Daniel R. Schlaepfer

Are invaders different? A conceptual framework of comparative approaches for assessing determinants of invasiveness

What determines invasiveness of alien organisms is among the most interesting and urgent questions in ecology. In attempts to answer this question, researchers compare invasive alien species either to native species or to non-invasive alien species, and this is done in either the introduced or native ranges. However, inferences that can be drawn from these comparisons differ considerably, and failure to recognize this could hamper the search for determinants of invasiveness. To increase awareness about this issue, we present a framework of the various comparisons that can be used to test for determinants of invasiveness, and the specific questions each comparison can address. Moreover, we discuss how different comparisons complement each other, and therefore should be used in concert. For progress in invasion biology, it is crucial to realize that different comparisons address different biological questions and that some questions can only be answered unambiguously by combining them.

A multi-species experiment in their native range indicates pre-adaptation of invasive alien plant species.

*To understand prerequisites of biological invasions, it is imperative to know whether species have traits that pre-adapt them to become invasive elsewhere. However, few experimental studies have explicitly tested this by comparing traits between invasive and noninvasive species in their native range instead of in the nonnative range. *We used native plant material of 14 European congeneric pairs of herbaceous species that were all introduced to North America, and of which one species per pair is invasive. *In our germination and common garden experiment with and without fertilizer addition, the invasive species germinated faster, produced more biomass and had a higher proportion of flowering plants than the noninvasive congeners. *Our results indicate that species traits, which lead to a high plant performance in the native range, can confer pre-adaptation to become invasive. We suggest that such traits may be especially relevant for use in risk-assessment protocols before introduction elsewhere.

Why only tetraploid Solidago gigantea (Asteraceae) became invasive: a common garden comparison of ploidy levels.

Many studies have compared the growth of plants from native and invasive populations, but few have considered the role of ploidy. In its native range in North America, Solidago gigantea Aiton (Asteraceae) occurs as a diploid, tetraploid and hexaploid, with considerable habitat differentiation and geographic separation amongst these ploidy levels. In the introduced range in Europe, however, only tetraploid populations are known. We investigated the growth performance and life history characteristics of plants from 12 European and 24 North American (12 diploid, 12 tetraploid) populations in a common garden experiment involving two nutrient and two calcium treatments. Twelve plants per population were grown in pots for two seasons. We measured 24 traits related to leaf nutrients, plant size, biomass production and phenology as well as sexual and vegetative reproduction. Native diploid plants had a higher specific leaf area and higher leaf nutrient concentrations than native tetraploids, but tetraploids produced many more shoots and rhizomes. Diploids grown with additional calcium produced less biomass, whereas tetraploids were not affected. European plants were less likely to flower and produced smaller capitulescences than North American tetraploids, but biomass production and shoot and rhizome number did not differ. We conclude that a knowledge of ploidy level is essential in comparative studies of invasive and native populations. While clonal growth is important for the invasion success of tetraploid S. gigantea, its potential was not acquired by adaptation after introduction but by evolutionary processes in the native range.

Terrestrial water fluxes dominated by transpiration: Comment

The fraction of evapotranspiration (ET) attributed to plant transpiration (T) is an important source of uncertainty in terrestrial water fluxes and land surface modeling (Lawrence et al. 2007, Miralles et al. 2011). Jasechko et al. (2013) used stable oxygen and hydrogen isotope ratios from 73 large lakes to investigate the relative roles of evaporation (E) and T in ET from their catchments and to estimate global T/ET. Two recent contributions have addressed data issues with their work highlighting uncertainty propagation (Coenders-Gerrits et al. 2014) and global interception estimates (Coenders-Gerrits et al. 2014, Schlesinger and Jasechko 2014). We address here the underlying assumptions of the model applied by Jasechko et al. (2013). They assumed that the isotope ratios of the lake water incorporate spatially integrated fractionation effects of total E and T over the entire catchment. This assumption is invalid and leads to substantial errors both for catchment-scale and global estimates of T/ET.

Climate change reduces extent of temperate drylands and intensifies drought in deep soils

Drylands cover 40% of the global terrestrial surface and provide important ecosystem services. While drylands as a whole are expected to increase in extent and aridity in coming decades, temperature and precipitation forecasts vary by latitude and geographic region suggesting different trajectories for tropical, subtropical, and temperate drylands. Uncertainty in the future of tropical and subtropical drylands is well constrained, whereas soil moisture and ecological droughts, which drive vegetation productivity and composition, remain poorly understood in temperate drylands. Here we show that, over the twenty first century, temperate drylands may contract by a third, primarily converting to subtropical drylands, and that deep soil layers could be increasingly dry during the growing season. These changes imply major shifts in vegetation and ecosystem service delivery. Our results illustrate the importance of appropriate drought measures and, as a global study that focuses on temperate drylands, highlight a distinct fate for these highly populated areas.

