Seed mucilage in temperate grassland species is unrelated to moisture requirements.

Myxospermy, the release of seed mucilage upon hydration, plays multiple roles in seed biology. Here, we explore whether seed mucilage occurs in a suite of temperate grassland species to test if the prevalence of species producing seed mucilage is associated with habitat type or seed characteristics. Seventy plant species found in wet or dry North American temperate grasslands were tested for the presence of seed mucilage through microscopic examination of seeds imbibed with histochemical stain for mucilage. Mucilage production was compared among species with different moisture requirements and seed mass. In this study, 43 of 70 of species tested produced seed mucilage. Seed mucilage did not differ based on habitat type, species moisture requirements, or seed mass. Most seed mucilage was non-adherent and did not remain stuck to the seed after extrusion. Seed mucilage was a common trait in the surveyed temperate grassland species and was observed in 61% of evaluated species. Surprisingly, seed mucilage was more common in temperate grasslands than in previous ecological surveys from arid/semiarid systems, which found 10%-31% myxospermous species. Given the high prevalence, seed mucilage may influence seedling ecology in temperate grasslands and requires further investigation.

Sixty years of community change in the prairie-savanna-forest mosaic of Wisconsin.

Biodiversity loss is a global concern, and maintaining habitat complexity in naturally patchy landscapes can help retain regional diversity. A mosaic of prairie, savanna, and forest historically occurred across central North America but currently is highly fragmented due to human land conversion. It is unclear how each habitat type now contributes to regional diversity. Using legacy data, we resurveyed savanna plant communities originally surveyed in the 1950s to compare change in savannas to that in remnant forests and prairies. Savanna community structure and composition changed substantially over the past 60 years. Tree canopy density nearly doubled and many prairie and savanna specialist species were replaced by forest and non-native species. All three habitats gained and lost many species since the 1950s, resulting in large changes in community composition from local colonizations and extinctions. Across all three habitats, regional species extinctions matched that of regional colonization resulting in no net change in regional species richness. Synthesis-Despite considerable species turnover within savannas, many species remain within the broader prairie-savanna-forest mosaic. Both regional extinctions and colonizations were high over the past 60 years, and maintaining the presence of all three community types-prairie, savanna and forest-on the landscape is critical to maintaining regional biodiversity.

Temporal heterogeneity increases with spatial heterogeneity in ecological communities.

Heterogeneity is increasingly recognized as a foundational characteristic of ecological systems. Under global change, understanding temporal community heterogeneity is necessary for predicting the stability of ecosystem functions and services. Indeed, spatial heterogeneity is commonly used in alternative stable state theory as a predictor of temporal heterogeneity and therefore an early indicator of regime shifts. To evaluate whether spatial heterogeneity in species composition is predictive of temporal heterogeneity in ecological communities, we analyzed 68 community data sets spanning freshwater and terrestrial systems where measures of species abundance were replicated over space and time. Of the 68 data sets, 55 (81%) had a weak to strongly positive relationship between spatial and temporal heterogeneity, while in the remaining communities the relationship was weak to strongly negative (19%). Based on a mixed model analysis, we found a significant but weak overall positive relationship between spatial and temporal heterogeneity across all data sets combined, and within aquatic and terrestrial data sets separately. In addition, lifespan and successional stage were negatively and positively related to temporal heterogeneity, respectively. We conclude that spatial heterogeneity may be a predictor of temporal heterogeneity in ecological communities, and that this relationship may be a general property of many terrestrial and aquatic communities.

Press-pulse interactions: effects of warming, N deposition, altered winter precipitation, and fire on desert grassland community structure and dynamics.

Global environmental change is altering temperature, precipitation patterns, resource availability, and disturbance regimes. Theory predicts that ecological presses will interact with pulse events to alter ecosystem structure and function. In 2006, we established a long-term, multifactor global change experiment to determine the interactive effects of nighttime warming, increased atmospheric nitrogen (N) deposition, and increased winter precipitation on plant community structure and aboveground net primary production (ANPP) in a northern Chihuahuan Desert grassland. In 2009, a lightning-caused wildfire burned through the experiment. Here, we report on the interactive effects of these global change drivers on pre- and postfire grassland community structure and ANPP. Our nighttime warming treatment increased winter nighttime air temperatures by an average of 1.1 °C and summer nighttime air temperature by 1.5 °C. Soil N availability was 2.5 times higher in fertilized compared with control plots. Average soil volumetric water content (VWC) in winter was slightly but significantly higher (13.0% vs. 11.0%) in plots receiving added winter rain relative to controls, and VWC was slightly higher in warmed (14.5%) compared with control (13.5%) plots during the growing season even though surface soil temperatures were significantly higher in warmed plots. Despite these significant treatment effects, ANPP and plant community structure were highly resistant to these global change drivers prior to the fire. Burning reduced the cover of the dominant grasses by more than 75%. Following the fire, forb species richness and biomass increased significantly, particularly in warmed, fertilized plots that received additional winter precipitation. Thus, although unburned grassland showed little initial response to multiple ecological presses, our results demonstrate how a single pulse disturbance can interact with chronic alterations in resource availability to increase ecosystem sensitivity to multiple drivers of global environmental change.

