ISSN 0798 1015

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Vol. 38 (Nº 28) Año 2017. Pág. 18

Floristic richness in a transitional area between Mixed and Semideciduous Forests in the middle Tibagi River region, southern Brazil

Riqueza florística em uma área de transição entre Florestas Mistas e Semidecíduas na região média do rio Tibagi, sul do Brasil

Eduardo ADENESKY-FILHO 1 ; Franklin GALVÃO 2; Paulo Cesar BOTOSSO 3

Recibido: 31/12/16 • Aprobado: 25/01/2017


Content

1. Introduction

2. Materials and Methods

3. Results

4. Discussion

Acknowledgements

Bibliographic references


ABSTRACT:

The vast forest that covered the state of Parana has been reduced to small forest fragments. The Tibagi River watershed has some of best fragments, but with little detailed information about this region is available. This study documented the tree and shrub vegetation found within the limits of the town of Telêmaco Borba, state of Parana. It recorded 221 species in 51 families and 138 genera, among which are one tree species previously unreported from that state and eight endangered species. The information obtained shows the relevance of forest fragments for the Tibagi River as well as that of floristic studies to preserve forest ecosystems.
Key words: Atlantic Forest, Ecotone, Ecological Group.

RESUMO:

A vasta floresta que cobria o estado do Paraná foi reduzida a pequenos fragmentos florestais. A bacia do rio Tibagi tem alguns dos melhores fragmentos, mas com pouca informação detalhada sobre esta região está disponível. Este estudo documentou a vegetação arbórea e arbórea encontrada dentro dos limites do município de Telêmaco Borba, estado do Paraná. Registou 221 espécies em 51 famílias e 138 gêneros, entre os quais uma espécie de árvore anteriormente não declarada desse estado e oito espécies ameaçadas de extinção. As informações obtidas mostram a relevância de fragmentos florestais para o rio Tibagi, assim como os estudos florísticos para preservar os ecossistemas florestais.
Palavras-chaves: Floresta Atlântica, Ecótono, Grupo Ecológico.

1. Introduction

Over the past thousands years, great transformations have shaped the Brazilian vegetation into its current physiognomic configuration. Palynological records on southern Brazil during the last glacial period indicate the predominance of grassland vegetation (grasslands and cerrado) in regions now covered with forests (Behling, 1995). During the late Quaternary, due to changes in climatic conditions and abundant rainfall, forests began to prevail over grasslands and cerrados on hillside slopes and in river valleys (Behling 2002). Until a few decades ago, the state of Parana had one of the richest forest covers in Brazil (Maack, 2012).

Nowadays, forest advance is no longer detained by climatic conditions, but by such anthropogenic pressures as agriculture, livestock, and wood exploitation. Forest which used to cover 83% of the state of Parana (Maack, 2012) only occupy 10.52% of its territory, most of it as fragments (SOS Mata Atlântica,2009).

Remnants of the Atlantic Forest domain, to which all the forests found in that state belong, are insufficiently sampled (Giulietti et al., 2005). This is particularly the case of the middle Tibagi River basin, which, nevertheless, is an area of high conservation priority according tothe Brazilian Ministry of Environment (MMA, 2007). Little detailed information about this region is available (Medri et al., 2002), although it has some of the largest and best remnants in that state (Torezan, 2002). Therefore, studies are urgently needed to understand the regional flora, redirect planning efforts, and preserve ecosystems.

The Tibagi River has its source in the Campos Gerais, to the west of the Devonian escarpment, and runs approximately 550 km before it empties into the Paranapanema River (Maack, 2012). Due to its extension, it flows through most of the lithologies found in the state of Parana (Pinese, 2002), which, in addition to the transition between Cfa and Cfb climates (Mendonça & Oliveira, 2002), promotes a great variety of soils and associations (Stipp, 2002). Together, such features imply the presence of different environments and distinct phytogeographic regions (IBGE, 2012) in this basin, whose main forest types are Mixed and Semideciduous Forests, (Dias et al., 2002; Cardoso & Sajo, 2004; Carmo & Assis, 2012).

Given the ecological importance of forest remnants in the middle Tibagi River watershed, Paraná, we intend, based on a floristic survey, to provide information on the tree and shrub species richness, the occurrence of endangered species, and the representation of its ecological groups.

