Bibliography - Grassland

To determine how grassland, woodland, and bordering forests respond to increased aridity, we used paleoecological methods to examine past responses along a transect of three sites at the eastern boundary of the Northern Plains of North America. Our study region corresponds to the confluence of three air streams that control central North American climates and, hence, should be sensitive to climate change. Sediment cores were analyzed for evidence of Holocene vegetation and fire from tall-grass prairie in eastern North Dakota (Moon Lake), from mixed forest near the prairie border in northwestern Minnesota (Deming Lake), and from mixed forest more remote from prairie in western Wisconsin (Dark Lake). Together with pollen and charcoal analysis, we present a new method for determining δ13C of terrestrial (charred) organic matter and, thus, the relative importance of C3 and C4 photosynthetic pathways in past vegetation. Paleorecords were supplemented with surface charcoal accumulation and δ13C from 21 North American lakes that span boreal, deciduous, pine, and mixed forest to tall- and mixed-grass prairie.Surface charcoal and δ13C follow vegetation and climate gradients, with high charcoal accumulation and δ13C (-20%25) in the Plains (Dakotas, Nebraska, and southwest Minnesota) and decreases to the east, west, and north. The δ13C pattern is consistent with observed patterns of C3:C4 dominance across the region. Sediment, pollen, charcoal, and terrestrial δ13C show that vegetation response to climate change varied substantially among tall-grass prairie, bordering woodland, and forest. During maximum aridity (8000;nd4000 yr BP) prairie vegetation in eastern North Dakota showed a demise of woody vegetation followed by a fluctuating dominance of grasses (40% C4) and forbs. Meanwhile, prairie expanded eastward into northwestern Minnesota, where it produced a shifting dominance between mostly C4 grasses and woody vegetation until more humid conditions and mixed forest developed after 4000 yr BP. Mixed forest in southwestern Wisconsin showed little response to mid-Holocene aridity. Elevated δ13C values from 5000 to 3000 yr BP suggest that composition of grasses changed (to increased C4), although pollen data indicate that the total abundance of grasses remained constant. The increase in C4 grasses at this time is consistent with previous studies suggesting a delayed dry interval in eastern Iowa. Reduced aridity of the last 2000 yr brought increased fire to tall-grass prairie as higher primary productivity led to increased fuel load. Meanwhile, forest expanded in northwestern Minnesota, leading to decreased ignition and fine fuels, in turn resulting in decreased fire at the woodland margin.

In water-limited ecosystems, discrete precipitation events trigger brief but important episodes of biological activity. Differential responses of above- and below-ground biota to precipitation may constrain biogeochemical transformations at the ecosystem scale. We quantified short-term dynamics of whole ecosystem response to 39 mm irrigation events (precipitation pulses) during June 2002 and 2003 using plant physiological and ecosystem gas-exchange measurements as state variables in a principal components analysis (PCA). Experimental plots consisted of either native (Heteropogon contortus L.) or non-native (Eragrostis lehmanniana Nees) bunchgrasses planted in monoculture on two distinct geomorphic surfaces in a semi-arid grassland. For 15 days, treatments followed similar, non-linear trajectories through state variable space with measurement periods forming distinct clusters; PCA axes 1 and 2 combined to explain 80.7% of the variation during both 2002 and 2003. During both years, bunchgrass species interacted with soil type such that there was a reduction in ecosystem functional resistance in plots planted with the non-native bunchgrass species on the fine-textured clay geomorphic surface. System-level hysteresis, emerging as a result of independent responses of photosynthesis, respiration and evapotranspiration to precipitation, indicated the potential for alternative functional states. Quantifying the frequency and duration of ecosystem alternative functional states in response to individual precipitation events within a season will provide insights into the controls of species, soils and climate on ecosystem carbon and water cycles.

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.

The prairie-forest transition in midcontinental North America is a major physiognomic boundary, and its shifts during the Holocene are a classic example of climate-driven ecotonal dynamics. Recent work suggests asymmetrical Holocene behavior, with a relatively rapid early Holocene deforestation and more gradual reforestation later in the Holocene. This paper presents a new synthesis of the Holocene history of the Great Plains prairie-forest ecotone in the north-central US and central Canada that updates prior mapping efforts and systematically assesses rates of change. Changes in percent woody cover (%WC) are inferred from fossil pollen records, using the modern analog technique and surface-sediment pollen samples cross-referenced against remotely sensed observations. For contemporary pollen samples from the Great Plains, %WC linearly correlates to percent arboreal pollen (%AP), but regression parameters vary interregionally. At present, %AP is consistently higher than %WC, because of high background levels of arboreal pollen. Holocene maps of the eastern prairie-forest ecotone agree with prior maps, showing a rapid decrease in %WC and eastward prairie advance between 10,000 and 8000 ka (1 ka = 1000 calibrated years before present), a maximum eastward position of the ecotone from 7 to 6 ka, and increased %WC and westward prairie retreat after 6 ka. Ecotone position is ambiguous in Iowa and southeastern Minnesota, due to a scarcity of modern analogs for early-Holocene samples with high Ulmus abundances and for samples from alluvial sediments. The northern prairie-forest ecotone was positioned in central Saskatchewan between 12 and 10 ka, stabilized from 10 to 6 ka despite decreases in %WC at some sites, then moved south after 6 ka. In both east and north, ecotonal movements are consistent with a dry early Holocene and increasing moisture availability after 6 ka. Sites near the ecotone consistently show an asymmetric pattern of abrupt early Holocene deforestation ( < 300 years) and gradual reforestation after 6 ka. Early Holocene decreases in %WC are faster than the corresponding drops in %AP, because the analog-based %WC reconstructions correct for the high background levels of arboreal pollen types that blur temporal variations in %AP. For example, at Elk Lake, the %AP decline lasts 1000 years, whereas the %WC decline occurs between adjacent pollen samples, approximately 300 years apart. Thus, early Holocene deforestation may have been even more abrupt than previously recognized. Rapid deforestation likely was promoted both by rapid climate changes around 8.2 ka and positive fire-vegetation feedbacks. Non-linear vegetational responses to hydrological variability are consistent with 1) other paleorecords showing rapid die-offs of some eastern tree species in response to aridity and 2) observations of threshold-type ecological responses to recent climate events. The 21st-century trajectory for the Great Plains prairie-forest ecotone is uncertain, because climate models differ over the direction of regional precipitation trends, but future drying would be more likely to trigger threshold-type shifts in ecotone position.

Aboveground net primary production (ANPP) is strongly correlated with annual precipitation (AP) in grassland ecosystems. However, the relationship between the interannual variation in ANPP and the variability in precipitation remains controversial. In this study, we used long-term data of biomass and precipitation from 118 sites across global grasslands to examine the relationship between variability in ANPP and AP. Our results showed that ANPP increased with precipitation, but leveled off in humid regions, and that increased variation in precipitation led to an increase in the variability in ANPP. The relative ANPP maxima significantly increased with relative AP maxima and the relative ANPP minima also positively correlated with relative AP minima. These suggest that the fluctuations in precipitation can alter the growth of grasslands, which should be incorporated into the prediction and modeling of climate changes.