Midpoint attractors and species richness: Modelling the interaction between environmental drivers and geometric constraints.

We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor - a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon-specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.

Impact of mycorrhization on the abundance, growth and leaf nutrient status of ferns along a tropical elevational gradient.

Mycorrhizal fungi are crucial for the ecological success of land plants, providing their hosts with nutrients in exchange for organic C. However, not all plants are mycorrhizal, especially ferns, of which about one-third of the species lack this symbiosis. Because the mycorrhizal status is evolutionarily ancestral, this lack of mycorrhizae must have ecological advantages, but what these advantages are and how they affect the competitive ability of non-mycorrhizal plants under natural conditions is currently unknown. To address this uncertainty, we studied terrestrial fern assemblages and species abundances as well as their mycorrhization status, leaf nutrient concentration and relative annual growth along an elevational gradient in the Ecuadorian Andes (500-4,000 m). We surveyed the mycorrhizal status of 375 root samples belonging to 85 species, and found mycorrhizae in 89% of the samples. The degree of mycorrhization decreased with elevation but was unrelated to soil nutrients. Species with mycorrhizae were significantly more abundant than non-mycorrhizal species, but non-mycorrhizal species had significantly higher relative growth and concentrations of leaf N, P, Mg, and Ca. Our study thus shows that despite lower abundances, non-mycorrhizal fern species did not appear to be limited in their growth or nutrient supply relative to mycorrhizal ones. As a basis for future studies, we hypothesize that non-mycorrhizal fern species may be favoured in special microhabitats of the forest understory with high soil nutrient or water availability, or that the ecological benefit of mycorrhizae is not related to nutrient uptake but rather to, for example, pathogen resistance.

Functional Diversity in Ferns Is Driven by Species Richness Rather Than by Environmental Constraints.

Functional traits determine how species interact with their abiotic and biotic environment. In turn, functional diversity describes how assemblages of species as a whole are adapted to their environment, which also determines how they might react to changing conditions. To fully understand functional diversity, it is fundamental to (a) disentangle the influences of environmental filtering and species richness from each other, (b) assess if the trait space saturates at high levels of species richness, and (c) understand how changes in species numbers affect the relative importance of the trait niche expansion and packing. In the present study, we determined functional diversity of fern assemblages by describing morphological traits related to resource acquisition along four tropical elevational transects with different environmental conditions and species richness. We used several functional diversity indices and their standardized effect size to consider different aspects of functional diversity. We contrasted these aspects of functional diversity with climate data and species richness using linear models and linear mixed models. Our results show that functional morphological trait diversity was primarily driven by species richness and only marginally by environmental conditions. Moreover, increasing species richness contributed progressively to packing of the morphological niche space, while at the same time decreasing morphological expansion until a saturation point was reached. Overall, our findings suggest that the density of co-occurring species is the fundamental driving force of morphological niche structure, and environmental conditions have only an indirect influence on fern resource acquisition strategies.

Alpha and beta diversity of plants and animals along a tropical land-use gradient.

Assessing the overall biological diversity of tropical rain forests is a seemingly insurmountable task for ecologists. Therefore, researchers frequently sample selected taxa that they believe reflect general biodiversity patterns. Usually, these studies focus on the congruence of alpha diversity (the number of species found per sampling unit) between taxa rather than on beta diversity (turnover of species assemblages between sampling units). Such approaches ignore the potential role of habitat heterogeneity that, depending on the taxonomic group considered, can greatly enhance beta diversity at local and landscape scales. We compared alpha and beta diversity of four plant groups (trees, lianas, terrestrial herbs, epiphytic liverworts) and eight animal groups (birds, butterflies, lower canopy ants, lower canopy beetles, dung beetles, bees, wasps, and the parasitoids of the latter two) at 15 sites in Sulawesi, Indonesia, that represented natural rain forest and three types of cacao agroforests differing in management intensity. In total, we recorded 863 species. Patterns of species richness per study site varied strongly between taxonomic groups. Only 13-17% of the variance in species richness of one taxonomic group could be predicted from the species richness of another, and on average 12-18% of the variance of beta diversity of a given group was predicted by that in other groups, although some taxon pairs had higher values (up to 76% for wasps and their parasitoids). The degree of congruence of patterns of alpha diversity was not influenced by sampling completeness, whereas the indicator value for beta diversity improved when using a similarity index that accounts for incomplete sampling. The indication potential of alpha diversity for beta diversity and vice versa was limited within taxa (7-20%) and virtually nil between them (0-4%). We conclude that different taxa can have largely independent patterns of alpha diversity and that patterns of beta diversity can be more congruent. Thus, conservation plans on a landscape scale need to put more emphasis on the high heterogeneity of agroforests and the overarching role of beta diversity shaping overall diversity patterns.

Natural and human-impacted diversity of bryophytes along an elevational gradient on an oceanic island (La Palma, Canarias).