Natural Regeneration Processes in Big Sagebrush (Artemisia tridentata)

Abstract Big sagebrush, Artemisia tridentata Nuttall (Asteraceae), is the dominant plant species of large portions of semiarid western North America. However, much of historical big sagebrush vegetation has been removed or modified. Thus, regeneration is recognized as an important component for land management. Limited knowledge about key regeneration processes, however, represents an obstacle to identifying successful management practices and to gaining greater insight into the consequences of increasing disturbance frequency and global change. Therefore, our objective is to synthesize knowledge about natural big sagebrush regeneration. We identified and characterized the controls of big sagebrush seed production, germination, and establishment. The largest knowledge gaps and associated research needs include quiescence and dormancy of embryos and seedlings; variation in seed production and germination percentages; wet-thermal time model of germination; responses to frost events (including freezing/thawing of soils), CO2 concentration, and nutrients in combination with water availability; suitability of microsite vs. site conditions; competitive ability as well as seedling growth responses; and differences among subspecies and ecoregions. Potential impacts of climate change on big sagebrush regeneration could include that temperature increases may not have a large direct influence on regeneration due to the broad temperature optimum for regeneration, whereas indirect effects could include selection for populations with less stringent seed dormancy. Drier conditions will have direct negative effects on germination and seedling survival and could also lead to lighter seeds, which lowers germination success further. The short seed dispersal distance of big sagebrush may limit its tracking of suitable climate; whereas, the low competitive ability of big sagebrush seedlings may limit successful competition with species that track climate. An improved understanding of the ecology of big sagebrush regeneration should benefit resource management activities and increase the ability of land managers to anticipate global change impacts.

Ecohydrology of Adjacent Sagebrush and Lodgepole Pine Ecosystems: The Consequences of Climate Change and Disturbance

Sagebrush steppe and lodgepole pine forests are two of the most widespread vegetation types in the western United States and they play crucial roles in the hydrologic cycle of these water-limited regions. We used a process-based ecosystem water model to characterize the potential impact of climate change and disturbance (wildfire and beetle mortality) on water cycling in adjacent sagebrush and lodgepole pine ecosystems. Despite similar climatic and topographic conditions between these ecosystems at the sites examined, lodgepole pine, and sagebrush exhibited consistent differences in water balance, notably more evaporation and drier summer soils in the sagebrush and greater transpiration and less water yield in lodgepole pine. Canopy disturbances (either fire or beetle) have dramatic impacts on water balance and availability: reducing transpiration while increasing evaporation and water yield. Results suggest that climate change may reduce snowpack, increase evaporation and transpiration, and lengthen the duration of dry soil conditions in the summer, but may have uncertain effects on drainage. Changes in the distribution of sagebrush and lodgepole pine ecosystems as a consequence of climate change and/or altered disturbance regimes will likely alter ecosystem water balance.

Ecohydrology of Dry Regions: Storage versus Pulse Soil Water Dynamics

Although arid and semiarid regions are defined by low precipitation, the seasonal timing of temperature and precipitation can influence net primary production and plant functional type composition. The importance of precipitation seasonality is evident in semiarid areas of the western U.S., which comprise the Intermountain (IM) zone, a region that receives important winter precipitation and is dominated by woody plants and the Great Plains (GP), a region that receives primarily summer precipitation and is dominated by perennial grasses. Although these general relationships are well recognized, specific differences in water cycling between these regions have not been well characterized. We used a daily time step soil water simulation model and twenty sites from each region to analyze differences in soil water dynamics and ecosystem water balance. IM soil water patterns are characterized by storage of water during fall, winter, and spring resulting in relatively reliable available water during spring and early summer, particularly in deep soil layers. By contrast, GP soil water patterns are driven by pulse precipitation events during the warm season, resulting in fluctuating water availability in all soil layers. These contrasting patterns of soil water—storage versus pulse dynamics—explain important differences between the two regions. Notably, the storage dynamics of the IN sites increases water availability in deep soil layers, favoring the deeper rooted woody plants in that region, whereas the pulse dynamics of the Great Plains sites provide water primarily in surface layers, favoring the shallow-rooted grasses in that region. In addition, because water received when plants are either not active or only partially so is more vulnerable to evaporation and sublimation than water delivered during the growing season, IM ecosystems use a smaller fraction of precipitation for transpiration (47%) than GP ecosystems (49%). Recognizing the pulse-storage dichotomy in soil water regimes between the IM and GP regions may be useful for understanding the potential influence of climate changes on soil water patterns and resulting dominant plant functional groups in both regions.

