Janneke Hille Ris Lambers

Productivity is a poor predictor of plant species richness

Standardized sampling from many sites worldwide was used to address an important ecological problem. For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters−2) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.

From selection to complementarity: shifts in the causes of biodiversity-productivity relationships in a long-term biodiversity experiment.

In a 10-year (1996–2005) biodiversity experiment, the mechanisms underlying the increasingly positive effect of biodiversity on plant biomass production shifted from sampling to complementarity over time. The effect of diversity on plant biomass was associated primarily with the accumulation of higher total plant nitrogen pools (N g m−2) and secondarily with more efficient N use at higher diversity. The accumulation of N in living plant biomass was significantly increased by the presence of legumes, C4 grasses, and their combined presence. Thus, these results provide clear evidence for the increasing effects of complementarity through time and suggest a mechanism whereby diversity increases complementarity through the increased input and retention of N, a commonly limiting nutrient.

Density-dependent mortality and the latitudinal gradient in species diversity.

Ecologists have long postulated that density-dependent mortality maintains high tree diversity in the tropics. If species experience greater mortality when abundant, then more rare species can persist. Agents of density-dependent mortality (such as host-specific predators, and pathogens) may be more prevalent or have stronger effects in tropical forests, because they are not limited by climatic factors. If so, decreasing density-dependent mortality with increasing latitude could partially explain the observed latitudinal gradient in tree diversity. This hypothesis has never been tested with latitudinal data. Here we show that several temperate tree species experience density-dependent mortality between seed dispersal and seedling establishment. The proportion of species affected is equivalent to that in tropical forests, failing to support the hypothesis that this mechanism is more prevalent at tropical latitudes. We further show that density-dependent mortality is misinterpreted in previous studies. Our results and evidence from other studies suggest that density-dependent mortality is important in many forests. Thus, unless the strength of density-dependent mortality varies with latitude, this mechanism is not likely to explain the high diversity of tropical forests.

EXPLOITING TEMPORAL VARIABILITY TO UNDERSTAND TREE RECRUITMENT RESPONSE TO CLIMATE CHANGE

Predicting vegetation shifts under climate change is a challenging endeavor, given the complex interactions between biotic and abiotic variables that influence demographic rates. To determine how current trends and variation in climate change affect seedling establishment, we analyzed demographic responses to spatiotemporal variation to temperature and soil moisture in the southern Appalachian Mountains. We monitored seedling establishment for 10 years in five plots located along an elevational gradient of five dominant tree species: Acer rubrum, Betula spp., Liriodendron tulipifera, Nyssa sylvatica, and Quercus rubra. A hierarchical Bayes model allowed us to incorporate different sources of information, observation errors, and the inherent variability of the establishment process. From our analysis, spring temperatures and heterogeneity in soil moisture emerge as key drivers, and they act through nonlinear population demographic processes. We found that all species benefited from warmer springs, in particular the species found on dry slopes, N. sylvatica, and those dominant at higher elevations, Betula spp. and Q. rubra. This last species also benefited from dry environments. Conversely, L. tulipifera, which is abundant on mesic sites, experienced highest establishment rates at high moisture. The mechanisms behind these results may differ among species. Higher temperatures are apparently more important for some, while dry conditions and reduced pathogenic attacks on their seeds and new seedlings have a large impact for others. Our results suggest that only communities found at higher elevations are in danger of regional extinction when their habitats disappear given the current climatic trends. We conclude that the recruitment dynamics of the communities where these species are dominant could be affected by minor changes in climate in ways that cannot be predicted using only climate envelopes, which use different variables and miss the nonlinearities.