2. Materials and Methods

2.1. Study Area

Study area is a 2.675 ha forested area located within the limits of the county of Telêmaco Borba, in the central-eastern part of the Second Parana Plateau, at coordinates 24º07′02″ S and 50º41′20″ W, in the middle Tibagi River watershed. Geologically, it corresponds to the sediment sequence of the Sedimentary Basin of the state of Parana, and outcropping rocks belong to the Itararé, Guatá, and Passa Dois groups (Milaniet al., 1994). Prevailing soils are Argisols, Latosols, Neosols, Nitosols, and Cambisols (EMBRAPA, 2006). According to Köppen’s classification, Cfb (humid with temperate summer) climate prevails in the region. Average annual temperature is 18°C (IAPAR, 1978) and average annual rainfall, 1.577 mm (SUDERHSA, 1998).

It is located on the Atlantic Forest Domain inserted at the Campos Gerais, according to Maack’s definition (2012). The geological, pedological, and climatic variations occurring in the Tibagi River watershed (Medri et al., 2002) allow the development of different plant types. Yet our study area is covered with both Semideciduous and Mixed Forests (Medri et al., 2009).

2.2. Data Collection

Our floristic survey follows the walk-through methodology proposed by Filgueiras et al., (1994). It consists in establishing parallel transects 100 m apart. Since our area is quite wide (2,675 ha), our collection efforts focused on approximately 2 ha sample blocks and, to encompass the most diverse environments, we walked along pre-established trails adjacent to the Tibagi River and its tributary, as well as on by- and main roads.

During our walks, we collected fertile samples of tree and shrub species with a diameter at breast height (DBH ~ 1.3 m above ground level) equal or superior to 15 cm. Data collection took place between December/2010 and February/2012. Botanical material was herborized and identified, with specialized bibliography and consultation to the Museu Botânico Municipal (MBM - City Botanical Gardens) of Curitiba, Paraná. Classification is based on APG III (2009), and botanical nomenclature was confirmed both by the Flora do Brasil (2016) and the Tropicos.org (2016). Collected samples and voucher specimens were deposited at the following herbaria: MBM, UPCB, HCF and RB, acronyms according to Thiers (2016).

To simplify the interpretation of the floristic composition, species were separated in ecological groups, defined according to Swaine & Whitmore (1988), and divided into three groups: pioneers (P), non-pioneer light-demanders (CL), and non-pioneer shade-tolerant (CS). Ecological information on species was found in the following bibliography: Gandolfi et al., (1995); Reis (1995); Ivanauskas et al.,(1999); Fonseca & Rodrigues (2000); Iserhagen et al.,(2002); Silva et al.,(2003); Mantovani et al.,(2005); Carvalho et al.,(2006); Guarantini et al.,(2008); Rodrigues et al., (2009); Lorenzi (2000; 2002; 2009); Carvalho (2003; 2006; 2008; 2010) and complemented with the authors’ knowledge.

3. Results

The survey documented 221 native and four exotic species: seven tree ferns (Cyatheaceae and Dicksoniaceae), one Gymnosperms (Araucariaceae), and 213 Angiosperms (Table 1) distributed in 51 families and 138 genera.

The richest families were Fabaceae (33 species), Myrtaceae (29), Euphorbiaceae (13), Meliaceae and Lauraceae (10 each), Rutaceae (9), Salicaceae (8) Rubiaceae, and Moraceae (7), totalizing 57% of all the species recorded. In the remaining families, approximately 8.1% (18) presented only one species. The richest genera were Eugenia L. (9 species), Machaerium (6), Casearia, Cyathea, Ficus, Solanum, and Trichilia (5 each).

Among tree species, Aspidosperma riedelii Müll. Arg. (Apocynaceae) had only been reported in the states of São Paulo and Santa Catarina in the Flora do Brasil (2016). It is thus a new record of occurrence to Paraná State. It was solely found on superficial fractures of diabase dikes in an area with Litholic Neosol.

The species Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae), Cedrela fissilis Vell. (Meliaceae), and Dicksonia sellowiana Hook. (Dicksoniaceae), are listed as endangered in Brazil (Martinelli et al., 2013), and Balfourodendron riedelianum (Engl.) Engl. (Rutaceae), Aspidosperma australe Müll. Arg., Aspidosperma polyneuron Müll. Arg. (Apocynaceae), Machaerium paraguariense Hassl., and Myrocarpus frondosus Allemão (Fabaceae) have been described as rare in this state (MMA, 2008).

