RESUMEN

Fluctuating asymmetry, defined as random differences between the two sides of a symmetrical structure, has been often related to development stress in both plants and animals. In plants, leaf fluctuating asymmetry has been related to stresses such as pollution and fire and may also be related to leaf growth and herbivory rates. We assessed whether leaf fluctuating asymmetry is related to plant and leaf size in Miconia albicans (Sw.) Triana (Melastomataceae), a common multi-stem Neotropical shrub, in a Brazilian savanna area. We collected 15 leaves from each of 70 individuals, and measured fluctuating asymmetry as the difference in area between the right and left sides of the leaves using the central vein as reference. To avoid spurious results due to measurement error, the division along the central vein was performed independently by three researchers. We also measured the basal area and height of each stem of the plant individuals. We used linear models to assess the relations between leaf fluctuating asymmetry, plant size and leaf size. No consistent relations were observed between leaf fluctuating asymmetry and plant size, as the analyses performed on the fluctuating asymmetry values obtained by the different researchers showed different results. However, relative fluctuating asymmetry values, obtained by dividing the fluctuating asymmetry by the total leaf area, tended to be smaller in larger leaves. It thus appears that, in the study species, fluctuating asymmetry is related to the developmental conditions faced by the individual leaves and not by the plant as a whole.

A assimetria flutuante, definida como diferenças aleatórias entre os dois lados de uma estrutura simétrica, tem sido frequentemente relacionada ao estresse de desenvolvimento em plantas e animais. Nas plantas, a assimetria flutuante foliar tem sido relacionada a estresses como poluição e fogo e também pode estar relacionada ao crescimento foliar e taxas de herbivoria. Nós avaliamos se a assimetria flutuante foliar está relacionada ao tamanho da planta e da folha em Miconia albicans (Sw.) Triana (Melastomataceae), um arbusto neotropical multicaule comum, em uma área de Cerrado. Coletamos 15 folhas de cada um dos 70 indivíduos e medimos a assimetria flutuante como a diferença de área entre os lados direito e esquerdo das folhas usando a nervura central como referência. Para evitar resultados espúrios devido ao erro de medição, a separação ao longo da nervura centra foi feita independentemente por três pesquisadoras/es. Também medimos a área basal e a altura de cada caule de cada planta. Usamos modelos lineares para avaliar as relações entre assimetria flutuante foliar, tamanho da planta e tamanho da folha. Não foram observadas relações consistentes entre a assimetria flutuante da folha e o tamanho da planta, pois as análises realizadas nos valores da assimetria flutuante obtidos pelas/os diferentes pesquisadoras/es mostraram resultados diferentes. No entanto, os valores de assimetria flutuante relativa, obtidos pela divisão da assimetria flutuante pela área foliar total, tenderam a ser menores nas folhas maiores. Assim, verifica-se que, na espécie estudada, a assimetria flutuante está relacionada às condições de desenvolvimento enfrentadas pelas folhas individuais e não pela planta como um todo.

Asunto(s)

RESUMEN

Fluctuating asymmetry, defined as random differences between the two sides of a symmetrical structure, has been often related to development stress in both plants and animals. In plants, leaf fluctuating asymmetry has been related to stresses such as pollution and fire and may also be related to leaf growth and herbivory rates. We assessed whether leaf fluctuating asymmetry is related to plant and leaf size in Miconia albicans (Sw.) Triana (Melastomataceae), a common multi-stem Neotropical shrub, in a Brazilian savanna area. We collected 15 leaves from each of 70 individuals, and measured fluctuating asymmetry as the difference in area between the right and left sides of the leaves using the central vein as reference. To avoid spurious results due to measurement error, the division along the central vein was performed independently by three researchers. We also measured the basal area and height of each stem of the plant individuals. We used linear models to assess the relations between leaf fluctuating asymmetry, plant size and leaf size. No consistent relations were observed between leaf fluctuating asymmetry and plant size, as the analyses performed on the fluctuating asymmetry values obtained by the different researchers showed different results. However, relative fluctuating asymmetry values, obtained by dividing the fluctuating asymmetry by the total leaf area, tended to be smaller in larger leaves. It thus appears that, in the study species, fluctuating asymmetry is related to the developmental conditions faced by the individual leaves and not by the plant as a whole.

