International Journal of Forestry Research

International Journal of Forestry Research / 2012 / Article

Research Article | Open Access

Volume 2012 |Article ID 537269 | 14 pages |

An Ecological Comparison of Floristic Composition in Seasonal Semideciduous Forest in Southeast Brazil: Implications for Conservation

Academic Editor: Frank Gilliam
Received30 Mar 2011
Revised06 May 2011
Accepted09 May 2011
Published06 Jul 2011


We examined floristic patterns of ten seasonal semideciduous forest sites in southeastern Brazil and conducted a central sampling of one hectare for each site, where we took samples and identified all individual living trees with DBH (diameter at breast height, 1.30 m) ≥4.8 cm. Arboreal flora totaled 242 species, 163 genera, and 58 families. Fabaceae (38 species) and Myrtaceae (20 species) were families with the largest number of species. Only Copaifera langsdorffii and Hymenaea courbaril occurred at all sites. Multivariate analysis (detrended correspondence analysis and cluster analysis) using two-way indicator species analysis (TWINSPAN) indicated the formation of a group containing seven fragments in which Siparuna guianensis was the indicator species. This analysis revealed that similarities between studied fragments were due mainly to the successional stage of the community.

1. Introduction

The extent of seasonal semideciduous forests (SSFs) in Brazil is underestimated because of its naturally fragmented distribution [1, 2]. Semideciduous seasonal forests occur along the contact zone between Atlantic forest and the diagonal of opened formations [35], comprising three different scenarios: (1) in northeastern Brazil, the SSFs form a marked belt (<50 km) in transition between the coastal rainforest and semiarid formations (Caatinga), but also occur in enclaves of montane forests, the altitude marsh [6, 7]; (2) the transition between the Cerrado and the coastal Atlantic forest in southeastern Brazil involves an extensive occurrence of SSFs to the south, up to the east of Paraguay and northeast of Argentina, forming a complex mosaic with the Cerrado vegetation in the west; (3) in southeastern Brazil, a large Araucaria forest confronts subtropical coastal Atlantic forest, and the SSF appear to the west and south as a transition to the Chaco forests, and to the southeast with fields or the southern pampas [8, 9], and beyond disjunct areas located in Mato Grosso and Tocantins states [10].

In southeast Brazil, SSFs are distributed widely in sites with a seasonal rainfall regime, characteristic of the Atlantic forest and Cerrado domains. In the Atlantic forest domain, SSF is the predominant typology, and in the Cerrado domain, SSF occurs in enclaves, associated with permanent or intermittent watercourses [12] and should be regarded as lato sensu Atlantic forest, since it presents a floristic-structural identity similar to forests of the Atlantic forest domain [12].

Semideciduous seasonal forests suffered the same degradation process as other Brazilian ecosystems. Since the 1970s, there was an accelerated replacement of natural vegetal formations to pasture and use for agriculture, transforming extensive areas into an important agriculture area for grain and fruit production and livestock [13]. This rural reorganization was determined by the II National Development Plan (NDP), which collaborated for a modern agricultural deployment. With increasingly intense adoption of mechanization and land use, SSFs were drastically reduced, with only a few remaining in Southeast Brazil.

Habitat fragmentation transforms the original landscape into different dynamic units that continually modify its structure [14]. Furthermore, the occurrence of disturbance histories on different scales and heterogeneity environmental landscapes can influence the species composition of forest fragments [15] and form species richness patterns according to the fragment successional stage [16, 17].

Therefore, owing to the similar process of disturbance in southeastern Brazilian forest formations and being located in the same catchment area, it is expected that the remaining SSF will have the same floristic pattern.

This work aims to analyze the floristic composition and regional richness in the arboreal species of ten fragments of SSF in southeastern Brazil, in order to answer the following questions: (1) what is the alpha diversity in sampled fragments? (2) is there a floristic pattern that represents the SSF in the studied region? (3) is the beta diversity in SSF similar to patterns found in other tropical forests?

2. Material and Methods

2.1. Floristic, Geographic, and Climate Data

We elaborate a floristic list from the tree species compilation in ten semideciduous seasonal forest fragments (according to Veloso classification [18]), distributed in five counties of southeastern Brazil (Table 1). Studied sites are located in the extreme west of Minas Gerais state, defined by geographic coordinates 18° 29′–19°40′ S and 47°30′– 49°53′ W (Figure 1).

