Who is who in the understory: the contribution of resident and
Contents
1. E B Almeida Jr 8 C S Zickel 2009 Riqueza de esp cies de sub bosque em um fragmento florestal urbano Pernambuco Brasil Biotemas 22 57 66 Souza A L G F Silva J F Chichorro amp R L Caraciolo 2006 Mata Nativa 2 Manual do usu rio Cientec Vi osa Minas Gerais Brasil Tabarelli M amp W Mantovani 1999 A regenera o de uma Floresta Tropical Montana ap s corte e queima S o Paulo Brasil Rev Bras Biol 59 239 250 The Internacional Plant Names Index 2010 Downloaded 10 January 2011 http www ipni org Trindade M A C B Lins e Silva H P Silva S B Figuei ra amp M Schessl 2008 Fragmentation of the Atlantic Rainforest in the northern coastal region of Pernam buco Brazil recent changes and implications for conservation Biorem Biodiv Bioavail 2 5 13 Triquet A M G A Mcpeek amp W C Mccomb 1990 Song bird diversity in clear cuts with or without a Riparian Buffer Strip J Soil Water Conserv 45 500 503 Tropicos 2010 Missouri Botanical Garden Missouri USA Downloaded 10 January 2011 http www tropicos org V lio I F M 2003 Seedling Growth of Understorey Spe cies of a Southeast Brazilian Tropical Forest Braz Arch Biol Techn 46 697 703 Vilela E A A T Oliveira Filho D A Carvalho amp M L Gavillanes 1995 Flora arbustivo arb rea de um fragmento de mata ciliar no Alto Rio Grande Itutin ga Minas Gerais Acta Bot Bras 9 87 100 Vol2. Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1029 estimated using rarefaction mean richness cal culated values were 34 4 12 93 median SD from the canopy and 47 69 18 09 from the understory not significantly different from observed values in both strata All of the frag ments except F3 were richer although not sig nificantly in the understory An average of 35 SD 16 76 of the species occurred exclusively in the understory while 15 2 10 18 of the species only occurred within the tree stratum of the riparian forests Of the total 75 4 of the species found in the tree stratum were present in the understory The similarity between the tree and understory strata averaged 0 54 0 09 Table 1 Fragment F3 had the highest similar ity 0 68 while fragments F4 and F5 were the least similar 0 46 and 0 45 respectively Families Myrtaceae 17 species 227 indi viduals Melastomataceae 13 327 Rubiace ae 10 247 Fabaceae Mimosoideae 10 234 Annonaceae 9 461 and Clusiaceae 7 266 had the highest species richness and added up to 40 50 of the understory s total species The lower understory was made up mainly of spe cies from families Rubiaceae and Melastomata ceae which totalled 52 17 of the species and 24 35 of individuals in this category while species from families Annonaceae Clusia ceae and Myrtaceae predominated in the upper understory 66 66 of the spec
3. respectively Fragment F5 differed from the others because it had a greater percentage of individuals in the lower understory 30 79 and a smaller per centage of individuals in the upper understory 12 22 in relation to fragments F1 G 29 30 p 0 05 F2 G 19 50 p lt 0 05 and F3 G 54 82 p lt 0 05 Fragment F4 was different due to its higher percentage transitory species 76 06 in comparison to the other fragments In this remnant the upper understory had the lowest value 12 23 and was different from fragments Fl G 16 10 p lt 0 05 F2 G 10 74 p lt 0 05 and F3 G 33 13 p lt 0 05 Significant differences were also found between fragments F2 and F3 G 13 06 p lt 0 05 which showed opposite trends in relation to the percentage of resident species F3 s lower understory had a smaller percentage 1 08 while the percent age of upper understory was larger 44 80 Among the 163 species recorded in the five fragments understory looking at the 10 with highest regeneration indexes TNR in each fragment 34 species are listed Table 2 These species include the most important in at least one fragment in which they added up to 10 75 31 25 of the total natural regeneration Transitory species were the majority with 21 recorded species 61 76 while the lower and upper understory had six 17 65 and seven species 20 59 respectively 1030 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 102
4. 2008 and Santos et al 2008 used Volpato s modified version for describing natural regeneration in Brazilian forests This latter incorporates a more detailed analysis per height class when calculating the index which considers the relative density of the species in each class In order to calculate the Natural Regen eration Index absolute and relative density and frequency parameters were estimated by height class for each species of each fragment The natural regeneration estimate by height class NRC was made based on these param eters according to Volpato s 1994 equation NRC RD RF 2 where NRC is the natu ral regeneration estimate of the i species of the j plant size class as a percentage RD is the relative density for the i species of the j natural regeneration size class RF is the relative frequency for the i species of the j natural regeneration size class i is the 1 2 3 p species sampled and j is the 1 2 3 and 4 classes The next step was to calculate the Total Natural Regeneration estimate TNR for each species by adding the natural regeneration indexes of each class according to the formula by Volpato 1994 TNR YNRC where TNR is the total natural regeneration estimate of the i species NRC is the natural regeneration estimate of the i species of the j plant size class i is the 1 2 3 p species sampled and jis the 1 2 3 and 4 classes RESULTS Species richness
5. 2008 who recorded an increase in the similarity between the canopy and the understory from a more disturbed area 0 45 edge to a more conserved one 0 59 forest interior in a 300ha mature Atlantic forest fragment In fragmented landscapes analyzing the understory s structure and identifying its func tional components has been crucial to diag nose the effects of disturbances Martins amp Rodrigues 2002 Ceccon et al 2006 fragmen tation and edge effects Benitez Malvido 1998 Laurance et al 2007 Bouroncle amp Finegan 2011 In the Brazilian Atlantic Forest where most areas have been substituted for cultivated land it is estimated that only 11 6 of the original cover remains Ribeiro et al 2009 Yet despite its status as a world conservation hotspot Mittermeier et al 2005 and the fact that many plants spend all or a considerable 1026 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 part of their lives in the understory Harms et al 2004 few studies have analyzed the under story vertical structure e g Silva et al 2007 Marangon et al 2008 described who is who in this stratum regarding plant habit Souza et al 2009 or successional category Tabarelli amp Mantovani 1999 Aquino amp Barbosa 2009 Onofre et al 2010 Sansevero et al 2011 or compared understory composition and struc ture among forests Gomes ef al 2009 Only 4 6 of the original vegetati