Classification into ecological groups revealed that 59.3% (131 species) are non-pioneer light-demanders, 24.9% (55) are non-pioneer shade-bearers and 15.8% (35) are pioneers.

Table 1 – Species checklist organized by ecological group and voucher numbers, documented in the forest fragment located within the limits of the county of Telêmaco Borba, PR. GE – Ecological Group (P = pioneer; CL = non-pioneer light-demander; CS = non-pioneer shade-tolerant; NC = not classified; I = Invasive-exotic; * identified in the field, but no fertile specimens were found.

FAMILY/SPECIES

GE

VOUCHER

ANACARDIACEAE

   

Astronium graveolens Jacq.

CL

*

Lithraea molleoides (Vell.) Engl.

P

UPCB 75667

Schinus terebinthifolius Raddi

P

MBM 375749

Rhus succedanea L.

I

*

ANNONACEAE

   

Annona cacans Warm.

CL

HCF 12239

Guatteria australis A.St.-Hil.

CL

*

Annona sylvatica   A.St.-Hil.

CL

*

Xylopia brasiliensis Spreng.

CL

MBM 385373

APOCYNACEAE

   

Aspidosperma australe Müll. Arg.

CS

MBM 387196

Aspidosperma polyneuron Müll. Arg.

CS

MBM 388165

Aspidosperma riedelii Müll. Arg.

CL

*

Rauvolfia sellowii Müll. Arg.

CL

HFC12316

Tabernaemontana catharinensis (A.DC.) Miers

P

*

AQUIFOLIACEAE

   

Ilex brevicuspis Reissek

CL

MBM 387456

ARALIACEAE

   

Dendropanax cuneatus (DC.) Decne. & Planch.

CL

*

Schefflera morototoni (Aubl.) Maguire et al.

CL

HCF 12250

ARAUCARIACEAE

   

Araucaria angustifolia (Bertol.) Kuntze

CL

*

ARECACEAE

   

Euterpe edulis Mart.

CS

*

Geonoma schottiana Mart.

CS

*

Syagrus romanzoffiana (Cham.) Glassm.

CL

*

Syagrus oleracea (Mart.) Becc.

CL

*

ASTERACEAE

   

Piptocarpha axillaris (Less.) Baker

P

*

BIGNONIACEAE

   

Cybistax antisyphilitica (Mart.) Mart.

CL

*

Jacaranda puberula Cham.

CL

MBM 385539

Jacaranda micrantha Cham.

P

MBM 388161

BORAGINACEAE

   

Cordia americana (L.) Gottschling & J.S.Mill.

P

*

Cordia ecalyculata Vell.

CS

MBM 388166

Cordia superba Müll. Arg.

CS

MBM 385598

Cordia trichotoma (Vell.) Arráb. ex Steud.

P

*

CANNABACEAE

   

Celtis iguanaea (Jacq.) Sarg.

P

MBM 389764

Trema micrantha (L.) Blume

P

UPCB 75684

CARDIOPTERIDACEAE

   

Citronella paniculata (Mart.) Howard

Cl

*

CARICACEAE

   

Jacaratia spinosa (Aubl.) DC.

CS

*

Vasconcellea quercifolia A. St.-Hil.

P

*

CELASTRACEAE

   

Maytenus aquifolia Mart.

CS

MBM 388163

Maytenus gonoclada Mart.

CL

*

COMBRETACEAE

   

Terminalia triflora (Griseb.) Lillo

P

*

CYATHEACEAE

   

Alsophila setosa Kaulf.

CS

*

Cyathea atrovirens (Langsd. & Fisch.) Domin

CS

HCF 12192

Cyathea corcovadensis (Raddi) Domin

CS

*

Cyathea delgadii Sternb.

CS

*

Cyathea hirsuta C.Presl

CS

*

Cyathea phalerata Mart.

CS

*

DICKSONIACEAE

   

Dicksonia sellowiana Hook.

CS

HCF 12417

EBENACEAE

   

Diospyros inconstans Jacq.

CL

HCF 12435

ELAEOCARPACEAE

   

Sloanea monosperma Vell.

CL

*

ERYTHROXYLACEAE

   

Erythroxylum cuneifolium (Mart.) O.E. Schulz

P

*

Erythroxylum deciduum A. St.-Hil.

P

MBM 385542

EUPHORBIACEAE

   

Actinostemum concolor (Spreng.) Müll. Arg.