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Abstract Fire is a recurrent disturbance in savanna vegetation and savanna species are adapted to it. Even so, fire may affect various aspects of plant ecology, including phenology. We studied the effects of a spatially heterogeneous fire on the reproductive phenology of two dominant woody plant species, Miconia albicans (Melastomataceae) and Schefflera vinosa (Araliaceae), in a savanna area in South-eastern Brazil. The study site was partially burnt by a dry-season accidental fire in August 2006, and we monitored the phenolology of 30 burnt and 30 unburnt individuals of each species between September 2007 and September 2008. We used restricted randomizations to assess phenological differences between the burnt and unburnt individuals. Fire had negative effects on the phenology of M. albicans, with a smaller production of reproductive structures in general and of floral buds, total fruits, and ripe fruits in burnt plants. All unburnt but only 16% of the burnt M. albicans plants produced ripe fruits during the study. Fire effects on S. vinosa were smaller, but there was a greater production of floral buds and fruits (but not ripe fruits) by burnt plants; approximately 90% of the individuals of S. vinosa produced ripe fruits during the study, regardless of having been burnt or not. The differences between the two species may be related to S. vinosa's faster growth and absence from the seed bank at the study site, whereas M. albicans grows more slowly and is dominant in the seed bank.

Resumo O fogo é uma perturbação recorrente em vegetação savânica e as espécies do cerrado são adaptadas a ele. Mesmo assim, o fogo pode afetar aspectos da ecologia vegetal, incluindo a fenologia. Nós estudamos os efeitos de um incêndio espacialmente heterogêneo sobre a fenologia reprodutiva de duas espécies lenhosas dominantes, Miconia albicans (Melastomataceae) e Schefflera vinosa (Araliaceae), em uma área de cerrado no Sudeste do Brasil. A área de estudo foi parcialmente queimada por um incêndio acidental na estação seca (agosto) de 2006. Nós acompanhamos a fenologia reprodutiva de 30 indivíduos queimados e 30 não-queimados de cada espécie, de setembro de 2007 a setembro de 2008, usando aleatorizações restritas para comparar os dois grupos. Em M. albicans, a produção de estruturas reprodutivas como um todo e de botões florais, frutos e frutos maduros foi maior em indivíduos queimados do que nos não-queimados. Todos os indivíduos não-queimados, mas apenas 16% dos queimados, produziram frutos maduros durante o estudo. Já em S. vinosa, a produção de botões florais e de frutos (mas não de frutos maduros) foi maior em indivíduos queimados; aproximadamente 90% dos indivíduos de S. vinosa produziram frutos maduros durante o estudo, independentemente de terem sido queimados ou não. As diferenças entre as duas espécies podem estar relacionadas com o fato de S. vinosa crescer mais rápido e estar ausente do banco de sementes, ao contrário de M. albicans, de crescimento mais lento e dominante no banco de sementes.

Asunto(s)

RESUMEN

Fire is a recurrent disturbance in savanna vegetation and savanna species are adapted to it. Even so, fire may affect various aspects of plant ecology, including phenology. We studied the effects of a spatially heterogeneous fire on the reproductive phenology of two dominant woody plant species, Miconia albicans (Melastomataceae) and Schefflera vinosa (Araliaceae), in a savanna area in South-eastern Brazil. The study site was partially burnt by a dry-season accidental fire in August 2006, and we monitored the phenolology of 30 burnt and 30 unburnt individuals of each species between September 2007 and September 2008. We used restricted randomizations to assess phenological differences between the burnt and unburnt individuals. Fire had negative effects on the phenology of M. albicans, with a smaller production of reproductive structures in general and of floral buds, total fruits, and ripe fruits in burnt plants. All unburnt but only 16% of the burnt M. albicans plants produced ripe fruits during the study. Fire effects on S. vinosa were smaller, but there was a greater production of floral buds and fruits (but not ripe fruits) by burnt plants; approximately 90% of the individuals of S. vinosa produced ripe fruits during the study, regardless of having been burnt or not. The differences between the two species may be related to S. vinosa's faster growth and absence from the seed bank at the study site, whereas M. albicans grows more slowly and is dominant in the seed bank.

