( PDF ) Decomposição foliar de espécies pioneiras e macrofauna de solo em ecossistemas degradados pela deposição de cinzas e extração do carvão

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Universidade Federal do Rio Grande do Sul

Instituto de Biociências

Programa de Pós-Graduação em Ecologia

Dissertação de Mestrado

DECOMPOSIÇÃO FOLIAR DE ESPÉCIES PIONEIRAS E

MACROFAUNA DE SOLO EM ECOSSISTEMAS DEGRADADOS PELA

DEPOSIÇÃO DE CINZAS E EXTRAÇÃO DO CARVÃO

Luciana Regina Podgaiski

Porto Alegre, abril de 2009

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DECOMPOSIÇÃO FOLIAR DE ESPÉCIES PIONEIRAS E

MACROFAUNA DE SOLO EM ECOSSISTEMAS DEGRADADOS PELA

DEPOSIÇÃO DE CINZAS E EXTRAÇÃO DO CARVÃO

Luciana Regina Podgaiski

Dissertação apresentada ao Programa de Pós-

Graduação em Ecologia, do Instituto de

Biociências, da Universidade Federal do Rio

Grande do Sul, como parte dos requisitos para

obtenção do título de Mestre em Ecologia.

Orientador: Dr. Gilberto Gonçalves Rodrigues

Comissão Avaliadora

Prof. Dr. Ademir Reis (UFSC)

Prof. Dr. Maria Luiza Porto (UFRGS)

Prof. Dr. Milton Mendonça Jr. (UFRGS)

Porto Alegre, abril de 2009

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À Odette de Mello Colombo, minha

super avó, por todo amor e carinho

dedicados a mim durante ¼ de século

iii

“Just to hear,

just to hear again:

Don’t matter how you go ....

Keep going!”

“Keep pray and trust the way is opening

Keep going!

Just keep going like me”

OS THE DARMA LOVERS

AGRADECIMENTOS

Ao meu noivo, André Luis Casara, pelo incentivo de continuar lutando,

trabalhando e crescendo profissionalmente, sempre com pensamentos

positivos e de sucesso. “Sucesso! Por que boa sorte é para quem não

está preparado”. Por toda segurança, apoio e dedicação, e é claro, por

todo amor que me transmites a cada dia. Te amo!

A minha mãe, Paula Cristina Colombo, por ser minha fortaleza. A minha

avó Odette de Mello Colombo, minha segunda-mãe, pelas orações e

dedicação.

Ao meu pai, Jorge Podgaiski, e minha terceira-mãe, Alice Kuzniar, por

todo carinho manifestado por mim, e pelas energias positivas que me

passam todas as vezes que nos encontramos.

Aos meus sogros, Sérgio e Berenice Casara (quarta-mãe), por escutarem

meus desabafos, pelas orações e força nos momentos difíceis e de

decisão.

A minha grande amiga Verônica Sydow, inestimável companhia em todas

as etapas do meu trabalho, sempre presente e de mãos dadas comigo

pro que der e vier. Valeu Vê!

A outra grande amiga, Aline Quadros, pesquisadora exepcional, que

muito me ajudou, ensinou, incentivou, e fortemente me encorajou a

seguir em busca do sucesso.

A professora Paula Araujo, fantástica orientadora, por me aceitar

alegremente de baixo de seus braços (laboratório), pela confiança

depositada em todos os momentos, e pelo reconhecimento.

Ao Alan Panata (Alão, in memorian), um anjo no céu, pela incrível

amizade e companheirismo. Deixaste a grande lição de que a vida é

efêmera, e temos que aproveitá-la intensamente (nem que para isso

não tenhamos tempo nem para dormir...). Tudo é muito rápido, e passa

voando pelos nossos cabelos... Um dia estamos aqui, no outro, podemos

já estar lá ...

A todas as pessoas que me axiliaram no projeto, campo e/ou

laboratório, Alan Panata, Aline Quadros, André Lima, André Castillo,

André Casara, André Frainer, Cíntia Beuvalet, Marisa Azzolini, Tamires

da Silva, Verônica Sydow e motorista do departamento Marcelo Saraiva.

A todos os colegas do PPG-Ecologia, em especial Ana Luiza Matte (Ziza),

André Lima, Marcelo Flores, Márlon Vasconscelos, Vagner Camiloti e

Verônica Sydow pela parceria, conversas especiais e papos furados.

A todos os taxonomistas que auxiliaram na identificação dos

macroinvertebrados terrestres: Aline Quadros (UFRGS), André Barbosa

(UFRJ), Erica Helena Buckup (FZBRS), Everton Rodrigues (FZBRS), Laura

Menzel (UFRGS), Letícia Schmidt (FZBRS), Luciano Moura (FZBRS), Maria

Aparecida Marques (Cida; FZBRS) e Profa. Paula Araujo (UFRGS).

A Marisa Azzolini, pela grande ajuda nas análises químicas e físicas, e é

claro pelas conversas, sugestões e positivo incremento dado ao

trabalho.

A Profa. Maria Luiza Porto, pela disponibilidade de uso do espaço e dos

equipamentos do Laboratório de Fitorremediação.

A Profa. Gislene Ganade e Prof. Adriano Melo por sugestões às análises

estatísticas.

A CGTEE, Companhia de Geração Térmica de Energia Elétrica – São

Jerônimo, em especial ao Januário e outros funcionários, pelo acesso à

área de estudo e auxílio na montagem dos experimentos.

A CRM, Companhia Rio-Grandense de Mineração, Minas do Leão, em

especial ao Eng. Müller, Baner e Pedro Paulo da Silva Batista, pelo

acesso a área de estudo, apoio logístico e incentivo ao trabalho.

Ao Programa de Pós-Graduação em Ecologia da UFRGS, por toda

logistica oferecida e contribuição financeira a análises químicas do

trabalho, e em especial aos coordenadores Prof. Adriano Melo e Profa.

Sandra Müller, por estarem sempre de portas abertas às necessidades

dos alunos, e a secretária Silvana Barzotto por toda assistência.

Ao Centro de Ecologia e Depto. de Ecologia do Instituto de Biociêncas,

pela infra-estrutura cedida ao Prof. Gilberto Rodrigues, para a execução

do projeto.

A equipe de trabalho de São Jerônimo, pela integração aos estudos

(Profa. Maria Luiza Porto, Dr. Mariza Azzolini, Profa. Maria Tereza Raya

Rodriguez e Prof. Gilberto Rodrigues).

Ao Prof. Gilberto Rodrigues pelo financiamento do projeto através de

projetos da CAPES/CNPq.

A C

APES pela concessão da bolsa de mestrado.

vii

RESUMO

Dois estudos envolvendo o processo de decomposição foliar de espécies

pioneiras e os organismos da macrofauna de solo foram realizados em

ecossistemas degradados pela deposição de cinzas e extração do carvão no Rio

Grande do Sul, Brasil. Os estudos resultaram em três artigos científicos. O

primeiro estudo (1° artigo) acessou a decomposição foliar e a colonização da

macrofauna nos folhiços do capim-bermuda (Cynodon dactylon- Poaceae), da

mamona (Ricinus communis- Euphorbiaceae), e da aroeira-vermelha (Schinus

terebinthifolius- Anacardiaceae), plantas participantes da sucessão natural

inicial, em áreas de deposição de cinzas leves e cinzas grossas/escória na

mata ciliar do Rio Jacuí, em São Jerônimo. O segundo artigo – originário deste

primeiro estudo-tratou exclusivamente sobre os padões de colonização de

espécies de tatuzinhos de solo (Isopoda), grupo de animais detritívoros com

alta abundância no local, no folhiço das mesmas três plantas. Ainda,

compararam-se as suas abundâncias e algumas características reprodutivas

entre os dois depósitos de cinzas. O segundo estudo (terceiro artigo) testou

uma técnica de manejo para acelerar o processo da decomposição foliar de

Pinus elliottii (Pinaceae) e enriquecer a fauna de macroartrópodes de solo em

uma floresta monodominate de pinus sobre solo minerado e reconstruído

topograficamente no município de Minas do Leão. Todos os estudos foram

realizados empregando-se a técnica de bolsas-de-folhiço, que consistiu em

bolsa de nylon de 30 X 20 cm e malha de 0.2 X 1.0 cm. A decomposição foi

medida calculando-se a perda de massa foliar seca nas datas amostrais (6, 35,

70 e 140 dias após a exposição das folhas– 1° e 2° artigo ; 3 e 6 meses– 3°

artigo ). Foram realizadas análises de macronutrientes do material foliar

remanescente. A macrofauna (indivíduos > 2 mm) foi retirada manualmente

das bolsas-de-folhiço em laboratório, ou com extrator de Berlese-Tüllgren

modificado (3° artigo), separada em ordens e morfoespeciada. Foi constatado

que a mamona apresentou decomposição foliar mais rápida (k= 20.7) e maior

densidade de indivíduos da macrofauna no seu folhiço, mas, no entanto,

viii

apresentou menor riqueza de espécies do que as outras plantas pioneiras. Os

folhiços do capim-bermuda e da aroeira-vermelha foram similares com relação

à decomposição e a diversidade da macrofauna. As espécies de tatuzinhos

colonizaram igualmente as três espécies de plantas ao longo do tempo. Apesar

da decomposição foliar não ter sido influenciada, a ocorrência de Atlantoscia

floridana (Isopoda), a abundância e fecundidade de fêmeas de Benthana

taeniata (Isopoda), e a composição total de espécies da macrofauna foram

afetadas pelos diferentes depósitos de cinzas de carvão. Como esperado, a

adição de folhas de plantas nativas sobre o solo homogêneo da floresta

monodominante de pinus em solo minerado modificou a composição química

do folhiço de pinus e aumentou a diversidade dos macroartrópodes de solo,

demonstrado ser uma estratégia potencial ao manejo e à restauração

ecológica. Estes estudos têm como metas contribur ao entendimento sobre a

ecologia (processos ecológicos e interação organismos – folhiço) de áreas

degradadas pela deposição de cinzas e áreas de extração e processamento do

carvão no sul do Brasil.

ABSTRACT

Two studies concerning the leaf decomposition of pioneer plant species and

the soil macrofauna organisms were accomplished in ecosystems degraded by

the deposition of ashes and coal extraction in Rio Grande do Sul, Brazil. The

studies result in three cientific articles. The first study (1° article) accessed

the decomposition and the macrofauna colonization among leaves of

bermuda-grasss (Cynodon dactylon - Poaceae), the castor oil plant (Ricinus

communis - Euphorbiaceae), and the Brazilian pepertree (Schinus

terebinthifolius - Anacardiaceae), plants that participate of the initial natural

succession in areas of deposition of fly ashes and boiler slag in Rio Jacuí's

ciliar forest, in São Jerônimo. The second article (also derivated from the 1°

study) treated exclusively about the colonization patterns of woodlice species

(Isopoda), detritivorous group with high abundance in the site, among the

leaf-litter of the same three plants. Still, their abundances and some

reproductive characteristics were compared between the two deposits of

ashes. The second study (3° article) tested a technique of management to

motivate the leaf decomposition of Pinus elliottii (Pinaceae) and to enrich the

soil macroarthropod fauna in a pinus monospecific forest with spontaneous

establishment on a mined and rebuilt soil in Minas do Leão. All of these

studies were accomplished using the litter-bags technique, that consisted in a

nylon’s bag of 30 X 20 cm and mesh of 0.2 X 1.0 cm. The decomposition was

measured calculating the leaf-litter mass loss through sampling dates (6, 35,

70 and 140 days after leaf expositions- 1° and 2° article; or 3 and 6 months -

3° article). Analyses of nutrient contents in remaining leaf-litter were

accomplished. The macrofauna (individuals> 2 mm) was removed manually

from the litter bags in laboratory, or with modified Berlese-Tüllgren extractor

(3° study), and it was separated in orders and morphospecies. We verified

that the castor oil plant presented faster decomposition (k=20.7) and larger

density of macrofauna individuals, but it presented smaller species richness

than the other pioneers plants. The leaf-litter of Bermuda-grass and Brazilian

pepeertree were similar concerning decomposition and macrofauna diversity.

The woodlice species colonized the three species of plants equally along the

time. Despite no effects on decomposition, the occurrence of Atlantoscia

floridana (Isopoda), the abundance and fecundity of Benthana taeniata

females (Isopoda), and the total macrofauna species composition was strongly

affected by the different deposits of coal ashes. As expected, the input of

leaves from native plants on the homogeneous soil of the pinus forest

modified the chemical composition of the pinus leaf-litter and it increased the

soil arthropod diversity, showing to be a potential strategy to management

and ecological restoration. Theses studies have goals to contribute to the

understanding about ecological processes and interaction between organisms-

litter in areas with extraction and processing of coal in southern Brazil.

SUMÁRIO

AGRADECIMENTOS..................................................................….v

ESUMO..........................................................................… viii

BSTRACT.............................................................................x

UMÁRIO.............................................................................xii

Introdução Geral

Introdução Geral...............................................................01

Artigos Científicos……………………………………………………………………….……….07

Métodos...............................................................…………….11

Artigo 1- Leaf-litter decomposition of pioneer plants and soil

macrofauna community on a coal ash disposal system undergoing

natural succession Podgaiski L.R. and Rodrigues G.G.

Summary..........................................................…...…………..15

Key words........................................................………………….15

Introduction..............................................................……….16

Methods..........................................................………………... 19

Study area.................................................................…. 19

Plant species ....................................…………………………………… 20

Physical and chemical characteristics of the ash disposal sites.….… 21

Litter bag experiments.............................……………............….22

Laboratory procedures .............................……………………......….22

Soil macrofauna ........................................……….........……….23

Data analyses .............................................………….....……….23

Results………………………………………………………….……………………………………..25

Leaf-litter decomposition.....…………….............................……...25

xii

Soil macrofauna diversity....................................…………….……26

Soil fauna in the treatments.............................……………………….26

Functional trophic groups in the treatments...…………………….........27

Detritivores........................................................………....27

Carnivores......................................................…...........28

Herbivores.............................................…......………………..28

Omnivores..............................................................…...28

Discussion.............................................................……….....28

Leaf-litter effects....................................……………………………..28

Coal ash disposal effects.........................………………………………….31

Final considerations and conclusions...........................….....…..32

Acknowledgements........................................……………………....33

References............................................................……….….34

Tables................................................................………………42

Figures..........................................................…………………...48

Appendix........................................................……………...…..55

Artigo 2 - Neotropical woodlice (Crustacea; Isopoda) colonizing early

successional plant leaf-litter in a coal ash disposal environment

Podgaiski L.R., Quadros, A.F., Araujo P.B. and Rodrigues G.G.

Abstract....................................………………………………………….……63

Key words.................................……………………………...........…….63

Introduction.......................................…………..........….........64

Material and methods...........................………………...........……...66

Study area....................………………………….........................…..66

Experiment design and laboratory procedures ..........……………….….67

Statistical analysis…………………………………….............……………………..68

Results……………..…………………………………………………………………………….....68

Discussion ................................………………………………..………….……69

Woodlice in coal ash disposal sites ………………………………………………..70

xiii

Woodlice in pioneer leaf-litter along decomposition.......……........71

Acknowledgements................................................…………....72

References...........................................…………………………..…...73

Tables............................………………………………………………………......79

Figures...............................………………..........................……..81

Artigo 3 - Does addition of mix leaves on the soil improve

macroarthropod diversity and litter decomposition in a post-mining

pinus forest? ………………………………………Podgaiski L.R. and Rodrigues G.G.

Summary.........................................................……….....….…85

Key words.........................................………………………............85

Introduction.......................................................……………....86

Methods.....................................................….........…………..88

Study area...............................................…..........………....88

Experimental design...................................................………89

Sample analysis……………………………………......……………………………….....90

Data analysis………….....................................……………….........91

Results………………………………………………………..………………………………….……93

Macroarthropods response to mix leaf-litter input...........……………93

Pinus elliottii leaf-litter response to mix leaf-litter........……..…...95

Discussion………………………………………………………………………………………………95

Macroarthropods response to mix leaf-litter input..………….…..………96

Pinus elliottii leaf-litter response to mix leaf-litter……………..……...97

Ecological application …………………………………………………………….……….99

Acknowledgements..........................…......…………………………….…100

References........………………………………................................…...100

Tables....................…………………..................................…....109

Figures...............................…………………………...........…..........113

Appendix.............................………………………...........…….......…119

xiv

Considerações Finais......................................................……122

Referências..........................................…....…..................128

NEXOS

Normas das revistas científicas escolhidas para publicação.......... 135

Forest Ecology and Management .......……………………………………...135

Studies on Neotropical Fauna and Environment…….................143

Ecology.....................................................................146

Procedimentos de publicação………………............………………….....……149

Atestado de submissão do Artigo 2 - Science of Total

Environment….............................................................149

Carta de rejeição do Artigo 2 - Science of Total

Environment…...........................................................…149

Atestado de submissão do Artigo 2- Studies on Neotropical Fauna

and Environment ………...................................................150

INTRODUÇÃO GERAL

A maioria dos ecossistemas terrestres no mundo apresenta áreas degradadas,

resultantes de atividades antrópicas tais como a agricultura e a mineração. Os

impactos ambientais oriundos da mineração do carvão são bastante complexos

nos ecossistemas, causando significativas alterações na paisagem, destruindo

a biota e gerando uma grande quantidade de resíduos. O consumo de energia

gerada pelo carvão mineral atualmente no Brasil alcança cerca de 6,2 % da

nossa oferta de energia interna (Brasil e EPE 2008). As maiores reservas de

carvão no Brasil estão situadas na região sul, apresentando o Rio Grande do

Sul cerca de 90 % das reservas. Todavia, o estado que contempla a maior

produção é Santa Catarina, uma vez que o minério do Rio Grande do Sul

apresenta elevado teor de impurezas e é pobre do ponto de vista energético

(ANEEL 2008).

A região carbonífera do Baixo Jacuí, depressão central do Rio Grande

do Sul, é constituída pelos municípios: São Jerônimo, Barão do Triunfo,

Charqueadas, Arroio dos Ratos, Butiá, Minas do Leão, General Câmara, Triunfo

e Eldorado do Sul (Fig. 1; Souza e Bittencourt 2000). A mineração do carvão

nesta região iniciou-se pela via subterrânea, mas ao passar dos anos, a lavra a

céu aberto passou a constituir a principal técnica de extração no estado

(Guerra 2000).

REGIÃO CARBONÍFERA DO RIO GRANDE DO SUL

Figura 1: Municípios constituíntes da região carbonífera do Baixo Jacuí, no Rio

Grande do Sul. Limites nos paralelos 29°37’ e 30°38’ de latitude sul e 51°15’

e 51°14’ de longitude oeste.