Beyond arctic and alpine: the influence of winter climate on temperate ecosystems.

Winter climate is expected to change under future climate scenarios, yet the majority of winter ecology research is focused in cold-climate ecosystems. In many temperate systems, it is unclear how winter climate relates to biotic responses during the growing season. The objective of this study was to examine how winter weather relates to plant and animal communities in a variety of terrestrial ecosystems ranging from warm deserts to alpine tundra. Specifically, we examined the association between winter weather and plant phenology, plant species richness, consumer abundance, and consumer richness in 11 terrestrial ecosystems associated with the U.S. Long-Term Ecological Research (LTER) Network. To varying degrees, winter precipitation and temperature were correlated with all biotic response variables. Bud break was tightly aligned with end of winter temperatures. For half the sites, winter weather was a better predictor of plant species richness than growing season weather. Warmer winters were correlated with lower consumer abundances in both temperate and alpine systems. Our findings suggest winter weather may have a strong influence on biotic activity during the growing season and should be considered in future studies investigating the effects of climate change on both alpine and temperate systems.

Grassland productivity limited by multiple nutrients.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited(1). Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)(2,3), the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized(4-8). However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+µ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+µ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

Above- and belowground responses to nitrogen addition in a Chihuahuan Desert grassland.

Increased available soil nitrogen can increase biomass, lower species richness, alter soil chemistry and modify community structure in herbaceous ecosystems worldwide. Although increased nitrogen availability typically increases aboveground production and decreases species richness in mesic systems, the impacts of nitrogen additions on semiarid ecosystems remain unclear. To determine how a semiarid grassland responds to increased nitrogen availability, we examined plant community structure and above- and belowground net primary production in response to long-term nitrogen addition in a desert grassland in central New Mexico, USA. Plots were fertilized annually (10 g N m(-2)) since 1995 and NPP measured from 2004 to 2009. Differences in aboveground NPP between fertilized and control treatments occurred in 2004 following a prescribed fire and in 2006 when precipitation was double the long-term average during the summer monsoon. Presumably, nitrogen only became limiting once drought stress was alleviated. Belowground NPP was also related to precipitation, and greatest root growth occurred the year following the wettest summer, decreasing gradually thereafter. Belowground production was unrelated to aboveground production within years and unrelated to nitrogen enrichment. Species richness changed between years in response to seasonal precipitation variability, but was not altered by nitrogen addition. Community structure did respond to nitrogen fertilization primarily through increased abundance of two dominant perennial grasses. These results were contrary to most nitrogen addition studies that find increased biomass and decreased species richness with nitrogen fertilization. Therefore, factors other than nitrogen deposition, such as fire or drought, may play a stronger role in shaping semiarid grassland communities than soil fertility.

Spatiotemporal dynamics of lianas during 50 years of succession to temperate forest.

Although they are important components of forest communities, the general ecology and spatiotemporal patterns of temperate lianas during forest regeneration are largely unknown. The dependence of lianas on other plants for physical support makes them a potentially important driver of community dynamics. We examined 50 years of vegetation data from an old-field succession study to determine the dynamics and community controls on liana expansion within the Piedmont region of New Jersey, USA. Four lianas, Lonicera japonica, Parthenocissus quinquefolia, Toxicodendron radicans, and Vitis spp., occurred in enough abundance for detailed analyses. In general, liana cover peaked during mid-succession (20-30 years post-abandonment) when community composition was mostly herbaceous with scattered trees and shrubs. Liana cover began to decrease as trees became dominant and the canopy closed. Temporal patterns of cover dynamics of abundant species indicated three early- and one late-successional liana species within the community. In contrast to cover, frequency of lianas increased throughout succession, indicating that liana populations persisted despite dramatic declines in cover for the three early-successional species. Temporal dynamics between native and nonnative lianas were similar but spatially distinct as cover of native species dispersed and expanded near the forest edge while the nonnative species preferentially grew far from the forest. These dynamics indicate that successional processes may ultimately lead to the decline of most lianas. However, the persistence of lianas as high numbers of suppressed individuals suggests that they may rebound quickly following canopy disturbance.

Simulated predator extinctions: predator identity affects survival and recruitment of oysters.

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.