6. J M Mello amp MLL Gavilanes 1994 Estrutura fitossociol gicas e vari veis ambientais em um trecho de mata ciliar do c rrego dos Vilas Boas Reserva Biol gica do Po o Bonito Lavras MG Rev Bras Bot 17 67 85 Onofre F F V L Engel amp H Cassola 2010 Regenera o natural de esp cies da mata atl ntica em sub bosque de Eucalyptus saligna Smith em uma antiga unidade de produ o florestal no Parque das Neblinas Bertio ga SP Sci For 38 39 52 Rayol P B M F F Silva amp F O Alvino 2006 Din mica da regenera o natural de florestas secund rias no munic pio de Capit o Po o Para Brasil Amaz nia Ci ncia e Desenvolvimento 2 93 109 Ribeiro M C J P Metzger A C Martensen F J Ponzoni amp M M Hirota 2009 The Brazilian Atlantic Forest How much is left and how is the remaining forest distributed Implications for conservation Biol Con serv 142 1141 1153 Richards P W 1996 The tropical rain Forest an ecological study Cambridge Cambridge England Salles J C amp I Schiavini 2007 Estrutra e composi o do estrato de regenera o em um fragmento florestal urbano implica es para a din mica e a conserva o da comunidade arb rea Acta Bot Bras 21 223 233 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1039 Sansevero J B B P V Prieto L F D Moraes amp P J Rodri gues 2011 Natural Regeneration in Plantati
7. Nativa 2 software Souza et al 2006 The botanical material collected was added to the Herbarium Geraldo Mariz UFP of the Federal University of Pernambuco Usina S o Jos Collection When possible samples were identified up to the species level with the aid of specific literature comparison with material from the UFP PEUFR and IPA herbaria that had been previously identified by specialists When necessary the material was sent to specialists for identification and or confirmation All of the species collected were listed alphabetically according to family the APG III 2009 classification system was used Species names were updated regarding synonymies based on the Missouri Botanical Garden website Tropicos 2010 author names and or abbreviations followed the International Plant Names Index IPNI 2010 Analysis of data structure and composi tion The woody flora of the studied understory was classified into functional stratification cat egories 1 transitory species TR represented by young individuals of the trees that make up the forest s canopy and 2 typical understory species i e resident species such as shrubs and small trees that develop under the canopy The typical understory species were also divided into a lower understory LU which included species that make up the lowest forest stratum and do not surpass 5m in height on average and b upper understory UU which included sciophilous species that a
8. abundance of these families do not seem to be good indicators of Atlantic Forest successional stages but yet dependent on the sampling criteria used and on complex characteristics of species life history Clark amp Clark 1992 Species classification into stratification categories In a manner similar to that already recorded by other authors Vilela et al 1995 Salles amp Schiavini 2007 Gomes et al 2009 the understory of the studied fragments is also composed primarily by species of the canopy which are passing through the lower strata Due to the different methodologies used for sampling this stratum widely varied percent ages for tree species have been recorded in the understory ranging for example from 54 54 Vilela et al 1995 to 89 18 Gomes et al 2009 Despite this variation all studies show that the understory is an important transitional environment that serves as an ecological filter for the species that will be part of the canopy i e an environment that selectively determines which individuals and species may survive under its conditions George amp Bazzaz 1999 Harms et al 2004 Few species were abundant and most had low density values this pattern is expected both for the understory L et al 2010 and for the tree stratum Richards 1996 These species classified as rare or locally rare gen erally occur in abundance percentages above 25 Nappo et al 2004 Oliveira amp Amaral 2005 di
9. and composition A total of 163 species distributed among 44 families were sampled from the understory of the five fragments analyzed fragment median 63 8 species SD 21 72 n 5 varying from 32 spe cies in F3 to 93 species in F1 Table 1 One hundred twenty six species of 44 families were sampled from the tree stratum The number of species in this stratum varied from 32 in F2 to 69 in F1 43 8 18 14 6 When richness was TABLE 1 Species richness for the total sample for the tree stratum and for the understory in riparian environments of five Atlantic Forest remnants in Pernambuco Brazil Fi E2 ES Understory density 736 643 558 Tree density 326 320 441 Total richness 117 72 50 Tree richness 69 32 44 Tree richness 51 24 30 Understory richness 93 66 32 Understory richness 74 48 24 Similarity Sorensen 0 56 0 53 0 68 Tree exclusives 24 6 18 Understory exclusives 48 40 6 Species with N gt 15 11 83 10 61 31 25 F1 F5 fragments 1 to 5 M mean SD standard deviation F4 FS Total M SD 981 630 3 368 710 164 40 218 162 1467 293 107 83 92 64 192 79 26 12 52 22 126 43 8 18 14 45 22 34 4 12 93 67 61 163 63 8 21 72 50 42 47 7 18 9 0 45 0 46 0 54 0 09 25 3 15 2 10 18 40 43 35 4 16 76 20 9 19 67 37 18 85 8 03 All species numbers are estimated for a sample of 160 tree individuals All species numbers are estimated for a sample of 531 understory individuals
10. regeneration classes based on a regeneration index The understory was richer in spe cies median 63 8 SD 21 72 n 5 fragments than the tree stratum 43 8 18 14 5 and the similarity between these components was relatively high median 0 54 SD 0 09 The results also showed that the studied under story in the forest fragments was mainly composed by transitory species median 67 01 SD 3 76 that were well distributed among height classes and had the highest densities which may favor their future presence in the canopy s structure and composition The typical understory species were grouped into two strata the lower understory made up of species that generally do not reach more than 4m in height mostly species from families Piperaceae Rubiaceae and Melastomataceae and the upper understory with intermediate heights between the lower understory and the canopy but with average heights that were not higher than 10m mainly of species from families Anonnaceae Clusiaceae and Myrtaceae These families richness was commonly used as an indi cator of the vegetation s successional stage however such results must be seen with caution as they show that these families co occurred and were highly important in different strata Studying the understory is fundamental because it represents a floristically rich stratum with a unique structure which promotes the natural regeneration of the tree stratum Rev Biol Trop 60 3 1025 1040 Epub 2012 Septem
