinção e verificamos um grande número

de espécies com ampla distribuição entre os domínios fitogeográficos do Brasil. Encontramos baixos valores

de similaridade florísticas entre as áreas analisadas, e a comunidade amostrada apresentou maior similaridade

com levantamentos realizados em área de cerradão e ecótonos. Em síntese, os resultados corroboram a grande

diversidade arbórea de áreas ecotonais, e o predomínio de espécies com ampla distribuição. Esse fato associado

ao adensamento da vegetação verificado em série fotográfica histórica, reforçam que a área do estudo pertence

a um ecótono em transformação.

Palavras-chave: ecótono; cerradão; floresta estacional semidecidual; fitogeografia; similaridade florísticas.

1. INTRODUCTION

Ecotones (gr. tonus = tension) are transitional areas

between ecological communities that commonly occur along

environmental gradients (ORCZEWSKA; GLISTA, 2005).

Studies have reported high species richness and abundance

tendency for ecotonal areas (e.g., KARK; van RENSBURG,

2006). Ecotones play an important role since environmental

gradients are important speciation and diversification sites

(SCHILTHUIZEN, 2000; SCHLUTER, 2000; SMITH et al.,

2001). The preservation of such areas can guarantee adaptive

responses of species to climate change (SMITH et al., 2001).

Savannah and forest ecotones play an important

ecological role in the tropics (HOFFMANN, 2005). Mosaics

of savannah-forest transitions occupy about 24% of the

Cerrado domain in Brazil (SILVA; BATES, 2002), and

several studies have pointed to edaphic factors as decisive

drivers of the establishment of Cerrado (savannah) and

Atlantic Forest physiognomies (FURLEY, 1992; RATTER,

1992; DURIGAN; RATTER, 2006; PINHEIRO;

MONTEIRO, 2008; BARROS et al. 2018; PINHEIRO et al.

2021). Furley (1992) concluded that soil moisture is a

determining factor in savannah-forest transitions. For Ratter

(1992), the soil is the main factor that influences the

geographic distribution of vegetation. Furthermore, fire

suppression can change the Cerrado vegetation types, since

Atlantic Forest meets the Cerrado: floristic, structure and species distribution of an ecotonal tree community

Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

472

forest vegetation types can replace open vegetations over the

years (DURIGAN; RATTER, 2006; PINHEIRO et al.,

2021). On the other hand, the presence of fire can facilitate

the invasion of savannah species in seasonal semideciduous

forests (PINHEIRO; MONTEIRO, 2008).

Sauer (1988) claims that there is a balance in savannah-

forest ecotones in South America. However, several studies

have shown a movement of these phytocenoses and an

expansion of forest formations over areas previously

occupied by Cerrado vegetation types (COUTINHO, 1990;

RATTER, 1992; PIVELLO; COUTINHO, 1996;

DURIGAN; RATTER, 2006; PINHEIRO; MONTEIRO,

2021).

Floristic and structural studies have been carried out on

Cerrado and seasonal semideciduous forests of southeastern

Brazil (DURIGAN et al., 1994; IVANAUSKAS et al. 1999;

DURIGAN et al. 2002; TEIXEIRA et al. 2004; NETO et al.,

2009; PINHEIRO & DURIGAN, 2012; HENCKER et al.,

2012; GARCIA et al., 2015), but few of them have

encompassed transitional areas (e.g., PINHEIRO;

MONTEIRO, 2008; GUILHERME; NAKAJIMA, 2007;

GOMES et al., 2004). Thus, we still do not understand this

complex mosaic completely. Data about transitional areas are

the basis for restoration projects, considering their

importance for legal activities for environmental adaptations

and licensing processes (e.g., Degraded Areas Recovery Plan

– PRAD). Furthermore, floristic relationships of these

ecotonal areas are important for characterizing and

contextualizing vegetations under its transitional condition.

In this study, we evaluate the composition, structure,

species distribution, and phytogeographic relationships of

the tree community of an ecotonal area to support recovery,

restoration and conservational actions in Cerrado-Atlantic

Forest ecotones in São Paulo state.

2. MATERIALS E METHODS

2.1. Study area

The study was conducted in an area with a cerradão

(CER) vegetation, a forested savannah, in transition with a

seasonal semideciduous forest (SSF) in Porto Ferreira State

Park (PFSP), municipality of Porto Ferreira, São Paulo state,

southeastern Brazil (Figure 1). Plots were placed in an area at

the north of “Lagoa do Cerrado” (21º50’49’’S, 47º25’39’’W),

where the average altitude was 600 m. The region’s climate is

classified as Cwa, or mesothermal dry winter, according to

the Köppen-Geiger scale (Alvares et al., 2013). The annual

mean temperature and precipitation are 22ºC and 1470 mm,

respectively, with a well pronounced dry winter (INMET,

2020).

The PFSP is composed of different vegetation types.