CS

MBM 385625

Alchornea glandulosa Poepp.

CL

MBM 385551

Alchornea sidifolia Müll. Arg.

CL

MBM 389778

Alchornea triplinervia (Spreng.) Müll. Arg.

CL

HCF 12284

Croton floribundus Spreng.

P

RB 606583

Croton urucurana Baill.

P

UPCB 75766

Manihot grahamii Hook.

P

MBM 389762

Phyllanthus sellowianus (Klotzsch) Müll.Arg.

P

*

Sapium glandulosum (Vell.) Pax

CL

*

Sebastiania brasiliensis (L.) Spreng.

CL

MBM 385615

Gymnanthes klotzschiana Müll.Arg.

P

HCF 12229

Sebastiania schottiana (Müll.Arg.) Müll.Arg.

P

*

Tetrorchidium rubrivenium Poepp. & Endl.

CL

*

FABACEAE

   

Albizia niopoides (Spruce ex Benth.) Burkart

P

MBM 385587

Albizia polycephala (Benth.) Killip ex Record

CL

*

Anadenanthera colubrina (Vell.) Brenan

P

*

Bauhinia forficata Link

CL

*

Bauhinia longifolia (Bong.) Steud.

P

*

Cassia leptophylla Vog.

CL

MBM 389766

Centrolobium tomentosum Guill. ex Benth.

P

HCF 12337

Copaifera langsdorffii Desf.

CL

MBM 385540

Dalbergia frutescens (Vell.) Britton

CL

MBM 385533

Enterolobium contortisiliquum (Vell.) Morong

CL

*

Erythrina crista-galli L.

P

*

Erythrina falcata Benth.

CL

*

Holocalyx balansae Mich.

CL

HCF 12382

Inga marginata Willd

CL

HCF 12291

Inga sessilis (Vell.) Mart.

CL

*

Inga striata Benth

CL

MBM 388152

Inga vera Willd

CL

*

Lonchocarpus campestris Mart.ex Benth.

CL

HCF 12386

Lonchocarpus cultratus (Vell.) A. M. G. Azevedo

CL

RB 607125

Lonchocarpus muehlbergianus Hassl.

CL

HCF 12375

Lonchocarpus subglaucescens Mart ex Benth.

CS

HCF 12432

Machaerium aculeatum Raddi

CL

UPCB 75685

Machaerium brasiliense Vog.

CL

MBM 385536

Machaerium nyctitans (Vell.) Benth.

P

*

Machaerium paraguariense Hassl.

CL

MBM 388151

Machaerium scleroxylon Tul.

CL

*

Machaerium stipitatum Vog.

CL

HCF 12228

Myrocarpus frondosus Allemão

CL

MBM 389740

Ormosia arborea (Vell.) Harms

CL

MBM 385604

Parapiptadenia rigida (Benth.) Brenan

CL

*

Peltophorum dubium (Spreng.) Taub.

CL

*

Piptadenia gonoacantha (Mart.) J.F.Macbr.

CL

*

Schizolobium parahyba (Vell.) Blake

P

*

LAMIACEAE

   

Aegiphila sellowiana Cham.

CL

*

Vitex megapotamica (Spreng.) Moldenke

CL

RB 607303

LAURACEAE

   

Cryptocarya aschersoniana Mez

CL

HCF 14249

Endlicheria paniculata (Spreng.) J. F. Macbr.

CL

*

Nectandra lanceolata Nees

CL

MBM 387459

Nectandra megapotamica (Spreng.) Mez

CL

MBM 385580

Nectandra oppositifolia Nees

CS

HCF 12388

Ocotea dyospirifolia (Meisn.) Mez

CL

RB 606199

Ocotea elegans Mez

CL

*

Ocotea corymbosa ( Meisn.) Mez

CL

*

Ocotea puberula (Rich.) Nees

CL

MBM 385581

Persea willdenovii Kosterm.

CL

MBM 389756

LAXMANNIACEAE

   

Cordyline spectabilis Kunth & C.DC. Bouché

CL

*

LECYTHIDACEAE

   

Cariniana legalis (Mart.) Kuntze

CL

*

LOGANIACEAE

   

Strychnos brasiliensis (Spreng.) Mart

CL

*

LYTHRACEAE

   

Lafoensia pacari A. St.-Hil.

CL

*

MAGNOLIACEAE

   

Magnolia ovata A.St.-Hil. (Spreng.)