SiteCountiesArea (ha)Latitude (S)Longitude (W)Altitude (m.a.s.l.)

1Araguari20018° 29′ 5048° 23′ 03680
2Ipiaçu4018° 43′ 3949° 56′ 22530
3Monte Carmelo11918° 44′ 5947° 30′ 56910
4Uberaba7019° 40′ 3548° 02′ 12790
5Uberlândia17,518° 40′ 2648° 24′ 32600
6Uberlândia3018° 57′ 0348° 12′ 22880
7Uberlândia22,319° 08′ 3948° 08′ 46930
8Uberlândia1619° 10′ 0448° 23′ 41800
9Uberlândia3518° 55′ 4048° 03′ 51890
10Uberlândia2018° 51′ 3548° 13′ 53890

The studied region is part of a relief global set, named the “Domain of Central Brazil Tropical Plateaus” by Ab'Saber [19], and the “Plateaus and elevated plains of the Paraná sedimentary basin” by the RADAM project [20]. The soils in the studied areas are predominantly red nonferric latossols (LV) [21]. The predominant climate in the studied region is tropical savanna (Aw Megathermic), characterized, according to Köppen classification [22], by rainy summers and dry winters. The clearly seasonal climate has two well-defined seasons, where the winter season (April to September) has approximately six months of drought, and the summer season (October to March) is warm and rainy. The average annual temperature lies between 23°C and 25°C, with July the month with the lowest average temperature (16°C). The annual rainfall ranges from 1160 to 1460 mm [23].

2.2. Data Collection and Analysis

We conducted a central sampling of one hectare from each fragment of SSF, in an attempt to exclude edge effect and to obtain uniform samples, avoiding the ecotones of the adjacent formations. Central sampling gave priority to sites without the influence of watercourses (gallery forest), savanna formations (cerrado sensu stricto), and others forestry formations (dry seasonal forest and cerradão), since these vegetation contact areas vary considerably between fragments and would lead to changes in the species listing of each area. In each central sampling, all individual living trees with a DBH (diameter at breast height, 1.30 m) of ≥ 4.8 cm were recorded and identified.

We identified the species using literature, queries in herbarium, and specialists. For specific binomial formation, we employed the w3 Tropicos database [24]. We deposited fertile material to the Uberlandia Federal University herbaria (Herbarium Uberlandensis: HUFU). Vegetative samples of all species were deposited at the Laboratory of Plant Ecology at the same university. Species were classified into families, according to the Angiosperm Phylogeny Group III system [25].

We compiled all species present in the inventories to undertake the floristic composition analysis. We carried out the analysis by preparing a binary database (presence/absence), containing the tree species listed in each fragment. We only considered individual trees identified at the species level in this database composition. For the Alpha diversity evaluation, we used the Shannon diversity index (H′) and the Pielou evenness index (J′).

We used the binary database for the measurement of richness analysis, and also for the total number of species projection, we used first- and second-order, nonparametric jackknife estimators, in addition to Chao and Chao 2 estimators, which were capable of projecting total species richness from the samples of species richness [26], with 5,000 randomizations. We conducted the analysis using the EstimateS 8.0 program [27].

2.3. Multivariate Analyses

We did similarity and sorting analysis utilizing the FITOPAC SHELL 1.6.4 program [28], using an absolute density matrix for the species of ten sites, considering only those species with two or more occurrences, since species that had just one occurrence did not contribute to the floristic site ordering for floristic similarity. For categorical floristic data (presence and absence), we analyzed the similarity between sites, using the same absolute density matrix, transformed into presence/absence. We used the Sørensen similarity coefficient [29] and the unweighted pair group method with arithmetic mean (UPGMA) for a dendrogram graphical representation.

We applied multivariate analysis to quantitative tree species floristic data (abundance) present in all the compared fragments. For this, we realized a data ordination by using the detrended correspondence analysis (DCA, [30]). In addition, to define predominantly the indicator and preferred species of the floristic groups, based on frequency and density (species with individuals), we used dichotomous hierarchical division using TWINSPAN (two-way indicator species analysis, [31]), from an absolute data matrix and its frequency at the 10 sites, with a cut level of 0, 2, 5, and 10. These analyses were made by PC-ord for Windows program version 4.0 [32].