CER is predominant in the north and SSF in the south of

PFSP (Rossi et al., 2005). At the southernmost limit of the

PFSP, parallel to the Mogi Guaçu River, the predominant

vegetation type is the riparian forest (KONOPCZYK, 2014).

CER occupies an area of about 180 ha, where the topography

is higher (Bertoni et al., 2001). The study area has a secondary

vegetation that has greatly recovered its structure, as

evidenced by the sequential aerial photographs taken in the

1960s, 1970s, and 1980s (Figure 2).

2.2. Sampling and species distribution

We sampled 64 plots with 10x10 m (totalizing 0.64 ha),

alternately placed along eight transects (with eight plots each).

The delimitation of transects and plots was made by using

timber stakes and cotton strings. We sampled all living trees

and palms with a perimeter at breast height (PBH) > 10 cm.

The botanical material was identified by consulting

specialized bibliography and taxonomic specialists and by

comparing it with materials deposited in the Herbario

Rioclarense (HRCB) collection. Fertile material was

deposited in the HRCB collection. Taxa classification

followed APG IV (2016) and the nomenclature was updated

according to Flora do Brasil (Flora do Brasil, 2020), The Plant

List (2020), and Tropicos (2020). We assessed the threatened

status of the species sampled following the Livro Vermelho

da Flora do Brasil (MARTINELLI; MORAES, 2013).

For phytosociological analysis, we recorded the PBH

values and the height of each individual sampled. We

assessed the natural distribution of the species in Brazilian

biomes according to Flora do Brasil 2020 (2020). Species

occurring in four or more biomes were classified as “widely

distributed”.

Figure 1. Location of Porto Ferreira State Park (PFSP), Porto

Ferreira, São Paulo state. A- study site in São Paulo state map; B-

PFSP; C- vegetation types of PFSP. Red ellipse indicates the site of

the study plots.

Figura 1. Localização do Parque Estadual de Porto Ferreira (PEPF),

Porto Ferreira, estado de São Paulo. A- Local de estudo no mapa do

estado de São Paulo; B - PEPF; C- fitofisionomias do PEPF. A

elipse vermelha indica a localização das parcelas do estudo.

Figura 2. Fotografias aéreas do Parque Estadual de Porto Ferreira.

A- de 1962; B- de 1971; C- de 1983; D- imagem de satélite de 2020

(Google Earth©). Retângulos vermelhos indicam a localização das

parcelas do estudo.

Figure 2. Aerial photographs of Porto Ferreira State Park. A- from

1962; B- from 1971; C- from 1983; D- satellite image from 2020

(Google Earth©). Red rectangles indicate the site of the study plots.

Sabino et al.

Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

473

We compiled 19 studies previously carried out on CER,

cerrado sensu stricto, SSF, and ecotones including one of these

vegetation types (Table 1). Only native shrub and tree taxa

identified at the species level were considered (if the study

included more life forms).

2.3. Data analysis

Phytosociological parameters such as relative frequency,

dominance, and importance value were calculated following

Shepherd (2008). We also estimated the Shannon diversity

index (H´) and Pielou equability (J´). Finally, to assess sample

sufficiency, we performed a species accumulation curve

(SAC) to compute the mean SAC and its standard deviation

from 999 random permutations of the data (GOTELLI;

COLWELL, 2001).

We conducted a cluster analysis to assess floristic

similarity among the compiled studies and the present study.

We built a binary matrix with the presence (1) and absence

(0) of all species and conducted an agglomerative hierarchical

cluster analysis (UPGMA) using the Jaccard dissimilarity

index (LEGENDRE; LEGENDRE, 2012). Then, we

verified exclusive and shared species among the studies

carried out on ecotones.

All analyses were conducted in R software (R

Development Core Team, 2020) using the ‘vegan’ package

(OKSANEN et al., 2013).

Table 1. Selected studies for the floristic similarity analysis. Co: Code; Ph: physiognomy; N: number of taxons included in similarity analysis;

Nt: total number of taxons; St. R. do P. Quatro: Santa Rita do Passa Quatro; S.B. do Campo: São Bernardo do Campo; Ssf: seasonal

semideciduous forest; Eco: ecotone; Cd: cerradão; C ss: cerrado sensu stricto.

Tabela 1. Estudos selecionados para a análise de similaridade florística. Co: Código; Ph: fisionomia; N: número de táxons utilizados na

análise de similaridade; Nt: Número total de táxons; St. R. do P. Quatro: Santa Rita do Passa Quatro; S.B. do Campo: São Bernardo do

Campo; Ssf: floresta estacional semidecidual; Eco: ecótono; Cd: cerradão; C ss: cerrado sensu stricto.

Co Study Site Ph. N / Nt REFERENCE

Porto Ferreira

–

Eco

87/ 101

Present study

Bauru

–

Eco

223/ 264

Pinheiro & Monteiro, 2008

Brotas

–

Eco

116/

125

Gomes et al., 2004

Uberlandia

–

Eco

121/ 141

Guilherme & Nakajima, 2007

Viçosa

–

Ssf

93/ 107

Ribas et al., 2003

Campinas

–

Ssf

132/ 362

Bernacci & Leitão, 1996

Campinas

–

Ssf

135/ 201

Guaratini et al., 2008

St. R. do P.

