# Secondary Forest Succession in the Mata Atlantica, Brazil: Floristic and Phytosociological Trends

Authors: Alexandre Siminski; Alfredo Celso Fantini; Raymond Paul Guries; Ademir Roberto Ruschel; Maurício Sedrez dos Reis
Journal: ISRN Ecology
 (2011)

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## Abstract

This study aimed at  understanding the dynamics of ecological processes and the use of
secondary forests in Santa Catarina state (Brazil). The data base for these studies was formed
through forest inventories carried out in the three forest types of the state. The results of this
study demonstrate that the patterns of diversity are very similar among the three forest types;
however, the species compositions among the types are quite different. A total of 343 woody
species belonging to 73 families were found in the 24,000 m2 sampling area, revealing the
potential role of secondary forest in the conservation of biodiversity at the landscape scale.
As expected, a small set of pioneer species dominates young secondary forests with shade-tolerant
species becoming structurally important after 30 years. The patterns of forest structure
and species diversity observed in study largely conform to the postagricultural secondary
succession observed for many tropical forests.


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## Body

## 1. Introduction

The continuing loss of primary tropical forests makes secondary forests increasingly important for maintaining biodiversity across large forested landscapes [1–4], while also providing for environmental services and sustainable economic development [5–7]. Today, less than 25% of the original forest area in Santa Catarina state (Brazil) remains, mostly as small fragments of secondary forest in a mosaic intermixed with other land uses [8]. Most secondary forests in Santa Catarina and throughout much of the Mata Atlântica are privately held and result from fallow rotations in a dynamic agricultural system [9]. The recovery potential of such secondary forests is influenced by many interacting factors including the length and intensity of past land use, parcel size, soil conditions and landscape position, local climate regimes, proximity to forest seed sources, the presence or absence of seed banks, and dispersal agents, and interactions among regenerating species [10–16].The important roles that secondary forests can play in conservation and economic development are poorly understood in Brazil. Even well-intentioned Brazilian laws governing forest protection and use can be counter-productive to conservation. Recent laws that define a “forest” in terms of arbitrary structural characteristics have led small farmers to shorten fallow periods so that secondary forest succession does not proceed to the point where it is recognized as “forest”. This avoids federal/state restrictions on forest harvesting or land conversion [17] with the result that secondary forests which could contribute to biodiversity conservation while providing a range of nontimber forest products are now less likely to develop. The landscape increasingly is held only in the earliest stages of forest succession before being cropped again.We focused on the successional dynamics of secondary forests in the youngest age classes that prevail across this landscape in order to suggest alternative definitions of “forest” that recognize their important values in conservation and economic development. In this paper we describe the regeneration patterns that occur in secondary forests of Santa Catarina on parcels having diverse cultivation histories and variable fallow periods. Anthropogenic disturbances such as forest clearing for agriculture are generally more intensive/severe than those arising from natural disturbances, but early stages of forest succession appear to be dominated in both instances by a small number of common woody species in well-defined stages typical of large gaps [16, 18, 19]. The specific objectives of this study were to examine the patterns of species diversity and abundance as well as the transitions during which woody species are added and removed during the early phases of secondary forest succession.


## 2. Methods

### 2.1. Study Areas

The study was carried out on 48 small farms in the counties of Anchieta, Garuva, Concórdia, Três Barras, Caçador and São Pedro de Alcântara, Santa Catarina, Brazil (Figure1). These areas included three different vegetation formations common to the Mata Atlântica of Santa Catarina, seasonal deciduous forest, mixed ombrophylous forest and dense ombrophylous forest [20] recognized and protected by Brazilian Law no. 11.426/2006.Figure 1
Santa Catarina phytogeographic map, adapted from Klein [25].


#### 2.1.1. Seasonal Deciduous Forest (SDF)

This forest type occurs in the Uruguay River basin at elevations ranging from 200 to 600 meters with an annual precipitation of 1.800 mm/yr [21–24]. It is characterized by a closed canopy dominated by Lauraceae and emergent deciduous species, mostly Fabaceae and a subcanopy dominated by Sorocea bonplandii, Gynnanthes concolorand Trichilia species; epiphytes are poorly represented in comparison with the dense ombrophylous forest [21].


#### 2.1.2. Mixed Ombrophylous Forest (MOF)

This forest type is distributed across plateaus at approximately 500 meters elevation upwards on slopes rising to 1600 meters above sea-level with annual precipitation ranging from 1,600 to 2,100 mm [20]. This moist forest is characterized by a supercanopy dominated by Araucaria angustifolia, with subcanopy layers rich in species of Lauraceae, Myrtaceae, and Fabaceae [25, 26].


#### 2.1.3. Dense Ombrophylous Forest (DOF)

This forest was formerly common in coastal regions at elevations below 500 meters. The forest is characterized by hot temperatures, heavy rainfall (annual precipitation ranges from 1,500 to 2,000 mm) and an evergreen canopy dominated by Lauraceae and Myrtaceae with an abundance of epiphytes and palms [18, 26].


