Laura M. Ladwig

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. 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 recognized4–8. 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.

Developing an ecological context for allelopathy

There has been a renewed interest in allelopathy as a plant–plant interaction as more plant ecologists have become involved in studying biological invasions. This resurgence highlights a major deficiency in our understanding of allelopathy—the lack of a well-developed ecological context for the interaction. In contrast to allelopathy, the plant–plant interaction of competition has a strong theoretical foundation as well as a large body of supporting empirical studies. We suggest that the plant-herbivore defense literature provides a mature and well-developed framework from which a broader ecological context for allelopathy can be developed. Here, we discuss three broad classes of questions, drawn from the herbivore defense literature, which may help to develop an appropriate ecological context for allelopathy. These questions focus on (1) variation in allelopathic expression within species, (2) community level variation in allelopathy across species, and (3) variation in the impacts of allelopathy on associated species. Addressing such broad population and community level themes in a variety of systems will be necessary to fully develop an ecological context for allelopathy and provide a theoretical basis for understanding its role in plant communities.

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.

Liana host preference and implications for deciduous forest regeneration 1

Abstract Lianas have the potential to shape forest communities and alter forest regeneration. However, impacts of lianas on forest regeneration, particularly in temperate forests, are largely unstudied. To understand potential liana impacts on the community we need to first know the location and intensity of liana burdens on host trees. We examined liana-tree host preferences within a series of young regenerating deciduous forests in the Piedmont region of New Jersey, USA. Established trees (≥ 5 cm dbh) and the lianas associated with each tree were surveyed in 2008. The five most abundant liana species were Celastrus orbiculatus, Lonicera japonica, Parthenocissus quinquefolia, Toxicodendron radicans, and Vitis species. Host preference for each liana species was measured in two ways, as colonization on tree trunks and coverage in the canopy. Host preferences based on tree species and tree size were compared among liana species. A total of 798 trees were measured and lianas occurred on 64% of them. Host preferences were generally consistent between colonization and canopy expansion, suggesting the same factors that regulate establishment also regulate liana growth. Most liana species had higher colonization and greater canopy cover on early successional trees, particularly Juniperus virginiana. In contrast, Vitis spp. were more abundant on canopy hardwood trees. Slight preferences based on tree size were seen for some species. The preference of lianas for early successional trees may make lianas a contributing factor to the acceleration of succession within this eastern deciduous forest. However, the continued expansion of some lianas at the site, particularly Vitis spp. and C. orbiculatus, may alter future liana-tree associations and forest trajectories.

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.

Conditional allelopathic potential of temperate lianas

Allelopathy is often treated as an innate characteristic of a species rather than a phenotypically plastic trait that can vary with environmental conditions. Lianas are a highly competitive, phenotypically plastic life form that typically occur in both shaded and unshaded environments. As such, we hypothesized that temperate lianas may conditionally change allocation to allelopathic chemicals in response to light availability though the expected direction of change is unclear. Shading may reduce resource availability and therefore reduce allocation to allelochemicals, induce allelopathy as a competitive mechanism, or may not be related to allelopathy. To test the conditionality of allelopathy, sun and shade leaves of five common liana species (Toxicodendron radicans, Parthenocissus quinquefolia, Celastrus orbiculatus, Lonicera japonica, and Vitisvulpina) were collected from a young deciduous forest in New Jersey, USA, and tested with laboratory bioassays to detect allelopathic potential. All liana species showed allelopathic potential, and three species exhibited induction of increased allelopathic potential in shaded environments. The two species that were not shade induced are late successional lianas that persist for long periods in forest canopies. In contrast, the inducible lianas were early successional species that typically decline with canopy closure. This research indicates that lianas have the potential to be allelopathic and allelopathic potential conditionally responds to shading only for species that would normally be excluded from the forest canopy. As early successional lianas are present throughout forest regeneration in a range of light environments, allelopathic plasticity may increase their success by differentially allocating resources based on environmental conditions.

Soil enzyme responses to varying rainfall regimes in Chihuahuan Desert soils

Extracellular enzyme activities (EEAs) are indicators of both soil microbial activity and nutrient availability for plants. However, it is unclear how EEAs change over the growing season in desert grasslands. We examined whether EEAs changed in response to the size and frequency of rain events during the summer monsoon in the northern Chihuahuan Desert, and if the response varied between plant and interspace associated soils. Potential EEAs were measured within a rainfall manipulation experiment at the Sevilleta National Wildlife Refuge in central New Mexico, USA. Rainfall treatments included either three 10-mm events or one 30-mm rain event per month throughout the three month summer monsoon (July–September). EEAs were measured immediately before and within hours after experimental rain events under plants and in unvegetated interspaces. Throughout the season hydrolase activities were higher under vegetation than in interspace soils. Potential activities of hydrolytic enzymes were similar for the two rai...

Salt marsh stabilization affects algal primary producers at the marsh edge

As sea level rise and human activities erode coastal wetlands, managers rebuild or preserve wetlands that can perform the ecosystem services of a natural system. One increasingly common mitigation activity is the construction of rock sills in the low marsh zone to stabilize marsh elevation. Sills dramatically alter the physical structure of marshes by changing elevation, adding hard substrate and potentially altering the spatial structure of benthic algal communities in and adjacent to the low marsh. We documented differences in benthic algal abundance at the seaward marsh edge in silled and unsilled marshes in North Carolina. We found that sills were associated with reduced standing stocks of benthic algal primary production and reduced macroalgal taxonomic richness, and this difference was driven primarily by differences in macroalgal abundance. We experimentally tested the effect of macroalgal abundance on cordgrass (Spartina alterniflora) growth in the low zone of an unmanipulated marsh, and found that macroalgal removal had no effect on final cordgrass abundance. Our study suggests that salt marsh management through the construction of sills in low marsh zones impacts benthic primary production in the low marsh zone, but that benthic algal production does not affect cordgrass growth over a growing season.

Chihuahuan Desert Grassland Responds Similarly to Fall, Spring, and Summer Fires During Prolonged Drought

Abstract Land managers frequently use prescribed burning to help maintain grassland communities. Semiarid grassland dynamics following fire are linked to precipitation, with increasing soil moisture accelerating the rate of recovery. Prescribed fires are typically scheduled to follow natural fire regimes, but burning outside the natural fire season could be equally effective and more convenient for managers, depending on their management objectives. We conducted a field experiment in desert grassland to determine if fire seasonality influenced plant community recovery. Experimental burn treatments occurred in fall, spring, and summer in replicate 0.24-ha plots to determine if fire seasonality affected the rate of recovery of an ungrazed Chihuahuan Desert grassland in central New Mexico. Plant communities were surveyed seasonally for 5 yr after the burns. Grassland community structure responded to fire but not fire seasonality. Grass cover in all burned treatments remained lower than unburned controls for 3 yr after the burns. Community change through time was largely influenced by low rainfall, as grass cover in burned and unburned communities converged during a year with severe drought. In conclusion, fire seasonality did not influence rate of community recovery, but extended drought was possibly more influential than fire on grassland dynamics.