Quatro

–

Ssf

56/ 56

Bertoni et al., 1988

St. R. do P. Quatro

–

Ssf

59/ 81

Vieira et al., 1989

Itatinga

–

Ssf

87/ 97

Ivanauskas et al., 1999

S. B. do Campo

–

Ssf

70/ 75

Pastore et al., 1992

São Carlos

–

Ssf

123/ 146

Silva &

Soares, 2003

Águas da Prata

–

Ssf

117/ 126

Toledo Filho et al., 1993

Uberlândia

–

73/ 74

Alves et al., 2013

St. R. do P. Quatro

–

81/ 148

Batalha & Mantovani, 2001

Luís Antonio

–

89/ 121

Silva et al., 2004

Bauru

–

126/ 135

Pinheiro & Monteiro, 2006

Mogi mirim

–

88/ 104

Toledo Filho et al., 1989

Uberlândia

–

Cd, C ss

103/ 107

Costa & Araújo, 2001

Paraopeba

–

Cd, C ss

81/ 91

Campos et al., 2006

3. RESULTS

We surveyed 1,755 trees and palms. We reported 101

species distributed in 37 botanical families and 69 genera

(Table 2). Euterpe edulis Mart. (Arecaceae) and Cedrela fissilis

Vell. (Meliaceae) were classified as “vulnerable” according to

the list of threatened species of the Livro Vermelho da Flora

do Brasil (Martinelli & Moraes, 2013). The Shannon-Wiener

diversity index was 3.65 nats/ind., and the Pielou equability

index was 0.79.

The richest species families were Myrtaceae (13 spp.),

Fabaceae (11 spp.), Annonaceae (6 spp.), Anacardiaceae (5

spp.), Melastomataceae, Rubiaceae, Euphorbiaceae, and

Lauraceae (4 spp. each). Altogether, these families comprised

50.4% of the species recorded and 54.3% of the individuals

sampled. Fifteen families were represented by a single

species, and 27 species were represented by a single

individual. Lauraceae was the family with the highest

importance value (IV= 28.9), followed by Anacardiaceae

(IV= 28.5), and Fabaceae (IV= 24.7).

Among the species, Tapirira guianensis Aubl. showed the

highest importance value (IV= 20), followed by Miconia affinis

DC. (IV= 18.7) and Siparuna guianensis Aubl. (IV= 18.4).

Siparuna guianensis was the most abundant species (188

individuals, 10.7% of the total amount of species) followed

by M. affinis (154 individuals, 8.7%), and Xylopia aromatica

(Lam.) Mart. (113 individuals, 6.4%). According to the

species accumulation curve, the sample size of our study was

sufficient to discuss differences among the areas (Figure 3).

The total basal area of the individuals was 11.65 m². The

mean height was 6.6 m (standard deviation= 2.9), with

minimum and maximum values of 1.9 and 22 m, respectively.

The mean diameter was 7.4 cm (standard deviation= 5.3),

with minimum and maximum values of 3.18 cm and 58.27

cm, respectively.

Out of the 101 species reported in this study, 87 (1,703

individuals) were identified at the species level and

classified based on their distribution among the Brazilian

biomes. Of this total, 38 species (44%) were classified as

“widely distributed”, and together they included 713

individuals (41.8%). Only five species (6%) and 31 individuals

(1.8%) are restricted to the Atlantic Forest. Likewise, six

species (7%) and 147 individuals (8.6%) are restricted to

Atlantic Forest meets the Cerrado: floristic, structure and species distribution of an ecotonal tree community

Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

474

Cerrado. Also, 31 species (36%) and 670 individuals (39.3%)

are reported for both Cerrado and Atlantic Forest.

The total amount of 644 species were sampled from the

20 lists chosen for the floristic similarity, of which 244

(37.8%) have been reported exclusively in a single study

(appendix 1). Only 26 species (4%) occurred in 50% or more

of the lists (Appendix 2), and no species were recorded for

all areas.

Among the ecotonal lists (A-D, Table 1), 341 species

were reported, of which 215 (63%) has exclusive occurrence

(appendix 3), 126 (36.9%) occur in 50% or more of the 4 lists,

and 19 species were reported for all areas (appendix 4).

The similarity analysis showed a high coefficient of

cophenetic correlation (0.91), and floristic similarities were

higher among closer areas. The cluster analysis segregated

two major floristic groups (Figure 4). The first group (1)

clustered the studies carried out on CER and ecotones,

including the present study; the second (2) clustered the

studies carried out on SSF.