### 2.2. Data Collection and Analysis

We inventoried a total of eighty10m×10m plots in the dense ombrophylous forest (Garuva and São Pedro de Alcântara sites), and forty 20m×10m plots in each of the seasonal deciduous forest (Anchieta and Concórdia sites), and mixed ombrophylous forest (Três Barras and Caçador sites) types (a total of 160 plots covering 24,000 m2). Based on physiognomic features, together with land ownership information, we sited 160 plots distributed across four successional stages (after Klein [18]: shrubby stage (0–8 years), small tree stage (8–15 years), Arboreal Stage (15–30 years) and advanced arboreal stage (30–60 years)).  All woody plants taller than 1.5 m were identified and measured for DBH (diameter at breast height = 1.3 m) and total height. Plant identifications were made with reference to the Angiosperm Phylogeny Group (APG) classification updated in APG II [27], and the expertise of Prof. Ademir Reis (Federal University of Santa Catarina and Curator of the Barbosa Rodriguez Herbarium) and Prof. Marcos Sobral (BHCB—Herbarium of the Botany Department at the Federal University of Minas Gerais).Species richness (number of species), Shannon-Wiener’s index of diversity (H′), evenness (J), Simpson’s diversity index (D), “importance value” (IV) and absolute frequency were estimated using the FITOPAC 1 software package [28]. Rarefaction curves (Sobs) constructed using 100 randomized orders and the Chao1, Jackknife and Bootstrap estimators were derived using EstimateS 6.0 software [29]. Differences in values of stem density and basal area among stages were tested by one-way ANOVA with Tukey’s multiple comparison tests using the STATISTICA 6.0 software package [30].


## 2.1. Study Areas

The study was carried out on 48 small farms in the counties of Anchieta, Garuva, Concórdia, Três Barras, Caçador and São Pedro de Alcântara, Santa Catarina, Brazil (Figure1). These areas included three different vegetation formations common to the Mata Atlântica of Santa Catarina, seasonal deciduous forest, mixed ombrophylous forest and dense ombrophylous forest [20] recognized and protected by Brazilian Law no. 11.426/2006.Figure 1
Santa Catarina phytogeographic map, adapted from Klein [25].


### 2.1.1. Seasonal Deciduous Forest (SDF)

This forest type occurs in the Uruguay River basin at elevations ranging from 200 to 600 meters with an annual precipitation of 1.800 mm/yr [21–24]. It is characterized by a closed canopy dominated by Lauraceae and emergent deciduous species, mostly Fabaceae and a subcanopy dominated by Sorocea bonplandii, Gynnanthes concolorand Trichilia species; epiphytes are poorly represented in comparison with the dense ombrophylous forest [21].


### 2.1.2. Mixed Ombrophylous Forest (MOF)

This forest type is distributed across plateaus at approximately 500 meters elevation upwards on slopes rising to 1600 meters above sea-level with annual precipitation ranging from 1,600 to 2,100 mm [20]. This moist forest is characterized by a supercanopy dominated by Araucaria angustifolia, with subcanopy layers rich in species of Lauraceae, Myrtaceae, and Fabaceae [25, 26].


### 2.1.3. Dense Ombrophylous Forest (DOF)

This forest was formerly common in coastal regions at elevations below 500 meters. The forest is characterized by hot temperatures, heavy rainfall (annual precipitation ranges from 1,500 to 2,000 mm) and an evergreen canopy dominated by Lauraceae and Myrtaceae with an abundance of epiphytes and palms [18, 26].


## 2.1.1. Seasonal Deciduous Forest (SDF)

This forest type occurs in the Uruguay River basin at elevations ranging from 200 to 600 meters with an annual precipitation of 1.800 mm/yr [21–24]. It is characterized by a closed canopy dominated by Lauraceae and emergent deciduous species, mostly Fabaceae and a subcanopy dominated by Sorocea bonplandii, Gynnanthes concolorand Trichilia species; epiphytes are poorly represented in comparison with the dense ombrophylous forest [21].


## 2.1.2. Mixed Ombrophylous Forest (MOF)

This forest type is distributed across plateaus at approximately 500 meters elevation upwards on slopes rising to 1600 meters above sea-level with annual precipitation ranging from 1,600 to 2,100 mm [20]. This moist forest is characterized by a supercanopy dominated by Araucaria angustifolia, with subcanopy layers rich in species of Lauraceae, Myrtaceae, and Fabaceae [25, 26].


## 2.1.3. Dense Ombrophylous Forest (DOF)

This forest was formerly common in coastal regions at elevations below 500 meters. The forest is characterized by hot temperatures, heavy rainfall (annual precipitation ranges from 1,500 to 2,000 mm) and an evergreen canopy dominated by Lauraceae and Myrtaceae with an abundance of epiphytes and palms [18, 26].