3. Results

A total of 242 species, distributed in 165 genera and 58 families, were found in the SSF (Table 2). Families with a higher species richness were Fabaceae (sensu lato) (38 species), subdivided into Fabaceae Faboideae (20), Fabaceae Mimosoideae (11), Fabaceae Caesalpinioideae (5) and Fabaceae Cercideae (2), Myrtaceae (20 species), Rubiaceae (13), Annonaceae (11), Moraceae (10), Lauraceae (9), Meliaceae (9), and Malvaceae (8). These eight families showed a great contribution to the regional tree diversity, covering 48.8% of the species. Twenty-six families were represented by only one species. Annonaceae, Apocynaceae, Combretaceae, Fabaceae, Mimosoideae, Fabaceae, Faboideae, Lauraceae, Myrtaceae, Meliaceae, Rubiaceae, Salicaceae, Sapindaceae, and Sapotaceae were found at all ten studied sites.




Most well-represented genera are in Fabaceae Faboideae (13), Rubiaceae (13), Myrtaceae (10), Annonaceae (8), and Euphorbiaceae, Fabaceae Mimosoideae, and Malvaceae, with seven genera each. Approximately 30% of families were represented by only one genus. The best represented genera in terms of species number were Aspidosperma, Machaerium, and Ficus, with seven species each, followed by Ocotea, Cordia, Inga, Trichilia, and Casearia with four species each.

The total species richness, designed by jackknife estimators 1 and 2 and Chao 1 and 2 estimators, showed a similar pattern (273, 284, 285, and 268 species, resp.). Results suggest high regional tree species richness. The 242 species found approximates of richness designed for estimators, demonstrating sampling sufficiency among the ten hectares studied (Figure 2).

The Shannon’s diversity index (H′) for each site showed diversity values between 2.92 and 3.97, with the Pielou’s evenness index (J′) between 0.73 and 0.87 (Table 2). However, when we considered the ten sites as a single sample, Shannon’s diversity index (H′) was 4.62, and Pielou’s evenness index was 0.84. This value can be attributed to the sampling covering quite heterogeneous sites, allowing differences in remnant structure and floristic composition and, consequently, increasing beta diversity.

Only Copaifera langsdorffii and Hymenaea courbaril were presented with the highest frequency in distribution, reaching 100% of occurrence at the sites (Table 3). However, 75 species occurred in at least five studied fragments, highlighting Cordiera sessilis, Apuleia leiocarpa, Casearia gossypiosperma, Cheiloclinium cognatum, Ixora brevifolia, Luehea grandiflora, Protium heptaphyllum, Sweetia fruticosa, and Terminalia glabrescens which occurred in nine sites, and can be indicated as characteristic SSF species (Table 3). On the other hand, 31.8% (77) of species occurred at only one site.

Families/SpeciesNIRF (%)