Table 2. Floristic list, structural parameters, and biomes of occurrence of an ecotonal tree community, Porto Ferreira State Park, municipality

of Porto Ferreira, São Paulo State, southeastern Brazil. Abd: abundance; RelDo: relative dominance; RelFr: relative frequency; IV:

importance value; WD: widely distributed; Cer: Cerrado; AF: Atlantic Forest; Am: Amazon; Pa: Pampa; Caa: Caatinga; Pn: Pantanal; “\”:

not evaluated; asterisks indicate threatened species according to “Livro Vermelho da Flora do Brasil” by CNCFlora (Martinelli & Moraes,

2013).

Tabela 2. Lista florística, parâmetros estruturais e ocorrência das espécies nos domínios fitogeográficos de uma comunidade arbórea

ecotonal, Parque Estadual de Porto Ferreira, cidade de Porto Ferreira, Estado de São Paulo, Sudeste do Brasil. Abd: abundância; RelDo:

dominância relativa; RelFr: frequência relativa; IV: valor de importância; WD: ampla distribuição; Cer: Cerrado; AF: Mata Atlântica; Am:

Amazônia; Pa: Pampa; Caa: Caatinga; Pn: Pantanal; “\”: não avaliado; asteriscos indicam espécies ameaçadas de acordo com o “Livro

Vermelho da Flora do Brasil”, CNCFlora (Martinelli & Moraes, 2013).

Family Species Abd RelDo RelFr IV Biomes

Anacardiaceae Astronium graveolens Jacq. 7 0.21 0.71 1.32 WD

Astronium urundeuva (M.Allemão) Engl. 1 0.04 0.1 0.2 WD

Lithraea molleoides (Vell.) Engl. 1 0.02 0.1 0.18 WD

Tapirira guianensis Aubl. 82 11.01 4.34 20.02 WD

Tapirira obtusa (Benth.) J.D.Mitch. 26 4.34 1.72 7.53 Cer, AF, Am

Annonaceae Annona sylvatica A.St.-Hil. 1 0.02 0.1 0.18 WD

Annona sp. 1 0.05 0.1 0.21 \

Duguetia lanceolata A.St.-Hil. 4 0.16 0.4 0.79 Cer, AF

Xylopia aromatica (Lam.) Mart. 113 4.13 4.34 14.91 Cer, Am

Xylopia brasiliensis Spreng. 2 0.11 0.2 0.42 AF

Xylopia sericea A.St.-Hil. 25 1.15 1.72 4.29 Cer, AF, Am

Apocynaceae Aspidosperma sp. 1 0.04 0.1 0.2 \

Araliaceae Dendropanax cuneatus (DC.) Decne. & Planch. 1 0.04 0.1 0.19 WD

Didymopanax morototoni (Aubl.) Decne. & Planch. 17 0.7 1.41 3.08 WD

Arecaceae Acrocomia aculeata (Jacq.) Lodd. ex Mart. 1 0.3 0.1 0.46 Cer, AF

Euterpe edulis Mart. * 1 0.03 0.1 0.19 Cer, AF

Syagrus romanzoffiana (Cham.) Glassman 27 5.34 1.92 8.8 Cer, AF, Pa

Asteraceae Piptocarpha macropoda (DC.) Baker 2 0.04 0.2 0.35 Cer, AF

Bignoniaceae Handroanthus ochraceus (Cham.) Mattos 3 0.07 0.2 0.44 WD

Handroanthus vellosoi (Toledo) Mattos 11 0.91 1.01 2.55 AF

Burseraceae Protium heptaphyllum (Aubl.) Marchand 4 0.17 0.2 0.6 WD

Calophyllaceae Calophyllum brasiliense Cambess. 1 0.02 0.1 0.17 WD

Clethraceae Clethra scabra Pers. 2 0.12 0.2 0.43 Cer, AF, Caa

Combretaceae Terminalia argentea Mart. & Zucc. 3 0.19 0.3 0.66 WD

Terminalia glabrescens Mart. 42 1.88 2.02 6.29 WD

Euphorbiaceae Alchornea glandulosa Poepp. & Endl. 2 0.7 0.2 1.02 Cer, AF, Am

Croton floribundus Spreng. 10 0.9 0.91 2.38 AF

Maprounea guianensis Aubl. 59 2.44 3.63 9.44 Cer, AF, Am

Sebastiania sp. 3 0.15 0.3 0.62 \

Fabaceae Andira anthelmia (Vell.) Benth. 4 0.13 0.2 0.56 Cer, AF

Andira humilis Mart. ex Benth. 1 1 0.1 1.15 Cer

Copaifera langsdorffii Desf. 61 9.27 2.72 15.47 WD

Dimorphandra mollis Benth. 1 0.21 0.1 0.36 Cer, Am, Pn

Inga striata Benth. 1 0.09 0.1 0.25 WD

Leptolobium dasycarpum Vogel 1 0.06 0.1 0.22 Cer

Machaerium aculeatum Raddi 6 0.3 0.61 1.25 Cer, AF, Pn

Machaerium villosum Vogel 5 0.29 0.5 1.08 WD

Peltophorum dubium (Spreng.) Taub. 1 0.01 0.1 0.17 WD

Platypodium elegans Vogel 9 1.88 0.71 3.09 WD

Senegalia polyphylla (DC.) Britton & Rose 12 0.22 0.81 1.72 WD

Lacistemataceae Lacistema hasslerianum Chodat 6 0.07 0.61 1.02 Cer

Lamiaceae Aegiphila integrifolia (Jacq.) Moldenke 3 0.2 0.3 0.67 WD

Sabino et al.