11. 20 25 A TNR 96 ji 20 40 60 80 100 120 140 160 180 o I 20 40 60 80 100 120 140 160 18 o pr o 20 40 60 80 100 120 140 160 180 80 120 160 200 240 280 oy 40 I 20 40 60 80 100 120 140 160 180 B Density 0 Fig 2 Species with the highest values of Total Natural Regeneration TNR A and or densities equal or higher than 15 individuals B in riparian environments of Atlantic Forest remnants in Pernambuco Brazil Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1033 categories SC 23 transitory species 62 16 six lower understory species 16 22 and seven upper understory species 18 92 Spe cies distribution between the different height classes was similar among the fragments The second regeneration class encompassed the largest number of species Fig 3A The transi tory species were the best represented in all of the classes followed by those from the upper and lower understory Fig 4 The average number of typical lower and upper understory species was similar for classes 1 and 2 In all of the forest remnants the species that occurred in only one class were concen trated in the second height class except for F3 where most occurred solely in class 4 Fig n mMm F2 E s ISS E a INSS o FS L OO 4 a Gi ai Cn ASS 2 aiL F3 ES Fa ANN FS AR E Gs u
12. 45 16 in Fl Transitory species obtained the greatest averages in all of the situations while the aver age number of species in the lower and upper understory varied from 1 3 DISCUSSION Species richness and composition In this study the understory generally contributed to an enhancement on plant richness in the for est fragments This confirms the pattern that is normally recorded in the literature on tropical forests Gentry amp Dodson 1987 Galeano et al 1998 Schnitzer amp Carson 2000 L et al 2010 as the lower stratum encompasses tran sitory and resident species In these forests the understory flora may represent 50 or more of the total species Schnitzer amp Carson 2000 and augment the list of tree species up to 30 Lins e Silva 2010 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 According to Jardim amp Hosokawa 1986 the floristic composition of tropical forests is very different for the upper stratum and under story Nevertheless the 54 average similarity among the understory and tree stratum of the different fragments studied here was higher than what is commonly found in the literature Alves amp Metzger 2006 It is important to point out that the herbaceous understory was not sampled and the upper understory was included which may have contributed to the high similarity found between the canopy and understory The high richness found by this s
13. 5 1040 September 2012 S pl IN Fragments o NS a NNE s NNE 40 60 80 100 mL Nem 2 D as SE BA UOV Soo Cs KWC CW CWT E T T 096 2096 4096 6096 8096 10096 Fig 1 Species stratification categories SC presented in the understory in riparian environments of five Atlantic Forest remnants Pernambuco Brazil A Number of species and B number of individuals in each SC Transitory Species white Lower Understory stripes Upper Understory grey and Non Classified black Regenerative capacity and vertical structure of the understory Most species with high total natural regeneration indices TNR were also well distributed among height classes and had the highest densities Fig 2 Only 37 species less than 21 of the total richness had densities equal to or higher than 15 individuals in the total sample Table 1 the highest percentage was recorded for F3 which stood out for housing 31 25 of the species with 15 or more individuals Species such as Inga thibaudiana DC Myrcia racemosa O Berg Kiaersk Miconia prasina Sw DC F2 and Pilocarpus cf giganteus Engl F3 all had less than 15 individuals each Nevertheless they were among those with the highest TNR as they were recorded in all of the height class es the exceptions were M prasina and P cf Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 giganteus which we
14. SP Biota Neotrop 6 1 10 APG III 2009 Angiosperm Phylogeny Group APG III An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants APG III Bot J Linn Soc 161 105 121 Aquino C amp L M Barbosa 2009 Classes sucessionais e s ndromes de dispers o de esp cies arb reas e arbus tivas existentes em vegeta o ciliar remanescente Conchal SP como subs dio para avaliar o potencial do fragmento como fonte de prop gulos para enrique cimento de reas revegetadas no rio Mogi gua u SP Rev rvore 33 349 358 Ara jo F S S V Martins J A A Meira Neto J L Lani amp LE Pires 2006 Estrutura da vegeta o arbus tivo arb rea colonizadora de uma rea degradada p s minera o de caulim em Br s Pires MG Rev rvore 30 107 116 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1037 Armesto J J amp J A Mart nez 1978 Relations between vegetation structure and slope aspect in the Medite rranean region of Chile J Ecol 66 881 889 Benitez Malvido J 1998 Impact of forest fragmentation on seedling abundance in a tropical rain forest Con serv Biol 12 380 389 Bohlman S amp S O Brien 2006 Allometry adult stature and regeneration requirement of 65 tree species on Barro Colorado Island Panama J Trop Ecol 22 123 136 Bouroncle C amp B Finegan 2011 Tree Regeneration and Under
15. Who is who in the understory the contribution of resident and transitory groups of species to plant richness in forest assemblages Juliana Silva Gomes Westphalen Ana Carolina Borges Lins e Silva amp Francisca Soares de Ara jo A Universidade Federal do Cear Campus do Pici Programa de P s Gradua o em Ecologia e Recursos Naturais Bloco 906 60455 760 Fortaleza CE Brasil juli ufrpe o gmail com tchesca ufc br Universidade Federal Rural de Pernambuco Departamento de Biologia rea de Ecologia 52171 900 Recife PE Brasil anacarol db ufrpe br Corresponding author Received 11 VIII 2011 Corrected 06 11 2012 Accepted 05 11 2012 Abstract The forest understory is made up of resident and transitory species and can be much richer than the canopy With the purpose to describe the contribution of these groups to the woody understory five Atlantic Forest fragments were selected and studied in Northeastern Brazil In order to analyze the understory s struc ture the sample included woody individuals with circumference at breast height CBH smaller than 15cm and circumference at ground level CGL greater than 3cm regardless of height The recorded species were quanti fied and classified into functional stratification categories resident and transitory and the floristic similarity between the understory and the tree stratum was calculated Species importance in the understory was analyzed by height and total natural