Astronium fraxinifolium Schott ex Spreng.1840
Astronium nelson-rosae Santin25760
Lithraea molleoides (Vell.) Engl.920
Myracrodruon urundeuva Allemão4240
Tapirira guianensis Aubl.3720
Tapirira obtusa (Benth.) J.D.Mitch.8550
Annona cacans Warm.2950
Annona montana Macfad.*110
Cardiopetalum calophyllum Schltdl.640
Duguetia lanceolata A. St.-Hil.22260
Guatteria australis A. St.-Hil.*2810
Porcelia macrocarpa (Warm.) R. E. Fr.*110
Rollinia sylvatica (A. St.-Hil.) Mart.530
Unonopsis lindmanii R. E. Fr.10240
Xylopia aromatica (Lam.) Mart.3160
Xylopia brasiliensis Spreng.3740
Xylopia sericea A. St.-Hil.110
Aspidosperma cuspa (Kunth) S. F. Blake ex Pittier1010
Aspidosperma cylindrocarpon Müll. Arg.2540
Aspidosperma discolor A. DC.*29450
Aspidosperma olivaceum Müll. Arg.810
Aspidosperma parvifolium A. DC.*2850
Aspidosperma polyneuron Müll. Arg.830
Aspidosperma subincanum Mart. ex A. DC.1950
Ilex cerasifolia Reissek310
Aralia warmingiana (Marchal) J. Wen1230
Dendropanax cuneatus (DC.) Decne. & Planch.1420
Schefflera morototoni (Aubl.) Maguire et al.2980
Acrocomia aculeata (Jacq.) Lodd. ex Mart.510
Piptocarpha macropoda Baker630
Handroanthus serratifolia (Vahl) Nicholson2270
Jacaranda cuspidifolia Mart. ex A. DC.220
Jacaranda macrantha Cham.*420
Tabebuia impetiginosus (Mart. ex DC.) Mattus220
Tabebuia roseoalba (Ridl.) Sandwith1320
Cordia alliodora (Ruiz & Pav.) Oken**110
Cordia sellowiana Cham.5250
Cordia superba Cham.1310
Cordia trichotoma (Vell.) Arrab. ex Steud.520
Protium heptaphyllum (Aubl.) Marchand28290
Celtis iguanaea (Jacq.) Sarg.3060
Trema micrantha (L.) Blume320
Citronella paniculata (Mart.) R.A.Howard410
Jacaratia spinosa (Aubl.) A. DC.420
Cheiloclinium cognatum (Miers.) A. C. Sm.40790
Maytenus floribunda Reissek*7160
Maytenus robusta Reissek410
Maytenus sp.510
Salacia elliptica (Mart. ex Schult.) G. Don110
Hirtella glandulosa Spreng.5140
Hirtella gracilipes (Hook. f.) Prance7250
Hirtella racemosa Lam.1110
Calophyllum brasiliense Cambess.110
Garcinia brasiliensis Mart.6460
Terminalia argentea (Cambess.) Mart.210
Terminalia glabrescens Mart.20290
Terminalia phaeocarpa Eichler5660
Lamanonia ternata Vell.1220
Diospyros hispida A. DC.10850
Sloanea monosperma Vell.1940
Erythroxylum daphnites Mart.610
Erythroxylum deciduum A. St.-Hil.310
Acalypha gracilis (Spreng.) Müll. Arg.**720
Alchornea glandulosa Poepp. & Endl.2030
Mabea fistulifera Mart.2510
Maprounea guianensis Aubl.3240
Micrandra elata Müll. Arg.11810
Pera glabrata (Schott) Poepp. ex Baill.1220
Sapium glandulosum (L.) Morong530
Fabaceae caesalpinoideae
Apuleia leiocarpa (Vogel) J. F. Macbr.11490
Cassia ferruginea (Schrad.) Schrad. ex DC.930
Copaifera langsdorffii Desf.155100
Hymenaea courbaril L.119100
Peltophorum dubium (Spreng.) Taub.*220
Sclerolobium paniculatum Benth.310
Fabaceae cercideae
Bauhinia rufa (Bong.) Steud.940
Bauhinia ungulata L.*2050
Fabaceae faboideae
Andira fraxinifolia Benth.720
Andira ormosioides Benth.*110
Centrolobium tomentosum Guillem. ex Benth.110
Dipteryx alata Vogel520
Lonchocarpus cultratus (Vell.) Az.-Tozzi & H. C. Lima1130
Machaerium acutifolium Vogel730
Machaerium brasiliense Vogel6970
Machaerium hirtum (Vell.) Stellfeld2530
Machaerium nyctitans (Vell.) Benth.210
Machaerium opacum Vogel110
Machaerium stipitatum (DC.) Vogel2060
Machaerium villosum Vogel6260
Myroxylon peruiferum L. f.110
Ormosia arborea (Vell.) Harms2370
Platycyamus regnellii Benth.5030
Platypodium elegans Vogel3270
Pterodon emarginatus Vogel210
Sweetia fruticosa Spreng.6990
Vatairea macrocarpa (Benth.) Ducke110
Zollernia ilicifolia (Brongn.) Vogel2140
Fabaceae Mimosoideae
Acacia polyphylla DC.3770
Albizia niopoides (Spruce ex Benth.) Burkart2330
Albizia polycephala (Benth.) Killip ex Record510
Anadenanthera colubrina (Vell.) Brenan6720
Calliandra foliolosa Benth.510
Enterolobium contortisiliquum (Vell.) Morong940
Inga laurina (Sw.) Willd.950
Inga marginata Willd.1010
Inga sessilis (Vell.) Mart.*4460
Inga vera Willd.8440
Piptadenia gonoacantha (Mart.) J. F. Macbr.16750
Lacistema aggregatum (P. J. Bergius) Rusby**320
Aegiphila sellowiana Cham.1150
Vitex polygama Cham.620
Cryptocarya aschersoniana Mez21270
Endlicheria paniculata (Spreng.) J. F. Macbr.310
Nectandra cissiflora Nees4430
Nectandra megapotamica (Spreng.) Mez2950
