Daniel L. Potts

Precipitation pulses and carbon fluxes in semiarid and arid ecosystems

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a “pulse” of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO2 is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.

Effects of seasonal drought on net carbon dioxide exchange from a woody‐plant‐encroached semiarid grassland

[1] Annual precipitation in the central and southern warm-desert region of North America is distributed climatologically between summer and winter periods with two prominent dry periods between them. We used energy and carbon dioxide (CO2) fluxes from eddy covariance along with standard meteorological and soil moisture measurements at a semiarid savanna in southern Arizona, United States, to better understand the consequences of warm or cool season drought on ecosystem CO2 exchange in these bimodally forced water-limited regions. Over the last 100 years, this historic grassland has converted to a savanna by the encroachment of the native mesquite tree (Prosopis velutina Woot.). During each of the 4 years of observation (2004–2007), annual precipitation (P) was below average, but monsoon (July–September) P was both above and below average while cool-season (December–March) P was always less than average by varying degrees. The ecosystem was a net source of CO2 to the atmosphere, ranging from 14 to 95 g C m � 2 yr � 1 with the strength of the source increasing with decreasing precipitation. When the rainfall was closest to the long-term average in its distribution and amount, the ecosystem was essentially carbon neutral. Summer drought resulted in increased carbon losses due mainly to a shortening of the growing season and the length of time later in the season when photosynthetic gain exceeds respiration loss. Severe cool season drought led to decreased spring carbon uptake and seemingly enhanced summer respiration, resulting in conditions that led to the greatest annual net carbon loss.

The sensitivity of ecosystem carbon exchange to seasonal precipitation and woody plant encroachment

Ongoing, widespread increases in woody plant abundance in historical grasslands and savannas (woody encroachment) likely will interact with future precipitation variability to influence seasonal patterns of carbon cycling in water-limited regions. To characterize the effects of woody encroachment on the sensitivity of ecosystem carbon exchange to seasonal rainfall in a semi-arid riparian setting we used flux-duration analysis to compare 2003-growing season NEE data from a riparian grassland and shrubland. Though less seasonally variable than the grassland, shrubland NEE was more responsive to monsoon rains than anticipated. During the 2004-growing season we measured leaf gas exchange and collected leaf tissue for δ13C and nitrogen content analysis periodically among three size classes of the dominant woody-plant, Prosopis velutina and the dominant understory species, Sporobolus wrightii, a C4 bunchgrass, present at the shrubland. We observed size-class and plant functional type independent patterns of seasonal plant performance consistent with greater-than-anticipated sensitivity of NEE in the shrubland. This research highlights the complex interaction between growing-season precipitation, plant-available alluvial groundwater and woody plant abundance governing ecosystem carbon balance in this semi-arid watershed.

SENSITIVITY OF MESQUITE SHRUBLAND CO2 EXCHANGE TO PRECIPITATION IN CONTRASTING LANDSCAPE SETTINGS

In semiarid ecosystems, physiography (landscape setting) may interact with woody-plant and soil microbe communities to constrain seasonal exchanges of material and energy at the ecosystem scale. In an upland and riparian shrubland, we examined the seasonally dynamic linkage between ecosystem CO2 exchange, woody-plant water status and photosynthesis, and soil respiration responses to summer rainfall. At each site, we compared tower-based measurements of net ecosystem CO2 exchange (NEE) with ecophysiological measurements among velvet mesquite (Prosopis velutina Woot.) in three size classes and soil respiration in sub-canopy and inter-canopy micro-sites. Monsoonal rainfall influenced a greater shift in the magnitude of ecosystem CO2 assimilation in the upland shrubland than in the riparian shrubland. Mesquite water status and photosynthetic gas exchange were closely linked to the onset of the North American monsoon in the upland shrubland. In contrast, the presence of shallow alluvial groundwater in the riparian shrubland caused larger size classes of mesquite to be physiologically insensitive to monsoonal rains. In both shrublands, soil respiration was greatest beneath mesquite canopies and was coupled to shallow soil moisture abundance. Physiography, through its constraint on the physiological sensitivity of deeply rooted woody plants, may interact with plant-mediated rates of soil respiration to affect the sensitivity of semiarid-ecosystem carbon exchange in response to episodic rainfall.