De acordo com o § 2.º do art. 225 da Constituição Federal de 1988

todas as áreas prejudicadas pela mineração deveriam obrigatoriamente ser

recuperadas no Brasil, mas, poucos estudos têm sido realizados enfocando a

recuperação ambiental destas áreas (Prochnow e Porto 2000). A reconstrução

de solos após a mineração necessariamente deveria seguir cuidados e regras

especiais, entre elas, a reposição das camadas de solo na seqüência original

de sua retirada, preservando seus horizontes. No entanto, os “solos

reconstruídos” são fundamentalmente antropogênicos, apresentando uma

série de desequilíbrios em seus atributos químicos, físicos e biológicos (Kämpf

et al. 2000), apesar da topografia recuperada. Kämpf et al. (2000) identificou

que solos reconstruídos podem apresentar acidificação, alta densidade, baixa

porosidade, baixa condutividade hidráulica e baixa retenção hídrica. Estes

parâmetros sugerem a existência de inúmeros fatores limitantes ao

estabelecimento da vegetação, assim como à restauração da paisagem e da

biodiversidade nestes locais.

O processo de combustão do carvão fóssil em usinas termelétricas

produz uma imensa quantidade de resíduos, oriundos do alto teor de matéria

mineral associada ao carvão (Guerra 2000; Rohde et al. 2006). Três centrais

termelétricas a carvão fóssil estão em atividade no Rio Grande do Sul, sendo

duas delas localizadas na região carbonífera tradicional: São Jerônimo e

Charqueadas. A termelétrica de São Jerônimo (Companhia de Geração

Térmica de Energia Elétrica, CGTEE) está situada às margens do Rio Jacuí, e

constitui a menor unidade geradora de eletricidade em operação no Brasil,

com potência de 20 MW (ANEEL 2008). Apesar da menor potência, esta usina

tem capacidade para gerar cerca de 55.000 toneladas de cinzas por ano (5.000

t – cinzas leves; 50.000 t - cinzas grossas/escória; Rohde et al. 2006).

As cinzas são materiais silico-aluminosos, sendo SiO

e Al

seus

principais componentes (Rohde et al. 2006). Cinzas leves, também

denominadas volantes (fly ash), são formadas por partículas finas (menores do

que 0,15 mm) que são arrastadas pelos gases de combustão nas fornalhas. As

cinzas grossas, ou escória (boiler slag), são retiradas do fundo das fornalhas,

apresentando altos teores de carbono não queimado, em granulometria

grosseira e blocos sintetizados (Rohde et al. 2006). A composição química das

cinzas varia de acordo com a qualidade do carvão parental e da tecnologia de

combustão, mas poluentes como metais pesados e compostos orgânicos são

geralmente associados a elas, em menor ou maior quantidade, dependendo do

tipo de cinza. Apesar do grande potencial do uso dos resíduos carboníferos na

indústria de construção civil (Rohde et al. 2006), grandes quantidades de

cinzas são depositadas nas cavas de extração do minério, ao fechamento das

mesmas, ou indiscriminadamente nos arredores das unidades geradoras

(Teixeira et al. 1999). Os depósitos de cinzas sobre o solo na região

carbonífera do Baixo Jacuí datam desde a década de 30 até os dias atuais

(Guerra 2000), e consistem em grave impacto ambiental contaminando o

ambiente, modificando aspectos estruturais, físicos e químicos do solo,

prejudicando a biota e influenciando, de uma maneira indireta, em todos os

processos do ecossistema (Carlson e Adriano, 1993).

As áreas afetadas por atividades relacionadas ao uso e processamento

do carvão constituem áreas degradadas por definição, uma vez que sua

capacidade de “retornar” ao estado original, através de seus meio naturais, é

drasticamente diminuída (Reis et al. 1999). Contudo, a colonização vegetal

destas áreas por espécies pioneiras agressivas e resistentes é um fato

(Prochnow e Porto 2000, Centro de Ecologia 2002, Shu et al. 2005, Azzolini

2008). As plantas pioneiras que se sujeitam a esta colonização primária,

partilham características ecofisiológicas peculiares, como o alto potencial de

dispersão, taxas de crescimento populacionais rápidas (Gotelli 2007), baixos

requerimentos fisiológicos e de substrato e, muitas vezes, resistência a metais

pesados e outros poluentes (Tordoff et al. 2000, Whiting et al. 2004). Como

conseqüência da colonização primária, as condições abióticas locais podem

ser modificadas ao longo do tempo, o solo pode ser estabilizado, sombreado,

aerado e fertilizado pela produção de matéria orgânica. Este processo pode se

refletir em uma facilitação, ou nucleação (Yarraton e Morrison 1974),

favorecendo a chegada de outras plantas com maiores requerimentos na

comunidade, bem como de animais em busca dos recursos disponíveis, e seus

predadores.

O aporte de matéria orgânica no solo é indispensável para a

manutenção da ciclagem de nutrientes dentro do ecossistema. Esta matéria

orgânica, liberada pelas plantas sob forma de folhas e galhos no solo (folhiço),

é passível de desintegração pela interação de agentes abióticos e biológicos

(Lavelle et al. 1993). Os agentes abióticos envolvidos neste processo são os

fatores climáticos - temperatura, luz, umidade; os bióticos são os organismos

detritívoros (invertebrados) e os decompositores (fungos e bactérias). Os

invertebrados detritívoros são responsáveis pela fragmentação do folhiço e

deposição de grandes quantidades de pelotas fecais no solo, o que estimula

enormemente a ação dos decompositores (Weeb 1977). A qualidade química

do folhiço, medida em termos de relação carbono-nitrogênio (C:N),

concentrações de macronutrientes (especialmente N), lignina e fenóis,

também é essencial para predizer a facilidade da ação dos organismos e da

decomposição (Zhang et al. 2008). O incremento da produtividade primária é

uma conseqüência positiva desta ciclagem.

O folhiço sobre o solo favorece a chegada e o estabelecimento da fauna

de invertebrados do solo. Os invertebrados do solo são muito diversos nos

ecossistemas, representando 23 % da diversidade total dos organismos vivos

descritos até o momento (Lavelle et al. 2006), contudo, eles são

reconhecidamente afetados por atividades antrópicas e pela contaminação

ambiental (Rusek et al. 2000). Os invertebrados participam de complexas

cadeias alimentares, pertencendo aos mais variados níveis tróficos (Coleman

et al. 2004). O folhiço oferece importante hábitat e alimentação a estes

organismos, e suas características podem ser determinantes à diversidade de

espécies que suporta. Como a alta heterogeneidade da superfície do solo é

intrínseca a muitos ecossistemas, seja pela diversidade do folhiço ou por

outras estruturas como rochas, é fundamental que estratégias de restauração

de hábitats levem estes fatores em consideração (Podgaiski et al. 2007). O

incremento na heterogeneidade de hábitats é altamente correlacionado ao

aumento na diversidade de microhábitats e microclimas, alimentos, refúgios e

sítios para atividades oferecidos aos invertebrados de solo, e desta forma vem

ao encontro da conservação destes organismos e ao enriquecimento de

hábitats.

A inserção de fundamentos ecológicos dentro das práticas de

restauração de áreas degradadas e conservação da biodiversidade é um

grande desafio. Antes de tomadas de decisões, se faz necessário um

entendimento mínimo sobre o funcionamento e a dinâmica dos sistemas a

serem contemplados (Engel e Parrota 2003), como, por exemplo, sobre seus

históricos de perturbação, composição de espécies vegetais, processos e

interações ecológicas (Eviner e Hawkes 2008). Informações de referência e

bases teóricas são essenciais ao sucesso das ações. Muitas vezes, a melhor

estratégia pode ser não manejar, e sim deixar com que a colonização natural

e espontânea dirija à sucessão e à restauração. Outras vezes, é necessário

subsidiar o sistema, oferecendo condições para que a biodiversidade e os

processos sejam encarecidos. De uma forma geral, estratégias nucleadoras

(Reis et al. 2003), que favoreçam a reconstrução da diversidade espacial dos

hábitats (Isaacs et al. 2009, Samways 2007) e facilitem a sucessão e as

interações interespecíficas (Reis et al. 1999, Silva 2003, Reis e Kageyama

2003) são importantes para manutenção da biodiversidade e a restauração de

ambientes degradados.

Artigos científicos

Nesta dissertação, foram realizados estudos contemplando assuntos

como o processo ecológico de decomposição de folhiço de plantas pioneiras e

interação da diversidade de organismos do solo em áreas altamente

degradadas pela ação do homem, servindo como subsídios à ecologia da

restauração. Dois estudos foram realizados em áreas de diferentes etapas do

processamento do carvão fóssil. Três artigos foram confeccionados: dois

referentes ao primeiro estudo, e um referente ao segundo. Os primeiros

artigos são entitulados respectivamente como:

 “Decomposição foliar de plantas pioneiras e a comunidade da macrofauna

de solo em um sistema com depósito de cinzas de carvão sofrendo sucessão

natural” e“Tatuzinhos neotropicais (Crustacea; Isopoda) colonizando o folhiço

de plantas pioneiras em um ambiente com depósitos de cinzas de carvão”

e apresentam como problemática a contaminação ambiental provinda em

longo prazo por depósitos irregurales de cinzas de carvão em uma area

adjacente à usina termelétrica (floresta ripária) de São Jerônimo, RS (Fig. 2).

Figura 2: Depósitos irregulares de cinzas de carvão em mata ripária do Rio

Jacuí no município de São Jerônimo, RS. Área de estudo dos artigos 1 e 2.

Mapa modificado de Centro de Ecologia

(2002).

O terceiro artigo da dissertação, entitulado “A adição de folhas mistas

no solo melhora a diversidade de macroartrópodes e a decomposição de

serapilheira em uma floresta de pinus pós-minerada?” é retratado em uma

floresta monodominate de pinus, estabelecida espontaneamente sobre um

solo minerado e reconstruído topograficamente, em Minas do Leão, RS (Fig.

3).

Figura 3: Mineração, reconstrução topografica do solo e estabelecimento

espontâneo de pinus, no município de Minas do Leão, RS. (Área de estudo do

artigo 3).

A abordagem do primeiro artigo refere-se especificamente ao processo

de decomposição foliar de três plantas pioneiras (Fig. 4) e a colonização deste

folhiço pela macrofauna de solo (organismos > 2 mm) no processo de

decomposição. As plantas pioneiras escolhidas para o trabalho foram: uma

gramínea exótica (capim-bermuda - Cynodon dactylon), um arbusto exótico

(mamona - Ricinus communis) e uma árvore nativa (aroeira-vermelha - Schinus

terebinthifolius). Todas são consideradas pioneiras e apresentam

representatividade na área de depósitos de cinzas, desempenhando um

importante papel no processo natural da sucessão ecológical destas áreas

(Azzolini, 2008). Ainda neste artigo, são comparados possíveis efeitos de dois

tipos de depósitos de cinzas (cinzas leves e cinzas grossas/ escória; Fig.2)

sobre a decomposição foliar e a macrofauna de solo.

Figura 4: Plantas pioneiras na área de depósito de cinzas, mata ripária do Rio

Jacuí, São Jerônimo, RS. Área de estudo dos artigo 1 e 2.

O segundo artigo se refere estritamente à colonização do folhiço destas

três plantas pelas espécies do grupo de artrópodes de solo mais abundante na

área degradada: os isópodas, ou mais comumente conhecidos, os tatuzinhos.

Estes organismos têm hábitos detritívoros, e uma imensa contribuição ao

processamento do folhiço (Quadros e Araujo 2008) e à ciclagem de nutrientes

nos ecossistemas. Mudanças em suas características de história de vida, como

crescimento e reprodução (Donker et al. 1993), são fortemente relatadas em

situações de contaminação ambiental por metais pesados. Desta forma, a

influência do tipo de depósito de cinza também é testada sobre a densidade e

características reprodutivas dos isopodas neste trabalho.

O terceiro e último artigo é um trabalho de teste de hipóteses

ecológicas, com especulações visando estratégias à restauração ecológica.

Visto forte impacto ambiental na área de estudo após a mineração, a adição

de folhas de espécies nativas mistas (Fig. 5) é testada como uma estratégia de

enriquecimento da fauna de solo e incentivo à decomposição de folhiço do

Pinus elliottii - a espécie pioneira e monodominante no sistema.

Figura 5: Adição de folhas de espécies nativas ao substrato da floresta

monodominante de pinus como uma estratégia para incrementar a diversidade

da fauna de solo e a ciclagem de nutrientes. (Artigo 3).

Métodos

Os experimentos foram delineados levando-se em consideração a

replicabilidade das bolsas-de-folhiço, princípios de aleatoriedade e a

utilização de blocos, para redução da heterogeneidade ambiental das áreas de

estudo (Gotelli e Ellison, 2004). Para os estudos de decomposição foliar e

colonização da fauna de solo foi utilizada uma técnica largamente conhecida

(Wieder e Lang, 1982): os “litter bags”, ou bolsas-de-folhiço (veja também

Rodrigues, 2006). Esta técnica consiste basicamente na adição de material

foliar seco, com massa conhecida, em uma bolsa que é disposta ao ambiente

para a decomposição. Após um período determinado, as bolsas-de-folhiço são

retiradas do ambiente e levadas ao laboratório onde o material foliar

remanescente é seco em estufa até temperatura constante, e pesado em

balança de precisão. Calcula-se a massa decomposta considerando-se a massa

inicial e a final (remanescente) em um determinado tempo. Uma abordagem

utilizada em dados de decomposição é o ajuste de um modelo matemático

que estima a constante que descreve a perda de massa ao longo do tempo (-

k). O modelo mais freqüentemente utilizado é o de decaimento exponencial

simples, considerado por Olson (1963), o qual se aproxima da biologia da

decomposição correspondente a uma perda de massa mais acentuada nos

primeiros dias (componentes solúveis e compostos fáceis de serem

degradados) e mais demorada em longo prazo (materiais recalcitantes)

(Wieder e Lang, 1982). A equação é assim descrita: X/ X

= e

(-kt)

, em que X

a massa inicial, X é a massa remanscente no tempo t (anos), “e” é a base de

logaritmo natural e –k é o coeficiente de processamento da decomposição.

Dependendo dos objetivos do trabalho, a malha da bolsa-de-folhiço

pode apresentar diferentes tamanhos. Malhas pequenas, de aproximadamente

1 mm

, são as mais utilizadas em ambientes terrestres (Gartner e Cardon,

2004), mas no entanto excluem organismos de solo de maior tamanho a

participarem da decomposição. Malhas largas, apesar de permitirem a

passagem da macrofauna (> 2 mm), por outro lado também facilitam maior

perda de fragmentos foliares. Tendo isto em vista, nesta dissertação, em

todos os experimentos, foram utilizadas bolsas-de-folhiço com tamanho 30 X

20 cm e malha de 10 x 2 mm, (Fig. 6). Esta malha permitiu a colonização da

macrofauna do solo.

Figura 6: Bolsa-de-folhiço utilizada nos experimentos do projeto.

Em campo, as bolsas-de-folhiço foram coletadas do solo e

condicionadas em sacos plásticos. Em laboratório, no caso do experimento de

São Jerônimo, as bolsas foram dispostas em bandeijas brancas, nas quais os

organismos representantes da macrofauna foram triados manualmente. No

caso do experimento de Minas do Leão, as bolsas foram inseridas em extrator

de Berlese-Tüllgren modificado, durante uma semana, e após, passaram por

uma rápida triagem manual. Este extrator consiste em uma estrutura com

forma de funil, que no topo apresenta uma fonte de calor e em baixo um

recipiente coletor. O gradiente de temperatura e umidade faz com que os

invertebrados migrem para baixo, e caiam nos coletores. Os invertebrados

foram acondicionados em potes contendo álcool 80 % e identificados em

morfoespécies e grupos tróficos funcionais. O tratamento dos dados e as

análises estatísticas foram realizadas de maneira independente e peculiar

para cada artigo da dissertação.

ARTIGO 1.

LEAF-LITTER DECOMPOSITION OF PIONEER PLANTS AND SOIL

MACROFAUNA COMMUNITY ON A COAL ASH DISPOSAL SYSTEM

UNDERGOING NATURAL SUCCESSION

Luciana Regina Podgaiski

and Gilberto Gonçalves Rodrigues

a,b

Programa de Pós-Graduação em Ecologia, Instituto de Biociências,

Universidade Federal do Rio Grande do Sul. Av: Bento Gonçalves, 9500, prédio

43422, Porto Alegre, RS, CEP 91501-970.

Departamento de Zoologia, Centro de Ciências Biológicas, Universidade

Federal de Pernambuco. Av: Professor Moraes Rego, S/N, Cidade

Universitária, Recife, PE, CEP 50670-420.

*Artigo a ser submetido para publicação na revista científica Forest Ecology

and Management (ISSN: 0378-1127; Elsevier; Impact factor: 1.579).

Summary

We studied leaf-litter decomposition of spontaneous pioneer plants and the diversity of

associated soil macrofauna community in a riparian forest affected by coal ash disposals

(fly ash and boiler slag) in Brazil. We conducted a litter bag experiment during 140 days

in the damaged area. We found that decomposition rate of Ricinus communis leaf-litter

was more than 80 % faster (k-value 20.7) than the other appraised species. This result

agrees with its low C:N ratio and high N (%), and increased detritivores. Whereas this

leaf-litter had supported the highest densities, it presented the lowest morphospecies

richness. Cynodon dactylon and Schinus terebinthifolius leaf-litter were similar in

decomposition rates and macrofauna diversity. Leaf-litter decomposition and

macrofauna densities were not affected by ash disposals type, on the other hand,

morphospecies composition was distinct in the different sites. Physical structure,

unfavorable pH-value and heavy metal concentration in the ash disposal types may be

selecting more adapted species to the different environmental conditions.

Key words: boiler slag, early successional plants; fly ash; soil invertebrates

Introduction

Anthropogenic habitat modifications are likely to have a major impact on the

composition and biodiversity of the Earth (Tilman, 1994). Alterations of biota have

modified ecosystem goods and services, which are very difficult to revert (Hooper at al.,

2005). In face of the global change phenomena, conservation and restoration of

biodiversity requires an immense increase in our knowledge, such as on environmental

constraints, drivers of biota diversity and ecological processes. Such knowledge is

essential to design ecological theory in order to understand and manage ecosystems,

communities, and species in a suitable way (Callaham et al., 2008).

Terrestrial ecosystems impacted by pollutant industrial activities generally

present altered biodiversity and ecosystem proprieties. Natural areas that received waste

deposits from coal combustion, for example, show leaching of potentially toxic trace

elements as heavy metals, lack of essential nutrients and inappropriate physical structure

of the substratum, which lead to: (1) reductions in plant establishment and growth, (2)

changes in plant elemental composition, and (3) increased cycling of toxic elements

through the food chain (Carlson and Adriano, 1993). Also, natural succession processes

in mine tailings is generally very slow, not changing for many years (Shu et al., 2005).