Nectandra membranacea (Sw.) Griseb.*8140
Ocotea corymbosa (Meisn.) Mez13670
Ocotea minarum (Nees) Mez410
Ocotea pulchella Mart.810
Ocotea spixiana (Nees) Mez5140
Cariniana estrellensis (Raddi) Kuntze5170
Lafoensia densiflora Pohl110
Byrsonima laxiflora Griseb.1240
Apeiba tibourbou Aubl.430
Ceiba speciosa (A. St.-Hil.) Ravenna1940
Eriotheca candolleana (K. Schum.) A. Robyns2850
Guazuma ulmifolia Lam.4950
Luehea divaricata Mart.620
Luehea grandiflora Mart. & Zucc.19990
Pseudobombax tomentosum (Mart. & Zucc.) A. Robyns320
Quararibea turbinata (Sw.) Poir.*510
Miconia cuspidata Mart. ex Naudin110
Miconia latecrenata (DC.) Naudin1030
Miconia minutiflora (Bonpl.) DC.*710
Cabralea canjerana (Vell.) Mart.740
Cedrela fissilis Vell.1640
Guarea guidonia (L.) Sleumer2350
Guarea kunthiana A. Juss.1920
Trichilia catigua A. Juss.16580
Trichilia claussenii C. DC.13020
Trichilia elegans A. Juss.5570
Trichilia pallida Sw.2870
Mollinedia widgrenii A. DC.630
Ficus clusiifolia Schott*210
Ficus guaranitica Chodat640
Ficus obtusiuscula (Miq.) Miq.110
Ficus pertusa L. f.*110
Ficus trigona L. f.320
Ficus sp1110
Ficus sp 2110
Maclura tinctoria (L.) Steud.430
Pseudolmedia laevigata Trécul520
Sorocea bonplandii (Baill.) W. C. Burger et al.940
Virola sebifera Aubl.12780
Calyptranthes clusiifolia O. Berg530
Calyptranthes widgreniana O. Berg720
Campomanesia guazumifolia (Cambess.) O. Berg110
Campomanesia velutina (Cambess.) O. Berg9370
Eugenia florida DC.19050
Eugenia involucrata DC.4240
Eugenia ligustrina (Sw.) Willd.2730
Eugenia subterminalis DC.*2020
Gomidesia lindeniana O. Berg110
Myrcia splendens (Sw.) DC.1160
Myrcia tomentosa (Aubl.) DC.940
Myrciaria glanduliflora (Kiaersk.) Mattos & D. Legrand**10640
Myrciaria tenella (DC.) O. Berg210
Psidium longipetiolatum D. Legrand**210
Psidium rufum DC.1240
Psidium sartorianum (O. Berg) Nied.4150
Siphoneugena densiflora O. Berg13360
Syzygium jambos (L.) Aston.*110
Myrtaceae 1110
Myrtaceae 2110
Guapira opposita (Vell.) Reitz2810
Guapira venosa (Choisy) Lundell5440
Neea hermaphrodita S. Moore**1210
Ouratea castaneifolia (DC.) Engl.1250
Heisteria ovata Benth.11360
Chionanthus trichotomus (Vell.) P. S. Green310
Agonandra brasiliensis Miers ex Benth. & Hook.2450
Margaritaria nobilis L. f.2770
Phyllanthus acuminatus Vahl210
Piper amalago L.410
Piper arboreum Aubl.210
Coccoloba mollis Casar.940
Ardisia ambigua Mez2530
Myrsine coriacea (Sw.) Roem. & Schult.210
Myrsine leuconeura Mart.220
Myrsine umbellata Mart.1520
Roupala brasiliensis Klotzsch3180
Rhamnidium elaeocarpum Reissek3450
Amaioua guianensis Aubl.3540
Chomelia pohliana Mull. Arg.1440
Cordiera sessilis (Vell.) Kuntze40490
Coussarea hydrangeifolia (Benth.) Müll. Arg.3260
Coutarea hexandra (Jacq.) K. Schum.2170
Faramea hyacinthina Mart.4350
Genipa americana L.110
Guettarda viburnoides Cham. & Schltdl.2950
Ixora brevifolia Benth.13190
Machaonia brasiliensis (Hoffmanss. ex Humb.) Cham. & Schltdl.*210
Rudgea viburnoides (Cham.) Benth.1150
Simira sampaioana (Standl.) Steyerm.*4770
Tocoyena formosa (Cham. & Schltdl.) K.Schum.110
Galipea jasminiflora (A. St.-Hil.) Engl.*14210
Metrodorea nigra A. St.-Hil.1020
Metrodorea stipularis Mart.610
Pilocarpus spicatus A. St.-Hil.*110
Zanthoxylum riedelianum Engl.1130
Casearia gossypiosperma Briq.17290
Casearia grandiflora Cambess.21840
Casearia rupestris Eichler810
Casearia sylvestris Sw.6560
Prockia crucis P. Browne ex L.520
Xylosma prockia (Turcz.) Turcz.*110
Allophylus edulis (A. St.-Hil., Cambess. & A.Juss.) Radlk.110
Allophylus racemosus Sw.830
Cupania vernalis Cambess.7880
Dilodendron bipinnatum Radlk.1020
Magonia pubescens A. St.-Hil.210
Matayba elaeagnoides Radlk.**3940
Matayba guianensis Aubl.10570
Chrysophyllum gonocarpum (Mart. & Eichler) Engl.6940
Chrysophyllum marginatum (Hook. & Arn.) Radlk.16120
Micropholis venulosa (Mart. & Eichler) Pierre6130
Pouteria gardneri (Mart. & Miq.) Baehni4270
Pouteria torta (Mart.) Radlk.12880
Siparuna guianensis Aubl.40770
Styrax camporum Pohl3550
Symplocos pubescens Klotzsch ex Benth.*810
Cecropia pachystachya Trécul1130
Urera baccifera (L.) Gaudich. ex Wedd.1010
Aloysia virgata (Ruiz & Pav.) A. Juss.320
Callisthene major Mart.6240
Qualea dichotoma (Mart.) Warm.530
Qualea jundiahy Warm.*3960
Vochysia magnifica Warm.11430
Vochysia tucanorum Mart.510