CL

*

MALVACEAE

   

Bastardiopsis densiflora (Hook & Arn) Hassl.

P

HCF 12317

Ceiba speciosa (A. St-Hil.) Ravenna

CL

RB 608779

Helicteres brevispira A.St.-Hil.

CL

UPCB 75769

Heliocarpus papayensis Kunth

P

*

Luehea divaricata Mart

CL

HCF 12319

Pseudobombax longiflorum (Mart. & Zucc.) A.Robyns

CL

*

MELASTOMATACEAE

   

Miconia cineracens Miq.

CS

MBM 385535

MELIACEAE

   

Cabralea canjerana (Vell.) Mart.

CL

HCF 12235

Cedrella fissilis Vell.

CL

*

Guarea guidonia (L.) Sleumer

CS

*

Guarea kunthiana A. Juss.

CL

*

Guarea macrophylla Vahl.

CL

HCF 12216

Trichilia casaretti C. DC.

CS

MBM 385562

Trichilia catigua A. Juss.

CS

*

Trichilia claussenii C. DC.

CS

*

Trichilia elegans A. Juss.

CS

*

Trichilia pallida Sw.

CS

MBM 387461

MORACEAE

   

Ficus enormis (Miq.) Miq.

CL

*

Ficus glabra Vell.

CL

*

Ficus guaranitica Chodat

CL

MBM 389774

Ficus insipida Willd.

CL

*

Ficus luschnathiana (Miq.) Miq.

CL

*

Maclura tinctoria L.

CL

MBM 387367

Sorocea bonplandii (Baill.) W.C.Burger et al.

CS

MBM 385557

MYRTACEAE

   

Calyptranthes concinna DC.

CS

MBM 385545

Campomanesia eugenioides (Cambess.) O. Berg

CL

MBM 387454

Campomanesia guaviroba (DC.) Kiaersk.

CL

RB 608037

Campomanesia guazumifolia (Cambess.) O.Berg

CS

*

Campomanesia xanthocarpa O. Berg

CS

MBM 385553

Eugenia blastantha (O.Berg) D. Legrand

CS

*

Eugenia brasiliensis Lam.

CS

*

Eugenia florida DC.

CL

*

Eugenia involucrata DC.

CL

*

Eugenia neoverrucosa Sobral

CS

*

Eugenia pluriflora DC.

CL

*

Eugenia pyriformis Cambess.

CL

*

Eugenia uniflora L

CL

*

Eugenia uruguayensis Cambess.

CL

*

Gomidesia palustris DC.

CL

*

Myrceugenia euosma (O. Berg) D. Legrand

CL

*

Myrcia anacardiifolia Gardner

CL

*

Myrcia laruotteana Cambess.

CS

*

Myrcia splendens (Sw.) DC.

CS

*

Myrcianthes pungens (O.Berg) D. Legrand

CL

*

Myrciaria cuspidata O.Berg

CS

*

Myrciaria deliculata (DC.) O. Berg

CL

*

Myrciaria floribunda (H.West ex Willd.) O. Berg

CL

*

Neomitranthes glomerata (D.Legrand) D.Legrand

CS

*

Plinia rivularis (Cambess.) Rotman

CL

RB 606744

Plinia trunciflora (O. Berg) Kausel

CS

*

Psidium cattleyanum Sabine

CL

*

Psidium guajava L.

CL

*

Siphoneugena crassifolia (DC.) Proença & Sobral

CS

*

NYCTAGINACEAE

   

Bougainvillea glabra Choisy

CL

UPCB 75689

Guapira hirsuta (Choisy) Lundell

CS

*

Guapira opposita (Vell.) Reitz

CS

HCF 12464

Pisonia ambigua Heimerl

CL

*

PHYTOLACCACEAE

   

Gallesia integrifolia (Spreng.) Harms

CL

*

Phytolacca dioica L.

CL

HCF 12234

PICRAMNIACEAE

   

Picramnia parvifolia Engl.

CS

*

Picramnia ramiflora Planch.

CS

MBM 389785

PRIMULACEAE

   

Myrsine coriacea (Sw.) R. Br.

P

HCF 12194

Myrsine gardneriana A.DC.

CL

*

Myrsine umbellata Mart.

CL

HCF 12430

PROTEACEAE

   

Roupala montana var. brasiliensis (Klotzsch) K.S.Edwards

CL

HCF 12380

RHAMNACEAE

   

Hovenia dulcis Thunb.