Environments with heavy metal contamination show decreases in litter decomposition

rates (Coughtrey et al., 1979; Giller at al., 1998; McEnroe and Helmisaari, 2001),

mainly due to inhibited abundance, diversity and feeding performance of soil

detritivores and microbial activity (Strojan, 1978; Van Wensem, 1997; Loureiro et al.,

2006; Filzek et al., 2004; Kools et al., 2008).

Current approaches to restoration of contaminated environments include the use

of technical reclamation, generally relating to use of covering systems and sowing or

planting target species (Tordoff et al., 2000; Dutta and Agrawal, 2003; Casselman et al.,

2005). However, lands are sometimes abandoned after use, allowing spontaneous biota

establishment through primary succession (Hodacová and Prach, 2003). Early

successional plants need to have suitable adaptive strategies (Shu et al., 2005), and over

time they might facilitate the arrival of other species by ameliorating harsh soil

conditions (Schulze, 2005). Although slower, this natural process may lead to a more

natural, richer vegetation cover (Hodacová and Prach, 2003).

As belowground and aboveground compartments of terrestrial ecosystems are

closely linked, the effects of one may be felt in the other (Hooper et al., 2000). Through

plant litter input, resources are provided to soil biota; which in turn is responsible for

most of the decomposition of the system. The decomposition process releases nutrients

for plants, which increases their productivity (Wardle, 1999; Hooper et al., 2000).

Impacts affecting plant community structure and abundance can induce changes in the

soil food webs and decomposition rates, by altering the decaying litter material entering

in soil. As plant species with different suites of traits differ in both quality and quantity

of resources offered, they might support soil animal communities that demand different

requirements of food and shelter (Wardle et al., 2004; Wardle, 2006). The litter

decomposition rates are greatly influenced by their chemical properties, specially

increasing in litters with low C:N ratio and high N, varying among species and across

habitats (Gartner and Cardon, 2004; Zhang et al., 2008). Yet, leaf-litter from herbaceous

plant species generally decomposes more easily than that from woody plant species

(Zhang et al., 2008), which could represent a trade-off between food and shelter for

animals from belowground communities (Hooper et al., 2000). Both soil invertebrates

and soil processes may be drove by forest dynamic and management (Salmon et al.,

2008; Lindsay and Cunningham, 2009).

Animals in soils are numerous, and constitute a diverse group of species,

organized into complex food webs (Coleman et al., 2004). They have fundamental role

in the delivery of ecosystem services and goods by the soil, contributing to water

storage and detoxification, nutrient cycling, soil formation, primary production, flood

and erosion control, and climate regulation (see Lavelle et al., 2006). Within soil fauna,

the macroinvertebrates represent large animals (body size > 2 mm; Swift et al., 1979)

that live in surface litter, in nests or burrows (Lavelle et al., 2006). This group supports

organisms that interact in different ways with their environment, belonging to several

trophic levels in the food web (Coleman et al., 2004), representing from detritivores

(e.g. Isopoda and Diplopoda) to true herbivores (e.g. some Hemiptera and Gastropoda),

predators (e.g., Araneae and Chilopoda) and omnivores (e.g., Opiliones and some

Hymenoptera).

In this study, we assess leaf-litter decomposition of early successional plants and

the associated soil macrofauna community structure in a riparian forest affected by

activities of a coal power plant. This ecosystem has received coal combustion waste

disposals (fly ash and boiler slag) for a long time, and after having disturbances

interrupted, it is undergoing a spontaneous succession process. This early succession

has been driving mainly by exotic plants, which are improving the substrate and

microsite conditions for other native species (Azzolini, 2008). For our experiments, we

chose three abundant plants, representing different life forms: the bermuda-grass

Cynodon dactylon (L.) Persoon (Poaceae; exotic), the shrub castor oil bean Ricinus

communis L. (Euphorbiaceae; exotic) and the Brazilian peppertree Schinus

terebinthifolius Raddi (Anacardiaceae; native). Specifically, we addressed four

questions about this system: (1) Which pioneer leaf-litter decay more quickly, releasing

nutrients to the ash substrate? (2) Does the leaf-litter identity influence the diversity of

colonizing soil macrofauna and its functional trophic groups? (3) Does the ash disposal

type affect leaf-litter decomposition and soil macrofauna?

Methods

Study area

The study area has about 9 ha and belongs to the riparian forest of the Jacuí river, in São

Jerônimo (29°57’55.6”S; 51°44’14.9”W), Central Depression region of Rio Grande do

Sul state, Brazil. The climate is temperate, with hot summer without dry season (Cfa

type, according to the Köppen-Geigen climate classification; Peel et al., 2007). Coal

combustion wastes from São Jerônimos’s thermal power plant were landfilled in the

study area for more than 30 years. Nowadays, the area stopped receiving additional

residues and natural succession is proceeding (Fig. 1A).

Two different kinds of coal wastes were disposed in the area: fly ash and boiler

slag (Fig. 1B, 1C). Fly ash is a fine-grained powder with spherical particles, and boiler

slag is molten grained with angular particles. Trace element affinities are similar for

both wastes, but fly ash is generally enriched by elements with calcium oxide-sulfate

affinities and boiler slag, by elements with iron oxide affinities (Querol et al., 1995).

The elemental concentrations of wastes vary according to parent coal composition and

combustion technology. In São Jerônimos’s thermal power plant, which burn coal from

Rio Grande do Sul, there are a production of fly ash and boiler slag. The most abundant

heavy elements from wastes are Sn, Ni and Mo, and in lower proportion Cr, As, Hg, Al,

Pb, Mn, V, Cd, Ba and Zn (Rohde et al., 2006).

Along decades, the waste disposals were accomplished in several times in a

heterogeneous way on the study area. Old and recent deposits are found one beside the

other. According to the landscape management history, we can classify two distinct

sites: one that received more fly ash and other that had more boiler slag disposals. The

sites are 200 m apart. Here forth, they will be called: fly ash site and boiler slag site.

Vegetation from early successional stages dominates both sites. However, as the

plant establishment was driven by the chronology of the ash deposits, the vegetation

physiognomy is also heterogeneous. Exotic and spontaneous plants such as Bermuda-

grass C. dactylon and castor oil plant R. communis assume prominent role in the

recovery of new disposals in both sites. Mimosa bimucronata (De Candolle) O. Kuntze

(Leguminosae) is found constituting patches, especially in the boiler slag site. Common

wood species in both sites are the Brazilian peppertree S. terebinthifolius, camboatá-

vermelho Cupania vernalis Cambessedes (Sapindaceae) and açoita-cavalo Luehea

divaricata Martius (Tiliaceae). Azzolini (2008) provides a list of plant species from the

area.

Plant species

For our study, we chose abundant three pioneer plants from the study area, representing

species from three different life forms and successional stages: a grass: (C. dactylon), a

shrub (R. communis), and a tree (S. terebinthifolius) (Fig. 1D, 1E). The three species are

broadly known by their spontaneous behavior in disturbed and natural habitats around

the world (GISP, 2005). Cynodon dactylon is a perennial grass, with both rhizomes and

stolons; its leaf blades measures are about 8.0 X 0.4 cm. Ricinus communis is a

perennial shrub, reaching 2–3 m height; with palmate leaves measuring around 15 X 45

cm and 7-11 lobes. Schinus terebinthifolius is a native tree from the study region, with

5-10 m height, has composite leaves with 3 to 10 pairs of leaflets measuring around 5 X

2.5 cm.

Physical and chemical characteristics of the ash disposals sites

We used twenty-four substrate core samples (approx. 10 cm deep and 5 cm diameter)

from the disposal sites (12 per site) to analyze the pH and the water retention capacity.

Sampling was done in eight plots; three samples were nested within each plot. In the

laboratory, the same dry weights of substrate (100 g) were conditioned into pots with

several homogeneous perforations in the bottom. We added water inside the pots until

saturation. Water retention was calculated by dividing the weight of the water retained

by the dry weight of the substrate. As the soil’s ability to retain water is related to

particle size, we expected that the fly ash site would retain more water than the boiler

slag site. However, the sites did not differ in their capacity of water retention (Nested

ANOVA, F

1,6

= 1.87, P= 0.22). The pH-values of both sites were acid, but we found that

the boiler slag disposal site was significantly more acid than the fly ash one (Nested

ANOVA; F

1,6

=32.38, P=0.001; Fig. 2).

Further, two samples of substrate from each site were analyzed for N, P, K, Ca,

Mg, S (SO

-2

) and B content at UFRGS Laboratory of Soil Analysis. Nitrogen was

accessed by distillation, after digestion in H

-H

. Phosphorous and Potassium

were analyzed with the Mehlich I method (Mehlich, 1953). Calcium and Magnesium

exchangeable were extracted in KCl mol L

-1

; Sulphur (SO

) was extracted in CaHPO

500 mg L

-1

of P; and Boron was extracted with hot water. The elemental concentrations

found on the sites are shown in the Table 1. The boiler slag site presented more N (%)

and B (mg/dm

) than the fly ash site, which in turn, had higher K (mg/dm

)

concentrations.

Litter bag experiment

For the experiments of leaf-litter decomposition and colonization by macrofauna we

constructed nylon litter bags measuring 30 X 20 cm, with coarse mesh size (10 x 2

mm). This mesh allows the entry of macrofauna (including small individuals). We filled

the litter bags with 20.3 ± 0.2 g of air-dried fresh leaves of the three pioneer plant

species, collected from the study region. In June 2007, we placed a total of 96 litter bags

in the field: 48 in the fly ash and 48 in the boiler slag site. Four blocks were settled per

site. Each plant species was replicated in four litter bags per block. At each sampling

occasion (6, 35, 70 and 140 days of leaf-litter exposure), one litter bag per plant species

was randomly removed from each block.

Laboratory procedures

In the laboratory, we dried (60°C; 72 h) and weighed the remaining leaf-litter from the

litter bags. The initial (t= 0) leaf contents of carbon, nitrogen, phosphorous and

potassium (% of dry mass) and of litter from 140 days of decomposition were analyzed

for each plant in each site. Carbon from the leaves was accessed by moisture

combustion/ Walkley-Black (Walkley and Black, 1934). Nitrogen was determined using

the Kjeldahl method (Kjeldahl, 1883). Phosphorous and potassium were analyzed using

nitric/ perchloric acid digestion followed by determination on an ICP-OES (Inducted

Coupled Plasma- Optical Spectrometer). All methods were performed at UFRGS

Laboratory of Soil Analysis.

Soil macrofauna

We collected the soil macrofauna by hand from the litter bags, before drying the litter.

We counted and separated the individuals into class, order, and family (whenever

possible). Within each group, we classified the individuals in species or morphospecies.

Immature insect (larvae) and spiders, and very small cockroaches and snails were not

classified in. The individuals were assigned to four functional trophic groups:

carnivores, detritivores, herbivores, or omnivores. Feeding assignments were based on

the literature (e.g., Marinoni et al., 2003) and personal communication with

systematists. Carnivores include spiders, cursorial hunters and haematophagous insects;

detritivores comprise saprophagous invertebrates, fungi feeders and litter grazers;

herbivores consist of sap feeders, and leaf chewers; and omnivores include species with

mix feeding habits.

Data analyses

We estimated the single constant (k), which describes the loss of mass over time, for the

three species, considering all litter bag data used in the study. The constant was

estimated fitting the single exponential decay model (Olson, 1963; Wieder and Lang,

1982):

X/ X

= e

(-kt)

where X

is the initial mass, X is the mass remaining at time t (year), “e” is the base of

natural logarithm, and –k is the processing rate coefficient. This approach was

performed just in terms of ecological knowledge (facilitating comparison with other

data sets; as in Zhang et al., 2008) and to obtain insights into the biology of the

decomposition process of these species (Wieder and Lang, 1982). To compare the

efficiency of decomposition process among the leaf-litter species and sites, we

expressed the decomposition as the percentage of initial leaf-litter mass remaining after

6, 35, 70 and 140 days.

We used a nested analysis of variance (ANOVA) with repeated measures for

testing the effects of site, leaf-litter species and time, in the leaf-litter mass remaining in

litter bags and soil macrofauna density (individuals.g

-1

d.w. litter). The blocks were

nested in the sites, and the time of leaf-litter exposure was used as a repeated measure.

Prior to analysis, data of fauna density were square root transformed to satisfy

parametric assumptions of normality of residuals and homogeneity of variances.

Comparisons of leaf-litter mass remaining of each specie between the sites were

performed with ANOVA, considering the time also as a repeated measure.

As species richness is largely influenced by individual’s abundance in samples

(Gotelli & Colwell, 2001), we used rarefaction curves with confidence intervals for

comparison of morphospecies richness among leaf-litter species and the two sites. To

compare the morphospecies composition between these same treatments, we performed

ordinations using non-metric multidimensional scaling (NMDS; Kruskal, 1964) with

Bray-Curtis distance measure as an index of dissimilarity. We used a matrix with

macrofauna morphospecie densities per plant specie within each block (temporal litter

bags data summed). To reduce the noise of rare species, we removed species that

occurred only once in the matrix improving interpretability of the ordination plot

(McCune and Grace, 2002). Analysis of similarity (ANOSIM; Clarke 1993) on a Bray-

Curtis distance matrix was performed to determine whether potential clustering patterns

identified

visually were statistically different. Both, rarefaction, NMDS and ANOSIM

were performed with PAST 1.9 (Hammer et al. 2001).

Results

Leaf-litter decomposition

Leaf nutrient concentration is shown in Table 2. The nitrogen and phosphorous contents

in R. communis leaves are more than 70 and 50 % higher than the other leaf species,

respectively, but their potassium content was low. The C: N ratio was lower for R.

communis (8.04) than for C. dactylon (26.7 %) and S. terebinthifolius (34.17 %).

Potassium content is highest in C. dactylon, and nitrogen and phosphorous contents are

lowest in S. terebinthifolius leaves.

The decomposition rate of leaf-litter over the time (single constant k) was more

than 80 % faster for R. communis (k=20.7 g g

-1

, R

=0.93), followed for C. dactylon

(k=3.5 g g

-1

, R

=0.38) and S. terebinthifolius (k=3.2 g g

-1

, R

=0.73). More than

90 % of R. communis leaves decayed in 35 days. C. dactylon and S. terebinthifolius

showed less of 70 % of their leaves decayed until the end of the experiment (140 days).

The leaf-litter mass remaining in litter bags was influenced by the interaction of leaf-

litter species and the time of leaf-litter exposure (Table 3; Fig. 3A). Ricinus communis

presented higher mass loss at 35, 70 and 140 days than the other leaf-litter species.

Cynodon dactylon presented more mass loss in 70 days than S. terebinthifolius, but in

the other times their leaf-litter mass remaining not differ.

The leaf-litter mass decaying was not different among sites (Table 3; Fig 3B),

neither when considering the individual leaf-litter species in interaction with time

(Table 4). Fly ash and boiler slag showed similar effects on the decomposition process.

However, after 140 days of leaf-litter exposure, some chemical elemental contents

showed to be loser in boiler slag site than in the fly ash site, such as carbon in C.

dactylon; potassium in R. communis; carbon, nitrogen and potassium in S.

terebinthifolius (Table 5).

Soil macrofauna diversity

We found a total of 2,573 individuals and 126 morphospecies of soil macroinvertebrates

belonging to the classes Arachnida, Chilopoda, Crustacea, Diplopoda Gastropoda,

Hexapoda and Oligochaeta colonizing the litter bags (Appendix 1). Rare species (≤ 2

indivíduos) were about 53 % of the total richness. Isopoda was the most abundant group

in terms of individuals (n= 987; 38 %), followed by Oligochaeta (n= 501; 19 %) and

Hymenoptera (n= 407; 16 %). Coleoptera and Hymenoptera presented the highest

morphospecies richness: 54 and 24 morphospecies, respectively. The invertebrates with

detritivorous habits represented 70.2 % of the total individuals and 28 morphospecies.

Omnivores represented 17.8 % of individuals and 26 morphospecies. Carnivores had 10

% of individuals and 51 morphospecies, and herbivores had 2 % of individuals and 21

morphospecies.

In one litter bag, from the 70 days period, we found a great abundance of one

morphospecie of ant (Formicidae; 258 individuals). As such insects have social

behavior, we considered this data as an outlier.

Soil macrofauna in the treatments

Total density of individuals presented significant effects from the interaction between

leaf-litter specie and time (Table 3, Fig. 3C). Ricinus communis sheltered more

individuals than the other species in 6, 35 and 140 days of decomposition. On the other

hand, rarefaction curves showed that this leaf-litter supported the smaller

morphospecies richness (Fig. 4A). Schinus terebinthifolius seemeed to have more

density of individuals in 70 days than the other species, but it can be the reflex of an

outlier produced by the increased ant abundance. In relation to density in other times

and number of morphospecies, C. dactylon and S. terebinthifolius did not differ.

The disposal sites did not influence the total individual colonization (Table 3),

although a tendence to be increased in the fly ash site (Fig. 3D). The morphospecies

richness did not differ between sites (Fig. 4B). As revealed by the ordination in two

dimensions (NMDS, stress=0.16) the overall community macrofauna composition was

responsive to site treatments, instead of leaf-litter species (Fig. 5). Analysis of similarity

showed weak but significant effects of the sites in the macrofauna (ANOSIM, R= 0.14,

P= 0.014). Mainly due the increased number of rare species, we did not use ordination

anlaysis for the functional throphic groups separately.

Functional trophic groups in the treatments

Detritivores

We found a significant interaction between leaf-litter species and time of leaf-litter

decomposition for detritivores (Table 6). Ricinus communis supported higher detritivore

density at the first days of litter decomposition (Fig. 6A), but it had the lower total

morphospecies richness (Fig. 7A). Cynodon dactylon and S. terebinthifolius seemed not

differ in in density and richness. The sites were similar concerning density (Table 6,

Fig. 6B) and richness of detritivores (Fig. 7B).

Carnivores

We found significant interaction between leaf-litter species and time, and effects of site

in carnivore density (Table 6). Schinus terebinthifolius had lower densities in 6 day, and

R. communis had higher densities at 140 days than the other species (Fig. 6C). The

plants did not differ in number of carnivore morphospecies (Fig. 7C). The fly ash site

had most density of individuals; on the other hand, the boiler slag site had most

morphospecies richness (Fig. 6D, Fig 7D).

Omnivores

We did not found effects of leaf-litter species on the omnivore densities and richness

(Table 6; Fig. 6E, Fig. 7E). The fly ash site had more morphospecies richness

colonizing litter bags than the boiler slag site (Fig. 7F), but no effects on density were

detected (Table 6, Fig. 6F).

Herbivores

The densities and richness of herbivores were similar concerning the treatments (Table

6; Fig. 6G, H, Fig. 7G, H).