The impact of invasion and subsequent removal of an exotic thistle, Cynara cardunculus, on CO2 and H2O vapor exchange in a coastal California grassland

Changes in vegetation structure and composition, particularly due to the invasion of exotic species, are predicted to influence biosphere-atmosphere exchanges of mass and energy. Invasion of Cynara cardunculus (cardoon or artichoke thistle), a perennial, non-native thistle in coastal California grasslands presently dominated by non-native annual grasses, may alter rates of ecosystem CO2 exchange and evapotranspiration (ET). During spring and summer 2006, we compared midday maximum net ecosystem CO2 exchange (NEE) and ET among adjacent grassland plots where Cynara was present and where it was absent. Measurements of NEE supported the prediction that deeply-rooted Cynara increase midday ecosystem C-assimilation. Cynara-mediated shifts in NEE were associated with increases in ecosystem photosynthesis rather than changes in ecosystem respiration. Furthermore, the presence of Cynara was associated with increased ET during the growing season. An increase in aboveground live biomass (a proxy for leaf area) associated with Cynara invasion may underlie shifts in ecosystem CO2 and water vapor exchange. Following mid-growing season sampling during April, we removed Cynara from half of the Cynara-containing plots with spot applications of herbicide. Three weeks later, midday fluxes in removal plots were indistinguishable from those in plots where Cynara was never present suggesting a lack of biogeochemical legacy effects. Similar to woody-encroachment in some semi-arid ecosystems, Cynara invasion increases midday ecosystem CO2 assimilation and evapotranspiration rates and has the potential to increase C-storage in California coastal grasslands.

Canopy structure, photosynthetic capacity and nitrogen distribution in adjacent mixed and monospecific stands of Phragmites australis and Typha latifolia

Shifts in canopy structure associated with nonnative plant invasions may interact with species-specific patterns of canopy resource allocation to reinforce the invasion process. We documented differences in canopy light availability and canopy resource allocation in adjacent monospecific and mixed stands of Phragmites australis and Typha spp. in a Great Lakes coastal wetland presently undergoing Phragmites invasion to better understand how light availability influences leaf nitrogen content (Nmass) and photosynthetic capacity (Amax) in these species. Due to their horizontally oriented leaves, light attenuates more rapidly in monospecific stands of Phragmites than in monospecific stands of Typha, where leaves are more vertically-oriented. Whereas Typha canopies followed our prediction that patterns of Nmass and Amax should closely parallel patterns of canopy light availability, Nmass and Amax were consistent throughout Phragmites’ canopies. Moreover, we observed overall greater Nmass and lower photosynthetic nitrogen use efficiency in leaves of Phragmites than in leaves of Typha. Improved understanding of the link between Nmass and Amax in these canopies should improve our understanding of carbon and nitrogen cycling consequences of Phragmites invasion in wetland ecosystems.

Stream Structural Limitations on Invasive Communities in Urban Riparian Areas

Urban riparian plant communities exist at the interface between terrestrial and aquatic habitats, and they are rich sources of species biodiversity and ecosystem services. The periodic floods that promote species diversity in riparian plant communities also increase their vulnerability to nonnative plant invasions. Plant invasions are constrained by seed and suitable habitat availability. However, how seed dispersal and establishment limitations interact to shape nonnative plant invasions in riparian communities is poorly understood. We use Stream Visual Assessment Protocol data to evaluate the hydrological and geomorphological parameters that influence the seeding and establishment of six common nonnative species in urban riparian habitats: garlic mustard, purple loosestrife, reed canarygrass, common reed, Japanese knotweed, and multiflora rose. To address this objective, we analyzed stream reach data collected during a basin-wide environmental assessment of the extensively urbanized upper Niagara River watershed. We found limited support for our prediction that propagule limitation constrains the distribution of nonnative riparian species, likely because these species are well established in the study area. Instead, we found that opportune stream reach characteristics better predict the distribution of the common invasive riparian species—most notably open tree canopy. Given that there is widespread investment in urban riparian forest restoration to improve water quality, increase stream-bank stability, enhance wildlife habitat and promote recreation, our data suggest that riparian forests may provide the additional benefit of reducing the abundance of some, but not all, invasive plants. Nomenclature: Garlic mustard, Alliaria petiolata (Bieb.) Cavara & Grande; purple loosestrife, Lythrum salicaria L.; reed canarygrass, Phalaris arundinacea L.; common reed, Phragmites australis (Cav.) Trin. ex Steud.; Japanese knotweed, Polygonum cuspidatum Sieb. & Zucc.; multiflora rose, Rosa multiflora Thunb. ex Murr. Management Implications: The progression from individual plant invasions to the establishment of widespread exotic communities requires that potential invaders are available and that there is suitable habitat in which they can establish. Nonnative riparian plant communities are widespread, particularly in urban areas, and often share common species. We evaluated the hydrologic and geomorphologic parameters that might influence the urban riparian distributions of six common nonnative riparian species in the urbanized upper Niagara River watershed: garlic mustard, purple loosestrife, reed canarygrass, common reed, Japanese knotweed, and multiflora rose. We found that four of these species are well established in the region so that seed availability did not appear limiting; instead, we found riparian canopy cover the strongest limiting factor on the invasive riparian communities. These data suggest that limiting the distribution of invasive species in urban riparian habitats requires a straightforward approach: restore riparian forests. Forest restoration in urban/riparian habitats is a widespread management goal expected to create many ecological benefits, including improved habitat and water quality. Still, the stream–forest interface is inherently edge habitat, and invasive species thrive along forest edges. Increasing forest canopy may reduce invasive species communities, but the widespread presence of well-established riparian invaders and the mitigating effects of stream-bank edge habitat suggest that full eradication is unlikely. Moreover, we also found conflicting species-specific associations and habitat responses, suggesting that managing individual invasive species may require specific interventions.

