Scale-dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity. 2. Here, we conduct an analysis of relationships between tree species richness, biomass and productivity in 25 forest plots of area 8-50ha from across the world. The data were collected using standardized protocols, obviating the need to correct for methodological differences that plague many studies on this topic. 3. We found that at very small spatial grains (0.04ha) species richness was generally positively related to productivity and biomass within plots, with a doubling of species richness corresponding to an average 48% increase in productivity and 53% increase in biomass. At larger spatial grains (0.25ha, 1ha), results were mixed, with negative relationships becoming more common. The results were qualitatively similar but much weaker when we controlled for stem density: at the 0.04ha spatial grain, a doubling of species richness corresponded to a 5% increase in productivity and 7% increase in biomass. Productivity and biomass were themselves almost always positively related at all spatial grains. 4. Synthesis. This is the first cross-site study of the effect of tree species richness on forest biomass and productivity that systematically varies spatial grain within a controlled methodology. The scale-dependent results are consistent with theoretical models in which sampling effects and niche complementarity dominate at small scales, while environmental gradients drive patterns at large scales. Our study shows that the relationship of tree species richness with biomass and productivity changes qualitatively when moving from scales typical of forest surveys (0.04ha) to slightly larger scales (0.25 and 1ha). This needs to be recognized in forest conservation policy and management.

Relief influence on tree species richness in secondary forest fragments of Atlantic Forest, SE, Brazil

(Relief influence on tree species richness in secondary forest fragments of Atlantic Forest, SE, Brazil). The aim of this work was to explore the relationship between tree species richness and morphological characteristics of relief at the Ibiuna Plateau (SE Brazil). We sampled 61 plots of 0.30 ha, systematically established in 20 fragments of secondary forest (2-274 ha) and in three areas within a continuous secondary forest site, Morro Grande Reserve (9,400 ha). At each plot, 100 trees with diameter at breast height > 5 cm were sampled by the point centered quarter method, and total richness and richness per dispersal and succession class were obtained. The relief was characterized by the mean and variance of slope, elevation, aspect and slope location. There was no significant relationship between relief heterogeneity and tree species richness. Relief parameters generally did not affect tree richness, but elevation was particularly important especially in the continuous forest. Despite the limited range of altitudinal variation (150 m), species richness increases with elevation. The highest areas were also those with the largest forest cover and the lowest disturbance degree, which should contribute to the greater richness of those sites. Our results suggest an indirect influence of relief, due to the fact that deforestation is less intense in higher regions, rather than a direct influence of abiotic factors related to the altitudinal gradient.

Relief influence on tree species richness in secondary forest fragments of Atlantic Forest, SE, Brazil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Species richness and composition of butterfly assemblages (Lepidoptera:Rhopalocera) along the altitudinal gradient of Serra da Estrela

Os gradientes altitudinais são importantes modelos em ecologia pois permitem, numa escala reduzida, estudar a influência da variação de diversas variáveis ambientais nos padrões de biodiversidade existentes. Estes padrões são influenciados por diversos factores, desde variáveis climáticas às relacionadas com a perturbação humana. Nesta tese é estudada a variação na composição das comunidades de borboletas ao longo do gradiente altitudinal na Serra da Estrela e são testadas duas hipóteses sobre os padrões de riqueza específica ao longo do gradiente altitudinal: a regra de Rapoport e o “Mid Domain Effect” (MDE). Foram assinaladas 70 espécies de borboletas neste estudo e constatou-se que a variação na riqueza de espécies de borboletas apresenta um padrão “mid peak” apoiando o MDE. Verificou-se também que as comunidades de borboletas do mesmo nível altitudinal são mais semelhantes entre si independentemente da vertente em que se encontram e são também mais semelhantes com as comunidades dos níveis altitudinais adjacentes. Finalmente, a análise da distribuição de algumas espécies ameaçadas de borboletas torna evidente a necessidade de realização de mais estudos sobre a biologia da conservação deste grupo de animais em Portugal; Abstract: Altitudinal gradients are important ecological models because they allow us to study the influence of diverse environmental variables in the existing biodiversity patterns. These patterns are influenced by various factors, from climatic variables to variables related to human disturbance. In this thesis we studied the variations in the composition of the butterfly assemblages along the altitudinal gradient of Serra da Estrela and we tested two hypotheses regarding the species richness patterns along the altitudinal gradient: Rapoport’s rule and the Mid Domain Effect (MDE). In this study 70 butterfly species were marked recorded and it was observed that the butterfly species richness presents a mid-peak pattern supporting the MDE. It was also noted that the butterfly assemblages in the same altitudinal level show more similarity within themselves regardless of the slope they are found in, and are also more similar to the assemblages present in the adjacent altitudinal levels. Finally, the distribution of some threatened butterfly species was analyzed and it became evident the need to do more studies regarding the conservation biology of this animal group in Portugal.