Discussion

Leaf-litter

From the three species of pioneer leaf-litter appraised by us, R. communis had the fastest

leaf-litter decay. The k value found for this plant (20.7 g g

-1

) is very high when

compared to other studies. For example, Zhang et al. (2008) reviewed 70 studies around

the world and found k values ranging from 0.006 to 4.993 g g

-1

. Besides R.

communis leaves present high N (%) and low C: N ration, they also supported highest

densities of detritivores in the first days of decomposition, which clearly explain its

mass loss rates (Gartner and Cardon, 2004). Some spontaneous exotic species, as R.

communis, often maintain higher leaf N concentrations, decomposing more rapidly and

releasing more nitrogen to the soil than native species (Levine et al., 2003; Ashton et al.,

2005). The increasing in decomposition rates and nutrient cycling may be negative

within the equilibrium of natural ecosystems (Ehrenfeld, 2003; Wolfe and Klironomos,

2005), but in our reclaiming system, the effects are certainly positive through

amelioration of nutrient availability into the coal ash substrate. Another aspect of R.

communis is that its leaves contain substances, such as ricin, ricinin and flavonoids,

which present insecticidal and antimicrobial actions (Okongkwo and Okoye, 1992; Bigi

et al., 2004). However, the amount of these substances may be reduced in the leaf-litter,

not seemed to directly affect the macrofauna, once the organisms had an increased

density in this leaf-litter and the morphospecies composition had negative response of

leaf-litter identity.

Leaf-litter is food for detritivores, which prefer to feed from palatable plants

with low C:N ratio and high N (e.g. Zimmer 2002). Leaf litter also is microhabitat for

organisms of all functional trophic groups of soil fauna. The decomposition process

may results in a trade-off between these two resources (food and microhabitat). A

greater comsumption of litter decreases the structural integrity that suits it for use as

microhabitat (Hooper et al 2000). The high-quality litter of R. communis, as expected,

was decomposed (consumed) very fast, at the same time that supported a soil

community with low species richness. This case may represent an example of such

trade-off, because the rapid loss of leaf-structure and its associated microhabitat

diversity can be related to a decrease in soil invertebrate diversity (Hooper et al 2000).

Despite markedly differences in the initial leaf nutrients content and leaf structure, C.

dactylon and S. terebinthifolius were similar in decomposition rates and macrofauna

diversity they supported. Cynodon dactylon is more palatable for detritivores and easier

to decompose than S. terebinthifolius. On the other hand, the broad leaves of S.

terebinthifolius offer greater resource heterogeneity (as role of habitat space to fauna)

than the narrow leaves of C. dactylon. Taking into account these characteristics, we

supposed that these species would differ in relation to the appraised ecological aspects.

However, they seem not to be distinguished in our study system.

In many ecosystems of the world, and especially in the system studied by us,

plant establishment, productivity and changes related to succession depend basically on

recycling of nutrients and amelioration of microsite conditions. Soil biota development

also requires adequate organic resources to be sustainable, which is provided by the

plant community by production of surface litter layers. Our results concerning litter

decomposition and associated soil fauna community complement the research of

Azzolini (2008) in some presuppositions about plant natural succession in this coal ash

contaminated area, according the following description. Firstly, C. dactylon have an

important role in rapidly establish a cover in recent ash disposals, stabilizing the

substrate. Plantules of R. communis occur together with C. dactylon, but in a little

number. With the grow-up of R. communis plantules, the cover of C. dactylon is

diminished due the shading, what favor the establishment of other R. communis plants.

In this time, leaves of C. dactylon (with high K) undergo decomposition process,

supplying resource for soil biota and nutrients to substrate. By falling, R. communis also

input their leaves (with high N and P) to the decomposer subsystem, increasing

macrofauna densities (detritivores in the beginning and carnivores in the end) and

quickly transferring nutrients to the soil. As such other shrubs, R. communis offer its

branches for perching, facilitating the arrival of birds dispersing native seeds. Due the

amelioration of soil conditions, S. terebinthifolius and other native plants get to

establish and grown in the substrate (around 4-5 years after ash disposals). Finally, with

the succession proceeding, litter of native trees would be inputted to the soil,

representing heterogeneous shelters for soil fauna.

Coal ash disposals effects

Major constraints to leaf-litter decomposition and soil macrofauna communities on the

two coal ash disposal sites studied could be particularly the high concentration of heavy

metals or other chemical elements, acidity and poor physic structure. In relation to

heavy metals, Rohde et al. (2006) found for this same thermal plant, that boiler slag has

increased concentrations of Sn, Mo, Cr, Al, and Pb than fly ash, which, in turn, had

increased concentrations of Ni, Hg, Cd and Zn. Concerning, other chemical elements,

we found that the site with major influence of boiler slag had more N and B

concentrations than the fly ash. The fly ash site, in its time, had increased K ions

concentrations, which could improve salinity and electrical conductivity. Related to the

pH-value, both disposal sites are acid, probably reducting microflora activities and

availability of major nutrients (such as N, P, K, Mg, B), as well as the increase of

availability of other toxic elements (such as Mn, Zn, Cu). These effects certainly are

higher in the boiler slag site, because it had the lowest pH-value. Substrate conditions

are also different, while the fly ash has fine-grained particles; boiler slag has larger size

particles with amount of unburned Carbon.

In view of all that, we found that the ash disposal type did not affected the

integrity of leaf-litter decomposition of the pioneer plants, in spite some chemical

elements showed to be mineralized faster in boiler slag site than in the fly ash. On the

other hand, the total morphospecies composition was clearly responsive. There are

several works that have been registering changes in survivorship, physiological and

morphological traits of some soil organisms when in contact with polluted food,

substrate, and not favorable pH-value (Jones and Hopkin, 1998; Rusek, 2000; Grumiaux

et al., 2007). There are recognized evidence of heavy metal adaptations in organisms

such Isopods (Donker et al., 1993) and Gastropods (Beeby and Richmond, 1989), which

always accompany altered life histories as part of the complex adaptation syndrome

(Posthuma and Van Straalen, 2002). Soil nutrient stoichiometry also can be explicitly

linked to the invertebrate litter fauna densities, explaining possible bottom-up regulation

of higher trophic levels (McGlynn et al., 2007). Thus, the difference in species

composition between the two disposal sites may be reflecting the selection of more

adapted or resistant species to the specific environmental conditions. As the level of

functional redundancy by a diverse range of taxa is significant into soil communities

(Wardle, 2006), integrity of leaf-litter decomposition process was not altered.

Conclusions

We examined fundamental aspects of ecology of an polluted environment:

decomposition of early successional plants and colonization by soil fauna in two distinct

sites of coal ash disposal. Similar works have not been yet conducted in Brazil relating

ecosystem process and soil fauna in the assessment of areas affected for coal residues.

We conclude that (1) the leaf-litter of R. communis decayed more quickly than the other

pioneer plants; (2) the leaf-litter identity influenced the diversity of colonizing soil

macrofauna, having the leaf-litter of R. communis supported more individuals density

but lower species richness than the other plants; (3) the coal ash disposal type did not

affect the leaf-litter decomposition, but selected distinct communities of soil

macroinvertebrates, showing the existence of singular environmental conditions and

suggesting effects of different constraints in the long term polluted environment.

According to Eviner and Hawkes (2008), efforts of understand feedbacks

between plants and soils have the potential to found a major tool for restoration (e.g.,

colonizing plants ameliorating poor soil conditions) or a major obstacle to restoration

(e.g., invasive species altering soil conditions to benefit themselves). We had evidence,

together with Azzolini (2008), that exotic plants have been ameliorating poor substrate

conditions through the litter decomposition, and benefiting the proceeding of natural

succession in a more long time. Also, the soil fauna development in the damaged area,

especially the detritivore animals, has bring clear benefits to ecological restoration

process, because these organisms greatly affect soil structure and chemistry and

facilitate the ecosystem processes (Snyder and Hendrix, 2008).

Acknowledgements

We thank M.A.L. Marques, E.H. Buckup and E.N.L. Rodrigues (FZBRS, Araneae); L.

Moura (FZBRS, Coleoptera); A.F. Quadros and P.B. Araujo (UFRGS, Isopoda); L.

Schmidt (FZBRS, Hemiptera) and A. Barbosa (UFRJ; Formicidae) for help in

macrofauna’s identifications; A.F.B. Lima, A. Castillo, A.L. Casara, T.B. da Silva and

V.G. Sydow for help in field and lab; G. Ganade for advise in statistical analysis; M.

Azzolini for advise in chemical/physical analysis procedures; Companhia de Geração

Térmica de Energia Elétrica (CGTEE) and its employees for allow access to the study

site and help in the fieldwork. L.R. Podgaiski received a scholarship from CAPES

(Brazil).

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Tables

Table 1. Mean ± SE (in brackets) of elemental concentration of substrate samples from

fly ash and boiler slag disposal sites.

Chemical elements Fly ash disposal Boiler slag disposal

N (%) 0.6 (0.11) 0.9 (0.08)

P (mg/dm

) 32.0 (10.61) 21.5 (4.60)

K (mg/dm

) > 313.0 185.0 (5.66)

Ca (cmol/dm

) 9.6 (1.70) 9.0 (0.64)

Mg (cmol/dm

) 1.4 (0.18) 1.2 (0.07)

S-SO

-2

(mg/dm

) 15.5 (1.77) 15.5 (0.35)

B (mg/dm

) 0.3 (0.03) 0.5 (0.03)

Table 2. Leaf nutrient content of the three pioneer plants.

Plant species C (%) N (%) P (%) K (%)

C. dactylon

40 1.5 0.18 1.9

R. communis

41 5.1 0.38 0.25

S. terebinthifolius

41 1.2 0.15 1

Table 3. Summary of results of nested ANOVA for repeated measures (F values with P

values in brackets) testing for effects of treatments on leaf-litter mass remaining (%)and

total soil macrofauna density (individuals g

-1

d.w. litter- square root transformed). All

statistically significant numbers (p<0.05) are in boldface.

Degrees of freedom are: site

(1, 6), blocks within site (6, 14), leaf-litter species (2, 14), time (3, 42), leaf-litter

species X time (6, 42).

Response variables Site

Blocks

within site

Leaf-litter

species (A)

Time (B) A x B

Leaf-litter mass 0.16 (0.70) 2.25 (0.10)

285.97 (< 0.01) 249.93 (< 0.01) 19.68 (< 0.01)

Total macrofauna 3.63 (0.105) 1.02 (0.45)

3.83 (0.05) 2.87 (0.05) 5.00 (< 0.01)

Table 4. Summary of results of ANOVA for repeated measures (F values with P values

in brackets) testing for effects of treatments on leaf-litter mass remaining (%) of leaf-

litter species. All statistically significant numbers (p<0.05) are in boldface.

Degrees of

freedom are: site (1, 6), blocks within site (6, 14), time (3, 42), leaf-litter species x time

(6, 42).

Leaf-litter species Site (A) Time (B) A x B

C. dactylon

3.46 (0.112)

138.39 (<0.001)

0.36 (0.786)

R. communis

0.04 (0.842)

196.25 (<0.001)

0.42 (0.743)

S. terebinthifolius

0.04 (0.857)

138.39 (<0.001)

0.36 (0.786)

Table 5. Percentage of nutrient loss of leaf-litter species after exposure (140 days) in fly

ash (F) and boiler slag (B) disposal sites. * Not detected.

Disposal sites/ C (%) N (%) P (%) K (%)

Leaf-litter species F B F B F B F B

C. dactylon

66.9 62 61.2 62.9 78.4 79.4 97.3 97.1

R. communis

* 97.1 99 98.4 99.2 98.8 98.3 96.2

S. terebinthifolius

71.4 67.4 51.2 45.7 78.5 72.6 94.7 94.5

Table 6. Summary of results of nested ANOVA for repeated measures (F values with p

values in brackets) testing for effects of treatments on functional trophic groups density

of soil macrofauna (ind. g

-1

d.w. litter – square root transformed). All statistically

significant numbers (p<0.05) are in boldface.

Degrees of freedom are: site (1, 6), blocks

within site (6, 14), leaf-litter species (2, 14), time (3, 42), leaf-litter species x time (6,

42).

Response

variables

Site

Blocks

within site

Leaf-litter

species (A)

Time (B) A x B

Detritivores 0.6.4 (0.45) 2.41 (0.08) 1.13 (0.35)

19.34 (<0.01) 5.37 (<0.01)

Carnivores

10.13 (0.02)

0.79 (0.59)

7.58 (0.01) 7.45 (< 0.01) 4.43 (< 0.01)

Omnivores 1.48 (0.27) 0.66 (0.68) 1.10 (0.36) 0.80 (0.50) 0.78 (0.59)

Herbivores 1.26 (0.30) 0.68 (0.67) 0.45 (0.64) 1.82 (0.16) 0.47 (0.83)

FIGURES

Figure 1. Study area with different ash disposal types and pioneer plants. A) Recent ash

deposit showing the plant community structure around; B) Boiler slag disposal site; C)

Fly ash disposal site; D) Cynodon dactylon (grass) and Ricinus communis (shrub); E)

Schinus terebinthifolius (tree).

Figure 2. Box plot for pH values of fly ash and boiler slag disposal sites.

C.dactylon

R.communis

S.therebinthifolius

C.dactylon

R.communis

S.therebinthifolius

C.dactylon

R.communis

S.therebinthifolius

C.dactylon

R.communis

S.therebinthifolius

0.0

0.6

1.2

1.8

2.4

6 35 70 140

Leaf-litter mass remaining (%)

Density of individuals

Time (days)

C. dactylon

S. terebinthifolius

R. communis

Fly ash

Boiler slag

0.0

0.6

1.2

1.8

2.4

0.0

0.4

0.8

1.2

1.6

Ash disposal site

Leaf-litter

Figure 3. Mean ± SE of leaf-litter mass remaining (A, B) and individuals.g

-1

d.w. litter

(square root transformed) from macrofauna (C, D) in response to effects of treatments:

leaf-litter species X sampling time (A, C), and ash disposal site(B, D).

0 300 600 900 1200

Boiler slag

Fly ash

0 250 500 750 1000

Abundance

Macrofauna species richness

S. terebinthifolius

C. dactylon

R. communis

B) Ash disposal site

A) Leaf-litter

Figure 4: Rarefaction curves of total soil macrofauna in response to leaf-litter species

and ash disposal sites. Error bars represent ± 1 confidence interval (C.I. 95%).

Figure 5. NMDS ordination in two dimensions of sample units showing the total soil

macrofauna community structure in response to leaf-litter species and ash disposal sites.

Circle= C. dactylon; triangle= R. communis, square= S. terebinthifolius; white symbols=

fly ash site; black symbols= boiler slag site.

Figure 6. Mean ±SE of individuals.g

-1

d.w.litter (square root transformed) per functional

trophic group of soil macrofauna under effects of treatments (leaf-litter species X

0.0

0.8

1.6

2.4

3.2

0.0

0.7

1.4

2.1

2.8

0.0

0.5

1.0

1.5

2.0

Density of individuals

0.0

0.4

0.8

1.2

1.6

Time (days)

6 35 70 140

0.0

0.2

0.4

0.6

0.8

0.00

0.25

0.50

0.75

1.00

Fly ash

Boiler slag

0.00

0.07

0.14

0.21

0.28

0.0

0.2

0.4

0.6

0.8

Ash disposal site

Leaf-litter

A) Detritivores

B) Detritivores

C) Carnivores

D) Carnivores

E) Omnivores

F) Omnivores

G) Herbivores

H) Herbivores

C. dactylon

S. terebinthifolius

R. communis

sampling time, and ash disposal site).

0350700

0 450 900

Ash disposal site

Leaf-litter

A) Detritivores

060120

0 60 120

C) Carnivores

01224

015

G) Herbivores

050100

0 50 100

E) Omnivores

S. terebinthifolius

C. dactylon

R. communis

Boiler slag

Fly ash

Abundance

B) Detritivores

D) Carnivores

H) Herbivores

F) Omnivores

Species richness of functional trophic groups

Figure 7: Rarefaction curves for functional trophic groups of soil macrofauna in

response to leaf-litter species and ash disposal sites. Error bars represent ± 1 confidence

interval (C.I. 95%).

position and abundance of soil macrofauna colonizing pioneer leaf litter (C-Cynodon dactylon, R-Ricinus communis,

) at 6, 35, 70 and 140 days of leaf-litter exposure, in two different coal ash disposal sites (fly ash and boiler slag). Functional

trophic groups (FTG) are: De

F ily /

A haen

I atur

G hosi

J ns

Ha iidae

H nida

Li hiid

L inac

yph

S ecur

I atur

Ly sidae

L cosidae

I atur

Oecobida

I atur

tritivores (D), Carnivores (C), Omnivores (O) and Herbivores (H). N.I.= not identified in morphospecie or throphic group.