16. alanaceae Hirtella racemosa Lam 4 85 0 23 1 09 0 53 TR Clusiaceae Symphonia globulifera L f 0 89 11 47 3 46 6 51 TR Tovomita mangle G Mariz 7 78 UU Erythroxylaceae Erythroxylum citrifolium A St Hil 3 79 0 70 1 86 0 45 TR Euphorbiaceae Pogonophora schomburgkiana Miers ex Benth 3 61 4 04 TR Fabaceae Mimosoideae Albizia pedicellaris DC L Rico 0 19 0 23 6 15 0 83 TR Albizia saman Jacq Merr 4 09 TR Inga thibaudiana DC 0 75 2 19 4 40 1 28 TR Lecythidaceae Eschweilera ovata Cambess Miers 8 56 10 38 2 19 20 61 2 40 TR Malvaceae Pavonia malacophylla Nees amp Mart Garcke 3 72 LU Melastomataceae Henriettea succosa Aubl DC 0 19 8 05 3 64 2 70 TR Leandra rufescens DC Cogn 345 LU Miconia minutiflora Bonpl DC 3 06 0 30 1 34 UU Miconia prasina Sw DC 1 08 2 1 3 42 10 92 TR Meliaceae Trichilia lepidota Mart 2 03 2 81 1 09 TR Myrtaceae Calyptranthes brasiliensis Spreng 0 38 0 70 8 30 0 31 2 52 TR Eugenia candolleana DC 3373 UU Myrcia racemosa O Berg Kiaersk 0 94 2 11 2 08 2 65 1 65 UU Nyctaginaceae Guapira opposita Vell Reitz 0 38 0 23 2 62 0 16 0 22 TR Phylantaceae Phyllanthus juglandifolius Willd 3 86 LU Piperaceae Piper marginatum Jacq 9 04 LU Rubiaceae Gonzalagunia dicocca Cham amp Schltdl 2 94 LU Psychotria carthagenensis Jacq 0 94 17 71 1 21 1 50 0 83 UU Rutaceae Pilocarpu
17. ber 01 Key words height classes floristic composition density natural regeneration richness functional groups Several life forms coexist in the forest understory which contributes to high species richness Gentry amp Dodson 1987 Schnitzer amp Carson 2000 There are two main approaches to view the regeneration dynamics of this stratum either as a physiognomic component defined by a maximum height or diameter including young trees seedlings saplings and shrubs regardless of their potential to occupy the forest vertically for example Campos amp Landgraf 2001 Lima Filho et al 2002 Rayol et al 2006 Silva et al 2007 Marangon et al 2008 Gomes et al 2009 L et al 2010 San severo et al 2011 or as functional groups of plants that occupy the lowest level of the forest and might include resident or transitory species Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1025 Tabarelli amp Mantovani 1999 Harms et al 2004 Ara jo et al 2006 Santos et al 2008 Aquino amp Barbosa 2009 Onofre et al 2010 The resident group of species which may be called the typical understory is made up of herbaceous and sprawling species as well as shrubs and small trees that remain in this stratum throughout their life cycle Conversely the transitory group also called the natural regeneration group includes species that have the potential to grow and reach the fo
18. biosfera da Mata Atl n tica em Pernambuco situa o atual a es e perspec tivas Caderno 12 Gr fica do Instituto Florestal S o Paulo S o Paulo Brasil Lima Filho D A J Revilla L S Coelho J F Ramos J L Santos amp J G Oliveira 2002 Regenera o natural de tr s hectares de Floresta Ombr fila Densa de Terra Firme na regi o do rio Urucu AM Brasil Acta Amaz 32 555 569 Lins e Silva A C B 2010 Influ ncia da rea e da hete rogeneidade de habitats na diversidade vegetal em fragmentos de floresta atl ntica Ph D Thesis Uni versidade Federal do Rio de Janeiro Rio de Janeiro Brasil Li X T J X Yin amp J W Tang 2010 Diversity and com position of understory vegetation in the tropical sea sonal rain forest of Xishuangbanna SW China Rev Biol Trop 59 455 463 Magurran A E 2004 Measuring Biological Diversity Blackwell Science Oxford England Marangon L C J J Soares A L P Feliciano amp C F Bran d o 2008 Regenerac o natural em um fragmento de floresta estacional semidecidual em Vigosa Minas Gerais Rev rvore 32 183 191 Martins S V amp R R Rodrigues 2002 Gap phase regene ration in a semideciduous mesophytic forest South eastern Brazil Plant Ecol 163 51 62 Mattei V L amp S J Longhi 2001 Avalia o da regenera o natural de Eucalyptus paniculata Smith Ci Fl 11 55 65 Metzger J P L C Bernacci amp R Goldenberg 1997 Pattern
19. e fragments were stud ied all from the extreme North of the state of Pernambuco Northeastern Brazil 07 41 05 07 54 17 S and 34 54717 35 05 07 W at 20 50m in elevation within properties of the Sao Jos sugarcane plantation processing plant For the purposes of this study the fragments were named Fl Piedade Forest 305 78ha F2 Macacos Forest 356 22ha F3 Zambana Forest 387 85ha F4 Vespas Forest 13 80ha and FS Pezinho Forest 29 89ha The S o Jos plant covers approximately 240km the landscape is dominated by sugarcane monocul ture The remnant forest vegetation 24 of the total area is inserted within the Atlantic Forest domain called Dense Ombrophilous Lowland Forest IBGE 1992 and covers 65 7km of this landscape divided among 106 fragments Trindade et al 2008 Following K ppen s classification climate is of type As tropical wet with an average precipitation of 1 689mm and rain concentrated from April August the average temperature is 25 1 C data from the S o Jos sugarcane processing plant for the period 1998 2006 The fragments are located on the Barreiras geological formation which dates back to the Plio Pleistocene and consists of unconsolidated sandy clay sediments of continental origin The terrain is made up of flattened tableland cut by deep narrow valleys with sides that slope at 30 declivity or higher CPRH 2003 The studied remnants house watercourses bet
20. est J Environ Qual 23 835 844 Gomes J S A C B Lins e Silva M J N Rodal amp H C H Silva 2009 Estrutura do sub bosque lenhoso em ambientes de borda e interior de dois fragmentos de floresta atl ntica em Igarassu Pernambuco Brasil Rodrigu sia 60 295 310 Harms K E J S Powers amp R A Montgomery 2004 Variation in small sapling density understory cover and resource availability in four Neotropical Forests Biotropica 36 40 51 IBGE 1992 Manual T cnico da vegeta o Brasileira IBGE Manuais T cnicos em Geoci ncias Rio de Janeiro Rio de Janeiro Brasil Ickes K S J Dewalt 8 S Appanah 2001 Effects of nati ve pigs Sus scrofa on woody understory vegetation in a Malasian lowland rain forest J Trop Ecol 17 191 206 Jardim F C S amp R T Hosokawa 1986 Estrutura da flores ta equatorial mida da esta o experimental de silvi cultura tropical do INPA Acta Amaz 16 411 508 1038 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 Kobe R K 1999 Light gradient partioning among tropical tree species through differential seedling mortality and growth Ecology 80 187 201 Laurance W F H E M Nascimento S G Laurance A Andrade R M Ewers K E Harms R C C Luiz o amp J E Ribeiro 2007 Habitat Fragmentation Varia ble Edge Effects and the Landscape Divergence Hypothesis PLoS ONE 2 1 8 Lima M L F C 1998 A reserva da