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475

Family Species Abd RelDo RelFr IV Biomes

Lauraceae Ocotea corymbosa (Meisn.) Mez 77 7.47 4.24 16.1 Cer, AF

Ocotea nutans (Nees) Mez 24 0.66 1.92 3.94 Cer, AF

Ocotea pulchella (Nees & Mart.) Mez 59 4.22 3.83 11.42 Cer, AF, Pa

Ocotea velutina (Nees) Rohwer 1 0.29 0.1 0.45 Cer, AF

Malpighiaceae Byrsonima intermedia A.Juss. 85 3.93 4.04 12.81 Cer

Byrsonima sp. 3 0.13 0.3 0.6 \

Malvaceae Luehea divaricata Mart. 2 0.03 0.2 0.35 WD

Luehea grandiflora Mart. 4 0.92 0.4 1.55 WD

Pseudobombax grandiflorum (Cav.) A.Robyns 7 0.42 0.4 1.22 AF

Melastomataceae Miconia affinis DC. 154 4.86 5.15 18.78 Cer, AF, Am

Miconia ligustroides (DC.) Naudin 3 0.12 0.3 0.6 Cer, AF, Caa

Miconia rubiginosa (Bonpl.) DC. 1 0.02 0.1 0.17 Cer, Am

Tibouchina sp. 1 0.01 0.1 0.17 \

Meliaceae Cedrela fissilis Vell. * 2 0.11 0.2 0.42 WD

Trichilia casaretti C.DC. 5 0.07 0.3 0.66 Cer, AF

Trichilia pallida Sw. 44 1.19 2.22 5.92 Cer, AF, Am

Moraceae Ficus guaranitica Chodat 6 0.14 0.61 1.08 Cer, AF

Maclura tinctoria (L.) D.Don ex Steud. 1 0.01 0.1 0.17 WD

Myristicaceae Virola sebifera Aubl. 17 0.73 1.41 3.11 Cer, AF, Am

Myrtaceae Eugenia florida DC. 6 0.09 0.61 1.04 WD

Eugenia sp. 1 0.42 0.1 0.58 \

Myrcia bella Cambess. 35 2.01 2.02 6.03 Cer

Myrcia guianensis (Aubl.) DC. 19 0.69 1.31 3.08 WD

Myrcia sp. 24 0.41 1.51 3.29 \

Myrcia splendens (Sw.) DC. 11 0.19 1.01 1.82 WD

Myrcia tomentosa (Aubl.) DC. 20 0.72 1.61 3.47 WD

Myrcia venulosa DC. 10 0.22 1.01 1.8 Cer, AF

Myrciaria sp. 2 0.03 0.2 0.34 \

Myrtaceae 1 1 0.01 0.1 0.16 \

Myrtaceae 2 2 0.02 0.1 0.23 \

Myrtaceae 3 1 0.02 0.1 0.18 \

Psidium rufum Mart. ex DC. 15 0.79 1.11 2.75 Cer, AF

Ochnaceae Ouratea castaneifolia (DC.) Engl. 2 0.2 0.2 0.51 WD

Peraceae Pera glabrata (Schott) Baill. 45 2.69 2.42 7.67 WD

Primulaceae Myrsine coriacea (Sw.) R.Br. ex Roem. & Schult. 16 0.5 1.61 3.03 Cer, AF, Pa