Shifts in plant functional types have time-dependent and regionally variable impacts on dryland ecosystem water balance

Summary 1. Terrestrial vegetation influences hydrologic cycling. In water-limited, dryland ecosystems, altered ecohydrology as a consequence of vegetation change can impact vegetation structure, ecological functioning and ecosystem services. Shrub steppe ecosystems dominated by big sagebrush (Artemisia tridentata) are widespread across western North America, and provide a range of ecosystem services. While sagebrush abundance in these ecosystems has been altered over the past century, and changes are likely to continue, the ecohydrological consequences of sagebrush removal and reestablishment remain unclear. 2. To characterize the immediate and medium-term patterns of water cycling and availability following sagebrush plant community alteration, we applied the SOILWAT ecosystem water balance model to 898 sites across the distribution of sagebrush ecosystems, representing the three primary sagebrush ecosystem types: sagebrush shrublands, sagebrush steppe and montane sagebrush. At each site, we examined three vegetation conditions representing intact sagebrush, recently disturbed sagebrush and recovered but grass-dominated vegetation. 3. Transition from shrub to grass dominance decreased precipitation interception and transpiration and increased soil evaporation and deep drainage. Relative to intact sagebrush vegetation, simulated soils in the herbaceous vegetation phases typically had drier surface layers and wetter deep layers. 4. Our simulations suggested that alterations in ecosystem water balance may be most pronounced in vegetation representing recently disturbed conditions (herbaceous vegetation with low biomass) and only modest in conditions representing recovered, but still grass-dominated vegetation. Furthermore, the ecohydrological impact of simulated sagebrush removal depended on climate; while short-term changes in water balance were greatest in wet areas represented by the montane sagebrush ecosystem type, medium-term impacts were greatest in dry areas of sagebrush shrublands and sagebrush steppe. 5. Synthesis. This study provides a novel, regional-scale assessment of how plant functional type transitions may impact ecosystem water balance in sagebrush-dominated ecosystems of North America. Results illustrate that the ecohydrological consequences of changing vegetation depend strongly on climate and suggest that decreasing woody plant abundance may have only limited impact on evapotranspiration and water yield.

Mid‐latitude shrub steppe plant communities: climate change consequences for soil water resources

In the coming century, climate change is projected to impact precipitation and temperature regimes worldwide, with especially large effects in drylands. We use big sagebrush ecosystems as a model dryland ecosystem to explore the impacts of altered climate on ecohydrology and the implications of those changes for big sagebrush plant communities using output from 10 Global Circulation Models (GCMs) for two representative concentration pathways (RCPs). We ask: (1) What is the magnitude of variability in future temperature and precipitation regimes among GCMs and RCPs for big sagebrush ecosystems, and (2) How will altered climate and uncertainty in climate forecasts influence key aspects of big sagebrush water balance? We explored these questions across 1980-2010, 2030-2060, and 2070-2100 to determine how changes in water balance might develop through the 21st century. We assessed ecohydrological variables at 898 sagebrush sites across the western US using a process-based soil water model, SOILWAT, to model all components of daily water balance using site-specific vegetation parameters and site-specific soil properties for multiple soil layers. Our modeling approach allowed for changes in vegetation based on climate. Temperature increased across all GCMs and RCPs, whereas changes in precipitation were more variable across GCMs. Winter and spring precipitation was predicted to increase in the future (7% by 2030-2060, 12% by 2070-2100), resulting in slight increases in soil water potential (SWP) in winter. Despite wetter winter soil conditions, SWP decreased in late spring and summer due to increased evapotranspiration (6% by 2030-2060, 10% by 2070-2100) and groundwater recharge (26% and 30% increase by 2030-2060 and 2070-2100). Thus, despite increased precipitation in the cold season, soils may dry out earlier in the year, resulting in potentially longer, drier summer conditions. If winter precipitation cannot offset drier summer conditions in the future, we expect big sagebrush regeneration and survival will be negatively impacted, potentially resulting in shifts in the relative abundance of big sagebrush plant functional groups. Our results also highlight the importance of assessing multiple GCMs to understand the range of climate change outcomes on ecohydrology, which was contingent on the GCM chosen.