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Pteridium is a cosmopolitan genus that acts as an invasive species in many parts of the world. Most research on this genus has occurred in Europe, and there is a lack of data on it from South America, in spite of causing considerable conservation problems. We compared the biomass allocation of P. esculentum subsp. arachnoideum in two ecosystems in Brazil - Atlantic forest and Brazilian savanna. We measured the biomass of fronds, rhizomes and above-ground litter. We also compared the density, length and biomass of fronds from this Brazilian study with similar data of P. esculentum subsp. arachnoideum derived from Venezuela and P. aquilinum from Europe. P. esculentum subsp. arachnoideum showed a wide response range. We found a negative relationship between frond and necromass, indicating a negative feedback effect, while a positive relationship was observed between frond and rhizome biomass. The continental comparison of relationships showed that Pteridium responds in a different way in both Brazil and Europe, and that in Brazil fronds tend to be longer and heavier, presumably as a result of the continuous growing season in South America while is shortened in Europe by frost. The paper shows the ability of Pteridium to adapt to different ecosystems.

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Pteridium is a cosmopolitan genus that acts as an invasive species in many parts of the world. Most research on this genus has occurred in Europe, and there is a lack of data on it from South America, in spite of causing considerable conservation problems. We compared the biomass allocation of P. esculentum subsp. arachnoideum in two ecosystems in Brazil - Atlantic forest and Brazilian savanna. We measured the biomass of fronds, rhizomes and above-ground litter. We also compared the density, length and biomass of fronds from this Brazilian study with similar data of P. esculentum subsp. arachnoideum derived from Venezuela and P. aquilinum from Europe. P. esculentum subsp. arachnoideum showed a wide response range. We found a negative relationship between frond and necromass, indicating a negative feedback effect, while a positive relationship was observed between frond and rhizome biomass. The continental comparison of relationships showed that Pteridium responds in a different way in both Brazil and Europe, and that in Brazil fronds tend to be longer and heavier, presumably as a result of the continuous growing season in South America while is shortened in Europe by frost. The paper shows the ability of Pteridium to adapt to different ecosystems.(AU)

Pteridium é um gênero cosmopolita que inclui espécies invasoras em várias partes do mundo. Os estudos sobre o gênero têm ocorrido principalmente na Europa, e existem poucas informações para as espécies que ocorrem na América do Sul. Nesse estudo comparamos a alocação de biomassa de P. esculentum subsp. arachnoideum em dois ecossistemas brasileiros Mata Atlântica e Cerrado - em cada um dos quais medimos a biomassa nos rametas, nos rizomas e na serapilheira. Comparamos, também, a densidade, comprimento e biomassa dos rametas com informações obtidas sobre P. esculentum subsp. arachnoideum na Venezuela e P. aquilinum na Europa. P. esculentum subsp. arachnoideum apresentou respostas distintas. Encontramos uma relação negativa entre a biomassa de rametas e da serapilheira, indicando uma retro-alimentação negativa, enquanto houve uma relação positiva entre a biomassa dos rametas e dos rizomas. A comparação das relações entre os continentes indicou que Pteridium responde diferentemente no Brazil e na Europa, e também que no Brasil os rametas são maiores e contêm mais biomassa, possivelmente devido à estação de crescimento contínua, enquanto na Europa o crescimento é limitado pelas baixas temperaturas. Esse estudo demonstra o sucesso adaptativo de Pteridium em diferentes ecossistemas.(AU)

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Palms are distinctive plants of tropics and have peculiar allometric relations. Understanding such relations is useful in the case of introduced species because their ability to establish and invade must be clarified in terms of their responses in the new site. Our purpose was to assess the survival and invasive capacity of an introduced palm species in the Atlantic rainforest, Euterpe oleracea Mart., compared to the native Euterpe edulis Mart. and to the hybrids produced between the two species. Considering this, we compared the allometry in different ontogenetic stages, the germination rates, and aspects of the initial development. The ontogenetic stages proposed for both Euterpe illustrated the growth patterns described for palm trees. E. oleracea and hybrids adjusted to the geometric similarity allometric model, while E. edulis presented a slope greater than would be expected considering this model, indicating a greater height for a given diameter. E. oleracea showed the same amount of pulp per fruit as E. edulis and a similar initial development of seedlings. The main differences observed were a lower germination rate and a faster height gain of E. oleracea seedlings. We conclude that E. oleracea, which is similar to E. edulis in aspects of allometry, development, seed and seedling morphology, may be an important competitor of this native palm tree in the Atlantic Forest.

