Shrub Expansion Research Articles

Effects of fragmentation on cattle in African savannas under variable precipitation

Cattle move to access patches that vary in forage quantity and quality. Fragmentation can prevent animals from reaching patches. I used an integrative ecosystem model applied to three African landscapes to explore the sensitivity of cattle populations to fragmentation (here, changes in populations as parcel areas decreased) under different precipitation patterns. I hypothesized that low and high precipitation would yield populations relatively insensitive to fragmentation, intermediate precipitation would yield more sensitive populations, and more variable inter-annual precipitation would reduce sensitivity to fragmentation. Precipitation data were altered to yield averages of 100–1,000 mm year−1 and inter-annual coefficients of variation of 0–60%. A 1,000 km2 landscape in each area was divided into progressively smaller parcels and simulations conducted for each parcel. Rainfall at 100 mm year−1 supported low populations that were insensitive to fragmentation. Populations peaked at rainfall levels similar to those observed, and declined under higher precipitation, due in-part to shrub expansion. Fragmenting landscapes caused up to a 62% decline in cattle. High inter-annual variation in precipitation reduced sensitivity to fragmentation when precipitation was above that observed. The pattern was opposite when precipitation was below what was observed. Cattle on the landscape with fine-scale heterogeneity were relatively insensitive to fragmentation, and those on the heterogeneous but coarse-grained landscape were extremely sensitive. Fragmentation in landscapes where populations are sensitive will require more intensive inputs to offset losses, and changes in the frequency of extreme weather associated with climate change will alter the sensitivity of some populations to fragmentation.

Mineralization and carbon turnover in subarctic heath soil as affected by warming and additional litter

Arctic soil carbon (C) stocks are threatened by the rapidly advancing global warming. In addition to temperature, increasing amounts of leaf litter fall following from the expansion of deciduous shrubs and trees in northern ecosystems may alter biogeochemical cycling of C and nutrients. Our aim was to assess how factorial warming and litter addition in a long-term field experiment on a subarctic heath affect resource limitation of soil microbial communities (measured by thymidine and leucine incorporation techniques), net growing-season mineralization of nitrogen (N) and phosphorus (P), and carbon turnover (measured as changes in the pools during a growing-season-long field incubation of soil cores in situ). The mainly N limited bacterial communities had shifted slightly towards limitation by C and P in response to seven growing seasons of warming. This and the significantly increased bacterial growth rate under warming may partly explain the observed higher C loss from the warmed soil. This is furthermore consistent with the less dramatic increase in the contents of dissolved organic carbon (DOC) and dissolved organic N (DON) in the warmed soil than in the soil from ambient temperature during the field incubation. The added litter did not affect the carbon content, but it was a source of nutrients to the soil, and it also tended to increase bacterial growth rate and net mineralization of P. The inorganic N pool decreased during the field incubation of soil cores, especially in the separate warming and litter addition treatments, while gross mineralized N was immobilized in the biomass of microbes and plants transplanted into the incubates soil cores, but without any significant effect of the treatments. The effects of warming plus litter addition on bacterial growth rates and of warming on C and N transformations during field incubation suggest that microbial activity is an important control on the carbon balance of arctic soils under climate change.

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.