Fly ash site Boiler slag site

C R S C R S

Class / Order am Species or Morphospecies FTG

6 35 70 140 6 35 70 140 6 35 70 140 6 35 70 140 6 35 70 140 6 35 70 140

Arachnida

Araneae nyp idae

mm es C 0 1 3 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 1 1 0

Araneida

mm es C 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Corinnidae

mm es C 0 0 0 0 1 0 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0

nap dae

ove C 0 2 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

ah e sp. C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0

nyp ae

Erigon sp. C 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 2 0 0 0 0 0 0 0 1

am auda sp. C 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

Lepth antes sp. C 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0

col a cambara Rodrigues, 2005 C 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0

mm es C 0 0 0 7 0 0 0 0 0 1 1 1 0 0 0 2 0 0 0 1 1 0 0 0

y sp. C 4 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 2 0 0 0 0 0 0 0

mm es C 5 1 0 2 2 0 0 5 6 4 0 1 3 1 1 0 2 1 0 0 0 0 0 1

mm es C 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

noopidae

onopinae sp. C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1

mmatures C 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Salticidae

Aphirape uncifera (Tullgreen, 1905) C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0

Appendix 1: Morphospecies com

S-Schinus terebinthifolius

Unidentatae sp.1 C 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Unidentatae sp.2 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Immatures C 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0

Scytodidae

Immatures C 0 0 1 10 0 0 0 1 0 1 3 0 0 0 0 0 0 0 0 0 0 0 0 1

0 0 0 0 0 0 0

Coleosoma sp. C 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Immatures C 0 0 0 2 1 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0

Thomisidae

Immatures C 0 0 4 0 1 0 0 0 0 0 0 0 0 0 1 1 0 2 1 0 1 1 0

Opiliones p. Opiliones s O 0 0 0 0 0 0 0 1 0 2 0 0 0 0 0 0 0 0 0 0 1 0 0

Chilopoda C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Chilopoda 0 1 1 0 2 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 4 0

Chilopoda C 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0

Amphipoda Talitridae

Talitroides sylvaticus (Haswell) O 0 0 0 0 0 2 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0

Isopoda Balloniscidae

Balloniscus sellowii (Brandt, 1833) D 11 6 0 0 2 0 3 0 1 0 1 5 1 1 2 1 2 2 0 0 0 8 0

Philosciidae

Atlantoscia floridana (van Name, 1940) D 7 1 4 0 20 9 0 0 27 32 14 2 0 0 0 0 0 0 0 0 0 0 0

Benthana taeniata Araujo & Buckup, 1994 D 81 28 13 8 88 11 0 1 159 72 39 16 85 24 12 1 22 2 0 54 16 17 0

Plathyarthridae

Trichorhina sp. D 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Diplopod 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0

Diplopoda sp. D 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 4 2 1

Soleolifera ae Veronicellid

Belocaulus angustipes (Heynemann, 1885) H 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Stylommatophora dae Agriolimaci

Deroceras laeve (Müller, 1774) D 0 1 0 1 2 0 0 0 0 0 0 3 0 0 1 2 2 0 0 0 0 4 1

Bradybaenidae

Bradybaena similaris (Férussac, 1821) H 0 0 0 6 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0

Bulimulidae

Scytodes sp. C 0 0 0 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Segestriidae

Immatures C 0 0 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Tetragnathidae

Immatures C 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0

Theridiidae

Thymoites sp. C 0

Chilopoda sp.1 0

sp.2 C 0

sp.3 0

Crustacea

Diplopoda a sp.1 D 0 0

2 0

Gastropoda

Continuação Appendix 1- Artigo 1

Drymaeus sp. H 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0

Euconulidae

Habroconus sp. D 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 6 0

Systrophiidae

Happia sp. C 0 0 1 0 2 0 0 0 5 0 1 2 0 0 0 0 0 0 0 0 0 0 2

aff. Drepanostomella sp. C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Gastropoda 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

N.I. 0 0 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Blattodea Blattodea sp.1 D 0 0 0 0 2 0 0 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0

Blattodea s 0 0 0 0 0 0 0 0 3 0 1 0 0 0 0 0 0 0 0 1 0 0

Blattodea sp.3 D 1 2 9 3 0 0 0 0 10 3 0 0 0 0 0 0 0 0 0 0 1 0 0

N.I. D 0 0 0 5 0 0 0 0 0 0 0 5 0 0 0 0 0 0 1 0 0 0 0

Coleoptera Anthicidae

Anthicidae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Chrysomelidae

Chrysomelidae sp.1 H 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0

Curculionidae sp.1 H 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Curculionidae sp.2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0

Curculionidae sp.3 H 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0

Curculionidae sp.4 H 1 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0

Elateridae

Elateridae sp. D 3 1 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Leiodidae

Leiodidae sp. D 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0

Nitidulidae

Nitidulidae sp.1 D 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Nitidulidae sp.2 D 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Nitidulidae sp.3 D 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Phalacridae

Phalacridae sp.1 0 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Phalacridae sp.2 D 4 0 0 0 6 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0

Ptilidae

sp. 0

Hexapoda

p.2 D 0 0

sp. C 0 0

0 0 0 0

Chrysomelidae sp.2 H 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Chrysomelidae sp.3 H 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Chrysomelidae H 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Diabrotica speciosa (Germar, 1824) H 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0

Colydiidae

Colydiidae sp. D 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Curculionidae

sp.4

H 1 0

D 0 0

Continuação Appendix 1- Artigo 1

Ptilidae sp. D 0 1 0 2 0 0 0 0 0 0 0 1 0 0 0 5 0 0 0 0 0 0 2 0

Scarabaeidae

Scarabaeidae sp. D 0 0 0 0 0 0 0 0 0 0 0 0 3 1 1 1 1 0 0 0 0 1 2 1

Scarabaeidae sp 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Scydmaenidae

Scydmaenidae sp. C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Staphilinidae

Staphilinidae s 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Staphilinidae sp.10 C 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae s p.11 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Staphilinidae s p.12 C 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae s C 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0

Staphilinidae s C 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae sp.15 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Staphilinidae s p.16 C 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0

Staphilinidae s 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Staphilinidae sp.18 C 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae sp.19 C 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae sp.2 C 0 0 0 0 0 0 0 0 0 0 0 0 6 0 0 0 0 0 0 0 0 0 0 1

Staphilinidae sp.20 C 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae sp.21 C 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1

Staphilinidae sp.3 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 3 0 0

Staphilinidae s C 0 0 0 0 0 0 0 0 0 0 0 0 8 0 0 0 1 0 0 0 1 0 0 0

Staphilinidae s 0 2 2 0 0 1 0 0 1 2 2 0 1 3 0 1 0 0 0 0 0 1

Staphilinidae sp.6 C 0 0 0 0 0 0 0 0 0 4 14 0 0 0 0 0 0 0 0 0 0 0 0 0

Staphilinidae sp.7 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0

Staphilinidae sp.8 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Staphilinidae sp.9 C 6 0 0 0 7 0 0 0 0 0 0 0 1 0 0 1 3 0 0 0 0 0 0 0

Pselaphinae

Pselaphinae sp C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0

Pselaphinae 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Pselaphinae s C 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0

Pselaphinae 3 6 1 0 2 0 0 0 0 3 3 1 0 0 3 0 0 0 0 0 0 0 1 0

Pselaphinae sp.5 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

Pselaphinae sp.6 C 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Pselaphinae sp.7 C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 1 0 0 0 0 0 0 1 0

Pselaphinae 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0

Scaphidiinae

Scaphidiinae sp. D 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0

Tenebrionidae

.2 D 1 0

p.1 C

p.13

p.14

p.17 C 0 0

C 16 14 0

p.4

p.5 C 0 0

sp.2 C

p.3

sp.4 C

sp.8 C

Continuação Appendix 1- Artigo 1

ae sp.1 Tenebrionid D 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Tenebronida 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

Dermaptera Dermaptera sp.1 O 0 0 0 0 0 0 0 0 4 1 3 0 0 0 0 0 0 0 0 0 0 0 0

Dermaptera sp.2 O 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Hemiptera Fulgoroidea

Fulgoroidea sp. H 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Cicadellidae

Cicadellidae sp.1 H 0 0 1 1 0 0 0 0 0 0 0 2 0 0 0 0

e sp.2 D 0 0 0 0

0 0 0 0 0 0 1 0

Cicadellidae sp.2 H 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0

Membracidae

Membracidae sp. H 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Cydnidae

Cydnidae sp. H 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0

Miridae

H 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Pentatomidae

0 0 0

Miridae sp. 0 0

Oebalus poecillus (Dallas, 1851) H 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

Reduviidae

0 0

Reduviidae sp. C 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Ryparochromidae

p. Ryparochromidae s H 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

NI H 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

Hymenoptera Formicidae

Dolichoderinae

Dorymyrmex sp. O 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Formicinae

0 0

Brachymyrmex sp.1 O 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0

Brachymyrmex sp.2 O 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 3 0

Camponotus sp.2 O 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0

Camponotus sp.3 O 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Paratrechina sp.1 2 O 3 0 1 0 0 0 1 0 58 0 3 0 0 0 0 0 1 0 2 0

Paratrechina sp.2 O 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0

Myrmicinae

0 0 0 0 0

0 0 0 0

0 0

0 0 0 0

Acromyrmex sp. D 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Camponotus sp.1

0 0 0 0 2

O 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Pheidole sp.1 O 0 0 0 2 0 0 0 0

0 0 0 0

1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Pheidole sp.2

O 6 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Pheidole sp. 0 0 0 7 0 0 0 0 0

Pheidole sp.4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Pyramica sp. O 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0

0 0 0 0

O 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

O 0 0 1

0 0 0 0

Continuação Appendix 1- Artigo 1

Solenopsis sp.1 0 3 0 1 0 4 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0

Solenopsis sp.2 0 0 0 0 0 0 0 0 1 1 2 0 3 4 2 0 1 1 2 10 6

Solenopsis sp.3 0 0 0 0 0 0 1 0 0

O 0 0 0

O 0 1 0

O 0 1 1 2 0 0 1 0 0 0 0 0 3 1 3

Solenopsis sp.4 O 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0

Solenopsis sp.5 O 0 0 0 0 1 0 0 0 0 0 0 0 6 0 0 0 0 0 0 1 0 0 0 0

Strumigeny sp. C 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 3 0

Wamannia sp.1 O 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Wasmannia auropunctata (Roger) O 1 0 0 0 0 0 0 0 0 0 2 7 0 1 1 0 0 0 0 0 1 0 0 0

Ponerinae

Hypoponera sp.1 O 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0

Hypoponera sp.2 O 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

N.I. 0 0 0 0 0 0 0 1 0 0 2 0 0 0 2 0 0 0 0 0 0 0 0

Orthoptera . Orthoptera sp H 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

Immatures (larvae) 2 2 42 N.I. - 6 6 0 0 5 0 1 0 7 6 5 5 4 1 5 4 3 0 0 0 9

Megascolecid

lifornica (Kinberg, 1867) 49 10 17 Metaphire ca D 17 20 19 13 0 2 2 3 8 22 36 27 8 156 16 0 0 6 49 21 0

55 16 67 82 82 42 5 17 250 166 67 76 162 80 71 46 72 49 8 7 72 97 150 34

Oligochaeta ae

TOTAL 1 1 1 3 2

Continuação Appendix 1- Artigo 1

ARTIGO 2.

Neotropical woodlice (Crustacea, Isopoda) colonizing

early successional plant leaf-litter in a coal ash disposal

a Podgaiski

, Aline Ferreira Quadros

, Paula Beatriz

Araujo

and Gilberto Gonçalves Rodrigues

a,c

I ncias, Universidade Federal do Rio Grande do Sul, Porto

Alegre, RS, Brazil. Fax: +55 51 33087626. E-mail: [email protected].

I to de Biociências, Universidade Federal do Rio Grande do Sul, Porto

pb [email protected].

Departamento de Zoologia, Centro de Ciências Biológicas, Universidade

Federal de Pernambuco. Av: Professor Moraes Rego, S/N, Cidade

Universitária, Recife, PE, CEP 50670-420. E-mail: [email protected].

environment*

Luciana Regin

stituto de Biociê

Alegre, RS, Brazil. Fax: +55 51 3308 7698. E-mail: [email protected];

nstitu

arau

*Artigo submetido em 26/09/2008 para a publicação na revista Science of

otal Environment (ISSN: 0048-9697; Elsevier; Impact factor: 2.182).

Rejeitado em 09/11/2008.

cal Fauna and Environment (ISSN: 1744-5140; Taylor &

Francis; Impact factor: 0.709).

Revisão da editora em 09/01/2009.

Artigo resubmetido em 15/01/2009.

Artigo reformulado submetido em 07/01/2009 para publicação na revista

Studies on Neotropi

Abstract

An experimental study was conducted to investigate the colonization patterns of

woodlice in leaf-litter from three spontaneous pioneer plants in an environment exposed

long-term coal ash pollution (fly ash and boiler slag). Three species were found:

sure (6, 35, 70 and 140 days). Occurrence of A.

f B. taeniata females was strongly influenced

discussed under ecological aspects.

Key words: Brazil, soil invertebrates; detritivores, litter decomposition, fly ash; boiler

slag.

Atlantoscia floridana (Philosciidae; n=116), Benthana taeniata (Philosciidae; n=817)

and Balloniscus sellowii (Balloniscidae; n=48). Woodlice colonized equally the plant

species along the time of leaf expo

floridana and abundance and fecundity o

by the ash disposal type. These results are

Introduction

rrounding

nvironment (Silva et al 2000).

e in terrestrial ecosystems is decline of plant establishment and growth (Carlson &

Adriano 1993). This decline results from changes in edaphic traits (Tordoff et al 2000;

Gupta et al 2002), that leads to unfavorable physical (compacted layers, reduced bulk

density) and chemical conditions (nutrient depletion, high toxicity and low pH-value)

and reduction in microbial and soil biota activity. Despite these challenges, early

successional plant species that show heavy metal tolerance or resistance are able to

colonize such environments (Whiting et al 2004). A positive feedback loop may then

occur between the pioneer plants and the substrate (Wilson & Agnew 1992) by the

improvement of the microclimate and resource availability through the stabilization of

coal wastes, shading, aerating, depositing litter and offering habitat and shelter for

animals (Carlson & Adriano 1993). In this way, early successional plants may pave the

way for subsequent species in a facilitation model (Connel & Slatyer 1977).

Coal has been the fastest-growing major fuel in the world (BP 2008). However, its

extraction and combustion implies in serious environmental impacts. The combustion of

coal in thermal power plants produces vast quantities of wastes, and instead of utilizing

those residues for the building industry (Scheetz & Earle 1998; Rohde et al 2006),

thermal power plants usually make landfills or minefills. Coal and associated coal

combustion products contain polycyclic aromatic hydrocarbons and trace elements of

“heavy metals” (Querol et al 1995; Teixeira et al 1999). These substances may contain

mobile toxic constituents with potentially genotoxic effects in the su

One of the major potential adverse impacts of the disposal of coal combustion

residu

Soil invertebrates play a crucial role in the early succession of the restoration

woodlice (Donker 1992). These changes are related to

increas

binthifolius

Raddi (Anacardiaceae), a native tree, is very abundant in South Brazil and it was found

process in polluted/damaged ecosystems. Woodlice (Crustacea, Isopoda) are commonly

found in metal-polluted environments (Jones & Hopkin 1996; Jones & Hopkin 1998;

Grelle et al 2000; Tajovský 2001). As abundant litter detritivores in terrestrial

ecosystems they participate in the leaf-litter processing (Quadros & Araujo 2008) by

breaking up the organic matter and returning nutrients to soil through their feces, which

strongly enhance microorganism activity (Hassall et al 1987). To avoid the toxic effects

from contaminated food, they are capable of immobilizing high levels of heavy metals

in their hepatopancreas (Raessler et al 2005). This may affect the life history traits by

changing resource allocation of

ed mortality, slower growth rates, reduced body size and reproduction effects

(Donker 1992; Donker et al 1993; Jones & Hopkin 1996; Jones & Hopkin 1998).

However, there is strong evidence for the occurrence of heavy-metal adaptation in

natural populations of isopods (Posthuma & Van Straalen 2002).

The southern region of Brazil is very rich in coal reserves, which have been

exploited, extracted and destined to generation of energy in thermal power plants for

decades (Pires & Querol 2004). Consequently, this region has a number of

environments exposed to long-term coal ash pollution (Teixeira et al 1999). A recent

research in one of those environments investigated the leaf decomposition of three

abundant early successional plant species and the diversity of associated soil

invertebrate macrofauna (Podgaiski & Rodrigues unpubl. data). Ricinus communis L.

(Euphorbiaceae) and Cynodon dactylon (L.) Persoon (Poaceae) are exotic plants in

Brazil, and they occurred in patches of recent ash deposits. Schinus tere

in olde

ifferent sites are found

in the a

r ash deposits. The research revealed woodlice as the most abundant group of soil

macrofauna in leaf-litter. In view of that, the present study investigates (1) the

colonization patterns of different woodlice species along leaf decomposition of these

plant species and (2) the woodlice abundance and some reproductive traits in different

types of coal ash disposals (fly ash and boiler slag).

Materials and methods

Study area

The study was conducted in the city of São Jerônimo in the state of Rio Grande do Sul,

Brazil (29°57’55.6”S; 51°44’14.9”W), in a site along a riparian forest of the Jacuí river

(Figure 1). The climate of this region is temperate (Cfa type of Köppen-Geiger climate

classification; Peel et al 2007) with hot summers but without a dry season.

The study area has been a coal combustion residue disposal site for more than 30

years, but has recently stopped receiving additional waste. Two d

rea: one that was more influenced by fly ash disposal and another that was more

influenced by boiler slag disposal (Figure 1). Fly ash and boiler slag are aluminosilicate

minerals, with SiO

and Al

being the predominant components (Rohde et al 2006),

and they differ in both physical and chemical properties. Fly ash is a fine-grained

powder, composed of spherical glassy and hollow particles captured by air pollution

control equipment in thermal power plants. Boiler slag is a specific type of bottom ash,

which is vitreous molten grained and composed of angular particles with high C content

from unburnt coal derived from wet ash removal of wet-bottom furnaces (ECOBA

2008). The elemental concentrations vary according to parent coal composition and

combustion technology. Rohde et al (2006) studied the results of leaching fly ash and

boiler slag from the São Jerônimo thermal power plant and found the most abundant

elements to be Sn, Ni and Mo, and in lower proportion Cr, As, Hg, Al, Pb, Mn, V, Cd,

Ba and Zn. Boiler slag had higher concentrations of Sn, Mo, Cr, Al, Pb and Mn than fly

ash, which, in turn, had higher concentrations of Ni, Hg, Cd and Zn. The boiler slag

disposal site has a lower pH (mean 5.1) than the fly ash disposal (mean 5.8) (Podgaiski

Rodrigues unpubl. data).

l design and laboratory procedures

ed by fly ash

disposa

Experimenta

Ninety-six nylon litter bags of 30 x 20 cm, made of coarse mesh (1.0 x 0.2 cm) were

filled with 20.3 ± 0.2 g of air-dried freshly fallen leaves of C. dactylon, R. communis

and S. terebinthifolius (32 litter bags per plant species). In June 2007, the litter bags

were placed on bare ground in eight blocks: four within the site influenc

l and four within the site influenced by boiler slag disposal (Figure 1). Each

block was composed of 12 litter bags (four litter bags per plant species), distant at least

2 m from each other (Figure 1). There were four successive sampling occasions (6, 35,

70 and 140 days after exposure), when one litter bag of each plant species was randomly

removed from each block (Figure 1).

The isopods from the litter bags were manually collected, identified at species

level and counted. The reproductive traits measured were the size of ovigerous females

(cephalothorax width; Araujo & Bond-Buckup 2004) and their fecundity (marsupial

content). After the inspection of the litter bags to sort out the animals, the remaining

leaf-litter in each bag was dried and weighed.

Keeping in mind that isopods prefer leaf-litter with a low Carbon-Nitrogen

(C:N) ratio (Zimmer 2002), the C and N content of green leaves of each plant leaf-litter

species was determined using the methods of moisture combustion/Walkley-Black

(Walkley & Black, 1934) and Kjeldahl (Kjeldahl 1883) with 0.01 % of detection limit,

respectively. The C:N ratio was lowest for R. communis (8.0), followed by C. dactylon

(26.7) and S. terebinthifolius (34.2) (Podgaiski & Rodrigues unpubl. data).

Statistical analysis

Isopod abundance in litter bags was standardized and expressed as individuals.g

-1

d.w.

for each isopod species) was

s were found, totaling 981 individuals (Table 1):

Bentha

with 63% of individuals occurring in the

fly ash

(dry weight) litter. The isopod abundance (total and

compared in plant leaf-litter species and sampling dates with repeated measures analysis

of variance (ANOVA) in blocks for each of two sites influenced by coal ash disposals.