Myrsine umbellata Mart. 100 3.33 4.54 13.57 WD

Proteaceae Roupala montana Aubl. 1 0.01 0.1 0.17 WD

Rosaceae Prunus myrtifolia (L.) Urb. 3 0.06 0.3 0.54 WD

Rubiaceae Amaioua guianensis Aubl. 21 0.47 1.72 3.38 Cer, AF, Am

Cordiera sessilis (Vell.) Kuntze 11 0.16 0.91 1.7 Cer, Caa

Ixora brevifolia Benth. 15 1.38 1.31 3.55 Cer, Caa

Rudgea viburnoides (Cham.) Benth. 1 0.02 0.1 0.18 Cer, Caa, Am

Rutaceae Zanthoxylum rhoifolium Lam. 2 0.14 0.2 0.46 WD

Salicaceae Casearia gossypiosperma Briq. 4 0.19 0.3 0.72 Cer, AF, Am

Casearia sp. 1 0.01 0.1 0.16 \

Casearia sylvestris Sw. 29 0.79 1.92 4.36 WD

Sapindaceae Allophylus semidentatus (Miq.) Radlk. 1 0.02 0.1 0.18 Cer, AF, Am

Cupania vernalis Cambess. 9 0.2 0.81 1.52 WD

Matayba sp. 10 0.77 0.81 2.15 \

Sapotaceae Chrysophyllum marginatum (Hook. & Arn.) Radlk. 9 0.41 0.71 1.63 WD

Pouteria sp. 1 0.03 0.1 0.19 \

Siparunaceae Siparuna guianensis Aubl. 188 2.18 5.55 18.44 WD

Styracaceae Styrax acuminatus Pohl 1 0.03 0.1 0.19 AF

Styrax camporum Pohl 16 0.71 1.51 3.13 Cer, AF, Caa

Styrax ferrugineus Nees & Mart. 2 0.06 0.2 0.38 Cer

Urticaceae Cecropia pachystachya Trécul 5 0.19 0.4 0.88 WD

Vochysiaceae Qualea cordata Spreng. 7 0.54 0.71 1.64 Cer, AF, Caa

Qualea grandiflora Mart. 17 1.67 1.41 4.06 Cer

Vochysia tucanorum Mart. 31 3.32 1.31 6.4 Cer, AF

Atlantic Forest meets the Cerrado: floristic, structure and species distribution of an ecotonal tree community

Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

476

Figure 3. Species accumulation curve generated by using the

sampling random method for the tree community surveyed in the

ecotonal area of Porto Ferreira State Park, Porto Ferreira, Brazil.

Figura 3. Curva de acumulação de espécies geradas para a

comunidade arbórea amostrada em área ecotonal, Parque Estadual

de Porto Ferreira, Porto Ferreira, Brasil.

Figure 4. Cluster analysis (UPGMA), using Jaccard’s index, of 20

studies carried out on Cerrado, Atlantic Forest, and ecotones. A –

the present study. A-T – previous selected studies. The list of the

selected studies is given in Table 1.

Figura 4. Análise de agrupamento (UPGMA) usando o índice de

Jaccard entre 20 estudos conduzidos no Cerrado, Mata Atlântica e

ecótonos. Letra A- presente estudo. Estudos (identificados pelas

letras A-T) estão disponíveis na tabela 1.

4. DISCUSSION

The tree diversity we found in this survey was similar

from what was found in previous studies carried out on

Cerrado (Toledo Filho et al.,1989; Costa & Araújo, 2001;

Silva et al., 2004), SSF (Vieira et al., 1989; Ivanauskas et al.,

1999), and ecotones (Gomes et al., 2004; Guilherme;

Nakajima, 2007). However, diversity comparisons should be

made with caution, given the differences in inclusion criteria

and sample sizes among the studies. Studies encompassing a

big range of life forms (e.g., Bernacci; Leitão, 1996; Batalha;

Mantovani, 2001; Guaratini et al., 2008; Pinheiro; Monteiro,

2008) naturally show a greater number of species.

The diversity value (3.65) found in the present study site

is higher than those of studies on CER (which range from

2.46 to 3.54) (Costa; Araújo, 2001; Alves et al., 2013) and

higher than the expected for SSF (ranging from 3.45 to 3.77)

(Ivanauskas, 1999; Silva; Soares, 2003), and ecotone sites

(ranging from 3.37 to 3.99) (Gomes et al., 2004; Guilherme;

Nakajima, 2007). This shows that the diversity in transitional

areas can be greater than what has been found in underlying

vegetation types.

The richest families of tree species corroborate the results

found for CER (Silva et al., 2004; Campos et al., 2006), SSF

(Bernacci; Leitão, 1996; Ivanauskas et al., 1999; Guaratini et

al., 2008), and ecotones (Gomes et al., 2004; Guilherme;

Nakajima, 2007; Pinheiro; Monteiro, 2008). In SSF surveys,

Fabaceae and Myrtaceae are often the richest families (Vieira

et al., 1989; Bernacci; Leitão, 1996; Ivanauskas et al., 1999;

Guaratini et al., 2008). Anacardiaceae is the only family of our

list rarely ranked among the most diverse family in other

studies. In Atlantic Forest, Myrtaceae, Fabaceae,

Melastomataceae, Rubiaceae, and Euphorbiaceae are among

the 10 richest families (BFG, 2015). These families are also

among the richest families in Cerrado, except for Myrtaceae

(BFG, 2015).

Lauraceae was the family with the highest importance

value (IV= 9.64%) due to the large dimensions of the

individuals and the abundance of Ocotea corymbosa and Ocotea

pulchella. Likewise, Anacardiaceae showed the second highest

IV (9.5%) due to the great number of individuals of Tapirira

guianensis and Tapirira obtusa surveyed and their big sizes.