EXPERIMENTAL WARMING, NOT GRAZING, DECREASES RANGELAND QUALITY ON THE TIBETAN PLATEAU

We investigated experimental warming and simulated grazing (clipping) effects on rangeland quality, as indicated by vegetation production and nutritive quality, in winter-grazed meadows and summer-grazed shrublands on the Tibetan Plateau, a rangeland system experiencing climatic and pastoral land use changes. Warming decreased total aboveground net primary productivity (ANPP) by 40 g x m(-2) x yr(-1) at the meadow habitats and decreased palatable ANPP (total ANPP minus non-palatable forb ANPP) by 10 g x m(-2) x yr(-1) at both habitats. The decreased production of the medicinal forb Gentiana straminea and the increased production of the non-palatable forb Stellera chamaejasme with warming also reduced rangeland quality. At the shrubland habitats, warming resulted in less digestible shrubs, whose foliage contains 25% digestible dry matter (DDM), replacing more digestible graminoids, whose foliage contains 60% DDM. This shift from graminoids to shrubs not only results in lower-quality forage, but could also have important consequences for future domestic herd composition. Although warming extended the growing season in non-clipped plots, the reduced rangeland quality due to decreased vegetative production and nutritive quality will likely overwhelm the improved rangeland quality associated with an extended growing season. Grazing maintained or improved rangeland quality by increasing total ANPP by 20-40 g x m(-2) x yr(-1) with no effect on palatable ANPP. Grazing effects on forage nutritive quality, as measured by foliar nitrogen and carbon content and by shifts in plant group ANPP, resulted in improved forage quality. Grazing extended the growing season at both habitats, and it advanced the growing season at the meadows. Synergistic interactions between warming and grazing were present, such that grazing mediated the warming-induced declines in vegetation production and nutritive quality. Moreover, combined treatment effects were nonadditive, suggesting that we cannot predict the combined effect of global changes and human activities from single-factor studies. Our findings suggest that the rangelands on the Tibetan Plateau, and the pastoralists who depend on them, may be vulnerable to future climate changes. Grazing can mitigate the negative warming effects on rangeland quality. For example, grazing management may be an important tool to keep warming-induced shrub expansion in check. Moreover, flexible and opportunistic grazing management will be required in a warmer future.

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

AbstractQuestion: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover?Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney.Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m2 size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen).Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa.Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump-backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m2 size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump-backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump-backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co-variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.

Late Glacial and Holocene history of Plešné Lake and its surrounding landscape based on pollen and palaeoalgological analyses

Pollen analysis has been carried out on a 549 cm thick sediment profile from lake Plesne jezero (Plesne Lake) in the Bohemian Forest (Sumava, Czech Republic; 1090 m a.s.l.; 48°47′ N; 13°52′ E). Analyses of 67 samples characterise the development of the lake biotope and the surrounding landscape during the last ca. 14,000 years. The pollen diagram shows a very distinct transition between the Late Glacial and the Holocene biostratigraphic units at a depth of ca. 312 cm. In the surroundings of Plesne Lake the vegetation was treeless during the entire Late Glacial. The alpine tree limit, formed by Betula and Pinus with undergrowth of shrubs, might have been at ca. 500 m a.s.l. Pollen transported from long distances was significant due to the openness of the landscape, coming from southern Europe and even Africa, and including high numbers of Artemisia, Poaceae, Chenopodiaceae, and some other herbs and shrubs from steppe and forest-steppe areas in southern Europe or Africa (likely Ephedra, certainly Lygeum spartum). The expansion of shrubs, particularly Juniperus, preceded the expansion of trees near the end of the Late Glacial. Afforestation of the region by thin stands of Betula and Pinus occurred during the Preboreal. Significant warming in the Boreal resulted in the expansion of Corylus, Quercetum mixtum (QM) trees, and probably also Picea and Alnus. Picea as well as QM trees were further expanding during the Early Atlantic. Picea was the dominant tree during the Late Atlantic and Fagus started to spread towards its end. Abrupt expansion of Abies marks the Subboreal. A high degree of afforestation (Abies, Fagus, Picea) was characteristic for the Early Subatlantic. During Late Subatlantic, pollen of synanthropic plants appears. Phases of the lake biotope development were defined on the basis of coccal green algae and Isoetes.