## 2.2. Data Collection and Analysis

We inventoried a total of eighty10m×10m plots in the dense ombrophylous forest (Garuva and São Pedro de Alcântara sites), and forty 20m×10m plots in each of the seasonal deciduous forest (Anchieta and Concórdia sites), and mixed ombrophylous forest (Três Barras and Caçador sites) types (a total of 160 plots covering 24,000 m2). Based on physiognomic features, together with land ownership information, we sited 160 plots distributed across four successional stages (after Klein [18]: shrubby stage (0–8 years), small tree stage (8–15 years), Arboreal Stage (15–30 years) and advanced arboreal stage (30–60 years)).  All woody plants taller than 1.5 m were identified and measured for DBH (diameter at breast height = 1.3 m) and total height. Plant identifications were made with reference to the Angiosperm Phylogeny Group (APG) classification updated in APG II [27], and the expertise of Prof. Ademir Reis (Federal University of Santa Catarina and Curator of the Barbosa Rodriguez Herbarium) and Prof. Marcos Sobral (BHCB—Herbarium of the Botany Department at the Federal University of Minas Gerais).Species richness (number of species), Shannon-Wiener’s index of diversity (H′), evenness (J), Simpson’s diversity index (D), “importance value” (IV) and absolute frequency were estimated using the FITOPAC 1 software package [28]. Rarefaction curves (Sobs) constructed using 100 randomized orders and the Chao1, Jackknife and Bootstrap estimators were derived using EstimateS 6.0 software [29]. Differences in values of stem density and basal area among stages were tested by one-way ANOVA with Tukey’s multiple comparison tests using the STATISTICA 6.0 software package [30].


## 3. Results

A total of 13,548 woody plants were identified on the 160 plots, representing 343 species from 73 families (the appendix). The most common families represented (number of species in parentheses) were the Myrtaceae (38), Fabaceae (33), Asteraceae (27), Lauraceae (27), Rubiaceae (20) and Melastomataceae (17). Forty-four species were common to all three forest types, representing around 13% of the species total, leading to a similarity estimate of about 10%.The dense ombrophylous forest (DOF) type contained 66% (230) of all the species recorded for all forest types. Our sampling design placed more but smaller plots in the DOF formation and this likely introduced a slight bias towards finding more species in this forest type. When the differences in plot number and area sampled are accounted for (DOF2 in Table2), we estimate that the dense ombrophylous forest type would still include 59% (204) of the species identified. The seasonal deciduous forest and mixed ombrophylous forest types were very similar in their estimated species richness but markedly less rich than the dense ombrophylous forest type.Table 1
Family and species richness and the number of species common to three forest types in the Mata Atlântica of Santa Catarina, Brazil.
SDFMOFDOFAll forestsFamilies46455573Species135135230343Common speciesSDF/MOFMOF/DOFDOF/SDFSDF/MOF/DOF14192044Table 2
Phytosociological indices estimated for four successional stages in three Mata Atlântica forest formations; SDF: seasonal deciduous forest, MOF: mixed ombrophylous forest, DOF: dense ombrophylous forest;H′ is Shannon-Wiener’s index of diversity, J′ is evenness, D is Simpson’s index.
Forest TypeSuccessional stagePlots (no.)Sampling area (m2)Species richness (no.)Family richness (no.)H′a (nats/ind)J′DSDFShrubby (0–8 y)10200045242.080.4000.335Small trees (8–15 y)10200072323.250.7600.076Arboreal (15–30 y)10200094403.980.8700.026Advanced arboreal (30–60 y)10200084333.700.8400.036All stages8000135464.000.8170.034MOFShrubby (0–8 y)10200046192.740.7190.122Small trees (8–15 y)10200051223.160.7970.069Arboreal (15–30 y)10200095413.760.8220.036Advanced arboreal (30–60 y)10200081343.570.8090.052All stages8000135453.920.7990.035DOFShrubby (0–8 y)20200080303.090.7050.073Small trees (8–15 y)202000116433.350.7000.058Arboreal (15–30 y)202000163514.240.8290.025Advanced arboreal (30–60 y)202000161484.420.8670.019All stages8000230554.280.7870.031DOF2Shrubby (0–8 y)10100075293.030.7010.081Small trees (8–15y)10100092353.320.7550.076Arboreal (15–30 y)101000132464.160.8520.026Advanced arboreal (30–60 y)101000134434.420.9020.016All stages4000204534.220.7940.031aEstimated using the natural logarithm (base e).The secondary succession process in all three forest types was characterized by an increase in (1) species richness, (2) evenness scores, and (3) Shannon-Wiener diversity index values. Only the Simpson’s diversity index values decreased over time during the formation of taxonomically more diverse and structurally more complex communities (Table2).Despite the relatively high species richness that our sampling revealed in these secondary forests, the rarefaction curves (Sobs) and other estimators of richness (Chao1, Jack2 and Bootstrap; Figure2) suggest that our sampling only found about three-fourths (75% in SDF, 71% in MOF and 78% in DOF) of the maximum number of species expected to be present in these forest types.Sample-based rarefaction curves (Sobs) and species richness estimators (Chao1, Jack2 and Bootstrap) for three forest types in the Mata Atlântica.
(a)(b)(c)Species composition for all three forest types changed with changes in the length of the preceding fallow period (Table3). Although all three forests are marked by high species diversity, a few species dominate each stage of succession as demonstrated by the large Importance Values (Table 3), but this tendency decreases with increasing succession. As an example, the shrubby stage of all three forest types is dominated by Baccharis dracunculifoliaand a few other species, while the small tree stage in the MOF and DOF types is dominated by Myrsine coriaceaeand a few other species Nectandra lanceolata (SDF), Mimosa scabrela (MOF) and Miconia cinnamomifolia (DOF) dominated the Arboreal Stage and species such as Nectandra megapotamica (SDF), Ocotea puberula (MOF) and Hyeronima alchorneoides (DOF) dominated the advanced arboreal stage of each type.Table 3
Woody species most characteristic of four secondary forest successional stages in the Mata Atlântica of Santa Catarina.
Successional stageDominan speciesaFive most important speciesIV valuebSuccessional positioncSeasonal deciduous forest (SDF)Shrubby6Baccharis calvensces, B. dracunculifolia, B. elaeagnoides, Anonna silvatica, Schinus terebinthifolius.137ESmall trees9Baccharis dracunculifolia, Dalbergia frutescens, Schinus terebinthifolius, Solanum mauritianum, Trema micrantha108EArboreal16Alchornea triplinervia, Aloysia virgata, Apuleia leiocarpa, Dahlstedtia pinnata, Nectandra lanceolata64MAdvanced arboreal12Alchornea triplinervia, Cupania vernalis, Nectandra lanceolata, N.  megapotamica, Parapiptadenia rigida.100M-LMixed ombrophylous forest (MOF)Shrubby6Baccharis dracunculifolia, B. semiserrata, Eupatorium vauthierianum, Piptocarpha angustifolia, Solanum mauritianum.137ESmall trees9Baccharis dracunculifolia, Matayba elaeagnoides, Mimosa scabrella, Myrsine coriaceae, Sapium glandulatum.100EArboreal11Clethra scabra, Matayba elaeagnoides, Mimosa scabrella, Ocotea puberula, O. pulchella.97MAdvanced arboreal11Clethra scabra, Cupania vernalis, Matayba elaeagnoides, Ocotea porosa, O. puberula.108M-L/UDense ombrophylous forest (DOF)Shrubby6Baccharis dracunculifolia, Dodonaea viscosa, Leandra dasytricha, Myrsine coriaceae, Tibouchina trichotona.115ESmall trees8Miconia cabucu, M. cinnamomifolia, M. rigidiuscula, Myrsine coriaceae, Tibouchina pulchra.133E-MArboreal17Hyeronima alchorneoides, Miconia cabucu, M. cinnamomifolia, Myrsine coriaceae, Tibouchina pulchra83MAdvanced arboreal20Euterpe edulis, Marlieria eugeniopsioides, Miconia cinnamomifolia, Psychotria longipes, Tibouchina pulchra.44M-L/UaDominant species are defined as those species whose summed importance values, when ranked from the highest to lowest, contained 50% of the total for a given stand, after Finegan [14].bIV: Importance Value, here we include the proportion (from a base of 300) accounted for by the five most important species.cSuccessional position (E: early successional, M: mid successional, L: late successional, U: understory) of the five most important species.During the early stages of secondary succession, species such asTrema micrantha (SDF) and Tibouchina trichona (DOF) had low absolute frequencies but they had high densities and/or dominance when they occurred.  In the advanced arboreal stages typical understory species such as Cupania vernalis, Allophylus edulis (SDF and MOF) and Euterpe edulis (DOF) all had high frequencies of occurrence. Most species with large Importance Values were common or abundant in only one or two successional stages but were quickly replaced by other species during succession. However, a few persisted, for example, Tibouchina pulchra was common in three successional stages in the DOF.