Species richness and abundance of low-trunk herb epiphytes in relation to host tree size and bark type, Eastern Amazonia

A composição e estrutura da comunidade epifítica herbácea de fuste baixo, assim como sua distribuição vertical, foram estudadas. O DAP de hospedeiros arbóreos e o tipo de casca influenciam a riqueza e abundância dessas espécies em um trecho de floresta de terra firme na Amazônia Oriental (1º57’36"S 51º36’55"W). Foram identificadas, no total, 37 espécies herbáceas epifíticas, sendo 60% delas Araceae. A riqueza de espécies e a abundância de herbáceas epifíticas mostraram tendência de correlação positiva com o tamanho de hospedeiros arbóreos e nenhuma relação com o tipo de casca. Correlação positiva baixa pode ser um subproduto da predominância de árvores de menor diâmetro na amostragem em vez de refletir relação neutra. A ausência de relações com o tipo de casca deve ser parcialmente explicada pelo grande número de hemiepífitas secundárias, generalistas, e também refletir a ausência de substratos adequados em árvores de menor diâmetro.

Ant community richness and composition across a gradient from Eucalyptus plantations to secondary Atlantic Forest

As florestas secundárias e plantações de espécies exóticas estão se expandindo nas paisagens tropicais. No entanto, nossa compreensão sobre o valor destas florestas para a conservação da biodiversidade de invertebrados ainda é incipiente. Neste trabalho, usamos a fauna de formigas de serapilheira para avaliar a diversidade desses insetos entre três florestas de Eucalyptus, sendo uma comercial (quatro anos de idade) e duas abandonadas em diferentes idades de regeneração (16 e 31 anos) e uma área de Mata Atlântica secundária. A riqueza total foi mais alta na floresta secundária e nos plantios de Eucalyptus abandonados há mais tempo. A densidade de espécies na floresta secundária foi significativamente maior quando comparado as plantações de Eucalyptus, mas não difere entre eucaliptais; análise de ordenação revelou diferenças na composição de espécies entre as plantações de Eucalyptus com subbosque ausente e com subbosque desenvolvido ou em desenvolvimento. Ainda, foi constatada uma sobreposição acentuada entre amostras de serapilheira das florestas de eucaliptos abandonadas há mais tempo e a floresta secundária. em geral, plantações de eucalipto foram caracterizadas pela presença de espécies generalistas e de ampla distribuição. Nossos resultados indicam que embora o subbosque de plantações de eucaliptos com maior idade de regeneração suporte um conjunto relativamente alto de espécies generalistas de formigas, é improvável que eucaliptais conservem a maioria das espécies de florestas primárias, especialmente predadores especializados, Dacetini e espécies nômades.

Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity?

Results from the humid tropics of Australia demonstrate that diverse plantations can achieve greater productivity than monocultures. We found that increases in both the observed species number and the effective species richness were significantly related to increased levels of productivity as measured by stand basal area or mean individual tree basal area. Four of five plantation species were more productive in mixtures with other species than in monocultures, offering on average, a 55% increase in mean tree basal area. A general linear model suggests that species richness had a significant effect on mean individual tree basal area when environmental variables were included in the model. As monoculture plantations are currently the preferred reforestation method throughout the tropics these results suggest that significant productivity and ecological gains could be made if multi-species plantations are more broadly pursued.

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers' residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers' residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m(2) along a transect and classified the contents by species. We observed 15.5-19.7 species per 0.01 m(2), which is high richness per 0.01 m(2) on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.

Tree species richness, does it play a key role on a forest restoration plantation?

The Brazilian Atlantic forest is considered one of the world's biodiversity conservation hotspot. Today there is less than ten percent remaining. Therefore it is necessary to restore these ecosystems. There are many ways of achieving restoration's main goals, but there is a lack of ecological studies that analyzes tree species richness as a variable. Thus, this study's goal is to investigate if there is a difference between a forest restoration in a gradient of tree species richness that varies from 20, 60 to 120 species, by using the litterfall as an indicator. Every month, for one year the forest litter was collected from litter traps that were previously installed. Results revealed that stands produced litterfall by the increasing gradient of species was of 5,370, 5,909 and 6,432 kg ha(-1) yr(-1). The statistical analyses revealed no significant difference among them. Therefore this six-year-old forest restoration plantation shows no difference on the litter production by the tree species richness.

Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity?

Results from the humid tropics of Australia demonstrate that diverse plantations can achieve greater productivity than monocultures. We found that increases in both the observed species number and the effective species richness were significantly related to increased levels of productivity as measured by stand basal area or mean individual tree basal area. Four of five plantation species were more productive in mixtures with other species than in monocultures, offering on average, a 55% increase in mean tree basal area. A general linear model suggests that species richness had a significant effect on mean individual tree basal area when environmental variables were included in the model. As monoculture plantations are currently the preferred reforestation method throughout the tropics these results suggest that significant productivity and ecological gains could be made if multi-species plantations are more broadly pursued. (c) 2006 Elsevier B.V. All rights reserved.

Impacts of level of utilisation by grazing on an Astrebla (Mitchell grass) grassland in north-western Queensland between 1984 and 2010. 2. Plant species richness and abundance

The occurrence of interstitial species in Astrebla grasslands in Australia are influenced by grazing and seasonal rainfall but the interactions of these two influences are complex. This paper describes three studies aimed at determining and explaining the changes in plant species richness and abundance of the interstitial species in a long-term sheep utilisation experiment in an Astrebla grassland in northern Queensland. In the first study, increasing utilisation increased the frequency of Dactyloctenium radulans (Button grass) and Brachyachne convergens (Downs couch) and reduced that of Streptoglossa adscendens (Mint bush). In the second study, seasonal rainfall variation between 1984 and 2009 resulted in large annual differences in the size of the seed banks of many species, but increasing utilisation consistently reduced the seed bank of species such as Astrebla spp. and S. adscendens and increased that of species such as B. convergens, D. radulans, Amaranthus mitchellii (Boggabri) and Boerhavia sp. (Tar vine). In the third study, the highest species richness occurred at the lightest utilisation because of the presence of a range of palatable forbs, especially legumes. Species richness was reduced as utilisation increased. Species richness in the grazing exclosure was low and similar to that at the heaviest utilisation where there was a reduction in the presence of palatable forb species. The pattern of highest species richness at the lightest grazing treatment was maintained across three sampling times, even with different amounts of seasonal rainfall, but there was a large yearly variation in both the density and frequency of many species. It was concluded that the maintenance of highest species richness at the lightest utilisation was not aligned with other data from this grazing experiment which indicated that the maximum sustainable wool production occurred at moderate utilisation.

Bird and mammal conservation in Gaoligongshan Region and Jingdong County, Yunnan, China: patterns of species richness and nature reserves

Elevational and latitudinal patterns of species richness for birds and mammals were compared with human population density in relation to nature reserve designation in two areas of Yunnan Province, China. Results suggest that species richness is not the same for the two areas. In Gaoligongshan Region, species richness is inversely correlated with elevation and altitude, while reserve designation is positively correlated with elevation and latitude. In Jingdong County, reserve designations are positively correlated with elevation, but species richness shows no clear trends. In general, the present situation is strongly influenced by human activities. It appears that reserve designation is mismatched with species richness in Gaoligongshan Region, while there is a better fit between the two in Jingdong County. In both areas, however, it appeared that reserves were located primarily in order to reduce conflict with humans rather than to maximize conservation of biodiversity, probably because humans were responsible for forest-especially primary forest-destruction and degradation in the low-lying areas.

Tree species-area and species-diameter relationships at three lowland rain forest sites in Sumatra

Data from three forest sites in Sumatra (Batang Ule, Pasirmayang and Tebopandak) have been analysed and compared for the effects of sample area cut-off, and tree diameter cut-off. An 'extended inverted exponential model' is shown to be well suited to fitting tree-species-area curves. The model yields species carrying capacities of 680 for Batang Ule, 380 species for Pasirmayang, and 35 for Tebopandak (tree diameter >10cm). It would seem that in terms of species carrying capacity, Tebopandak and Pasirmayang are rather similar, and both less diverse than the hilly Batang Ule site. In terms of conservation policy, this would mean that rather more emphasis should be put on conserving hilly sites on a granite substratum. For Pasirmayang with tree diameter >3cm, the asymptotic species number estimate is 567, considerably higher than the estimate of 387 species for trees with diameter >10cm. It is clear that the diameter cut-off has a major impact on the estimate of the species carrying capacity. A conservative estimate of the total number of tree species in the Pasirmayang region is 632 species! In sampling exercises, the diameter cut-off should not be chosen lightly, and it may be worth adopting field sampling procedures which involve some subsampling of the primary sample area, where the diameter cut-off is set much lower than in the primary plots.