IMPLICATIONS OF SEED BANKING FOR RECRUITMENT OF SOUTHERN APPALACHIAN WOODY SPECIES

Seed dormancy is assumed to be unimportant for population dynamics of temperate woody species, because seeds occur at low densities and are short lived in forest soils. However, low soil seed densities may result from low seed production, and even modest seed longevity can buffer against fluctuating seed production, potentially limiting density-dependent mortality and ensuring that seeds are available for germination when recruitment success is likely. To investigate whether seed banking affects woody seedling dynamics in the southern Appalachians, we monitored seed rain, seed bank, and seedling densities to (1) determine the prevalence of seed banking among southern Appalachian woody species, (2) quantify annual seed mortality rates for three seed-banking species using a Bayesian statistical approach, and (3) assess whether or not the ability to seed bank affects recruitment rates. We found that the seeds of eight woody taxa ( Acer rubrum, Betula spp., Liriodendron tulipifera, Nyssa sylvatica, Robinia pseudoacacia, Rubus spp., Sassafras al- bidum, and Vitis sp.) remain viable in the soil for more than one year. Seeds of six taxa (Amelanchier spp., Acer pennsylvanicum, Carya spp., Quercus prinus, Quercus rubra, and Tsuga canadensis) were never found in the soil seed bank, despite high seed production and germination. For three species, a substantial proportion of seeds available for germi- nation came from dispersal events two or more years in the past (Acer rubrum 12-37%, Betula spp. 59-73%, Liriodendron tulipifera 40-76%), even though annual seed mortality was high (Acer rubrum 70-98%, Betula spp. 21-81%, Liriodendron tulipifera 12-59%). In years when no seeds fall in local microsites (approximately one in five years), seed banks are the only source of seedling recruitment for these species. Comparing our results to those of previous studies led to valuable insights: first, that seeds of Acer rubrum and Betula spp. suffer high mortality while being incorporated into the seed bank; and second, that seed decay varies greatly over relatively small spatial scales (i.e., within a watershed). Taken together, these results demonstrate that seed banking may play a critical role during woody seedling recruitment in temperate forests.

‘Natural experiment’ Demonstrates Top-Down Control of Spiders by Birds on a Landscape Level

The combination of small-scale manipulative experiments and large-scale natural experiments provides a powerful approach for demonstrating the importance of top-down trophic control on the ecosystem scale. The most compelling natural experiments have come from studies examining the landscape-scale loss of apex predators like sea otters, wolves, fish and land crabs. Birds are dominant apex predators in terrestrial systems around the world, yet all studies on their role as predators have come from small-scale experiments; the top-down impact of bird loss on their arthropod prey has yet to be examined at a landscape scale. Here, we use a unique natural experiment, the extirpation of insectivorous birds from nearly all forests on the island of Guam by the invasive brown tree snake, to produce the first assessment of the impacts of bird loss on their prey. We focused on spiders because experimental studies showed a consistent top-down effect of birds on spiders. We conducted spider web surveys in native forest on Guam and three nearby islands with healthy bird populations. Spider web densities on the island of Guam were 40 times greater than densities on islands with birds during the wet season, and 2.3 times greater during the dry season. These results confirm the general trend from manipulative experiments conducted in other systems however, the effect size was much greater in this natural experiment than in most manipulative experiments. In addition, bird loss appears to have removed the seasonality of spider webs and led to larger webs in at least one spider species in the forests of Guam than on nearby islands with birds. We discuss several possible mechanisms for the observed changes. Overall, our results suggest that effect sizes from smaller-scale experimental studies may significantly underestimate the impact of bird loss on spider density as demonstrated by this large-scale natural experiment.

Spatial Heterogeneity in Ecologically Important Climate Variables at Coarse and Fine Scales in a High-Snow Mountain Landscape

Climate plays an important role in determining the geographic ranges of species. With rapid climate change expected in the coming decades, ecologists have predicted that species ranges will shift large distances in elevation and latitude. However, most range shift assessments are based on coarse-scale climate models that ignore fine-scale heterogeneity and could fail to capture important range shift dynamics. Moreover, if climate varies dramatically over short distances, some populations of certain species may only need to migrate tens of meters between microhabitats to track their climate as opposed to hundreds of meters upward or hundreds of kilometers poleward. To address these issues, we measured climate variables that are likely important determinants of plant species distributions and abundances (snow disappearance date and soil temperature) at coarse and fine scales at Mount Rainier National Park in Washington State, USA. Coarse-scale differences across the landscape such as large changes in elevation had expected effects on climatic variables, with later snow disappearance dates and lower temperatures at higher elevations. However, locations separated by small distances (∼20 m), but differing by vegetation structure or topographic position, often experienced differences in snow disappearance date and soil temperature as great as locations separated by large distances (>1 km). Tree canopy gaps and topographic depressions experienced later snow disappearance dates than corresponding locations under intact canopy and on ridges. Additionally, locations under vegetation and on topographic ridges experienced lower maximum and higher minimum soil temperatures. The large differences in climate we observed over small distances will likely lead to complex range shift dynamics and could buffer species from the negative effects of climate change.