I

*

ROSACEAE

   

Eriobotrya japonica Lindl.

I

*

Prunus myrtifolia (L.) Urb.

CL

MBM 385830

RUBIACEAE

   

Bathysa australis (A.St.-Hil.) K.Schum.

CL

HCF 12336

Psychotria carthagenensis Jacq.

CS

MBM 389642

Psychotria suturella Müll. Arg.

CS

HCF 12242

Psychotria vellosiana Benth.

CS

*

Randia cf. armata (Sw.) DC.

CS

*

Rudgea jasminoides (Cham.) Müll. Arg.

CS

RB 575620

Rudgea parquioides (Cham.) Müll. Arg.

CS

MBM 389759

RUTACEAE

   

Balfourodendron riedelianum (Engl.) Engl.

CL

HCF 12391

Citrus limon (L.) Osbeck

I

*

Esenbeckia febrifuga (A. St.-Hil.) A. Juss. ex Mart.

CL

HCF 12233

Esenbeckia grandiflora Mart.

CL

*

Pilocarpus pennatifolius Lem.

CL

*

Zanthoxylum caribaeum Lam.

P

*

Zanthoxylum fagara (L.) Sarg.

P

MBM 385623

Zanthoxylum petiolare A. St.-Hil.

P

*

Zanthoxylum rhoifolium Lam.

P

HFC12414

SALICACEAE

   

Banara tomentosa Clos

CS

*

Casearia decandra Jacq.

CS

*

Casearia gossypiosperma Briq.

CS

*

Casearia lasiophylla Eichler

CL

*

Casearia obliqua Spreng.

CS

HCF 12387

Casearia sylvestris Sw.

CL

*

Prockia crucis P. Browne ex. L.

CL

*

Xylosma ciliatifolia (Clos) Eichler

CL

*

SAPINDACEAE

   

Allophylus edulis (A. St.-Hil. et al.) Hieron. ex Niederl.

CL

*

Cupania vernalis Cambess.

CL

HCF 12279

Dianopteryx sorbifolia Radlk.

CL

MBM 385541

Matayba elaeagnoides Radlk.

CL

MBM 388179

SAPOTACEAE

   

Chrysophyllum marginatum (Hook. & Arn.) Radlk.

CL

HCF 12196

Chrysophyllum gonocarpum (Mart. & Eichl.) Engl.

CS

UPCB 75775

Pouteria beaurepairei (Glaz. & Raunk.) Baehni

CL

*

SOLANACEAE

   

Solanum swartzianum Roem. & Schult.

CL

*

Solanum granulosoleprosum Dunal

CL

*

Solanum mauritianum Scop

CL

*

Solanum pseudoquina A. St.-Hil.

CL

*

Solanum sanctaecatharinae Dunal

CL

*

STYRACEAE

   

Styrax acuminatus Pohl

CL

*

Styrax leprosus Hook. & Arn.

CL

*

SYMPLOCACEAE

   

Symplocos tenuifolia Brand

CS

MBM 389761

THEACEAE

   

Laplacea fruticosa (Schrad.) Kobuski

CS

*

URTICACEAE

   

Boehmeria caudata Sw.

P

MBM 385538

Cecropia glaziovii Snethl.

CL

HCF 12221

Cecropia pachystachya Trécul

P

HCF 12205

Urera baccifera (L.) Gaudich. ex Wedd.

CL

*

VERBENACEAE

   

Aloysia virgata (Ruiz & Pav.) Juss.

CL

HCF 12385

Citharexylum myrianthum Cham.

CL

MBM 385544

 

4. Discussion

The high species richness of Fabaceae and Myrtaceae, as found in this study, is recurrent in forest formations from southern and southeastern Brazil (Ivanauskas & Rodrigues 2000; Colonetti et al., 2009; Higuchi et al., 2012; Kurtz et al., 2009; Ribeiro et al., 2013), as well as in fragments located in the Tibagi River watershed (Dias et al.,1998; Bianchini et al., 2003; Costa et al., 2011; Carmo & Assis, 2012), especially in its middle course (Silva et al.,1995; Dias et al., 2002).