Net Carbon Exchange Across the Arctic Tundra-Boreal Forest Transition in Alaska 1981–2000

Shifts in the carbon balance of high-latitude ecosystems could result from differential responses of vegetation and soil processes to changing moisture and temperature regimes and to a lengthening of the growing season. Although shrub expansion and northward movement of treeline should increase carbon inputs, the effects of these vegetation changes on net carbon exchange have not been evaluated. We selected low shrub, tall shrub, and forest tundra sites near treeline in northwestern Alaska, representing the major structural transitions expected in response to warming. In these sites, we measured aboveground net primary production (ANPP) and vegetation and soil carbon and nitrogen pools, and used these data to parameterize the Terrestrial Ecosystem Model. We simulated the response of carbon balance components to air temperature and precipitation trends during 1981–2000. In areas experiencing warmer and dryer conditions, Net Primary Production (NPP) decreased and heterotrophic respiration (R H ) increased, leading to a decrease in Net Ecosystem Production (NEP). In warmer and wetter conditions NPP increased, but the response was exceeded by an increase in R H ; therefore, NEP also decreased. Lastly, in colder and wetter regions, the increase in NPP exceeded a small decline in R H , leading to an increase in NEP. The net effect for the region was a slight gain in ecosystem carbon storage over the 20 year period. This research highlights the potential importance of spatial variability in ecosystem responses to climate change in assessing the response of carbon storage in northern Alaska over the last two decades.

The evidence for shrub expansion in Northern Alaska and the Pan‐Arctic

AbstractOne expected response to climate warming in the Arctic is an increase in the abundance and extent of shrubs in tundra areas. Repeat photography shows that there has been an increase in shrub cover over the past 50 years in northern Alaska. Using 202 pairs of old and new oblique aerial photographs, we have found that across this region spanning 620 km east to west and 350 km north to south, alder, willow, and dwarf birch have been increasing, with the change most easily detected on hill slopes and valley bottoms. Plot and remote sensing studies from the same region using the normalized difference vegetation index are consistent with the photographic results and indicate that the smaller shrubs between valleys are also increasing. In Canada, Scandinavia, and parts of Russia, there is both plot and remote sensing evidence for shrub expansion. Combined with the Alaskan results, the evidence suggests that a pan‐Arctic vegetation transition is underway. If continued, this transition will alter the fundamental architecture and function of this ecosystem with important ramifications for the climate, the biota, and humans.

Response of understory plant community of boreal mixedwood stands to fire, logging, and spruce budworm outbreak

The understory plant community of 63 boreal mixedwood stands in southeastern Manitoba, Canada, that were disturbed in the 1980s (21 by crown fire, 20 by logging, and 22 by severe spruce budworm ( Choristoneura fumiferana ) outbreak) were investigated. Understory plant communities that developed after fire, logging, and spruce budworm outbreak shared a large number of species (47 out of 81 identified). However, compared with fire and spruce budworm outbreak, logging promoted the rapid expansion of tall shrubs (52.7% vs. 17.2% and 29.75% coverage, respectively), particularly Corylus cornuta . Significant differences were also found between disturbance types when species coverage was compared on the basis of shade tolerance and regeneration strategy. Lower shrub coverage following fire is attributed to greater disturbance severity on the forest floor, affecting in situ propagules and competition from dense trembling aspen ( Populus tremuloides Michx.) regeneration. Lower shrub coverage following spruce budworm outbreak is attributed to slow opening of the canopy coupled with retention of a residual canopy of nonhost trees. Uniformly high shrub coverage following logging resulted in less heterogeneity and lower species diversity at both the stand and the disturbance level compared with natural disturbances. High shrub coverage may negatively impact conifer recruitment and have significant implications for future stand composition and productivity. Consideration of a natural insect outbreak model, rather than fire, for management of boreal mixedwood stands is recommended, particularly if stands are to be left to regenerate naturally.