## 4. Discussion

The sampling method used identified those species most common to the early successional stages of secondary forest regeneration in three forest types of the Mata Atlântica. The high number of species (230) found in the dense ombrophylous forest was not unexpected, as this forest type represents some 82% of all arboreal species found in Santa Catarina [26].Diversity and especially species richness increased over time since abandonment, similar to what has been found in other studies of swidden fields, abandoned plantations and pasture [13, 31–34].The peak in the numbers of species found in Arboreal Stages of all forest types is consistent with the “intermediate disturbance hypothesis” [35], that during the early stages of secondary forest succession a few pioneer species dominate but are replaced over time by species better adapted to the increasingly competitive environments that develop. Species diversity is often greatest during mid-successional stages that contain both early and late successional species. In all three forest types studied here, the Arboreal Stage (15–30 years) had a richness that was greater than for any other stage in the SDF and MOF types and equal in richness to the advanced arboreal stage of the DOF type.Species richness is only one component of diversity but because it gives the same weight to all species irrespective of their relative abundance it is strongly influenced by the number of rare species. Evenness, another component of diversity, is strongly influenced by the relative frequencies of dominant species [36, 37].  We use different measures of diversity to emphasize different diversity patterns that reflect different ecological processes.Our Shannon-Wiener diversity estimates were similar to, or slightly higher than, those of other studies of comparably aged Mata Atlântica forests. For example, Oliveira [34] and Torezan [38] estimated H′=2.51 nats/ind in five-year-old early successional stands; Pessoa et al. [39] estimated H′=3.66 nats/ind in a thirty-year-old pole stand; and Oliveira [34] estimated H′=3.33 nats/ind in a twenty-five-year-old stand and H′=3.10 nats/ind in a fifty-year-old area stand. All these studies were conducted on sites with land use histories characterized by shifting cultivation.The Shannon index is a common diversity estimate reported in the literature ranging from 3.26 to 4.36 for the mature forests of these types [40–46]. The higher values reported here for all stages of our study in relation to values reported in the literature may be due to the sampling method and criteria used for plant inclusion. Also our use of more plots, especially noncontiguous plots, likely raised the levels of beta diversity.Values of the Shannon index can be influenced by the number of species with intermediate values of relative abundance [47], and may introduce some variation in the estimates of species richness of communities [48]. As the value of Simpson’s D has an inverse relationship with the indices of Shannon and evenness [49], the value of D decreases along successional stages.The floristic composition of secondary forest formations described in this study was very similar with respect to genera reported for other studies in the Mata Atlântica of southeastern Brazil (Table4).Table 4
Plant genera with a high frequency of occurrence in secondary forests of the Mata Atlântica in Southern and Southeastern Brazil.
Forest FormationFrequent botanic genusAuthorsSeasonal deciduous forestBaccharis, Alchornea, Casearia, Inga, Solanum, Nectandra, Ocotea, Ilex, Allophylus, Apuleia, Cedrela, Cupania, Lonchocarpus, Luehea, Machaerium, Sorocea and Trema.Vaccaro and Longhi [50]; Rondon-Neto et al. [51]; Andreis et al. [52]; Hack et al. [53].Mixed ombrophylous forestBaccharis,Casearia, Chethra, Ilex, Solanum, Nectandra, Ocotea, Myrsine, Matayba, Piptocarpha, Sapium, Vernonia, Allophylus, Zanthoxylum and Capsicodendron.Rondon-Neto et al. [54]; Pezzatto et al. [55]; Narvaes et al. [56]; Ramos and Boldo [57].Dense ombrophylous forestBaccharis, Tibouchina, Myrsine, Cecropia, Alchornea, Solanum, Miconia, Nectandra, Ocotea, Jacaranda, Chethra, Ilex,Cedrela, Cupania, Psychotria, Euterpe, Guarea and Matayba.Klein [18]; Tabarelli and Mantovani [33]; Oliveira [34]; Oliveira-Filho et al. [58]; Mantovani et al. [59]; Schorn and Galvão [60]; Liebsh et al. [61].Table 5