According to Torezan (2002), the Tibagi River watershed is composed of grassland and forest phytophysiognomies whose proximity favors the growth of a higher number of species typical of each. Thus, the high species richness of Myrtaceae and Fabaceae found in this study area is linked to its closeness to the lower Tibagi River, covered by semideciduous seasonal forests, in which these families, especially Fabaceae, are very species-rich (Jarenkow & Waechter, 2001; Leite et al., 2013; França & Stehmann, 2013).

Eugenia and Machaerium, which are frequent in this survey, are also part of the main species growing in this watershed (Bianchini et al., 2003; Carmo & Assis, 2012) and, exceptionally, in the middle Tibagi River area (Dias et al., 2002; Yamamoto et al., 2005). Studies carried out in forests of southern Brazil report that both genera present the highest number of species (Leite et al., 2013) and are distributed on the different forest layers (Dias et al., 2002).

Based on floristic and phytosociological surveys and on the consultation herbaria, 547 tree and shrub species have so far been mentioned as occurring in the (25.000 km2) Tibagi River watershed (Dias et al., 2002). Since our study area shelters 221 species, i.e. 40% of the total species of the whole watershed, it is a taxon rich habitat.

The high species richness probably is related to both the geo-pedological (Pinese, 2002; Stipp, 2002) and climatic (Mendonça & Oliveira, 2002) conditions and their local associations watershed, which allow environment diversification and, consequently, the presence of more species (Torezan, 2002). Another relevant factor is the species mix of typical of distinct phytogeographic regions: mixed ombrophilous forests, characterized by the presence of Araucaria angustifolia, and semideciduous seasonal forest, identified by the occurrence of Aspidosperma polyneuron. Such results indicate that the middle Tibagi River area is an ecotone region (Carmo & Assis, 2012; Dias et al., 2002). According to Durigan et al., (2008), transition areas tend to present high species richness and diversity because taxa from distinct phytogeographic regions co-occurs.

The presence of species in the different ecological groups allows us to infer that this fragment constitutes a mosaic of different successional stages, where areas sheltering non-pioneer shade-bearers or light-demanders are adjacent to disturbed ones occupied by early successional species. The alternation of ecological groups, forming a dynamic component of overcoming among species, was also observed by Silva et al., (2003) and Mantovani et al., (2005), who pointed out changes in plant communities over time as possible causes.

Non-pioneer light-demanders were the prevailing ecological group, followed by non-pioneer shade-bearers. According to Mantovani et al., (2005), this denotes conditions are better suited to a successful recruitment of late successional species. However, the predominance of light-demanders suggests that the area has not yet reached its full development or that another factor may have modified its successional status.

According to Peixoto (2004), among the factors that can delay succession are fires and selective logging. Guarantini et al., (2008) also noted species richness in early ecological groups, after fires in semideciduous seasonal forests. They stated that natural or anthropic disturbance originates small gaps that can be colonized by light-demanding species, which can thus be quite numerous within the forest.

Furthermore, with regard to climax species, it is worth stressing that emrgent tree species Annona cacans, Araucaria angustifolia, Aspidosperma polyneuron, Machaerium scleroxylon, and Myrocarpus frondosus, were distributed within the whole survey area. Similar results were obtained by Ivanauskas & Rodrigues (2000) and Costa et al., (2011) who studied the successional character of tree species in forest fragments of Southern Brazil and acknowledged species typical of climax and/or better preserved forests.

Information on the richness of the regional flora including endangered species and a new tree species in this state proves the relevance of forest fragments in the Tibagi River watershed as well as the importance of floristic studies to preserve forest ecosystems and maintain their ecological functions.

Acknowledgements

The authors thank the Companhia Paranaense de Energia (COPEL), and more particularly Murilo Lacerda Barddal, who allowed data collection and provided the necessary infrastructure; Osmar dos Santos Ribas, curator at the Museu Botânico de Curitiba (MBM), for his help in the identification of species; and the Cooperativa Interdisciplinar de Serviços Técnicos (INTERCOOP), for their help in collecting and identifying species.

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1. Universidade Federal do Paraná – UFPR, Doctoral candidate in Forest Enginnering, Curitiba, PR, Brasil. E-mail: eduardo_adenesky@yahoo.com.br 

2. Universidade Federal do Paraná – UFPR, Departamento de Ciências Florestais, Curitiba, PR, Brasil.

3. Empresa Brasileira de Pesquisa Agropecuária – Embrapa Florestas, Estrada da Ribeira, CEP 83411-000, Colombo, PR, Brasil.


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