A projection of severe near‐surface permafrost degradation during the 21st century

The current distribution and future projections of permafrost are examined in a fully coupled global climate model, the Community Climate System Model, version 3 (CCSM3) with explicit treatment of frozen soil processes. The spatial extent of simulated present‐day permafrost in CCSM3 agrees well with observational estimates – an area, excluding ice sheets, of 10.5 million km2. By 2100, as little as 1.0 million km2 of near‐surface permafrost remains. Freshwater discharge to the Arctic Ocean rises by 28% over the same period, largely due to increases in precipitation that outpace increases in evaporation, with about 15% of the rise directly attributable to melting ground ice. Such large changes in permafrost may provoke feedbacks such as activation of the soil carbon pool and a northward expansion of shrubs and forests.

Role of Land-Surface Changes in Arctic Summer Warming

A major challenge in predicting Earth's future climate state is to understand feedbacks that alter greenhouse-gas forcing. Here we synthesize field data from arctic Alaska, showing that terrestrial changes in summer albedo contribute substantially to recent high-latitude warming trends. Pronounced terrestrial summer warming in arctic Alaska correlates with a lengthening of the snow-free season that has increased atmospheric heating locally by about 3 watts per square meter per decade (similar in magnitude to the regional heating expected over multiple decades from a doubling of atmospheric CO2). The continuation of current trends in shrub and tree expansion could further amplify this atmospheric heating by two to seven times.

Changes in a hemlock-dominated forest following woolly adelgid infestation in southern New England1

Small, M. J., C. J. Small (Department of Botany, Connecticut College, New London, Connecticut 06320), and G. D. Dreyer (Goodwin-Niering Center for Conservation Biology and Environmental Studies, Connecticut College, New London, Connecticut 06320). Changes in a hemlock-dominated forest following woolly adelgid infestation in southern New England. J. Torrey Bot. 132: 458–470. 2005.—The hemlock wooly adelgid (HWA; Adelges tsugae), a small aphid-like insect introduced from Japan, has caused widespread hemlock (Tsuga canadensis) mortality throughout the mid-Atlantic and southern New England region over the last twenty years. We examined long-term (1952–2002) changes in hemlock-dominated stands before and after the appearance of the HWA in 1987 in the Connecticut College Arboretum, southeastern Connecticut. With HWA infestation, basal area of T. canadensis declined dramatically, dropping 70% from 1982 to 2002. Forest communities responded to the elimination of the dominant species by quickly filling various sized gaps. Black oaks (Quercus velutina, Q. coccinea, Q. rubra) increased from 28% of canopy basal area in 1982 to 41% in 2002. Sapling density increased markedly following HWA infestation, from 80 stems/ha in 1982 to nearly 5600 in 2002, with greatest increase in Sassafras albidum (0 to 1900 stems/ha) and Acer rubrum (4 to 1100 stems/ha). Ledge and ravine communities, formerly dominated by T. canadensis, became more compositionally distinct, with greater importance of black oaks on more xeric ledge sites and mixed canopy dominance in mesic ravine sites. Major trends associated with the decline of T. canadensis included a shift in canopy dominance to oak and mixed hardwoods, considerable understory development, including greater herb richness and abundance and increased density of clonal saplings, and expansion of several invasive shrubs and woody vines.

Beringia as a glacial refugium for boreal trees and shrubs: new perspectives from mapped pollen data