21. fferently from abundant species these are more likely to be substituted by others as the forest develops either for natural reasons or due to disturbances in the area Campos amp Landgraf 2001 Regenerative capacity and vertical structure of the understory Regarding spe cies vertical distribution most were between 1 2m high similar to what was found by Silva et al 2007 and Marangon et al 2008 The species that were found in all height classes despite their lower occurrence percentages must have their growth accompanied and their successional characteristics observed because these are the species with the greatest potential to establish themselves and assure their pres ence in the forest s structure and composition Clark amp Clark 1992 Kobe 1999 Silva et al 2007 The exceptions are the species that due to particular characteristics are resident and never surpass the lower stratum Finol 1971 Species that scored high in relative density and were well distributed vertically also had the highest regeneration rates as other authors had already recorded Volpato 1994 Silva et al 2007 Marangon et al 2008 with few excep tions Some species remained among those with high TNR despite low relative density because of their homogeneous vertical distribu tion and higher relative frequency in relation to the others Nevertheless some species did not have a high regeneration rate despite their high relative de
22. ies and 56 95 of individuals In addition to the families cited above despite its lower richness the family Lecythi daceae 3 species 447 individuals was high in density Anaxagorea dolichocarpa Sprague amp Sandwith Annonaceae Eschweilera ovata Cambess Miers Lecythidaceae and Sym phonia globulifera L f Clusiaceae stood out due to the number of individuals 24 10 of the total and for being among the ten species with the highest natural regeneration indexes TNR in three of the five remnants studied Species classification into stratification categories The percentage of transitory spe cies was greater than that of typical under story species Fig 1A An average of 67 01 SD 3 76 transitory species were recorded while lower and upper understory species added up to 12 42 4 33 and 18 4 79 of the total species sampled respectively Between frag ments differences in species distribution into stratification categories were only detected for F3 which was different from fragments F2 G 14 63 p lt 0 05 F4 G 10 37 p lt 0 05 and FS G 9 38 p lt 0 05 because it had the small est percentage of lower understory species 6 25 and the greatest percentage of upper understory species 25 Transitory species averaged 60 73 9 26 of the understory sampled individuals Fig 1B Conversely the lower and upper under story species represented 11 52 11 33 and 27 06 14 50 of the individuals
23. justed the classification applied by Oliveira Filho et al 1994 and Marangon et al 2008 Marangon et al 2008 used the minimum Im height limit and justified that individuals with such a height have better morphological characteristics and thus allow for a more reliable identification In this study we also included plants that were less than 1m tall as Oliveira Filho et al 1994 have stated that this makes it possible to sample more individuals Individuals were grouped into the following height classes class 1 height lt 1m class 2 height gt 1m and lt 2m class 3 height gt 2m and 3m and class 4 height 23m with CBH less than 15cm Species importance in the understory was analyzed based on the Natural Regeneration 1028 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 Index by height class and Total Natural Regen eration suggested by Finol 1971 and modified by Volpato 1994 The natural regeneration index includes parameters related to horizontal and vertical structure density frequency and distribution of individuals by height class Over the last years the natural regeneration index has been used with the purpose of car rying out more complete diagnoses on the dynamics and development of forest succes sion For example Armesto amp Mart nez 1978 used the index by Finol for describing Mediter ranean vegetation in Chile whereas Silva et al 2007 Marangon et al
24. l dynamics of a near natural temperate deciduous forest in Denmark Forest Ecol Manage 106 83 95 Felfili J M J F Ribeiro C W Fagg amp J W B Machado 2000 Recupera o de Matas de Galeria EMPRAPA Cerrados S rie T cnica 21 1 45 Finol H U 1971 Nuevos parametros a considerarse en el analisis structural de las selvas virgenes tropicales Rev Forest Venez 14 29 42 Galeano G S Su rez amp H Balslev 1998 Vascular plant species count in a wet forest in the Choc area on the Pacific Coast of Colombia Biodivers Conserv 7 1563 1575 Gama J R V S A Botelho M M Bentes Gama amp J R S Scolforo 2003 Estrutura e potencial futuro de utili za o da regenera o natural de floresta de v rzea alta no munic pio de Afu Estado do Par Ci FI 13 71 82 Gentry A H amp C Dodson 1987 Contribuition of nontrees to species richness of a tropical rain forest Biotropica 19 149 156 George L O amp F A Bazzaz 1999 The fern understory as an ecological filter emergence and establishment of canopy tree seedlings Ecology 80 833 845 Gilliam F S N L Turril amp M B Adams 1995 Herba ceous layer and overstory species in clear cut and mature central Appalachian hardwood forests Ecol Appl 5 947 955 Gilliam F S N L Turrill S D Aulick D K Evans amp M B Adams 1994 Herbaceous layer and soil response to experimental acidification in a central Appalachian hard wood for
25. nsity which is due to their low rela tive frequency and absence from at least one height class In fact since the TNR index 1s based upon three species descriptors it showed to be adequate for ranking understory spe cies according to their importance in the plant assemblage Furthermore the species ranking based on the index maintained the same bal ance between transitory and understory typical species thus allowing an appropriate inter pretation of the data for understanding the contribution of each group to the richness of this stratum The results of this study reinforced the idea that the understory of forest fragments 1036 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 is made up mainly of transitory species that are well distributed among the height classes and have higher densities which favors their future presence in the structure and composi tion of the canopy Typical understory spe cies however occur in two strata the lower understory which includes the species that don t generally surpass four meters such as those from families Piperaceae Rubiaceae and Melastomataceae and the upper understory intermediate between the low understory and the canopy and made up of species from fami lies Anonnaceae Clusiaceae and Myrtaceae which do not surpass an average of ten meters in height Our findings support the importance of studying the understory s vegetation becau