The floristic similarity analysis between sites showed a similar pattern in seven of the ten sites examined (Figure 3). Accordingly, there was the formation of a floristically cohesive group (Group 1), where sites 3, 6, 7, 8, 9, and 10 were presented with values exceeding 0.5, indicating sites that were floristically similar. Also, a second group was formed with sites 1 and 4, whereas site 2 was floristically different from all the other sites (Figure 3).

Detrended correspondence analysis, DCA, presented eigenvalues of 0.61 and 0.25 in the two first ordination axes, explaining about 85% of the total data variation (Figure 4). The diagram demonstrated a large group formed from seven sites. Structural ordination, on the left, contemplates the same sites forming Group 1 by grouping analysis. These seven sites indicate the floristic pattern representative of the studied SSF. The second DCA axis was associated strongly with the sampled fragments’ conservation degree. The group formed on the center left is characterized by intermediate conservation stages, while other sites on the top right are characterized by more advanced conservation stages (Group 2). On the lower right is the initial stage (site 2). Therefore, the three fragments on the right do not form a cohesive group. In addition, these three sites are those that are more distant from each other. Thus, the structural floristic similarity of the seven areas above may have been established on a geographical position basis. The division made by TWINSPAN corroborated the ordination results by DCA and the Sørensen coefficient (Figure 3).

The first division by TWINSPAN separated the seven SSF representative sites in this region from the three other fragments, owing to the absence of Siparuna guianensis, a species regarded as an indicator, which only occurs in the group formed on the other side of the dichotomy (Figure 5). In the second division, sites 3 and 8 were separated from the other fragments (Figure 5). The lower levels of division by TWINSPAN do not reveal new groups that make sense.

4. Discussion

Studies conducted in areas of SSF in Brazil presented Fabaceae, Myrtaceae, Rubiaceae, Annonaceae, Lauraceae, and Meliaceae as the families with greater species richness [3336]. In São Paulo state SSF, as well as the greater richness of families found in this study, there is also an emphasis on Melastomataceae and Solanaceae [37, 38]. In northeastern Brazil, the most prominent families in terms of species number in SSF sites are Fabaceae Mimosoideae, Euphorbiaceae, Fabaceae Faboideae, Myrtaceae, and Rubiaceae [39]. The pattern in families and genera richness found here corroborates that described by Oliveira Filho and Fontes [40] for SSF domains in southeastern Brazil, with significant tree species richness in Fabaceae (all subfamilies), Lauraceae, Myrtaceae, Moraceae, and Rubiaceae and genres Aspidosperma, Ficus, Machaerium, and Ocotea.