Many species were represented by a few individuals,

corroborating the distribution pattern of individuals of

tropical forest species (HUBBEL, 2013). In that sense, 45

species were represented by three or fewer individuals, 27 of

which were represented by a single individual (e.g., Andira

humilis, Acrocomia aculeata, Calophyllum brasiliense, and Ocotea

velutina). The presence of rare or naturally low-density species

indicates that new taxa tend to be found as the sample area

increases, as demonstrated by Condit et al. (1996).

Regarding the most important species, Tapirira guianensis,

has been reported in several studies carried out on different

vegetation types and is considered a generalist species

(Oliveira-Filho; Fontes, 2000). Siparuna guianensis, the most

abundant species in the present study, has been frequently

sampled in CER vegetation (e.g., Fina; Monteiro, 2009;

Pereira-Silva et al., 2004) and reported in studies carried out

on SSF (e.g., Joly, 1994; Silva et al., 2004; Yamamoto et al.,

2005).

The basal area of the individuals we found was low (e.g.,

Toledo Filho et al., 1989; Vieira et al., 1989; Ivanauskas et al.,

1999; Costa; Araújo, 2001; Guilherme; Nakajima, 2014). This

fact may be related to anthropic disturbances suffered in the

past, such as selective cutting, as showed in Figure 2 for the

area of the present study. This supports that low basal area

values are found in areas with selective cutting history

(ALVES et al., 2013).

Many species of our study have wide distribution,

occurring in different Brazilian biomes, which was also

reported by Gomes et al. (2004) for another ecotonal area.

Durigan et al. (2012) highlighted that generalist species with

high ecological plasticity are generally abundant in ecotonal

Sabino et al.

Nativa, Sinop, v. 9, n. 4, p. 471-480, 2021.

477

areas. The predominance of species with wide distribution

and the presence of species occurring in both Cerrado and

Atlantic Forest reinforce that the area we studied is, in fact,

an ecotone. On the other hand, few species are restricted to

the Cerrado domain, e.g., Qualea grandiflora, Byrsonima

intermedia, and Myrcia bella; and to the Atlantic Forest, e.g.,

Xylopia brasiliensis, Handroanthus vellosoi, and Pseudobombax

grandiflorum (Flora do Brasil, 2020).

The shared species among ecotonal studies (Appendix 4),

such as Copaifera langsdorffii, Ocotea corymbosa, Platypodium

elegans, Terminalia glabrescens, and Vochysia tucanorum, can be

indicator species of Cerrado-Atlantic Forest ecotones from

southeastern Brazil (DURIGAN et al., 2012).

Changes in the vegetation type of the study site over time

(e.g., Figure 2), with a putative replacement of species of SSF

over those of CER, indicate that fire suppression and soil

conditions allowed the development of both floristics unities.

The ecotonal area, was composed of savannah

physiognomies with species of Cerrado (BERTONI et al.,

2001; MATTOS; ROCHA, 2002). Currently, it is mainly

composed of CER and SSF. Studies have indicated that fire

is an important modelling agent in the physiognomies of

Cerrado (HENRIQUES, 2005; HOFFMAN; MOREIRA,

2002; PINHEIRO et al., 2021). Fire suppression can replace

areas initially occupied by open forms of Cerrado by CER

and SSF over the years (DURIGAN et al., 1987; DURIGAN;

RATTER 2006). A common action taken in protected areas

in Brazil is to extinguish fire events (Monteiro, personal

communication), which naturally occur in Cerrado

(FIDELIS, 2020).

The analysis of floristic relationships among the

compared areas showed a marked distinction between

Cerrado and SSF, but placed ecotonal areas closer to the

Cerrado or as a subgroup. Although, species of the SSF can

colonize areas with fire suppression in Cerrado, the

transformation of the vegetation type can be constrained by

the soil type (HARIDASSAN, 1992; ROSSI et al., 2005;

RODRIGUES et al., 2016; BARROS et al., 2018).

Furthermore, we observed a great floristic heterogeneity

among the analyzed areas, which included many exclusive

and few shared species. Even though communities were

sampled in geographically close areas and had the same

vegetation types, they had few shared species. Our results

support the hypothesis that tropical tree communities have a

great floristic heterogeneity over short geographic distances

(HUBBEL, 2013; BUENO et al., 2018). In addition, our

results indicate that ecotonal tree communities may shelter a

heterogeneous species composition and have species from

different vegetation types. However, geographically closer

communities share more species.

5. CONCLUSIONS

In this study, we provided important data about a poorly

known vegetation type, corroborating the importance of

preserving ecotonal areas for maintaining the high diversity

of tropical tree communities. Our data can also help to

recognize ecotones between Cerrado and Atlantic Forest

from southeastern Brazil and can be used as a floristic

reference for future conservation and restorative measures.

6. ACKNOWLEDGMENTS

We would like to thank Marco Antonio de Assis and

Alessandra Fidelis for all suggestions; and the PFSP

employees for the technical support.