The relationship between isopod ovigerous female size (mm) and fecundity was

assessed between sites using analysis of covariance (ANCOVA). The size of the

ovigerous females in both sites was compared with ANOVA. Residual analyses were

carried out to check all models used in this study.

Results

Three native neotropical isopod specie

na taeniata Araujo & Buckup, 1994 (Philosciidae), Atlantoscia floridana (van

Name, 1940) (Philosciidae) and Balloniscus sellowii (Brandt, 1833) (Balloniscidae).

The species behave differently regarding the two sites of disposal. Atlantoscia floridana

occurred exclusively in the fly ash site (116 ind.), B. taeniata was found in a much

higher abundance than the others (817 ind.),

. Balloniscus sellowii, on the other hand, was rarer than the others (48 ind.) and

showed no differences between the two sites (Table 1).

According to results of repeated measures ANOVA, the total abundance of

isopods as well as the abundance of each isopod species was not significantly different

among leaf-litter species (Table 2; Figure 2) in both sites. There was no interaction

between treatments (leaf-litter species) and time (sampling dates). However, a tendency

wards a higher abundance of all isopod species in R. communis leaf-litter could be

f leaf exposure (Figure 2). The time of leaf exposure only had a

1, 50

= 10.79; P= 0.002; Figure 3).

evertheless, ovigerous female size was not significantly different between sites (F

1, 51

.88).

ion (Araujo et al.

observed at 35 days o

significant effect in the abundance of B. taeniata in the fly ash disposal site. In this case,

B. taeniata was more abundant in 6 days than in 70 days of leaf-litter decomposition

(Tukey test; P=0.04).

Benthana taeniata was the only species that presented a sufficient number of

ovigerous females for the analysis of reproductive traits (53 individuals: 34 in the fly

ash site and 19 in the boiler slag site). ANCOVA indicates that fecundity in relation to

female size (F

1, 50

= 107.36; P< 0.001) was lower in females from the boiler slag disposal

site than in females from the fly ash disposal site

0.02; P=0

Discussion

The woodlice species found in the coal waste disposal sites studied are native to the

Neotropical region. Among them, A. floridana is the most common and abundant

species in southern Brazil. Its populations can reach up to 1040 ind. m

-2

(Araujo &

Bond-Buckup 2005), with an average biomass of 1 kg ha

-1

(Quadros & Araujo 2008).

Balloniscus sellowii is also commonly found in southern Brazil (Araujo et al. 1996;

Lopes et al. 2005). Benthana taeniata occurs exclusively in this reg

1996)

seems to have different levels of tolerance to the environments

created

ts, such as monocultures of exotics plants (Eucaliptus spp. and Pinus

p.) and urban parks. Moreover, B. sellowii shows an increased fecundity, which is a

at enhances its colonization ability (Quadros et al. 2008).

and no data on its biology and ecology is available. The fragment of riparian

vegetation situated between the disposal sites and the adjacent river probably serves as a

source for these colonizing populations. Interestingly, synantropic woodlice as

Armadillidum spp., Porcellio spp. and Porcellionides spp., which are very abundant in

urban areas in Brazil (Araujo et al. 1996), were not present in the area of the thermal

power plant and disposal sites.

Woodlice in coal ash disposal sites

The woodlice species

by the coal ash disposal. The boiler slag is clearly less suitable for isopods,

especially A. floridana. The increased amount of contaminants such as Se, Mo and Mn

in boiler slags (Rohde et al 2006) probably contributed to the low abundances verified

in this site, if they imply in higher mortality and/or slower growth rates, then decreasing

or inhibiting colonization by woodlice. Compared to A. floridana and B. taeniata, B.

sellowii seems to be more tolerant to this habitat, since it is commonly found in human

managed habita

key reproductive trait th

Fecundity of B. taeniata females was differently affected in the two sites.

Previous studies have shown evidence of changes in life history traits of woodlice

inhabiting contaminated environments (Donker et al 1993; Jones & Hopkin 1996; Van

Brummelen et al 1996) due to changes in resource allocation (Donker 1992). The lower

fecundity of B. taeniata females in the most polluted site was may be an example of

such trade-off, but since there are no other studies concerning the ecology and

reproduction of B. taeniata, especially in non-polluted sites, further research needs to be

conducted.

Woodlice in pioneer leaf-litter along decomposition

Terrestrial isopods prefer to feed from leaf litter of decayed, dicotyledonous (Rushton &

Hassall 1983; Hassall et al 1987) plants with low C:N ratio (Zimmer 2002). Considering

at they may be attracted to a litter patch that offers more palatable food (Tuck &

expected in R. communis bags (based on its

ource,

howev

Hassall 2005), a higher abundance was

lower C:N ratio) and in leaf-litter from the latest days of decomposition (more decayed

leaves). However, our predictions were not supported as woodlice used the three plant

species in all times of leaf decomposition. Another possibility is that woodlice were

attracted to the litter bags for sheltering and protection from direct light, high

temperatures (Hassall & Tuck 2007) and potential predators. For organisms that utilize

the litter both as food and habitat, there is a trade-off between these two resources,

because the decomposition process that increases the palatability of the litter (Hassall et

al 1987) at the same time decreases the structural integrity that makes it best suited for

use as persistent shelter (Hooper et al 2000). The nitrogen-rich leaf-litter of R.

communis is an example of such a trade-off. It may constitute a valuable food s

er, it decomposes much faster than the others (Podgaiski & Rodrigues unpubl.

data). If the litter was not suitable for feeding in the beginning of the experiment,

woodlice may have been attracted to the litter bags mainly for sheltering and, along the

decaying process and after considerable microbial degradation (Wolters 2000), they

may have fed from the litter.

The woodlice populations studied here inhabit a contaminated and highly

modified riparian ecosystem of southern Brazil. Considering that they are abundant

etritivores, they influence the soil restoration processes in this area by the acceleration

utrient availability for the

of humus-forming processes that contribute to an efficient n

establishment of the plant community. On the other hand, as bioaccumulator organisms

that are predated by a wide range of animals, both invertebrates and vertebrates

(Sunderland & Sutton 1980), they are likely to take part in the process of

biomagnification of heavy metals through the food chain (Paoletti & Hassall 1999).

Despite some studies have been conducted with woodlice and other invertebrates in

polluted-environments in Europe and other regions of the world (e.g., Majer et al 2007;

Tajovský 2001; Grelle et al 2000), these kinds of studies are not yet found in Brazil with

neotropical invertebrates. Studies relating plants and invertebrate detritivores,

ecosystem processes (such as litter decomposition) and effects of contaminants on

invertebrate biology would be very important to the understanding of the ecology of

polluted environments. This knowledge would be essential to meet restoration and

diversity conservation objectives.

Acknowledgements

To André F.B. Lima, André Castillo, André L. Casara, Tamires B. da Silva and

Verônica G. Sydow for helping with fieldwork and/or lab procedures; to Adriano S.

Melo and Gislene Ganade for statistical analysis advice; to CAPES for the scholarship

awarded to Luciana R. Podgaiski and Aline F. Quadros; to CGTEE for access to the

study site.

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Tables

Table 1. Total abundance of isopod species colonizing leaf-litter from three pioneer

tere

Fly ash disposal site Boiler slag disposal site Total

plant species (C- Cynodon dactylon, R- Ricinus communis and S- Schinus

binthifolius) in two coal ash disposal sites. N= 96 litter bags.

Isopod species

Total

osciidae

ntoscia floridana

12 29 75 116 0 0 0 0 116

thana taeniata

130 100 286 516 122 92 87 301 817

loniscidae

Phil

Atla

Ben

Bal

Balloniscus sellowii

2 4 18 24 6 4 14 24 48

Total iso

ods 144 133 379 656 128 96 101 325 981

easures ANOVA (F and P-valu fo rs cti the al ab ance o op a B. tae ta,

sellowii and A. floridana abundances for each of two coal ash disp tes ou of riatio re: trea nt af er species:

dactylon, R. communis and S. terebinthifolius), time (6, 35, 70 a 14 aft eaf po ) and cks (n= pe site).

Fly ash disposal site Boiler slag disposal site

es)

r fa

osa

0 d

cto

l si

ays

affe

. S

er l

rces

tot

sure

und

n a

blo

ime

1.5 (0.256)

0.3 (0.817) 1.4 (0.267)

* 1) 2.9 (0.120)

ent ime, Treatment x Tim and

f is ods nd

tme (le -litt

4 r

A x B

1.2 (0.346) 0

1.2 (0.365) 0

1.7 (0.169)

ck, respectively. *

nia

Table 2. Results of repeated m

Response variable

Treatment (A) Time (B) A x B Block Treatment (A) T (B) Block

Total abundance 1.4 (0.306) 2.3 (0.110) 1.0 (0.442) 1.4 (0.319) 0.9 (0.455) .8 (0.556)

B. taeniata

0.6 (0.567) 3.7 (0.031) 0.6 (0.730) 0.9 (0.448) .8 (0.543)

A. floridana

1.0 (0.417) 1.4 (0.280) 1.0 (0.46 * *

B. sellowi

0.4 (0.695) 2.2 (0.128) 0.6 (0.705) 0.7 (0.565) 0.4 (0.677) 2.5 (0.093) 0.2(0.908)

P-values are in brackets. Degrees of freedom are 2, 3, 6 and 3 for Treatm , T e Blo A.

floridana was not found at the boiler slag disposal site.

Figures

Figure 1. Study area and sites in São Jerônim

distance of the two sites

site had four blocks with four litter bags of

inside the triangle (B). T

exposure).

). Each study

sposed as illustrated

o, Rio Grande do Sul, Brazil, showing the

the coal power plant and to the Jacuí river (A

each plant species, di

n ers in the l tte g icate sa

from

he umb i r ba s ind mpling dates (days after

Figure 2. Mean ± standard error of abundances (individuals.g

-1

d.w. litter) of terrestrial

isopods in different plant species in litter bags and in four sampling occasions (6, 35, 70

and 140 days) in two coal ash disposal sites.

Figure 3. Relationships between fecundity (marsupial content) of Benthana taeniata

females and female size (cephalothorax width) in two coal ash disposal environments

(fly ash and boiler slag). Each point represents one ovigerous female (n= 53).

ARTIGO 3.

Does enhancement of mix leaves on the soil improve

macroarthropod diversity and litter decomposition in a

post-mining pinus forest?*

Luciana Regina Podgaiski

and Gilberto Gonçalves Rodrigues

a,b

Departamento de Zoologia, Centro de Ciências Biológicas, Universidade

Federal de Pernambuco. Av: Professor Moraes Rego, S/N, Cidade

Universitária, Recife, PE, CEP 50670-420.

rtigo a ser submetido para publicação na revista científica Ecology (ISSN

0012-9658; ESA; Impact factor: 4.78).

Programa de Pós-Graduação em Ecologia, Instituto de Biociências,

Universidade Federal do Rio Grande do Sul. Av: Bento Gonçalves, 9500, prédio

43422, Porto Alegre, RS, CEP 91501-970.

Summary

In ecological systems, the diversity of one component may promote the diversity in

other components (e.g., belowground resources vs. aboveground biota). In the case of

native leaves on the floor of a mined Pinus

elliottii monospecific forest in southern Brazil would increase the soil macroarthropod

community diversity. Further, as high quality and structured litters may promote the

decomposition of associated litters by interactions between them and due increase in

biota diversity, we also hypothesized that our experiment could alter the decomposition

dynamics of P. elliottii litter. We show that mix leaf-litter plots attracted higher

individual density, 25 % more species and distinct macroarthropods species

composition when compared to P. elliottii leaf-litter plots. Moreover, considering

spiders separately, the increased in species richness reach up to 58 %. Mass and nutrient

loss of P. elliottii leaf-litter were not altered when in contact with mixed leaf-litter. On

the

of less C (%).

rces on a damaged monospecific forest

can improve soil biodiversity and alter the dynamic of an ecosystem process. These

results may have implications for forest management and restoration of damaged

system

Key words: Brazil, leaf-litter decomposition, litter mixing, soil fauna

plant litter vs. soil invertebrates, the diversity of the last would be promoted manly due

to microhabitat spatial heterogeneity and a diverse set of food resources. In this study,

we hypothesized that enhancement of mix

the other hand, the nutrient content composition of litter was responsive to

treatment, where we found higher P (%) and K (%) and an indicative

Thus, it appears that the input of diverse resou

Introduction

of interespecific competition, and possibilities of more species coexistence in

greater resource use (MacArthur 1970, McKane et al 2002, Finke and Snyder 2008).

obes, nematodes

and mites to effects of plant species diversity (reviewed in Wardle 2006). Although the

The human effects on biodiversity are worldwide known. Changes in land use through a

variety of activities increase rates of deforestation, soil exhaustion, alien invasion, and

species extinction. Habitat degradation and changes in biotic functional diversity are

usually accompanied by changes in ecosystem properties (Hopper et al. 2005). The

impact of conversion of natural to agricultural systems, for example, reflects drastically

in the decrease on plant diversity, which in turn reduces the range of invertebrates and

the functions that they drive, such as related to decomposition (Swift and Anderson

1994, Hooper et al. 2000). Because biodiversity influences and is influenced by the

system components at many scales, an enhancing of our understanding about diversity

drivers are required in front of environmental changes (Tilman 1994, Wardle et al.

2004), seeking best taking decisions about ecosystem management and conservation.

In terrestrial ecosystems, diversity of one component may promote the diversity

in other components (Siemann et al. 1998, Tews et al. 2004, Armbrecht et al. 2004).

This linkage is clearly recognized, mainly when we look at the influences of

aboveground communities in the belowground communities. High diversity of plants

can input high litter diversity on the soil, which favors a diversity of decomposers,

detritivores and organisms from the higher trophic levels in the soil food web (Hooper

et al. 2000). Several litter qualities and litter types offer both diverse food and

microhabitats to the soil biota, favoring more niches and resource-use differences,

reduction

Several recent studies had emphasized specially responses of soil micr

large representativeness of macroarthropod groups in the soil diversity, great

(Hätten

importance in ecological roles (Seastedt and Crossley 1984, Giller 1996, Lavelle et al.

2006) and potential to be linked with plant communities (Siemann et al. 1998, Shaffers

et al. 2008), to our knowledge they have been rarely addressed in studies concerning

relationships between litter resource diversity and belowground diversity (e.g.,

Armbrecht et al. 2004).

Litter decomposition is a key process to determine soil fertility and quality in

ecosystems; it is governed by interactions among physical parameters (such as climate

and soil proprieties), resource quality and organisms (Swift et al. 1979, Lavelle et al.

1993). Decomposer invertebrates directly affect litter decomposition through litter

fragmentation and modifications of the structure and activity of the microbial

community, which in turn is responsible for most of the carbon and energy flo

schwiler et al. 2005). Chemical properties of leaves, especially nitrogen, lignin

and polyphenols are important predictors for biota action and litter decomposition (e.g.,

Palm and Sanchez 1991, Zhang et al. 2008). Therefore, litter decomposition of different

plant species has the potential to influence each other, by affecting dynamics of litter

mass loss, litter nutrient content and decomposer abundance and activity (Gartner and

Cardon 2004). A pattern is that high quality litters may promote the decomposition of

associated litters, while poor quality litters have negative effects on decomposition of

other litters (Seasted 1984, Quested et al. 2002, Wardle et al. 2003, Gartner and Cardon

2004).

Despite many natural ecosystems have high diversity of plants in the

aboveground community, damaged anthropized systems with monospecific plant

dominance are easily found in silvicultures, constituting single homogeneous litter

layers. If located in infertile conditions, these systems may undergo slow decomposition

rates, in which nutrients are conserved and soil carbon sequestration is promoted

(Wardle et al. 2004). Litter of monocultures, and soil under infertile conditions, may

present soil biodiversity reduced, food webs altered and negative feedbacks in linkages

between aboveground and belowground compartments (Swift and Anderson 1994,

Hansen 2000, Wardle et al. 2004, Haase et al. 2008).

Study

A common assumption about suitable management practices is that goals to

restore and rebuild landscape diversity/heterogeneity within anthropized systems will

help to maintain the biota diversity and preserve a range of ecosystem functions (Isaacs

et al. 2009, Samways 2007). In the present work we added an amount of mix leaves

from native trees on the floor of a post-mining monospecific forest of Pinus elliottii

Engelm (Pinaceae) in Brazil, with goal of enhance soil fauna diversity and the P.

elliottii leaf litter decomposition. Specifically, we experimentally tested the hypothesis

that (1) different native leaves species would create soil substrates with different

structural, chemical, physical and microbial community properties, and that these

differences would in turn increase soil macroarthropod community diversity; and (2) the

decomposition dynamics of P. elliottii litter could be altered both directly by the

interactions with mix native litters and indirectly through the interaction with enhanced

soil biota community.

Method

area

The experiment was performed during December 2007-July 2008 at a monodominate

forest of P. elliottii (approx. 20 ha; 30°09’44. 2”S; 52°00’17. 2”W), in the city of Minas

do Leão, Rio Grande do Sul state, Brazil. The landscape in this region is characterized

by a mosaic of land uses, including coal mining activities and agricultural systems

(specially exotic silvicultures). Native forests mainly remaining along streams, as

riparian vegetation. The climate represents the Cfa type of Köppen-Geigen climate

classification, temperate, with hot summer and precipitation in all months (Peel et al.

2007).

Coal was exploited by opencast mining from the study area around the year 1980

and between 2001-2002. The cave was filled with soil, coal wastes and coal combustion

residues. Whereas revegeted with grasses, spontaneous P. elliottii colonized itself the

land (Fig. 1A), quickly establishing a monodominate forest (personal communication,

Pedro Paulo da Silva Batista- mining work chief of CRM, Companhia Riograndense de

Mineração). Plants from this tree genus are exotic and invasive in the South America,

threatening natural ecosystems and advancing in deforested areas (GISP 2005).

We obtained data of litterfall production of this forest, at the same time of our

experiments. Starting from 12 litterfall collectors (1 m

; mesh 1 X 3 mm), we found an

average production of 2005.3 kg.ha

-1

of P. elliottii leaf-litter in the summer period

(December 2007 to March 2008) and 1531.8 kg.ha

-1

in the autumn period (March to

July 2008).

Experimental design

d sixteen 90 X 60 cm plots on the forest floor in pairs to either a mix leaf-

tment (8 replicates/treatment; Fig. 2). The pairs were distant more

We assigne

litter or control trea

than 100 meters each other, and the plots from a same par were distant two meters each

other. Each plot was composed by nine leaned litter bags (30 X 20 cm length; mesh 10

X 2 mm; Fig. 1B). Control plots presented all litter bags belonging to P. elliottii leaves.

Mix leaf-litter plots had one litter bag belonging to P. elliottii leaves in the center of the

plot, and eight surrounding litter bags with native leaves from four different tree

species. Of these, four litter bags had single species, and four had mix of native species.