List of families, species and number of individuals in four successional stages (SR: shrubby, ST: small trees, AR: arboreal and AA: advanced arboreal) in the Mata Atlântica forest formations, Brazil; SDF: seasonal deciduous forest, MOF: mixed ombrophylous forest, DOF: dense ombrophylous forest.
Botanic family/speciesSDFMOFDOFSRSTARAASRSTARAASRSTARAAAdoxaceae (Caprifoliaceae)Sambucus australis Cham. & Schltdl.1AnacardiaceaeLithraea brasiliensis March.1138Schinus molle L.353Schinus terebinthifolius Raddi2151822404111Tapirira guianensis Aubl.4AnnonaceaeAnnona glabra L.34Duguetia lanceolata A.St.-Hil.125Guatteria australis A.St.-Hil.1113Annona rugulosa Schl.633112110Annona sericea R.E.Fries16920Annona sylvatica A.St.-Hil.23294796Annona sp.12Xylopia brasiliensis Spreng.3427ApocynaceaeAspidosperma parvifolium A.DC.524126Rauvolfia sellowii Müll.Arg.137Tabernaemontana catharinensis A. DC.235AquifoliaceaeIlex brevicuspis Reissek1114Ilex dumosa Reissek61Ilex microdonta Reissek231Ilex paraguariensis A.St.-Hil.224357327Ilex theezans Mart. ex Reissek271197AraliaceaeSchefflera angustissima (Marchal) Frodin111143Schefflera morototoni (Aubl.) Maguire, Stey. & Frod353AraucariaceaeAraucaria angustifolia (Bertol.) Kuntze223ArecaceaeBactris setosa Mart.52819Euterpe edulis Mart.18131Geonoma gamiova Barb.1641Geonoma schottiana Mart.222Syagrus romanzoffiana (Cham.) Glassman2AsteraceaeAsteracea sp11Asteracea sp21Baccharis calvensces A.P. Candole1610142179232Baccharis dentata (Vell.) G.M.Barroso1Baccharis dracunculifolia DC.3077212031141181221Baccharis elaeagnoides Steud.14112328124Baccharis erioclada DC.31Baccharis semiserrata DC.385Baccharis sp116101Baccharis sp218132Baccharis sp311Baccharis trimera  (Less.) DC.4Baccharis uncinella DC.5Eupatorium serratum Spreng.39Eupatorium sp23272Eupatorium sp35Eupatorium sp121Eupatorium vauthierianum DC.1325113Gochnatia polymorpha (Less.) Cabrera.2Piptocarpha angustifolia Dusén ex Malme14415112013Piptocarpha tomentosa Baker.16121Vernonia discolor (Spreng.) Less.615252546373Vernonia petiolaris DC.27Vernonia puberula Less.131Vernonia sp11422Vernonia sp27Vernonia sp31BignoneaceaeJacaranda micrantha Cham.1526422153473Jacaranda puberula Cham.113133712Tabebuia sp.62Tabebuia umbellata (Sond.) Sandw.14823BoraginaceaeCordia ecalyculata Vell113912Cordia americana (L.) Gottschling & J.S. Mill.3235BurseraceaeProtium kleinii Cuatrec.31Cannabaceae (Ulmaceae)Celtis triflora (Klotszch.)Miq.4Trema micrantha (L.) Blume8186411771CanellaceaeCapsicodendron dinisii (Schwacke) Occhioni23231Cardiopteridaceae (Icacinaceae)Citronella paniculata (Mart.) R.A.Howard214CaricaceaeCarica quercifolia (A.St.-Hil.) Hieron2CelastraceaeMaytenus muelleri Scheacke3141Maytenus robusta Reiss.122ChloranthaceaeHedyosmum brasiliensis Mart.1917ChrysobalanaceaeHirtella hebeclada Moric. ex DC.1238ClethraceaeClethra scabra Pers.1214242922210607ClusiaceaeCalophyllum brasiliense Cambess.41Clusia parviflora (Saldanha) Enfler9247Garcinia gardneriana (Planch. & Triana) Zappi158CombretaceaeTerminaliacf.australis Cambess.21CunoniaceaeLamanonia speciosa (Camb.) L.B. Smith.3357Lamanonia ternata Vell.2Weinmania humilis Engl.12Weinmannia paulliniifolia Pohl ex Ser.341CyatheaceaeAlsophila sp.121Cyathea schanschin Mart.2233Cyathea vestita Mart.82435DicksoniaceaeDicksonia sellowiana Hook32EbenaceaeDiospyros inconstans Jacq.1031ElaeocarpaceaeSloanea guianensis (Aubl.) Benth.11ErythroxylaceaeErythroxyllum deciduum A.St.