26. of tree species diversity in riparian forest fragments of different widths SE Brazil Plant Ecol 133 135 152 Mittermeier R A P R Gil M Hoffman J Pilgrim T Brooks C G Mittermeier J Lamoreux amp G A B Fonseca 2005 Hotspots Revisited Earth s Bio logically Richest and Most Endangered Terrestrial Ecoregions Chicago Chicago Illinois USA Naiman R J H D camps amp M E McClain 2005 Riparia ecology conservation and management of stream side communities Elsevier San Diego California USA Nappo M E J J Griffith S V Martins P Marco J nior AL Souza amp A T Oliveira Filho 2004 Din mica da estrutura fitossociol gica da regenerag o natural de esp cies arb reas e arbustivas no sub bosque de povoamento de Mimosa scabrella Bentham em rea minerada em Pogos de Caldas MG Rev rvore 28 811 829 Nascimento L M 2010 Sucess o secund ria em rea de Mata Atl ntica de Pernambuco mudan as flor sticas e estruturais Ph D Thesis Universidade Federal Rural de Pernambuco Pernambuco Brasil Oliveira A N amp LL Amaral 2005 Aspectos flor sticos fitossociol gicos e ecol gicos de um sub bosque de terra firme na Amaz nia Central Amazonas Brasil Acta Amaz 35 1 16 Oliveira E C L amp J M Felfili 2005 Estrutura e din mica da regenerag o natural de uma mata de galeria no Distrito Federal Brasil Acta Bot Bras 19 801 811 Oliveira Filho A T R J Almeida
27. on remains in the extreme North of the Atlantic Forest s distribution area Lima 1998 in the form of small irregular fragments most less than 500ha in size Trindade et al 2008 Such forests are located mainly at the bottom of deep valleys areas where small springs and watercourses can be found and have closed canopies They are thus riparian forests Naiman et al 2005 made up of assemblages that border watercourses in areas where the interfluvial vegetation is also forest Metzger et al 1997 In addition to the typically ripar lan species species from the upper forests can also be found in these habitats since the riparian vegetation serves as an important source of diaspores for forest remnants natural regeneration and recolonization processes Tri quet et al 1990 With the purpose of describing the phys iognomic and functional structure of the understory in Atlantic Forest fragments five differently sized remnants were selected from a highly fragmented landscape The study had five goals 1 to uncover species richness and composition 2 to discover how many and which canopy species are found in the understory 3 to functionally classify species as resident or transitory 4 to investigate the existence of a pattern in the understory s struc ture among the different fragments and 5 to discover the present and potential regenerative capacity of the remnants studied MATERIAL AND METHODS Study area Fiv
28. ons of Native Trees in Lowland Brazilian Atlantic Forest Community Structure Diversity and Dispersal Syn dromes Restor Ecol 19 379 389 Santos R V Citadini Zanette L S Leal Filho 8 W T Hennies 2008 Spontaneous Vegetation on Overbur den Piles in the Coal Basin of Santa Catarina Brazil Restor Ecol 16 444 452 Santos S L amp LF M V lio 2002 Litter accumulation and its effects on seedling recruitment in a Southeast Brazilian Tropical Forest Rev Bras Bot 25 89 92 Scariot A 2000 Seedling mortality by litterfall in Amazo nian forest fragments Biotropica 32 662 669 Schnitzer S A amp W P Carson 2000 Have we forgotten the forest because of the trees Trends Ecol Evol 15 375 376 Schorn L A amp F Galv o 2006 Din mica da regenera o natural em tr s est gios sucessionais de uma floresta ombr fila densa em Blumenau SC Floresta 36 59 74 Silva H C H A C B Lins e Silva J S Gomes amp M J N Rodal 2008 The effects of internal and external edges on vegetation physiognomy and structure in a remnant of atlantic lowland rainforest in Brazil Biorem Biodiv Bioavail 2 47 55 Silva W C L C Marangon L C Ferreira A L P Feliciano amp R F Costa J nior 2007 Estudo da regenera o natural de esp cies arb reas em fragmento de floresta ombr fila densa Mata das Galinhas no munic pio de Catende Zona da mata sul de Pernambuco Ci FI 17 321 331 Souza A C R
29. pato M M L 1994 Regenerac o natural em uma flo resta secund ria no dom nio de Mata Atl ntica uma an lise fitossociol gica Master Thesis Universidade Federal de Vigosa Minas Gerais Brasil 1040 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012
30. r est canopy such as trees and vines Gilliam et al 1994 1995 The transitory group is generally made up of tree individuals with dendrometric values lower than those defined for the tree stratum normally diameter at breast height DBH gt 5cm Finol 1971 Since these two groups concurrently occur Gilliam et al 1995 the understory s composition may include much more species than the canopy Galeano et al 1998 L et al 2010 Future canopy composition in turn is dependent on the density of transitory species in the under story as well as on the recruitment of these species from the lower to the highest height classes Clark amp Clark 1992 Volpato 1994 Kobe 1999 Species that occur in all of the height classes have greatest survival chances as natural mortality is greater in the lower height classes Volpato 1994 Felfili et al 2000 caused by physical damage leaf litter vertebrates Scariot 2000 Ickes et al 2001 Santos amp V lio 2002 or by biological damage such as predation and parasitism Cadenasso amp Picket 2000 The physiognomic and functional analy ses when added to the analysis of the under story s vertical structure help understanding the forest s future development and composi tion Gama et al 2003 and allow us to better estimate the richness and state of conservation Richards 1996 diagnose the dynamics of natural forest fragments Finol 1971 measure responses to soil and climatic
31. re found in three classes yet had high relative frequencies in relation to the others Tapirira guianensis Aubl F1 E ovata F3 Henriettea succosa Aubl DC M racemosa Erythroxylum citrifolium A St Hil Coccoloba sp16 F4 Calyptranthes brasilien sis Spreng and Piper caldense C DC F5 although not being among the species with the highest regeneration represent the regen eration potential of the fragment in which they occur as they stand out in number of individu als These species had low relative frequency and with the exception of T guianensis and M racemosa were absent from one of the height classes The percentages of species with densities of 15 or more individuals had the follow ing distribution among functional stratification 1031 TABLE 2 Estimate of the Total Natural Regeneration TNR of the 34 most important species samples in riparian environments of Atlantic Forest remnants in Pernambuco Brazil Family Scientific name Fl E2 F3 F4 FS SC Anacardiaceae Tapirira guianensis Aubl 2 30 1 17 0 60 2 55 4 65 TR Annonaceae Anaxagorea dolichocarpa Sprague amp Sandwith 12 17 13 47 4 95 0 22 UU Annonaceae 7 11 02 TR Guatteria spl 0 19 14 18 TR Xylopia frutescens Aubl 0 38 0 23 0 30 0 62 6 00 TR Araliaceae Schefflera morototoni Aubl Maguire Steyerm amp Frodin 0 75 0 47 3 47 TR Burseraceae Protium heptaphyllum Aubl Marchand 6 89 1 49 0 99 2 90 0 22 TR Chrysob