We based the experiment on leaves cut from living trees, collected from the study

region. The leaves were placed in litter bags after oven-drying. Each litter bag was filled

with 40

m and 5

X 2.5 c

We sampled the litter bags after three (March - Summer) and six months (July -

of the experiments. At each sampling time, we removed four

± 0.016 g of leaves, and each plot contented around 360 g. Mix litter bags were

standardized by volume basis of the four native species.

The native species were: Cupania vernalis Cambessedes (Sapindaceae), Inga

marginata Willdenow (Fabaceae), Luehea divaricata Martius (Tiliaceae) and Schinus

terebinthifolius Raddi (Anacardiaceae). These species represent common trees in forests

of the study region (Reitz et al. 1983). Leaves of C. vernalis (Fig. 1C) and S.

terebinthifolius (Fig. 1D) are composts, and their leaflets have about 10 x 3.5 c

m, respectively. Leaves of L. divaricata (Fig. 1E) and I. marginata (Fig. 1F) are

generally found completely curly in the forest floor; they have mean of 10.5 X 5 and 7.5

X 2.5 cm, respectively. Leaf nutrient contents (%) of the plants are presented in Table 1.

Sample analyses

Autumn) of the exposure

pairs of plots randomly. Soil macroarthropods (> 2 mm; Swift 1979) were immediately

extracted from litter bags by hands, and using modified Berlese-Tüllgren funnels (6

days).

The macroarthropods were classified in morphospecies. Immature spiders and

insect larvae were not classified in morphospecies. The mass remaining in each litter

bag was determined after oven-drying the leaf-litter (60°C; 72 h). Contents of carbon

and macronutrients (nitrogen, phosphorous, potassium, calcium and magnesium) were

obtained for the P. elliottii litter from the central litter bag of the plots. Carbon content

was determined by moisture combustion/ Walkley-Black (Walkley and Black 1934);

nitrogen was analyzed by the Kjeldahl method (Kjeldahl 1883). Phosphorous and

potassium were assessed by nitric/ perchloric acid digestion followed by determination

on an ICP-OES. Calcium and Magnesium were extracted in KCl mol L

-1

. All analyses

were conducted at the Laboratory of Soil Analysis at UFRGS.

Data analysis

We assessed the effects of mix leaf-litter addition on the macroarthropods diversity by

comparing the plots of both treatments, considering data of all litter bags from the plots.

For the assessment of effects on decomposition and nutrients dynamics of P. elliottii

leaf-litter, we take into account data of the central litter bag inside each plot.

Arthropod’s abundance was standardized to a density measure in each plot:

eight (d.w.) litter. We used analysis of covariance (ANCOVA) in

blocks

MANCOVA yielded significant result, we performed univariate ANCOVAs to

test for difference among each variable separately. Data from these analyses were

individuals.g

-1

dry w

to test effects of treatments in the total macroarthropod density. The sampling

dates (time) were used as covariate factor in analysis. Also, we evaluated the effects of

treatments in the density of the six most abundant macroarthropod groups (> 3% of total

abundance) using multivariate analysis of covariance (MANCOVA) in blocks. In cases

where

square

parately, and the other

maining groups together) was responsive to treatments. Rare species that occurred

were deleted from analyses to improve interpretability of the ordination

root transformed to satisfy parametric assumptions of normality of residuals and

homogeneity of variances.

We compared the morphospecies richness of macroarthropods (considering all

groups together, the two richest groups separately, and the other remaining groups

together) between the leaf-litter treatments using rarefaction curves, with confidence

intervals (95%). We carried out this analyze in order to remove the effects of sample

abundance on morphospecies richness (Gotelli and Colwell 2001, Vasconcelos and

Melo 2008). We use nonmetric multidimensional scaling analyses (NMDS, Kruskal

1964) to ordinate sample units (plots) and assess whether morphospecies composition

(considering all groups together, the two richest groups se

only in one plot

plot (McCune and Grace, 2002). We used the Bray-Curtis distance metric in the data of

species density. To determine whether the treatments were statistically different we used

a two-way analysis of similarity (ANOSIM; Clarke 1993) on a Bray-Curtis measure

matrix. Both rarefaction, NMDS and ANOSIM were performed with PAST 1.9

(Hammer et al. 2001).

To test P. elliottii mass loss in treatments, we used analysis of covariance

(ANCOVA) in blocks. The mass loss of entire plots (considering all litter bags from the

treatments) also was assessed. We evaluated the nutrient content proportions (%), and

the nutrient content loss in P. elliottii litter using multivariate analysis of covariance

(MANCOVA) in blocks.

Results

A total of 2,291 individuals from the orders Araneae, Blattodea, Chilopoda, Coleoptera,

Diplop

ponse to mix leaf-litter input

As we

atment (ANOSIM, R=0.72; P< 0.001) and time of samplings

(ANOSIM, R=0.75; P< 0.001).

oda, Hemiptera, Hymenoptera, Isopoda, Opiliones, Orthoptera and

Pseudoscorpiones were collected (Table 2, Appendix 1). Predators (spiders, centipedes,

and pseudoscorpions) were dominants in number of individuals (49 %) in our

experiment, being 98 % of this represented by spiders. Ants, beetles and harvestman,

that have species with a range of feeding strategies inside the major group, represented

26 % of total individuals. Herbivores, as hemipterans and crickets, were around 14 %;

and groups with detritivorous habits, as millipedes, cockroaches and woodlice had 11 %

of the individuals. A total of 109 morphospecies were identified, being 75 % of these

represented by beetles (45 %) and spiders (30 %; Table 2, Appendix 1). About 58% of

the total species were rare, being 50% singletons (represented only by one individual)

and 8% doubletons (represented only by two individuals). For beetles, 79% were rare

species (67% singletons and 12% doubletons), and for spiders, 53% were rare (47%

singletons and 6% doubletons).

Macroarthropods res

hypothesized, addition of mix native leaves on the forest floor showed positive

effects on the macroarthropod diversity. Mixed leaf-litter plots supported more

individuals (ANCOVA, F

1,7

= 24.48, P< 0.001), and attracted 25 % more morphospecies

(Fig. 3A) than control plots. The morphospecie composition (Fig. 4A) was strongly

responsive to leaf-litter tre

From the eleven classified groups of macroarthropods, ten and seven groups had

is, we decided to analyse the composition data of the

eetles (8 species remaining) together with the other arthropod groups.

od groups analysed together

respectively their total abundances and morphospecies richness increased in at least one

sampling time in the mix leaf-litter plots (Table 2). We found a significant response

concerning the densities of the six most abundant groups (Table 2) in the leaf-litter

treatments (MANCOVA, Pillai Trace= 0.99, F

1,7

= 50.72, P= 0.019). Separated analyzes

yielded significant positive effect of mix litter in spider, cockroach, beetle, millipede,

and ant densities, but did not for hemipteran densities (Table 3).

Spiders had 58 % more species richness in the mix leaf-litter than in control

plots (Fig. 3C), and their morphospecie composition (Fig. 4B) was clearly different in

treatments (ANOSIM, leaf-litter R=0.72, P= 0.001; sampling time R=0.54, P= 0.002).

Rarefied species richness of beetles in the mix leaf-litter plots did not differ from the

control, indicating that the increase in the total species richness (show in the Table 2)

was a sampling artifact resulting from increased sample abundance (Gotelli and Colwell

2001). Mainly due the increased number of species that occurred only in one plot (84%)

and were excluded from analys

The rarefied species richness of the other arthrop

(excepting the most rich orders-spiders and beetles) did not differ between leaf-litter

treatments. On the other hand, their species composition (in this case, only excepting

spiders) was significantly responsive to treatments (Fig. 4B; ANOSIM, leaf-litter

R=0.45, P< 0.001; sampling time R=0.69, P< 0.001).

Pinus

ed by the contact with

mix lea

The ma

such as microhabitat spatial

eterogeneity, and a range of food resources could be attracted diverse organisms to the

ix native litter patches. Our experiment also shows that, unlike the expected, input of

ative mix leaf-litter did not change P. elliottii litter decomposition rates. On the other

elliottii leaf-litter response to mix leaf-litter

Pinus elliottii needles have lower litter quality compared to the native broad leaves

(Table 1). Thus, mass loss of entire plots (all litter bags per plot analyzed together) was

very high in the mix leaf-litter plot (F

1,7

= 135.05, P<0.001; Fig. 5A) compared to

control plots.

Not as expected, P. elliottii mass loss (F

1,7

= 0.51, P= 0.50; Fig. 5B) and nutrients

loss (Pillai Trace= 0.84, F

1,7

= 1.83, P= 0.39) were not influenc

f-litter. However, the proportions of nutrients content (%) exhibited responses of

treatments (Pillai Trace= 0.98, F

1,7

= 17.56, P= 0.05). Univariate analysis reveled

significant differences in K (%) and P (%) and marginally differences in C (%) (Table

4; Fig. 6). Potassium and P concentrations were higher in P. elliottii leaf-litter when in

contact with mix leaf-litter. On the other hand, C concentrations seemed to be increased

in the control plots.

Discussion

in conclusion of this paper is that the input of diverse resources upon a damaged

soil in a monospecific forest favored an increase in soil arthropod diversity and

diferential species composition. Our experimental technique strongly influenced

diversity of many groups such as spiders, cockroachs, beetles, harvestman, millipedes,

ants and woodlice. In particular, characteristics

hand, our strategy altered P. elliottii litter nutrients content, probably trough nutrient

er experimental studies with resource manipulation (Chen and Wise 1999,

Leroy

habitat space, and then can offer more suitable shelters against

arsh climatic conditions and predators, microsites for foraging, mating, oviposition,

ay, all trophic levels of soil fauna could be influenced positively by the

crease in microhabitat complexity (Stevenson and Dindal 1982, Kaneko and

transferation or differential microbial activity.

Macroarthropods response to mix leaf-litter input

As oth

et al. 2007), we have overlap in the factors (micromicrohabitat and food

enhancement) enriching soil fauna diversity in mixed litter plots. Concerning

microhabitat enhancement, it is know that a diverse litter layer can support higher

spatial structure heterogeneity, through different leaf-sizes, leaf shapes and leaf-surface

structures than monospecific litter layers (Hättenschwiler et al. 2005). This prediction is

yet more convincing when we compare the structure of needle litter of P. elliottii with

the native broadleaf litters studied. Uetz (1974) related that more spatial structured litter

layers produce more

etc. In this w

Salamanca 1999, Hansen 2000, Tews et al. 2004, Lassau et al. 2005). On the other hand,

habitat preferences also reflect foraging habits. Bottom-up controls propagated through

trophic levels are very common in detritus food webs, and are related in experiments

excluding enhancement in microhabitats (Scheu and Schaefer 1998). The input of mix

leaves, as a role of food with diverse qualities and chemical compositions, may lead to

diversity of detritivores through food selectivity and resource partitioning (Hooper et al.

2000, Wardle 2006). Perhaps, the microenvironment created by the mix of different

species of trees influenced in a differentiate water retention and decomposition, favoring

growth of different types of fungi and bacteria (Armbrecht et al. 2004, Hättenschwiler et

al. 2005) which conceivably constitute part of a diet of detritivores. Still, diversity of

redators may be responded to diversity of preys (Seetle et al. 1996, Chen and Wise p

1999, Scheu and Schaefer 1998, Miyashita et al. 2003).

Interaction between species through top-down controls in the soil food web can

limit the influence of aboveground diversity in the belowground species (Wardle and

Yeates 1993, Hooper et al. 2000). Spiders were about 48 % of the total macroarthropod

individuals found, and they had a significant enhanced density in mixed leaf-litter plots.

This result reflects a consistent pattern from literature, in which spiders tends to increase

their abundances in a more intensive way then did other groups in front of increased

habitat complexity (reviewed in Langellotto and Denno 2004). Structurally rich habitats

can diminish antagonistic interactions among predators, such as intraguild predation,

and have effective implications to prey suppression (Finke and Denno 2002, Langellotto

and Denno 2006, Sanders et al. 2008). Thus, the increased densities of spiders may lead

to negative effects in diversity of other invertebrate groups (their preys; Wise et al,

1999), for example in the case of hemipterans density and beetles richness, therefore

hiding some positive correlations between the preys and the enhancement of mix litter.

Pinus elliottii leaf-litter response to mix leaf-litter

Despite many works demonstrated additive effects of mixes in decomposition rates of

individual litters (Chadwick 1998, Gartner and Cardon 2004), we did not find direct

effects on the litter mass and nutrients loss of P. elliottii. Indirectly, we show significant

changes in its chemical composition, related to differential concentrations of

phosphorous, potassium and carbon. We identify three plausible explanations. First, as

nutrient concentrations usually increase in the lower quality while decreasing in the

higher quality litter thought nutrient transfer (Wardle et al. 1997, Gartner and Cardon

2004), the greater concentrations of P and K may had leached from the surrounding

high quality native litters. Second, in view of the increased diversity of soil

macroa

xperiment we still had more than 50% of initial remaining leaf-litter in the plots.

ophic cascades induced by

rthropods in the more diverse litter layer, we suppose that a greater production of

diverse faecal materials may had occurred in this treatment. As assimilation efficiency

is low in the food digestion by soil organisms (Wolters 2000), increased nutrients may

be coming from the animal’s defecation. Third, as soil carbon and energy flow is mainly

driven by microbial activities (Coleman et al. 2004), which are substantially influenced

by the two previous topics, the changed chemical composition of P. elliottii in mix litter

treatment may be reflecting the action of differential microbial communities. Infertile

soils tend to support fungal-based energy channel (Wardle et al. 2004). Increased

potassium and decreased carbon concentrations may be related to greater fungal

activities and respiration (Coleman et al. 2004, Sayer 2006).

Then, why we did not find major mass loss in P. elliottii litter when surrounding

by heterogeneous native litter? One plausible possibility is that our experimental time

was short to find an effect in the decomposition rates, because in the end of the

Another speculative reason is the existence of top-down tr

predators (Wise et al 1999). Spiders are evolved in decomposition food webs and have

the potential to generate positive or negative impacts on rates of litter mineralization.

Wise et al. (1999) hypothesized that in a four-level food chain, an increase in spiders

should lead to a decrease in detritivores/fungivores (e.g. Collembola). Through relieve

in the predation pressure in fungi, the nutrients may became immobile in senescent

fungal hyphae, not entering in the soil.

Ecological implications

We find that the input of diverse resources on a damaged monospecific system

can improve soil macroarthropod biodiversity and alter the dynamic of an ecosystem

process. These results certainly have great implications in conservation biology and

restoration ecology. Isaacs et al. (2009), attempting to promote conservation of

beneficial aboveground arthropods in agricultural landscapes, show that the increase in

native plants provides local adaptation, habitat permanency, and support of native

biodiversity, therefore maximizing arthropod-mediated ecosystem services, such as crop

pollination and pest control. Here, we show that the input of native leaves on the floor

of an exotic monospecific forest benefits belowground arthropods, probably

invigorating biotic and abiotic relationships and also maximizing soil arthropod-

mediated ecosystem services (Lavelle et al. 2006). Moreover, the technique of

incorporate diverse organic matter in poor anthropized systems have clear benefits to

the soil nutrient enrichment, and consequently, to primary productivity (Neher 1999).

Finally, based in our data and taking into account the increased rates of habitat

degradation and changes in biotic diversity and ecosystem properties around the world,

we suggest two aspects related to ecological restoration practices. First, it should to

maintain quality landscape heterogeneity, simulating condictions of natural ecosystems

(Samways 2007, Podgaiski 2007) to benefits fauna diversity permanency. Second, it

should to seek soil fertility maintenace, which will in turn have positive consequences

for above–belowground linkages and their effects on natural communities (Haase et al.

2008).

We tha

Armbrecht, I., I. Perfecto, J. Vandermeer. 2004. Enigmatic biodiversity correlations: ant

diversity responds to diverse resources. Science 304:284-286.

Clarke, K. R. 1993. Non-parametric multivariate analyses of changes in community

structure. Australian Journal of Ecology 18: 117–143.

Acknowledgements

nk taxonomists that help in arthropod’s identifications (M.A.L. Marques, E.H.

Buckup and E.N.L. Rodrigues –FZBRS, Araneae; A.F. Quadros -UFRGS, Isopoda; L.

Menzel -UFRGS; Formicidae); field and lab assistants (A. Panatta, A.L. Casara and

V.G. Sydow); Companhia Rio-Grandense de Mineração (CRM) for allow access to the

study site and its employees for help in the fieldwork. L.R. Podgaiski received a

scholarship from CAPES (Brazil).

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Tables

Table 1. Initial leaf nutrient content (%) of the plants.

Plants C N P K Mg Ca

Pinus elliottii

44.75 1.02 0.09 0.32 0.18 0.52

Inga m

Luehe

2.4 0.20 0.93 0.28 0.66

Schinus terebinthifolius

43 1.2 0.12 0.65 0.36 1.8

pania vernalis

47 2.8 0.29 1.7 0.24 0.42

arginata

44 3.7 0.24 0.87 0.15 0.42

a divaricata

109

Table 2. Total abundance and morphospecies richness (in brackets) of soil

acroarthropods in mix leaf-litter plots and control plots in two sampling times.

Note: Each column represents diversity values from four plots.

Mix leaf-litter plot Control plot

Class Order

3 months 6 months 3 months 6 months

l Tota

Araneae 6(16 538(2 6 2 ) 1 3) 29 ) 4) 3(3) 13(11 110(3

rachnida

Pseud s 0 0 1(1)

rustacea Isopoda ) 0 0 21(1)

Opiliones 0 23(1) 0 1(1) 24(1)

oscorpione 1(1) 0

Chilopoda 0 12(2) 0 6(2) 18(2)

C 8(1) 13(1

Diplopoda 8(1) 128(2) 0 16(2) 152(2)

Blattodea 29(3) 46(2) 3(1) 3(1) 81(3)

Coleoptera 13(10) 72(31) 6(5) 29(13) 120(49)

Hemiptera 6(3) 187(4) 3(1) 118(2) 314(5)

Hymenoptera 58(5) 271(3) 32(6) 88(3) 449(11)

Hexapoda

Orthoptera 0 0 0 1(1) 1(1)

Total 419(40) 1290(70) 107(16) 475(36) 2291(109)

110

Table 3. F and P-values (in brackets) from ANCOVA in block results for square root

transformed densities (individuals.g-1 d.w. litter) of five macroarthropod groups.

Response variable Treat cking (df= e (df=1,7) ment (df=1,7) Blo 6,7) Tim

Araneae

5 01

.51

002)

4.93 (<0.0 )

3 (0.06)

21.79 (0.