Copaifera langsdorffii and H. courbaril are wide-ranging adaptive species, occurring in various forest formations of the Cerrado region, as well as in other geographical provinces in Brazil and are, therefore, considered to be habitat generalists [41]. Apuleia leiocarpa and Luehea grandiflora are also among the most frequent species of SSF, according to a study presented by Ferreira Júnior et al. [42], which brought together 15 studies in southeastern Brazil. The tree species composition studied in SSF showed a relationship with lowland seasonal forests of Brazil’s eastern Minas Gerais state, based on the indicator species analysis made by Oliveira-Filho and Fontes [40]. Of the 57 species identified by these authors as eastern Minas Gerais region indicators, 36 (63.1%) were recorded in the sample sites presented here. This data corroborates the idea that interior SSF fragments may have a floristic connection with the Atlantic forest that is able to expand its distribution in sites with strong seasonality by means of gallery forests [43].

The presence of unique species to each location highlights the heterogeneity between samples (sites), reflected in increased beta diversity. We should point out that the limited number of individuals in some populations may impair the biological conservation of many species, causing serious difficulties for their preservation. However, some species, such as Galipea jasminiflora and Micrandra elata, occurred in only one fragment, but with a high density. This observation acquires relevance in terms of plant population conservation. The conservation of the remaining forest, with different structures and consequent higher beta diversity, can be a relevant parameter to the decision for choosing priority areas for conservation in SSF [44].

Diversity values are among those frequently found for SSF and Atlantic rainforest, which generally vary between 3.2 and 4.2 [45]. However, they are lower than those found in other tropical forests [46]. The lower values for evenness found in fragments indicate a relatively high concentration in density of a small number of species, which dominate the tree community. The predominance of a few species, in number or biomass in a community (also known as ecological dominance), is not uncommon in tropical forests [47].

A variation in similarity values occurs as a result of different conditions in the use and historic occupation of each fragment and acts as an important force capable of modifying plant communities through spatial and temporal heterogeneity, thus determining the community composition and structure [16, 48]. The separation of site 2 from other areas, along with Group 2, can be related to different stages of forest succession. Site 2 presents in an initial successional stage owing to an intense human processes. This fragment, although situated in the countryside, reveals recent historical disturbance as a result of selective logging and the presence of cattle, and the surrounding matrix is formed entirely by agricultural sites [49]. Selective logging enables the creation of gaps within the fragment and the establishment of pioneering species. As Whitmore states [47, 50], pioneer trees and shrubs need high light and temperature intensities for seed germination and seedling establishment and growth. Group 2 (sites 1 and 4) is formed by the more well-preserved fragments among those studied and is characterized by the presence and dominance of individuals with a large basal area. In fragment 4, for example, Micrandra elata was sampled with a density of 118 ind·ha−1 and a basal area of 24.51 m2·ha−1 [51], whereas in fragment 1, Psidium sartorianum was the species with the highest dominance [52].

Although the floristic similarity between the fragments can be related to the geographical proximity, as already reported by MaCdonald et al. [53], we verified that similarity relations between studied sites are established mainly as a result of the community successional stage. This same pattern was also found by Durigan et al. [17]; that is, within the same vegetal formation, communities at similar stages of the successional process tend to have similar flora.

This analysis showed high beta diversity of the studied SSF arboreal flora, mainly as a result of the coverage of heterogeneous areas in relation to fragments at a successional stage. The largest SSF sampling in the Cerrado domain led to the registration and distribution of several plant populations, which had previously only been described for the Atlantic forests domain, demonstrating not only a lack of studies like this, but also corroborating the idea of strong floristic connections with rainforests in the east of the country [40].

The fragmentation process started in the 1970s in the studied region of Brazil, especially as a result of the incentive given to landowners by the public authority for the advancement of occupied areas for agricultural and livestock activities, which had a direct consequence for the formation of many smaller forest fragments. Fragmentation possibly created similar processes in forests’ successional development, which has conditioned a floristic pattern as found for seven of the ten sites surveyed.


Thanks are due to all those who assisted fieldwork and Professor Dr. Glein Monteiro de Araújo for aid in the field and in species identification. The authors would like to thank Fundo de Amparo a Pesquisa do Estado de Minas Gerais (FAPEMIG) for financial support for this Project (Process no. APQ-00694-08).


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