Can local adaptation explain varying patterns of herbivory tolerance in a recently introduced woody plant in North America

Abstract Patterns of woody-plant mortality often reflect tradeoffs associated with resource allocation. Plants that allocate a high proportion of carbon acquired from photosynthesis to non-structural carbohydrate storage may be buffered from the synergistic effects of climate change and episodic disturbance.

Rural land use bifurcation in the urban-rural gradient

The classic urban-rural gradient concept assumes a decline in land use intensity from an intensively developed urban core outward to residential suburbs, culminating in lightly developed rural areas. The concept provides a common framework directing urban socio-ecological research. Given that rural land use includes woodlands and croplands, and croplands appear as ecologically depauperate as urban lands, we investigated land-use patterns along urban-rural gradients for 30 large metropolitan areas in the eastern United States. We predicted a bifurcation at the rural end of the gradient between woodland and cropland land use that does not correspond with human population density (expected to be relatively low in rural areas regardless of land use type). Our data indicated that ‘rural’ was a poor substitute for ‘natural’ as the rural end of the gradients bifurcated at the rural end between woodland and cropland – croplands being demonstratively poor ecological habitats. Indeed, we found that when defined by habitat quality, the habitat known to be biotically homogenized (urban and cropland) remained steady along the urban-rural gradient. Our results do not undermine the utility of the urban-rural gradient framework, but do suggest that the gradient and/or human population density do not necessarily indicate shifts in habitat quality.

Do novel weapons that degrade mycorrhizal mutualisms promote species invasion

Non-native plants often dominate novel habitats where they did not co-evolve with the local species. The novel weapons hypothesis suggests that non-native plants bring competitive traits against which native species have not adapted defenses. Novel weapons may directly affect plant competitors by inhibiting germination or growth, or indirectly by attacking competitor plant mutualists (degraded mutualisms hypothesis). Japanese knotweed (Fallopia japonica) and European buckthorn (Rhamnus cathartica) are widespread plant invaders that produce potent secondary compounds that negatively impact plant competitors. We tested whether their impacts were consistent with a direct effect on the tree seedlings (novel weapons) or an indirect attack via degradation of seedling mutualists (degraded mutualism). We compared recruitment and performance using three Ulmus congeners and three Betula congeners treated with allelopathic root macerations from allopatric and sympatric ranges. Moreover, given that the allelopathic species would be less likely to degrade their own fungal symbiont types, we used arbuscular mycorrhizal (AMF) and ectomycorrhizal (ECM) tree species to investigate the effects of F. japonica (no mycorrhizal association) and Rhamnus cathartica (ECM association) on the different fungal types. We also investigated the effects of F. japonica and R. cathartica exudates on AMF root colonization. Our results suggest that the allelopathic plant exudates impact seedlings directly by inhibiting germination and indirectly by degrading fungal mutualists. Novel weapons inhibited allopatric seedling germination but sympatric species were unaffected. However, seedling survivorship and growth appeared more dependent on mycorrhizal fungi, and mycorrhizal fungi were inhibited by allopatric species. These results suggest that novel weapons promote plant invasion by directly inhibiting allopatric competitor germination and indirectly by inhibiting mutualist fungi necessary for growth and survival.