32. re part of the inter mediate stratum and do not reach the canopy The subdivision of typical species into the low understory and upper understory catego ries was based on the works of Vilela et al 1995 Oliveira amp Amaral 2005 Gomes et al 2009 Souza et al 2009 and Onofre et al 2010 Upper limit was fixed at 10m based on the average height of the maximum heights of each species recorded in 370 plots studied by the Fragments Project undergoing at S o Jos Sugarcane Property since 2003 The number and percentage of species in each functional stratification category SC was determined for each fragment As density varied amongst samples richness of canopy and understory strata where estimated through rarefaction 100 permutations using the software EstimateS 8 2 Cowel 2009 for the minimal density value recorded in each stratum 160 trees 531 under story plants Observed and estimated values were then compared using T test Homogene ity of species and individuals distribution was compared among the SCs using the G Test and the floristic similarity between the under story and the tree stratum was calculated using Sorensen s similarity index with the purpose of defining which species recorded in the understory have regeneration potential for the overlying tree stratum For test descriptions see Magurran 2004 Studies on natural regeneration usually stratify vegetation into height classes In this study we ad
33. s 2 a ss E L ESSEN e DNS 5 Mil o F4 L sss F5 E O xe GM we GM mi Fig 3 Percentage distribution of species among stratification categories SC A and exclusive of each SC B Class 1 white Class 2 stripes Class 3 grey and Class 4 black In C percentage of species that occurred in one white two stripes three grey or all classes black presented in the understory in riparian environments of five Atlantic Forest remnants Pernambuco Brazil 1034 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 40 E 35 E LU 304 m UU 2 25 pea 9 204 2 15 4 5 2 104 57 0 Class 1 Class 2 Class 3 Class 4 Height classes Vertical bars standard deviation Fig 4 Average number of transitory species TR and species typical of the lower LU and upper UU understory distributed by height classes among riparian environments of Atlantic Forest remnants in Pernambuco Brazil 3B The distribution of transitory and typical understory species exclusively in one height class revealed that these species do not gener ally occur in classes 3 2 3m height and 4 height gt 3m and CBH lt 15cm Few species occurred in all of the height classes percentages of such species varied from 16 67 in F2 to 28 13 in F3 Fig 3C Most species only occurred in one class with percentages varying from 25 in F3 to
34. s cf giganteus Engl 4 69 4 80 TR Sapindaceae Cupania oblongifolia Mart 2 88 3 44 TR Siparunaceae Siparuna guianensis Aubl 4 59 8 62 LU Violaceae Paypayrola blanchetiana Tul 1 14 13 73 0 16 UU SC stratification category TR transitory species LU typical of the lower understory UU resident species of the upper understory Species with the maximum TNR per fragment 1032 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 Cupania oblongifolia Miconia minutiflora Eugenia candolleana Tapirira guianesis Cupania oblongifolia Miconia minutiflora Eugenia candolleana Erythroxylum citrifolium o yt yi Erythroxylum citrifolium EF Siparuna guianensis n a o Siparuna guianensis o Hirtella racemosa Hi Y i irtella racemosa Protium heptaphyllum Protium heptaphyllum Tovomita mangle Tovomita mangle Eschweilera ovata Eschweilera ovata Anaxagorea dolichocarpa Anaxagorea dolichocarpa F1 0 5 10 15 20 25 Myrcia racemosa Miconia prasina Inga thibaudiana o Trichilia lepidota Em Trichilia lepidota Ro Leandra rufescens mmm Leandra rufescens U E J Pilocarpus cf giganteus Pilocarpus cf giganteus u Henriettea succosa EE Henriettea succosa Eschweilera ovata eg Eschweilera ovata Symphonia globulifera Symphonia globulifera Psychotria carthagenensis EI Psychotria carthagenensis F2 0 5 10 15 20 25 Pilocarpus cf giganteus Esch
35. se it is floristically rich structurally unique and promotes the natural regeneration of both the lower and tree strata and the recovery from natural or anthropic disturbances The funda mental aspects of understory assemblages must be prioritized in order to correctly understand the role of transitory and resident species in this stratum ACKNOWLEDGMENTS This study is part of the Fragments Project Phase II Sustainability of Atlantic Forest Remnants in Pernambuco and their Implications for Local Conservation and Development a Brazil Germany scientific collaboration pro gram Science and Technology for the Atlantic Forest financed by CNPq 590039 2006 7 and BMBF 01 LB 0203 A1 with the support of the Sao Jos S A Sugarcane Mill Caval canti Petrib Group The authors thank the support received from the teams of the UFRPE Plant Ecology and Phytosociology Laboratory and the UFPE Plant Morphotaxonomy Labora tory the Ceara Foundation for Scientific and Technological Development FUNCAP and the Brazilian National Council for Scientific Development CNPq for the research scholar ship provided to the first author We also thank Lenilson Santos and Marcos Chagas for their indispensable aid in the field RESUMEN El sotobosque forestal est compuesto por especies residentes y transitorias Con el objetivo de describir la contribuci n de esos grupos en el sotobosque lefioso cinco fragmentos de Bosque Atlantico f
36. story Woody Plants Show Diverse Responses to Forest Pasture Edges in Costa Rica Biotropica 43 562 571 Cadenasso M L amp S T A Pickett 2000 Linking forest edge structure to edge function mediation of herbi vore damage J Ecol 88 31 44 Campos J C 8 P R C Landgraf 2001 An lise da regene ra o natural de esp cies florestais em matas ciliares de acordo com a dist ncia da margem do lago Ci FI 11 143 151 Cardoso Leite E T B Covre R G Ometto D C Caval canti amp M I Pagani 2004 Fitossociologia e caracte riza o sucessional de um fragmento de mata ciliar em Rio Claro SP como subs dio recupera o da rea Rev Inst Flor 16 31 41 Ceccon E P Huante amp E Rinc n 2006 Abiotic factors influencing tropical dry forests regeneration Braz Arch Biol Techn 49 305 312 Clark D A amp D B Clark 1992 Life history diversity of canopy and emergent trees in a neotropical rain forest Ecol Monogr 62 315 344 Cowel R K 2009 EstimateS 8 2 Statistical estimator of species richness and shared species from samples Department of Ecology amp Evolutionary Biology University of Connecticut Storrs USA Down loaded 20 October 2011 http viceroy eeb uconn edu estimates CPRH 2003 Diagn stico s cio ambiental do Litoral Norte de Pernambuco CPRH Recife Pernambuco Brasil Emborg J 1998 Understorey light conditions and rege neration with respect to the structura