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8. APPENDIX

Appendix 1. Numbers of species reported exclusively in this and

each one of the previous studies. Co: code; Vt: vegetation type; E

ssp.: number of exclusive species; Ssf: seasonal semideciduous

forest; Cd: cerradão; C ss: cerrado sensu stricto.

Apêndice 1. Número de espécies exclusivas entre os estudos

selecionados para a análise de similaridade florística. Co: código; Vt:

Fitofisionomia; E spp: número de espécies exclusivas; Ssf: floresta

estacional semidecidual; Cd: cerradão; C ss: cerrado sensu stricto.

Reference Vt E spp.

A Present study Ecotone 5

B Pinheiro & Monteiro 2008 Ecotone 30

C Gomes et al. 2004 Ecotone 10

D Guilherme & Nakajima 2007 Ecotone 14

E Ribas et al. 2003 Ssf 32

F Bernacci & Leitão 1996 Ssf 21

G Guaratini et al. 2008 Ssf 22

H Bertoni et al. 1988 Ssf 1

I Vieira et al. 1989 Ssf 0

J Ivanauskas et al. 1999 Ssf 16

K Pastore et al. 1992 Ssf 24

L Silva & Soares 2003 Ssf 18

M Toledo Filho et al. 1993 Ssf 20

N Alves et al. 2013 Cd 1

O Batalha & Mantovani, 2001 Cd 6

P Silva et al. 2004 Cd 4

Q Pinheiro & Monteiro 2006 Cd 1

R Toledo Filho et al. 1989 Cd 7

S Costa & Araújo 2001 Cd, C ss 5

T Campos et al. 2006 Cd, C ss 7

Appendix 2. Species occurring in 10 (50%) or more lists among the

compared studies and PFSP. N: number of lists in which the species

occurs.

Appendix 2. Espécies com ocorrência igual ou superior a 10 listas

(50%) entre os estudos selecionados para a análise de similaridade

florística. N: número de listas com ocorrência.

Species N %

Roupala montana Aubl. 18 90

Copaifera langsdorffii Desf. 16 80

Qualea multiflora Mart. 16 80

Tapirira guianensis Aubl. 14 70

Casearia sylvestris Sw. 13 65

Myrcia splendens (Sw.) DC. 13 65

Ocotea corymbosa (Meisn.) Mez 13 65

Vochysia tucanorum Mart. 13 65

Siparuna guianensis Aubl. 12 60

Dimorphandra mollis Benth. 11 55

Lafoensia pacari A.St.-Hil. 11 55

Miconia albicans (Sw.) Triana 11 55

Myrcia tomentosa

(Aubl.) DC. 11 55

Myrsine umbellata Mart. 11 55

Pera glabrata (Schott) Baill. 11 55

Plathymenia reticulata Benth. 11 55

Platypodium elegans Vogel 11 55

Qualea grandiflora Mart. 11 55

Rudgea viburnoides (Cham.) Benth. 11 55

Xylopia aromatica (Lam.) Mart. 11 55

Bowdichia virgilioides Kunth 10 50

Brosimum gaudichaudii Trécul 10 50

Caryocar brasiliense Cambess. 10 50

Croton floribundus Spreng. 10 50

Protium heptaphyllum (Aubl.) Marchand 10 50

Terminalia glabrescens Mart. 10 50

Appendix 3. Numbers of exclusive species among the studies

carried out on ecotonal vegetation. Co: code; Vt: vegetation type; E

ssp.: number of exclusive species; Ssf: seasonal semideciduous

forest; Cd: cerradão; C ss: cerrado sensu stricto.

Apêndice 3. Número de espécies exclusivas entre os estudos

selecionados para a análise de similaridade florística. Co: código; Vt:

fitofisionomia; E spp: número de espécies exclusivas; Ssf: floresta

estacional semidecidual; Cd: cerradão; C ss: cerrado sensu stricto.

REFERENCE Vt E spp.

A Present study Ecotone 18

B Pinheiro & Monteiro 2008 Ecotone 125

C Gomes et al. 2004 Ecotone 30

D Guilherme & Nakajima 2007 Ecotone 46

Appendix 4. Shared species among ecotonal vegetation lists.

Apêndice 4. Espécies compartilhadas nas listas de vegetação

ecotonal.

Species

Casearia sylvestris Sw.

Copaifera langsdorffii Desf.

Dimorphandra mollis Benth.

Myrcia tomentosa (Aubl.) DC.

Myrsine umbellata Mart.

Ocotea corymbosa (Meisn.) Mez

Pera glabrata (Schott) Baill.

Platypodium elegans Vogel

Protium heptaphyllum (Aubl.) Marchand

Qualea grandiflora Mart.

Roupala montana Aubl.

Rudgea viburnoides (Cham.) Benth.

Siparuna guianensis Aubl.

Styrax camporum Pohl

Tapirira guianensis Aubl.

Terminalia glabrescens Mart.

Vochysia tucanorum Mart.

Xylopia aromatica (Lam.) Mart.

Zanthoxylum rhoifolium Lam.