AbstractAim Beringia, far north‐eastern Siberia and north‐western North America, was largely unglaciated during the Pleistocene. Although this region has long been considered an ice‐age refugium for arctic herbs and shrubs, little is known about its role as a refugium for boreal trees and shrubs during the last glacial maximum (LGM,c. 28,000–15,000 calibrated years before present). We examine mapped patterns of pollen percentages to infer whether six boreal tree and shrub taxa (Populus,Larix,Picea,Pinus,Betula,Alnus/Duschekia) survived the harsh glacial conditions within Beringia.Methods Extensive networks of pollen records have the potential to reveal distinctive temporal–spatial patterns that discriminate between local‐ and long‐distance sources of pollen. We assembled pollen records for 149 lake, peat and alluvial sites from the Palaeoenvironmental Arctic Sciences database, plotting pollen percentages at 1000‐year time intervals from 21,000 to 6000 calibrated years before present. Pollen percentages are interpreted with an understanding of modern pollen representation and potential sources of long‐distance pollen during the glacial maximum. Inferences from pollen data are supplemented by published radiocarbon dates of identified macrofossils, where available.Results Pollen maps for individual taxa show unique temporal‐spatial patterns, but the data for each taxon argue more strongly for survival within Beringia than for immigration from outside regions. The first increase ofPopuluspollen percentages in the western Brooks Ranges is evidence thatPopulustrees survived the LGM in central Beringia. Both pollen and macrofossil evidence supportLarixsurvival in western Beringia (WB), but data forLarixin eastern Beringia (EB) are unclear. Given the similar distances of WB and EB to glacial‐age boreal forests in temperate latitudes of Asia and North America, the widespread presence ofPiceapollen in EB andPinuspollen in WB indicates thatPiceaandPinussurvived within these respective regions.Betulapollen is broadly distributed but highly variable in glacial‐maximum samples, suggesting thatBetulatrees or shrubs survived in restricted populations throughout Beringia.Alnus/Duschekiapercentages show complex patterns, but generally support a glacial refugium in WB.Main conclusions Our interpretations have several implications, including: (1) the rapid post‐glacial migration rate reported forPiceain western Canada may be over estimated, (2) the expansion of trees and shrubs within Beringia should have been nearly contemporaneous with climatic change, (3) boreal trees and shrubs are capable of surviving long periods in relatively small populations (at the lower limit of detection in pollen data) and (4) long‐distance migration may not have been the predominant mode of vegetation response to climatic change in Beringia.

Spatio-temporal changes of a nature protection area in the agricultural landscape of Jaeren, South-West Norway

A nature protected area within agricultural landscapes is analysed based on studies of landscape metrics, vegetation and soil, as well as measurements of the water and matter balance. Landscape metrics analyses show the influences of agriculture in shaping the landscape over the past 65 years. Increased separation between intensively used agricultural land and low-productive wet fen areas is found. Both individual patches and the total area of arable land have increased in size. Changes restricted to the nature reserve are identified as to sedimentation and vegetation succession. Expansion of reed and Salix shrubs profits from nutrient fluxes from arable land and drainage canals. Plant compositions differ with regard to the complex nutrient balance which is affected by hydrologic conditions. Scenarios are drawn which indicate that extensive land use and landscape management neither prevent sedimentation processes nor reverse the successive development. We conclude that future changes in bird species composition and occurrence will have to be anticipated. Nature protection in agricultural landscapes based on structural stability must implement management measures. The authors suggest an approach including dynamic systems to support decisions within the frame of a sustainable development of changing landscapes.

Remote sensing of vegetation and land-cover change in Arctic Tundra Ecosystems

The objective of this paper is to review research conducted over the past decade on the application of multi-temporal remote sensing for monitoring changes of Arctic tundra lands. Emphasis is placed on results from the National Science Foundation Land–Air–Ice Interactions (LAII) program and on optical remote sensing techniques. Case studies demonstrate that ground-level sensors on stationary or moving track platforms and wide-swath imaging sensors on polar orbiting satellites are particularly useful for capturing optical remote sensing data at sufficient frequency to study tundra vegetation dynamics and changes for the cloud prone Arctic. Less frequent imaging with high spatial resolution instruments on aircraft and lower orbiting satellites enable more detailed analyses of land cover change and calibration/validation of coarser resolution observations. The strongest signals of ecosystem change detected thus far appear to correspond to expansion of tundra shrubs and changes in the amount and extent of thaw lakes and ponds. Changes in shrub cover and extent have been documented by modern repeat imaging that matches archived historical aerial photography. NOAA Advanced Very High Resolution Radiometer (AVHRR) time series provide a 20-year record for determining changes in greenness that relates to photosynthetic activity, net primary production, and growing season length. The strong contrast between land materials and surface waters enables changes in lake and pond extent to be readily measured and monitored.