37. tudy for the understory associated with an intermediate similarity between the understory and the canopy may also indicate the absence of large scale disturbances in the past Bohl man amp O Brien 2006 since even higher simi larities are expected in early secondary areas Nascimento 2010 With the exception of Family Clusiaceae the high richness and abundance of individu als recorded for families such as Rubiaceae Melastomataceae Annonaceae and Myrtaceae seems to be a standard for flora understory in more humid forests Schorn amp Galv o 2006 Gomes et al 2009 Onofre et al 2010 L et al 2010 as well as in seasonal forests and savan nah Vilela et al 1995 Cardoso Leite et al 2004 Oliveira amp Felfili 2005 Although these 1035 families richness in the understory is common ly used as an indicator of succession stage our results show that these families co occur and are highly important in distinct strata Thus in studies with less inclusive size criteria the fam ily Myrtaceae which has a higher richness of shade tolerant species Tabarelli amp Mantovani 1999 is indicative of more mature succes sional stages When individuals of the smaller height classes are sampled Melastomataceae and Rubiaceae are cited as representative of pioneer species Tabarelli amp Mantovani 1999 i e of forests in an initial successional stage Thus in a more detailed analysis of forest stra ta the richness or
38. ueron seleccionados en el nordeste de Brasil El muestreo incluy individuos con circunferencia a la altura del pecho CPA lt I5cm y con circunferencia a la altura del suelo CAS gt 3cm Las especies fueron cuantificadas y clasificadas en residentes o transitorias y la similitud flor stica entre el sotobosque y el dosel fue calculada El sotobosque se mostr m s rico en especies mediana 63 8 DS 21 72 n 5 fragmentos que el dosel 43 8 18 14 5 y la similitud entre esos com ponentes fue relativamente alta 0 54 0 09 El sotobosque est compuesto principalmente de especies transitorias mediana 67 01 DS 3 76 las cuales est n bien distri buidas entre las clases de altura y presentan las mayores densidades Las especies residentes fueron agrupadas en dos estratos el sotobosque bajo lt 4m de altura la mayor a Piperaceae Rubiaceae y Melastomataceae y el sotobos que superior altura media lt 10m principalmente Anonna ceae Clusiaceae y Myrtaceae El estudio del sotobosque es fundamental porque es un estrato flor sticamente rico con un estructura nica lo cual promueve la regeneraci n natural del estrato arb reo Palabras clave clases de altura composici n flor stica densidad regeneraci n natural riqueza de especies Bos que Atl ntico grupo funcional REFERENCES Alves L F amp J P Metzger 2006 A regenera o florestal em reas de floresta secund ria na Reserva Florestal do Morro Grande Cotia
39. variations and to environmental stress Harms et al 2004 plan management actions forestry practices and vegetation restoration activities Emborg 1998 Mattei amp Longhi 2001 Gama et al 2003 Silva et al 2007 Sansevero et al 2011 In a mature tropical forest the floristic composition of the canopy and understory is expected to be distinct from each other Jardim amp Hosokawa 1986 due to the low height of herbs and shrubs which contributes to an increase of richness in the understory and also to the dynamics of tree species in different successional groups As shown by Nascimento 2010 in a 12 year secondary area regener ated after clearcutting the similarity between woody species in the canopy and in the under story was of 76 and diminished to 48 with successional maturity in 20 year secondary forests For a more mature stage Alves amp Metzger 2006 recorded 30 similarity in a secondary 80 year forest Such low similarity in intermediate regeneration stages is due to the concentration of secondary species in the understory and pioneer species in the canopy since the pioneer species grow faster than the secondary species until they reach maturity V lio 2003 Bohlman amp O Brien 2006 Simi larity may start increasing again during a more advanced successional stage as the secondary species that once occupied the regeneration stratum become part of the tree stratum This pattern was found by Silva et al
40. ween one and three meters wide that are part of the Botafogo River water basin which allows them to be classified as riparian forest sensu Naiman et al 2005 Composition and structure data sam pling In order to analyze the understory 20 non contiguous 5x5m parcels were installed in each fragment at the apex of larger 10x10 plots that had already been used for a study on trees available from Lins e Silva 2010 as part of the databank of the Fragments Project The par cels were arranged to guarantee that both forest edge and interior environments be sampled as well as both sides of the watercourse at a minimum distance of 5m between the units To sample to tree stratum we included all of the trees with CBH circumference at breast Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 3 1025 1040 September 2012 1027 height greater or equal to 15cm In the under story we included the woody individuals with CBH lesser than 15cm and circumference at ground level CGL greater than 3cm regard less of height All of the individuals included in the tree stratum or understory had their CBH or CGL taken with a measuring tape and their height estimated with the aid of an 8m graduat ed ruler Additionally a fertile or sterile sample of each species branches was collected Field work was undertaken from 2007 2009 The data CGL CBH height botanical family and species was organized into a database using the Mata
41. weilera ovata Guapira opposita Guapira opposita Symphonia globulifera Symphonia globulifera o Pogonophora schomburgkiana Pogonophora schomburgkiana y Henriettea succosa Henriettea succosa a Calyptranthes brasiliensis Calyptranthes brasiliensis Annonaceae 7 Annonaceae 7 Anaxagorea dolichocarpa Anaxagorea dolichocarpa Paypayrola blanchetiana Paypayrola blanchetiana Guatteria sp 1 Guatteria sp 1 F3 0 5 10 15 20 25 Coccoloba sp 16 Erythroxylum citrifolium Protium heptaphyllum Myrcia racemosa Himatanthus phagedaenicus Henriettea A Miconi gt Protium heptaphyllum Iconia prasina Himatanthus phagedaenicus E Phyllanthus juglandifolius Miconia prasina o Pogonophora schomburgkiana Phyllanthus juglandifolius ro Inga thibaudiana Pogonophora schomburgkiana Inga thibaudiana Anaxagorea dolichocarpa 5 NE Anaxagorea dolichocarpa Albizia pedicellaris Albizia pedicellaris Symphonia globulifera Symphonia globulifera Eschweilera ovata Eschweilera ovata F4 0 5 10 15 20 25 Gonzalagunia dicocca Piper caldense Cupania oblongifolia Calyptranthes brasiliensis Scheffera morototoni Gonzalagunia dicocca n Pavonia malacophylla Cupania oblongifolia o Albizi Scheffera morototoni 9 d OH Pavonia malacophylla 8 Tapirira guianensis Albizia saman nm Xylopia frutescens Tapirira guianensis Siparuna guianensis Xylopia frutescens Pi A Siparuna guianensis per marginatum d M i Piper marginatum Miconia prasina f Miconia prasina F5 0 5 10 15
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