-Hil.2Erythroxylumcf.cuneifolium (Mart.) O.E. Schulz3Erythroxylum myrsinites Mart.42EuphorbiaceaeAlchornea iricurana Casar.11Alchornea sidifolia Müll.Arg.322224Alchornea triplinervia (Spreng.) Müll. Arg.15232811571276Gymnanthes concolor Spreng.1202462114Manihot grahami Hook.6114110215Pausandra morisiana (Casar.) Radlk.33Pera glabrata (Schott) Baill145325Ricinus communis L. (exótica)2Sapium glandulosum (L.) Morong11162018111711Sebastiania commersoniana (Baill.) L.B. Smith & R.J.Downs1343Tetrorchidium rubrivenium Poepp. & Endl.1202127PhylanthaceaeHyeronima alchorneoides Fr. Allem.32587839Richeria australis Mull.Arg.1FabaceaeAbarema langsdorfii (Benth.) Barneby & J.W. Grimes25Albizia polycephala (Benth.) Killip5Andira anthelmintica Benth.151Andira fraxinifolia Benth.1Apuleia leiocarpa (Vog.) Macbr.22511Bauhinia forficata Link.211519Calliandra selloi (Spreng.) Macbr.361Copaifera trapezifolia Hayne3Dahlstedtia pinnata (Benth.) Malme736Dalbergia frutescens Britton177319362Enterolobium contortisiliquum (Vell.) Morong.1Holocalyx balansae Micheli2Inga marginata Willd.262077362Inga sessilis (Vell.) Mart.4221Inga vera Willd.1171Inga virescens Benth.113181Lonchocarpus campestris Mart. ex Benth54391029171112Lonchocarpus guilleminianus (Tul.) Malme312710121Lonchocarpus sp.911Machaerium hirtum (Vell.) Stell Feld2Machaerium paraguariensis Hassler62221221Machaerium sp.149Machaerium stipitatum (DC.) Vog.343324131322Mimosa bimucronata (DC.) O. Ktze.61623Mimosa scabrella Benth813752Myrocarpus frondosus Allem.23Ormosia arborea (Vell.) Harms.11Papilionaceae sp.1Parapiptadenia rigida (Benth.) Brenan5111537Peltophorum dubium (Spreng.) Taub.1Piptadenia gonoacantha (Mart.) Macbr.63Schizolobium parahyba (Vell.) Blake3Zollernia ilicifolia Vog.194LamiaceaeAegiphila sellowiana Cham.5235Vitex megapotamica (Spreng.) Moldenke231LauraceaeCinnamomum amoenum (Nees) Kosterm.11Cryptocarya aschersoniana Mez3Cryptocaryacf. moschata Nees et Mart. ex Nees10Endlicheria paniculata (Spreng.) Macbride338Lauraceae sp111Lauraceae sp21Nectandra lanceolata Nees et Mart. ex Nees2286773163961211Nectandra leucothyrsus Meissn.123Nectandra megapotamica Mez.14189064111220Nectandra membranacea (Sw.) Griseb.610Nectandra oppositifolia Nees121013Nectandra rigida (Kunth) Nees12Nectandra sp11Nectandra sp21Nectandra sp36Ocotea catharinensis Mez1Ocotea diospyrifolia (Meisn.) Mez141329221Ocotea odorifera (Vell.) Rohwer55Ocotea porosa (Nees & C. Mart.) Barroso14119Ocotea puberula (Rich.) Nees24716533812Ocotea pulchella Mart.21017331810Ocotea sp13Ocotea sp23Ocotea teleiandra (Meissn.) Mez1Persea americana Mill.4Persea major (Nees) L. E. Kopp1Persea sp.21LoganiaceaeStrychnos brasiliensis (Spreng.) Mart.1365LythraceaeLafoensia pacari A.St.-Hil.1MagnoliaceaeMagnolia ovata (A.St.-Hil.) Spreng.21MalpighiaceaeBunchosia maritima (Vell.) J. F. Macbr.2Byrsonima ligustrifolia A.Juss.36MalvaceaeBastardiopsis densiflora (Hook. & Arn.) Hassler531Luehea divaricata Mart.114821171Sida rhombifolia L.2MelastomataceaeHuberia semiserrata DC.1029Leandra australis (Cham.) Cogn.3Leandracf. dasytricha (A. Gray) Cogn.217313Leandra sp1631166Leandra sp24139411Miconia cabucu Hoehme27610931Miconiacf.latecrenata(DC.) Naudin48Miconia cinnamomifolia (DC.) Naudin744419723Miconia cubatanensis Hoehne245Miconia flammea Cesar.5Miconia ligustroides (DC.) Naudin17Miconia rigidiuscula Cogn.1163973Miconia sp1371Mouriri chamissoniana Cogn.32Tibouchinacf.trichopoda Baill.1202127Tibouchina pulchra Cogn.4421112643Tibouchina sellowiana (Cham.) Cogn.63510MeliaceaeCabralea canjerana (Vell.) Mart.2613411424Cedrela fissilis Vell.1219191554321Guarea macrophylla Vahl11142026Trichilia catigua A.Juss.42Trichilia elegans A.Juss.71325175Trichilia lepidota Mart.46MonimiaceaeMollinedia sp.1315Mollinedia triflora (Spreng.) Tul231326MoraceaeBrosimum lactescens (S. Moore) C.C. Berg1148Ficuscf.insipida Willd.2Ficus organensis (Miq.) Miquel12Ficus sp.1Maclura tinctoria (L.) D.Don ex Steud.144Sorocea bonplandii (Bail.) Burg., Lanj & Boer42294MyristicaceaeVirola bicuhyba Schott1924MyrsinaceaeMyrsinecf. lancifolia Mart.1Myrsine coriacea (Swartz) R. Brown ex Roemer & Schultz13321355212116277717Myrsine parviflora A. DC.1165Myrsine umbellata Mart ex. DC.221176227219128MyrtaceaeBlepharocalyx salicifolius (Kunth) O. Berg12Calyptrantes sp.21Calyptranthescf.grandifolia O. Berg.9Campomanesia guaviroba (DC.) Kaiaerskou12Campomanesia guazumifolia (Cambess.) O. Berg32Campomanesia reitziana Legr.1113Campomanesia sp.4Campomanesia xanthocarpa  O. Berg143724Eugenia burkartiana (Legr.) Legr.1Eugenia chlorophylla O. Berg2Eugenia involucrata DC.2212Eugenia neoaustralis Sobral1Eugenia playtsema