C3shrub expansion in a C4grassland: positive post‐fire responses in resources and shoot growth

Changes in land management and reductions in fire frequency have enabled woody species to increase in grasslands worldwide. Nevertheless, fire is rarely eliminated from grasslands, and for shrubs to survive, they must be able cope with fire and replace aboveground structures. Because new shoots may have more available solar radiation, greater root : shoot ratios, and thus more resources available belowground after fire compared to undisturbed shrub communities, we hypothesized that carbon, nutrient, and water relations may be enhanced in stems compared to those in an undisturbed grassland. However, this same post-fire resource pulse stimulates the grasses and may intensify competitive interactions between shrubs and grasses. To test these predictions, we measured seasonal patterns in net photosynthesis (A), predawn xylem pressure potentials (XPP), leaf nitrogen (N) content, and productivity of Cornus drummondii shoots from shrub patches (islands) of different sizes in mesic grasslands burned annually, burned infrequently, and protected from fire. Seasonal average A was 20% higher (P = 0.016) in burned than in unburned shrubs, regardless of island size. Shrubs in burned sites also produced shoots with higher leaf N than unburned shrubs, and N content was higher in leaves from small islands compared to large islands (P < 0.0001). Burning caused a decrease in late summer predawn XPP in small islands (-3.1 MPa), whereas burned large islands did not differ from unburned shrubs. Post-fire productivity of new shoots was significantly greater compared to shoots in unburned sites. These results indicate that a transient period of high resource availability after fire allows for increased growth and rapid recovery of grassland shrubs. Thus, although fire has a negative effect on aboveground biomass of shrubs, the post-fire increases in resource availability, which enhance growth in the dominant grasses, are also important for recovery of woody species.

Paleovegetation of marine isotope stages 4 and 3 in northern new zealand and the age of the widespread rotoehu tephra

Paleolake sediment, constrained by tephrochronology, from Onepoto basin volcanic crater in Auckland, Northern New Zealand (36° 48′S), provides one of the few uninterrupted records of paleovegetation for marine oxygen isotope stages (MIS) 4 and 3 (76,000–26,000 yr B.P.) in the region. This period was characterized by cool temperate conifer-hardwood forest that lacked some of the warmer taxa typical of the Holocene. The period 64,400–60,500 yr B.P. was marked by opening of forest canopy and expansion of small trees and shrubs, and correlates to the thermal minima of MIS 4. However, the landscape was never as open as the forest-shrubland mosaic of the MIS 2. The beginning of MIS 3 (60,500–50,500 yr B.P.) was marked by the dramatic expansion and then decline of conifer-hardwood forest dominated by Dacrydium cupressinum, a species that prefers wetter conditions. This forest was succeeded by the typically montane Nothofagus at 50,500 yr B.P., corresponding to a thermal decline. Thus, MIS 3 began with an abrupt change to moist cool conditions that lasted about 5000 yr, followed by gradual cooling and dryer conditions. This supports some interpretations from other parts of the southwest Pacific region, that MIS 3 was a period of increased precipitation. The widespread and stratigraphically important Rotoehu tephra, erupted from Okataina Volcanic Centre, has been variously dated at 45,000–65,000 yr B.P. At Onepoto, sedimentation rate and paleovegetation reconstruction imply an age of c. 44,300 yr B.P. The tephra provides a correlation horizon in the marine and terrestrial realms during a period (MIS 3) difficult to date by radiometric methods.

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract.In the mesic grasslands of the central United States, the shrubCornus drummondiihas undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused byC. drummondiiexpansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C4grassAndropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency inA. gerardiiand lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain inA. gerardiiin the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state ofC. drummondiidominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect ofC. drummondiidominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability.C. drummondiiremoval facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.

Long‐term patterns of shrub expansion in a C4‐dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.