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Studies of allometry are important in explaining effects of fire and herbivory, for estimating biomass in forests, and so on. There has been extensive research on plant allometry in temperate and tropical forests, showing that plant architecture often adjusts to the elastic similarity model, but not in Brazilian savannas (cerrado). We studied allometry of Dalbergia miscolobium, Diospyros hispida, Erythroxylum suberosum, Miconia albicans, M. ligustroides, Schefflera vinosa, and Xylopia aromatica in a cerrado sensu stricto area that was affected by a fire in August 2006. We expected that the study species would not adjust to any of the allometric models considered common for forest species ("constant stress", "elastic similarity", and "geometric growth"), and that there would be differences in allometry in burnt and unburnt patches. We sampled two species in 60 5 × 5 m contiguous plots placed in three transects, and five species in 100 5 × 5 m contiguous plots placed in five transects, where we measured the diameters at soil level (DSL) and the heights of all shoots. We used standardized major axis regressions on log-transformed data. The regression slope between the height and DSL was higher than 1.0 (p < 0.05) for four species, showing a greater height than would be expected under geometric growth, not predicted by theoretical models. We found significant differences (p < 0.05) in regression slopes and/or correlation coefficients between burnt and unburnt plots for five species, indicating that fire may influence plant allometry in the Brazilian cerrado, and that such a response is highly variable between species.

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Studies of allometry are important in explaining effects of fire and herbivory, for estimating biomass in forests, and so on. There has been extensive research on plant allometry in temperate and tropical forests, showing that plant architecture often adjusts to the elastic similarity model, but not in Brazilian savannas (cerrado). We studied allometry of Dalbergia miscolobium, Diospyros hispida, Erythroxylum suberosum, Miconia albicans, M. ligustroides, Schefflera vinosa, and Xylopia aromatica in a cerrado sensu stricto area that was affected by a fire in August 2006. We expected that the study species would not adjust to any of the allometric models considered common for forest species ("constant stress", "elastic similarity", and "geometric growth"), and that there would be differences in allometry in burnt and unburnt patches. We sampled two species in 60 5 × 5 m contiguous plots placed in three transects, and five species in 100 5 × 5 m contiguous plots placed in five transects, where we measured the diameters at soil level (DSL) and the heights of all shoots. We used standardized major axis regressions on log-transformed data. The regression slope between the height and DSL was higher than 1.0 (p < 0.05) for four species, showing a greater height than would be expected under geometric growth, not predicted by theoretical models. We found significant differences (p < 0.05) in regression slopes and/or correlation coefficients between burnt and unburnt plots for five species, indicating that fire may influence plant allometry in the Brazilian cerrado, and that such a response is highly variable between species.(AU)

Estudos de alometria são importantes para interpretar o efeito de fogo e da herbivoria, para estimar biomassa em florestas, entre outros. Há diversos estudos em florestas temperadas e tropicais, mostrando que as plantas desses ecossistemas frequentemente ajustam-se ao modelo de similaridade elástica; mas quase não há estudos realizados no cerrado. Estudamos a alometria de Dalbergia miscolobium, Diospyros hispida, Erythroxylum suberosum, Miconia albicans, M. ligustroides, Schefflera vinosa e Xylopia aromatica em um fragmento de cerrado sensu stricto, afetado de maneira heterogênea por um incêndio em agosto de 2006. Esperávamos que as espécies não se ajustassem a quaisquer dos modelos considerados comuns para espécies florestais ("estresse constante", "similaridade elástica" e "crescimento geométrico"), e que haveria diferenças significativas entre manchas atingidas pelo fogo e as não atingidas. Amostramos duas espécies em 60 parcelas contíguas de 5 × 5 m dispostas em três transectos e cinco espécies em 100 parcelas contíguas de 5 × 5 m dispostas em cinco transectos, dentro das quais medimos o diâmetro à altura do solo (DAS) e a altura de todos os caules. Usamos regressões do tipo eixo principal reduzido sobre logaritmos dos dados. A inclinação da regressão foi maior do que 1,0 (p < 0,05) para quatro espécies, mostrando uma altura maior do que seria esperada sob crescimento geométrico. Esse padrão não é previsto por modelos teóricos. Encontramos diferenças significativas (p < 0,05) nos coeficientes de inclinação e/ou de correlação entre parcelas queimadas e não queimadas para cinco espécies; generalizamos que o fogo pode influenciar a alometria de plantas do cerrado, sendo que essa resposta pode variar entre espécies.(AU)