O.Berg12Eugenia pyriformis Camb.135Eugenia uniflora L.121Gomidesia palustris (DC.) D.Legrand11Gomidesia schaueriana O.Berg1Gomidesia spectabilis (DC.) O. Berg251319Gomidesia tijucensis (Kiaerskov) Legr.11Marlieriacf.eugeniopsioides (Kaus. & Legr.) Legr.32538Marlierea parviflora O. Berg.4Marlierea tomentosa Cambess.112023Mosiera prismatica (D. Legrand) Landrum317Myrceugenia sp.1Myrcia gracilis O.Berg22Myrcia guianensis (Aubl.) DC.12Myrcia multiflora (Lam.) DC.12Myrcia palustris DC.1Myrcia rostrata DC.173332210Myrcia splendens (Sw.) DC.1Myrcia tenuivenosa Kiaersk.3Myrcianthes pungens (O. Berg) Legr.1Myrciaria tenella (DC.) O. Berg.1222Myrtaceae sp1220Myrtaceae sp2211Myrtaceae sp32Myrtaceae sp4210Psidium cattleyanum Sabine1932104NyctaginaceaeGuapira hirsuta(Choisy) Lundell14Guapira opposita(Vell.) Reitz32430Pisonia ambiguaHeimerl14145OchnaceaeOuratea parviflora(DC.) Baill.1916OleaceaeChionanthus filiformis(Vell.) P.S. Green1OlacaceaeHeisteria silvianiiSchwacke112PhytolaccaceaePhytolacca dioicaL.62Seguieria glazioviiBriq.1Picramniaceae (Simaroubaceae)Picramnia parvifoliaEngl.2Picramnia sp.3PiperaceaePiper arboreumAubl55Piper cernuumVell.533Piper gaudichaudianumKuntze42411228PodocarpaceaePodocarpus lambertiiKlotzsch18PolygonaceaeCoccoloba warmingiiMeisn.131Ruprechtia laxifloraMeisn.15122ProteaceaeRoupala asplenioidesSleumer9Roupala cataractarumSleumer7Roupalacf.brasiliensisKlotzsch1QuiinaceaeQuiina glazioviiEngl.1211RhamnaceaeHovenia dulcisThumb. (exótica)30161Rhamnaceae sp131Scutia buxifoliaReissek.157RosaceaePrunus brasiliensis(Cham. & Schlechtd) D. Dietr.141Prunus myrtifolia(l.) Urb.5Prunus sellowiiKoehne12113Prunus sp.11RubiaceaeAlibertia concolor (Cham.) K. Schum.122Amaioua guianensis Aubl.994Bathysa australis (A.St.-Hil.) Benth. & Hook.F.432415Chomelia pedunculosa Bentham15Posoqueria latifolia (Rudge) Roem. & Schult.1256Psychotria carthagenensis Jacq.24Psychotria leiocarpa Cham.197Psychotria longipes Muell.Arg.1186243Psychotria nuda (Cham. and Schltdl.) Wawra2810Psychotria officinalis (Aubl.) Raeusch. ex Sandwith1514Psychotria sp1312Psychotria sp2136Psychotria sp323Psychotria sp413445Psychotria stenocalyx Müll.Arg.2921Psychotria suterella Müll. Arg.2211522Randia armata (Sw.) DC.5122Rubiaceae sp1312Rubiaceae sp25Rudgea jasminioides (Cham.) Müll.Arg.132125Ruscaceae (Agavaceae)Cordyline dracaenoides Kunth317RutaceaeBalfourodendron riedelianum (Engl.) Engl.45Esenbeckia grandiflora Mart.55Helietta apiculata Benth.2Pilocarpus pennatifolius Lem.12Zanthoxylumcf. astrigera Cowan1Zanthoxylum hiemalis A.St.-Hil.131Zanthoxylum petiolare A.St.-Hil. & Tul.3Zanthoxylum rhoifolia (Lam.) Engl.137811126912173SabiaceaeMeliosmacf.sellowii Urban.32Salicaceae (Flacourtiaceae)Banara parviflora (A. Gray) Benth.257Banara tomentosa Clos.15Casearia decandra Jacq.65427212223331Casearia sylvestris Sw.21192948856723Xylosma pseudosalzmanniiSleumer.11SapindaceaeAllophylus edulis (A.St.-Hil. & et al.) Radlk1493919232015Allophylus guaraniticus (A.St.-Hil.) Radlk33626Cupania oblongifolia Mart.22Cupania vernalis Camb.21334654924752210517Diatenopteryx sorbifolia Radlk.5Dodonaea viscosa (L.) Jacq.127181Matayba elaeagnoides Radlk.61525283093140Matayba guianensis Aubl.43218SapotaceaeChrysophyllum gonocarpum (Mart. & Eichl.)Engl.168Chrysophyllum marginatum (Hook. & Eichler) Engl.1310111Pouteria venosa (Mart.) Baehni283SolanaceaeCapsicumcf. flexuosum Sendtn2Cestrum strigilatum Ruiz & Pav.14Solanaceae sp14Solanum diflorum Vell.9292111341692Solanum mauritianum Scopoli13642Solanum pseudoquina A.St.-Hil.2711Solanum sanctae-catharinae Dunal301Solanum sp22Solanum sp31336111248StyracaceaeStyrax leprosus Hook. & Arn.1122SymplocaceaeSymplocos celastrina Mart.2Symplocos lanceolata (Mart.) A.DC.1Symplocos sp1543Symplocos tenuifolia Brand325Symplocos uniflora (Pohl) Benth.12TheaceaeLaplacea fruticosa (Schrad.) Kobuski.48UrticaceaeBoehmeria caudata Sw13307Cecropia glaziovii Snethl.3328102Urera baccifera (L.) Gaudich. ex Wedd3VerbenaceaeAloysia virgata (Ruiz & Pav.) Juss.12311Citharexylum myrianthum Cham.31Lantana camara L.21WinteraceaeDrimys brasiliensis Miers89UndeterminedNonidentified36110237615DeadDead651215971361