Bryophytes have been proposed as ideal indicators of ecosystem change, because they are important components of forest integrity, and considerable research indicates that some groups are sensitive to the changes associated with specific human disturbances. Bryophyte richness and abundance have been found to vary predictably along elevational gradients, but the role of human impacts on these distribution patterns remains unclear. The aim of this study is to explore the impact of human disturbance on the elevational patterns of bryophyte diversity, along an elevational gradient. Along the gradient we collected three datasets in the following sites: preserved (P), forest track roadsides (R) and disturbed by agriculture/silviculture practices (D). Two survey plots of 100 m2 were established at every 200 m elevational step for each sites P, R, D, and in each plot bryophytes were sampled in a stratified manner. At each plot we recorded all species on available substrates and estimated their percentage cover. Our results showed that species number did not differ among studied sites, but that species diversity pattern differs among the three gradient types and species life strategy composition along the elevational gradient showed a clear response to the disturbance of mature communities. We conclude that human impact has strongly changed the elevational pattern of diversity, and that these changes vary depending on the ecological and taxonomical group considered.

Selliguea kachinensis (Polypodiaceae), a new fern species of uncertain affinity from Northern Myanmar.

We describe Selliguea kachinensis as a new species from Northern Myanmar and discuss its generic placement in either Selliguea or Arthromeris. The conservation status is assessed as Data Deficient. In addition, we make the new combination Selliguea erythrocarpa (Mett. ex Kuhn) Hovenkamp, S. Linds., Fraser-Jenk.

Elevational patterns of Polylepis tree height (Rosaceae) in the high Andes of Peru: role of human impact and climatic conditions.

We studied tree height in stands of high-Andean Polylepis forests in two cordilleras near Cuzco (Peru) with respect to variations in human impact and climatic conditions, and compared air and soil temperatures between qualitatively defined dry and humid slopes. We studied 46 forest plots of 100 m(2) of five Polylepis species at 3560-4680 m. We measured diameter at breast height (dbh) and tree height in the stands (1229 trees in total), as well as air and soil temperatures in a subset of plots. The data was analyzed combining plots of given species from different sites at the same elevation (±100 m). There was no elevational decrease of mean maximum tree height across the entire data set. On humid slopes, tree height decreased continuously with elevation, whereas on dry slopes it peaked at middle elevations. With mean maximum tree heights of 9 m at 4530 m on the humid slopes and of 13 m at 4650 m on the dry slopes, we here document the tallest high-elevation forests found so far worldwide. These highest stands grow under cold mean growing season air temperatures (3.6 and 3.8°C on humid vs. dry slopes) and mean growing season soil temperatures (5.1 vs. 4.6°C). Mean annual air and soil temperature both decreased with elevation. Dry slopes had higher mean and maximum growing season air temperatures than humid slopes. Mean annual soil temperatures did not significantly differ and mean annual air temperatures only slightly differed between slopes. However, maximum air temperatures differed on average by 6.6 K between dry and humid slopes. This suggests that the differences in tree height between the two slopes are most likely due to differences in solar radiation as reflected by maximum air temperatures. Our study furthermore provides evidence that alpine Polylepis treelines grow under lower temperature conditions than global high-elevation treelines on average, suggesting that Polylepis species may have evolved special physiological adaptations to low temperatures.

Can joint carbon and biodiversity management in tropical agroforestry landscapes be optimized?

Managing ecosystems for carbon storage may also benefit biodiversity conservation, but such a potential 'win-win' scenario has not yet been assessed for tropical agroforestry landscapes. We measured above- and below-ground carbon stocks as well as the species richness of four groups of plants and eight of animals on 14 representative plots in Sulawesi, Indonesia, ranging from natural rainforest to cacao agroforests that have replaced former natural forest. The conversion of natural forests with carbon stocks of 227-362 Mg C ha(-1) to agroforests with 82-211 Mg C ha(-1) showed no relationships to overall biodiversity but led to a significant loss of forest-related species richness. We conclude that the conservation of the forest-related biodiversity, and to a lesser degree of carbon stocks, mainly depends on the preservation of natural forest habitats. In the three most carbon-rich agroforestry systems, carbon stocks were about 60% of those of natural forest, suggesting that 1.6 ha of optimally managed agroforest can contribute to the conservation of carbon stocks as much as 1 ha of natural forest. However, agroforestry systems had comparatively low biodiversity, and we found no evidence for a tight link between carbon storage and biodiversity. Yet, potential win-win agroforestry management solutions include combining high shade-tree quality which favours biodiversity with cacao-yield adapted shade levels.

The influence of climatic seasonality on the diversity of different tropical pollinator groups.

Tropical South America is rich in different groups of pollinators, but the biotic and abiotic factors determining the geographical distribution of their species richness are poorly understood. We analyzed the species richness of three groups of pollinators (bees and wasps, butterflies, hummingbirds) in six tropical forests in the Bolivian lowlands along a gradient of climatic seasonality and precipitation ranging from 410 mm to 6250 mm. At each site, we sampled the three pollinator groups and their food plants twice for 16 days in both the dry and rainy seasons. The richness of the pollinator groups was related to climatic factors by linear regressions. Differences in species numbers between pollinator groups were analyzed by Wilcoxon tests for matched pairs and the proportion in species numbers between pollinator groups by correlation analyses. Species richness of hummingbirds was most closely correlated to the continuous availability of food, that of bees and wasps to the number of food plant species and flowers, and that of butterflies to air temperature. Only the species number of butterflies differed significantly between seasons. We were not able to find shifts in the proportion of species numbers of the different groups of pollinators along the study gradient. Thus, we conclude that the diversity of pollinator guilds is determined by group-specific factors and that the constant proportions in species numbers of the different pollinator groups constitute a general pattern.