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In order to identify the effect of P. arachnoideum, we studied 11 native tree species commonly used in reforestation projects. Bioassays were conducted in laboratory to evaluate the effect of bracken leachate on the germination and morphology of seedlings. Juveniles of some of these species were planted in two adjacent but contrasting areas in relation to the dominance of P. arachnoideum. The evaluation of growth and survivorship was performed after six and twelve months. This study reveals that for some pioneer and secondary trees P. arachnoideum leachate exerted an inhibitory effect on seed germination and seedling morphology. Field experiments revealed that pioneers are apparently more resistant to P. arachnoideum leachate than secondary species.

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Since cerrado fires may impede the growth of seedlings into trees, they may shape the population of woody species. In this study, we assessed the effects of a severe fire on the population structure and spatial distribution of Zanthoxylum rhoifolium, a widespread cerrado tree. We were interested to know the importance of the resprouting and sexual reproduction in regenerating the population. The study area had been for about six years protected from fire, before a severe fire at the end of the dry season in 2006. We sampled and measured all individuals of Z. rhoifolium found in 80 plots of 25 m(2). We found 149 individuals before the fire and 112 after the fire, of which 77 were resprouts from burned seedlings and saplings. We did not find significant differences between the population structure before and after the fire. The spatial distribution of the population remained clumped after the fire. Thus, the Z. rhoifolium population was very resilient to a severe fire. We did not find any new seedlings. As a consequence, resprouting seems to be more important than sexual reproduction in promptly regenerating the Z. rhoifolium populations.

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Since cerrado fires may impede the growth of seedlings into trees, they may shape the population of woody species. In this study, we assessed the effects of a severe fire on the population structure and spatial distribution of Zanthoxylum rhoifolium, a widespread cerrado tree. We were interested to know the importance of the resprouting and sexual reproduction in regenerating the population. The study area had been for about six years protected from fire, before a severe fire at the end of the dry season in 2006. We sampled and measured all individuals of Z. rhoifolium found in 80 plots of 25 m². We found 149 individuals before the fire and 112 after the fire, of which 77 were resprouts from burned seedlings and saplings. We did not find significant differences between the population structure before and after the fire. The spatial distribution of the population remained clumped after the fire. Thus, the Z. rhoifolium population was very resilient to a severe fire. We did not find any new seedlings. As a consequence, resprouting seems to be more important than sexual reproduction in promptly regenerating the Z. rhoifolium populations.

Uma vez que as queimadas no cerrado podem impedir o crescimento de plântulas, elas podem moldar a população de espécies arbóreas. Neste estudo, avaliamos os efeitos de uma queimada severa na estrutura e na distribuição espacial da população de Zanthoxylum rhoifolium, uma espécie de árvore comum no cerrado. O objetivo era saber a importância relativa do rebrotamento e da reprodução sexuada na regeneração da população. A área de estudo, há aproximadamente seis anos sem queimar, sofreu uma queimada severa no final da estação seca de 2006. Amostramos todos os indivíduos de Z. rhoifolium encontrados em 80 parcelas de 25 m². Encontramos 149 indivíduos antes do fogo e 112 depois dele, dos quais 77 eram rebrotas de plântulas e jovens queimados. Não encontramos diferenças significativas entre a estrutura da população antes e após o fogo. A distribuição espacial da população permaneceu agregada após a queimada. Portanto, a população de Z. rhoifolium foi resiliente ao fogo. Não foi encontrada nenhuma nova plântula. Consequentemente, o rebrotamento parece ser mais importante que a reprodução sexuada na regeneração imediata das populações de Z. rhoifolium.

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