Note: Sources of variation are: treatment (mix leaf-litter and control plots), blocks

(n=8), t plings (3 and 6 mo ll stati y sign num

(P<0.05) are in boldface.

ime of sam nths). A sticall ificant bers

Blattodea

20.49 (0.003)

0.65 (0.689) 0.42 (0.537)

Coleoptera

8.12 (0.024)

0.85 (0.567)

16.26 (0.005)

Diplopo

6.24 (0.041)

1.25 (0.385)

10.19 (0.015)

Hemiptera 1.25 (0.301) 1.49 (0.304)

24.02 (0.002)

Hymenoptera

5.73 (0.048)

2.80 (0.102)

8.89 (0.020)

111

Table 4. F and P-values (in brackets) from ANCOVA in block results for nutrients

content (%) of P. elliottii leaf-litter.

R Trea l Time (df=1,7) esponse variable tment (df=1,7) B ocking (df=6,7)

(%) 9 (0.684) 5 (0.264)

11.487 (0.012)

(%)

6 (0.025) 31 (0.029)

(%)

7 (0.015)

00 (0.332)

a (%) (0.918) 02 (0.063)

g (%) (0.785) 03 (0.077) 0.438 (0.529)

(%) (0.079) 96 (0.367) 1.296 (0.292)

N 0.17 1.64

7.98 4.8 16.662 (0.005)

10.28

1.4

32.200 (<0.001)

C 0.011 3.5

36.505 (<0.001)

M 0.080 3.2

C 4.200 1.2

Note: Sources of variation are: treatment (mixed leaf-litter and control plots), blocks (n=

8), time of samplings (3 and 6 months). All statistically significant numbers (P< 0.05)

are in boldface.

112

IGURES

Figure 1. Plantule of Pinus elliottii (A) growing up in front of a land with mining

activities in the study region, and experimental plot (B) in the post-mining P. elliotti

forest. Leaves of trees: Cupania vernalis (C), Schinus terebinthifolius (D), Luehea

divaricata (E) and Inga marginata (F) showed in the same scale.

113

Figure 2. Experimental design testing the effects of enhancement of leaves from native

trees on the floor of a post-mining P. elliottii forest in the soil macroarthropod diversity

and in the P. elliottii litter decomposition.

114

C) Araneae

0 150 300

B) Coleoptera

Number of individuals

04590

Number of species

A) Total arthropods

0 275 550 825 1100

Number of species

100

Mix leaf-litter plot

Control plot

D) Other arthropods

0400800

Figure 3. Rarefaction curves for species richness of total soil arthropod abundances (A),

Coleoptera (B), Araneae (C) and other arthropods (D) in relation to effects of mix leaf-

litter and control plot treatments. Error bars represent ± 1 conficence interval (CI).

115

Figure 4: NMDS ordinations of mix leaf-litter (Mix) and control plots from two sample

dates (3 and 6 months) in the first two principal axes, considering total soil arthropods

(A), Araneae (B) and other arthropods (C).

116

Figure 5: Leaf-litter mass loss of entire plots (A), and single litter bags of P. elliottii

from the center of the plots.

117

Figure 6: Nutrients concentrations (%) dynamics in of P. elliottii leaf litter in mix leaf-

litter and control plots.

118

Appendix 1: Morphospecies composition and abundance of soil macroarthropods in mix

leaf-litter and control plots in 3 and 6 months of exposure.

Mix leaf-litter plot Control plot

Class / Order Family / Specie or Morphospecie

3 months 6 months 3 months 6 months

Arachnida

Araneae Amphinectidae

Immature 0 4 0 0

Araneidae

Immature 0 1 0 2

Caponiidae

Nops meridionalis Keyserling, 1891 6 4 0 0

Immature 3 5 0 0

Corinnidae

Meriola mauryi Platnick & Ewing, 1995 5 0 0 0

Orthobula sp.1 21 7 0 3

Orthobula sp.2 0 1 0 0

Orthobula sp.3 0 1 0 0

Orthobula sp.4 0 1 0 0

Immature 8 0 0 0

Ctenidae

Isoctenus sp. 0 1 0 0

Immature 9 20 1 4

Gnaphosidae

Apopyllus silvestrii (Simon, 1905) 0 1 0 0

Gnaphosidae sp. 1 0 0 0

Immature 23 23 0 3

Hahniidae

Hahniidae sp.1 2 30 0 10

Hahniidae sp.2 0 1 0 0

Immature 7 35 4 11

Linyphiidae

Lygarina sp. 0 55 0 42

sp. 0 1 0 0

Neomaso arundicolaMillidge, 1991 1 0 0 0

Neomaso damocles Miller, 2007 0 15 0 10

Neomaso sp.1 0 1 0 0

Neomaso sp.2 0 0 0 1

Scolecura cambara Rodrigues, 2005 1 1 0 5

Smermisia vicosana (Bishop & Crosby, 1938) 2 3 1 6

Sphecozone sp. 1 5 0 1

Immature 22 86 1 38

Lycosidae

Immature 0 3 1 0

Migalomorphae

Immature 1 0 0 0

Mysmenidae

Itapua sp. 0 2 0 0

Immature 0 2 0 1

Oecobidae

Oecobius navus Blackwall, 1859 0 0 1 0

Immature 1 0 1 0

Erigone sp. 0 2 0 0

Gigapassus octarine Miller, 2007 1 5 0 0

Mermessus

Oonopinae sp. 4 37 0 2

15 4 1

Immature 0 1 0 0

Salticidae

Aphirape sp.

Salticidae sp.1 3 0 0 0

Salticidae sp.2

54 98 10 23

Scytodiidae

7 1 0

Theridiidae

p.1

Opiliones

Pseudoscorpiones sp.

hilopoda

Oonopidae

Immature 4

Oxyopidae

0 1

1 2

0 0

Breda sp.

Unidentatae sp. 0 0 0 1

1 0 0 0

Immature

Immature 0

Thymoites s 4 1 0 0

Thymoites sp.2 44 16 23 23

Immature 59 49 14 25

Thomisidae

Immature 0 1 1

Opiliones sp. 0 23 0 1

Pseudoscorpiones 1 0 0 0

C Chilopoda sp.1 0 11 0 4

Chilopoda sp.2 0 1 0 2

Crustacea

Isopoda e

ridana (van Name, 1940)

iplopoda .1

Philoscida

Atlantoscia flo 8 13 0 0

D Diplopoda sp 8 99 0 9

Diplopoda sp.2 29

exapoda

0 0 7

Blatodea

Coleoptera

p.2

0 0

1 0

0 0

Scarabaeidae

sp.

Staphilinidae

e sp.5

ae sp.6

ae sp.9

Blatodea sp.

1 20

0 Blatodea sp

Blatodea sp.3 1 0 0 0

Carabidae

sp.1

Carabidae 0

0 Carabidae s

Crisomelidae

Crisomelidae sp.

Curculionid

Curculionidae sp 0 0 0 2

Curculionidae sp.2 1

0 Curculionidae sp.3

Disticidae

Disticidae sp. 0

Nitidulidae

Nitidulidae sp.1 0 1 0 0

Nitidulidae sp.2 1 0

0 Scarabaeidae 0 1 0

0 Staphilinidae sp

ae sp Staphilinid 0

0 Staphilinidae sp.3

ae sp.4 Staphilinid

0 1 0 0

Staphilinid

0 Staphilini

Staphilinidae sp.7

ae sp.8

0 0 0 1

Staphilinid

Continuação Appendix 1- Artigo 3 120

Staphilinidae sp.10

.11

ae sp.12

ae sp.13

ae sp.14

dae sp.15

ae sp.1

ae sp.4

oleoptera sp.13

.14

oleoptera sp.18

Hemiptera 178 115

.3 2

Hymenoptera

ufipes (Fabricius, 1775)

llax (Mayr, 1870) 2

.2 10

rhoptrum) sp.

victa Buren, 1972 41 57 77

Orthoptera 0

OTAL 4 1290 108 474

0 2 0 0

Staphilinidae sp 0 1 0 0

Staphilinid 0 2 0 0

Staphilinid 0 1 0 0

Staphilini 0 0 1 0

Pselaphineae

Pselaphine 0 1 0 0

Pselaphineae sp.2 0 1 0 0

Pselaphineae sp 0 1 0 0

Pselaphineae sp 0 4 0 0

Pselaphine 0 0 0 1

Coleoptera sp 0 0 0 1

Coleoptera sp 0 2 0 1

Coleoptera sp 0 0 0 1

Coleoptera sp.4 1 0 0

Coleoptera sp.5 0 1 0 0

Coleoptera sp 0 1 0 0

Coleoptera sp 0 4 0 3

Coleoptera sp 2 2 0 4

Coleoptera sp.9 0 23 0 4

Coleoptera sp.10 0 0 0 1

Coleoptera sp.11 0 1 0 0

Coleoptera sp.12 0 5 0 8

C 1 0 0 0

Coleoptera sp 1 0 0 0

Coleoptera sp.15 1 0 0 0

Coleoptera sp.16 2 0 1 0

Coleoptera sp.17 0 1 2 1

C 1 0 0 0

Coleoptera sp.19 2 0 0 0

Hemiptera sp.1 0 0

Hemiptera sp.2 1 6 0 3

Hemiptera sp 2 0 0

Hemiptera sp.4 0 1 0 0

Hemiptera sp.5 3 3 0

Formicidae

Formicinae

Camponotus r 4 0 0 0

Camponotus sp. 0 0 6 0

Myrmicinae

Crematogaster sp. 0 0 1 0

Pheidole fa 2 05 0 0

Pheidole sp.1 7 0 0 0

Pheidole sp 0 0 6

Pheidole sp.3 0 0 1 0

Solenopsis (Diplo 4 9 0 0

Solenopsis in 3

Solenopsis sp.1 0 0 13 0

Solenopsis sp 0 0 1 0

Solenopsis sp.3 0 0 1 0

Ponerinae

Pachycondyla sp. 0 0 1 0

Orthoptera sp. 0 0 1

T 19

Continuação Appendix 1- Artigo 3 121

CONSIDERA

Estudos ecológicos sobre decomposição foliar de plantas pioneiras e a

macrofauna de solo associada em ecossistemas degradados pela deposição de

cinzas e extração do carvão a céu aberto foram elaborados e realizados em

curto prazo. Apesar de vários estudos terem sido realizados em ambientes

extremamen na Europa e outras regiões do mundo (e.g., Rodrigu

2001; McEnroe et al 2001; Majer et al 2007), abordagens similares a es

estudos, relacionando estas variáveis ecológicas em eco istemas xtrem

ainda são in Brasil. Há carência de informações tradicionais sobre

o funcionam stemas degradados, o que prejudica a tomada de

decisões acerca do manejo, restauração e conservação da biodiversidade

destes sistem

Em São Jerônimo (RS), o local exato onde cinzas de carvão fóssil (cinz

leves e cinzas grossas) foram despejadas irregularmente durante longos anos

constitui área integrante da mata ripária do Rio Jacuí, que por lei (Código

Florestal, nº 4771, de 1967) é considerada área de preservação permanente

(APP). De acordo com a resolução do CONAMA nº 303 (2002), a função

ambiental das APPs é de preservar os recursos hídricos, a paisagem, a

estabilidade a biodiversidade, o fluxo gênico de fauna e flora,

proteger o s r o bem estar das po lações humanas. Com base

nestas funções essenciais, dever-se-ia fazer o possível para manter e

preservar as características originais destas áreas, não subjugando nenhum

ÇÕES INAISF

te poluídos es

tes

ss e os,

cipientes no

ento dos si

as.

geológica,

olo e assegura pu

122

uso antrópico degradante a elas. As cinzas de carvão fóssil produzidas na

Usina Termelétrica de São Jerônimo (CGTE – Companhia de Geração Térmica

de Energia Elétrica) são classificadas como “resíduos sólidos– não inertes”

(NBR 10004), e desta forma, seu despejo em áreas naturais deveria ser

altamente restrito, uma vez que contêem elementos com propriedades

químicas e toxicológicas que podem causar danos à saúde e ao ambiente

(Rohde et al. 2006). Sua disposição no solo (NBR 8419), deveria ser

necessariamente em um aterro sanitário, local em que apresentaria mínimos

impactos ambientais. Segundo a constituição Federal (1998), compete ao

gerador de resíduos sólidos a sua inteira responsabilidade sobre eles,

compreendendo desde as etapas de acondicionamento, disponibilização para

coleta, coleta, tratamento, até sua disposição final ambientalmente

adequada. Desta forma, a CGTE, como produtora de resíduos sólidos

altamente poluidores (cinzas), e agente de disposição imprópria dos resíduos

em APP (mata ripária), apresentou graves irregularidades com a lei.

Esta mata ripária em São Jerônimo, que felizmente não vem mais

recebendo os depósitos de cinzas de carvão, está sofrendo o importante

processo ecológico da sucessão natural a partir de plantas nucleadoras

(Azzolini, 2008). A partir do nosso estudo, foi constatado que a mamona (R.

communis) –abundante planta pioneira na área - apresentou decomposição

foliar muito mais rápida do que outras plantas pioneiras avaliadas (capim-

bermuda C. dactylon e aroeira-vermelha S. terebinthifolius). Isto significa que

esta espécie está disponibilizando nutrientes mais rapidamente ao substrato

(cinzas), e desta forma, pode estar contribuindo ao melhoramento das

123

condições bióticas locais de uma maneira mais eficiente do que as outras

espécies. No entanto, é importante reconhecer o papel de cada espécie

dentro da sucessão ecológica na área (Azzolini, 2008). Apesar das três

espécies avaliadas apresentarem caráter pioneiro, elas participam em

momentos distintos do processo de sucessão, apresentando uma certa ordem

na qual ocorrem facilitações e desfavorecimentos, e assim devem ser julgadas

com importâncias singulares na transformação do ambiente degradado. A

partir dos resultados de Azzolini (2008), podemos supor que as duas espécies

exóticas avaliadas (mamona e capim-bermuda) alterem as condições do

substrato (cinzas), beneficiando elas mesmas em um primeiro momento, mas

também as espécies nativas que se estabelecem no andamento da sucessão,

como a aroeira-vermelha. Nesta área, percebe-se o processo de restauração

ecológica ocorrendo naturalmente.

O folhiço das três espécies de plantas pioneiras (mamona, capim-

bermuda e aroeira-vermelha) avaliadas no estudo de São Jerônimo foram

largamente colonizados pela macrofauna do solo nos experimentos, o que

demonstra claramente a importância desta camada orgânica para os

organismos. Os folhiços da aroeira-vermelha e do capim-bermuda foram

similares com relação à diversidade de espécies que suportaram. A mamona,

por sua vez, apesar de ter apresentado maiores densidades de indivíduos,

apresentou a menor colonização por diferentes espécies, visto a sua rápida

desintegração foliar.

Embora a decomposição de folhiço não tenha sido influenciada por

diferentes depósitos de cinzas (cinzas leves X cinzas grossas), a diversidade de

124

invertebrados carnívoros, a ocorrência de A. floridana (Isopoda), a

abundância e fecundidade de fêmeas de B. taeniata (Isopoda), e as

composições totais de espécies da macrofauna foram fortemente afetadas

pelos depósitos. A estrutura física do substrato, o pH não favorável, as

diferentes concentrações de metais pesados ou até mesmo características

peculiares de cada área, como a composição da vegetação, podem estar

influenciando e selecionando as diferentes espécies nos ambientes. Assim

como outros experimentos naturais realizados em campo, nosso estudo tem

limitações em apontar os mecanismos estritamente responsáveis pelos

padrões encontrados. Seriam necessários estudos manipulativos - que

padronizassem condições físicas/químicas – mais análises químicas e réplicas

amostrais para melhor compreender os efeitos dos tratamentos sobre a

decomposição foliar e a macrofauna de solo. Mas, em geral, apesar da não

replica

do que vem ocorrendo nos depósitos de cinzas.

Benefí

ção do fator cinza no experimento, parece que as cinzas grossas

apresentam mais fatores condicionantes e limitaram mais a macrofauna, o

que poderia ser levado em consideração na recuperação destas áreas.

Com relação à área de estudo em Minas do Leão (RS), que teve seu solo

minerado e reconstruído topograficamente após a extração do carvão, pode se

considerar que apresenta uma série de desequilíbrios nos atributos químicos,

físicos e biológicos do seu solo (Kämpf et al. 2000). A colonização espontânea

da exótica P. elliottii neste sistema pós-minerado não permite sucessão

natural, ao contrário

cios sobre o ambiente pretérito, sem dúvida, são intrínsecos ao

estabelecimento desta floresta monodominante de pinus, mas intervenções de

125

manejadores e restauradores, neste caso, são bem vindas em ordem a

estimular processos carentes e incrementar a biodiversidade.

Tendo isto em vista, a adição de folhiço de plantas nativas sobre as

camadas homogêneas de acículas de pinus no solo da floresta de pinus

demonstrou ser uma prática de manejo viável, uma vez que modificou a

composição química do folhiço de pinus, representando uma diferenciação na

ciclagem de nutrientes. Esta prática também enriqueceu a fauna de

artrópodes de solo nesta área, provavelmente pelo aumento na

heterogeneidade de recursos oferecidos a fauna, aumento de espaço de

hábitat e qualidade de alimento.

Com base nos resultados deste estudo, e certamente em um forte

embasamento da literatura (ver artigo 3), nós pudemos propor dois aspectos a

serem levados em conta ao sucesso de práticas de restauração ecológica:

1) almejar a manutenção da heterogeneidade estrutural da

paisagem, simulando as condições de ecossistemas naturais, e

2) auxiliar na manutenção da fertilidade do solo, a qual trará

positivas conseqüências a todas os outros elementos do

sistema.

Com relação a macrofauna encontrada nestes dois projetos de pesquisa

pertencentes à dissertação, observou-se a existência de um grande número de

indivíduos e espécies nas áreas degradadas estudadas. Duas extensas listas de

espécies são apresentadas, e constituem informação importante ao inventário

de biodiversidade do Rio Grande do Sul. Tatuzinhos (Isopoda) e aranhas

126

(Araneae) foram os dois grupos mais representativos em indivíduos nos

depósitos de cinzas e na floresta de pinus pós-minerada, respectivamente.

Besouros (Coleoptera) foram os mais ricos em espécies nas duas áreas.

Visto todas estas questões na presente dissertação, esperamos estar

contribuindo um pouco mais ao entendimento de processos ecossistêmicos e a

interação dos organismos do solo em áreas degradadas pelo carvão no sul do

Brasil, principalmente no Rio Grande do Sul. A decomposição foliar e a

interação fauna – serapilheira servem como ferramentas e podem auxiliar em

estratégias e o desenvolvimento prático-teórico da ecologia da restauração.

127

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Winter, M., Gaskell, P. 1998. The Agenda 2000 debate and CAP reform in Great Britain. Is th

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Rasmussen, L., Wright,

European environmental policy, 1998. In: Rasmussen, L., Wright, R.F., (Eds.), The Whole

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101, 353-363.

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