## 5. Conclusion

The floristic diversity observed for the forests we surveyed largely agree with patterns of post-agricultural secondary forest succession observed for many other neotropical forests. The chronosequence covers a relatively large range of successional ages (0–60 years) that results in large changes in species diversity and composition despite a limited geographical sampling. This result suggests that the patchy mosaic of secondary forests in Santa Catarina has a high potential for biodiversity conservation. Whether it can also provide for economic development in the way of nontimber forest products would depend upon the life histories of individual species and any rules instituted to guide their sustainable management.The results of this study demonstrate that the patterns of secondary succession appear very similar among the three forest types with respect to changes in species richness and other measures. However, the species composition among the types, while quite similar in the earliest shrubby stage, diverges during succession, with the largest differences noted between the DOF and the other two types. As expected, and despite high species richness, a small set of “pioneer species” dominates the shrubby and small tree stages until about age 15. After that time, more shade-tolerant species increase during the Arboreal and advanced arboreal stages of forest succession, but only a small number become structurally important after 25–30 years.A chronosequence approach only allows us to infer successional changes because we do not analyze the underlying processes mediating these changes on our sites (growth, mortality, and recruitment).  Thus, we suggest that long-term permanent plots be established in this region, to improve our understanding of secondary forest dynamics, while also creating a framework for future comparative studies of the role of ecological processes and mechanisms in different successional stages.



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*Source: 759893-2011-04-28.xml*