Restoration Ecology of an Endangered Plant Species: Establishment of New Populations of Cirsium pitcheri

We determined the effects of shade, burial by sand, simulated herbivory, and fertilizers on the survival and growth of artificially planted population of Cirsium pitcheri—an endangered plant species of the sand dunes along Lake Huron. Sand burial experiments showed that greenhouse grown plants should optimally be transplanted into areas receiving 5 cm of sand deposition: burial at this depth maximized emergence, survivorship, and below‐ground biomass. Under field conditions, simulated herbivory of up to 50% of the plant height produced a slight increase in biomass after one year of growth. Field observations showed that when white‐tailed deer removed more than 50% of the transplant's leaf tissue, the plant died. The application of a 20:20:20 (N:P:K) water‐soluble fertilizer produced a significant increase in the dry leaf biomass, total leaf area, and total dry biomass relative to control plants. We also tested for the presence or absence of a persistent seed bank. Few seeds were recovered from soil samples collected from Pinery Provincial Park and Providence Bay. However, C. pitcheri has the ability to form a persistent seed bank under field conditions but only at soil depths of 15 cm. Cirsium pitcheri seeds are able to germinate and seedlings can emerge from a burial depth of up to 6 cm. Thus, seeds planted in open, sunny areas will probably maximize emergence, growth, and survivorship of seedlings. Populations of C. pitcheri can be restored by planting seeds at shallow depths, transplanting greenhouse‐grown plants, applying water soluble fertilizers, and protecting plants from herbivores.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.     

Patch identity and the spatial heterogeneity of woody encroachment in exotic-dominated old-field grasslands

Most grassland communities in agricultural landscapes comprise a mix of exotic and native plants, where grasses and forbs are disposed in low diversity patches conforming a heterogeneous matrix of vegetation. Within these “novel” ecosystems, woody encroachment is one of the principal causes of ecosystem degradation. Here, we examined the resistance to exotic woody establishment (Gleditsia triacanthos) into four different monospecific patches characteristics of old-field grasslands in Inland Pampa: an annual forb (Conium maculatum), an annual grass (Lolium multiflorum), and two perennial grasses (Cynodon dactylon and Festuca arundinacea). We evaluated the filter to tree recruitment by rodent seed removal and survival and growth of Gleditsia seedlings transplanted into undisturbed and disturbed microsites, within each patch. Beneath intact vegetation seed removal was an important biotic filter to woody establishment whereas disturbances facilitated seed survival in patches of perennial grasses. Patch identity affected tree growth, and Cynodon reduced the final biomass compared to forbs. Disturbance enhanced tree performance independently of patch type. After 2 years, tree survival was independent of disturbance and patch identity. As patch identity may regulate granivory and growth of tree saplings, community susceptibility or resistance to woody invasion rather than representing a static community attribute could vary according to the dynamic changes in the proportion of susceptible-resistant patches. Broadly, our work reinforces the concept that mechanisms regulating vegetation heterogeneity add a component of stochasticity to biotic resistance to community plant invasion.     

A native nitrogen-fixing shrub facilitates weed invasion

Invasions by exotic weedy plants frequently occur in highly disturbed or otherwise anthropogenically altered habitats. Here we present evidence that, within California coastal prairie, invasion also can be facilitated by a native nitrogen-fixing shrub, bush lupine (Lupinus arboreus). Bush lupines fix nitrogen and grow rapidly, fertilizing the sandy soil with nitrogen-rich litter. The dense lupine canopy blocks light, restricting vegetative growth under bushes. Heavy insect herbivory kills lupines, opening exposed nitrogen-rich sites within the plant community. Eventual re-establishment of lupine occurs because of an abundant and long-lived seed bank. Lupine germination, rapid growth, shading and fertilization of sites, and then death after only a few years, results in a mosaic of nutrient-rich sites that are available to invading species. To determine the role of bush lupine death and nitrogen enrichment in community composition, we examined nutrient dynamics and plant community characteristics within a site only recently colonized by lupine, comparing patches where lupines had recently died or were experimentally killed with adjacent areas lacking lupine. In experimentally killed patches, instantaneous pool sizes of exchangeable ammonium and nitrate nitrogen were higher than in adjacent sites free of lupine. Seedlings of the introduced grass Bromus diandrus accumulated 48% greater root biomass and 93% more shoot biomass when grown in a greenhouse in soil collected under experimentally killed lupines compared to B. diandrus seedlings grown in soil collected at least 1 m away from lupines. At the end of the spring growing season, total above-ground live plant biomass was more than twice as great in dead lupine patches as in the adjacent lupine-free grassland, but dead lupine patches contained 47% fewer plant species and 57% fewer native species. Sites where lupines have repeatedly died and reestablished during recent decades support an interstitial grassland community high in productivity but low in diversity, composed of mostly weedy introduced annual plants. In contrast, at a site only recently colonized by bush lupines, the interstitial grassland consists of a less productive but more diverse set of native and introduced species. We suggest that repeated bouts of lupine germination, establishment, and death can convert a rich native plant community into a less diverse collection of introduced weeds.     

Linking the pattern to the mechanism: How an introduced mammal facilitates plant invasions

Non‐native mammals that are disturbance agents can promote non‐native plant invasions, but to date there is scant evidence on the mechanisms behind this pattern. We used wild boar (Sus scrofa) as a model species to evaluate the role of non‐native mammals in promoting plant invasion by identifying the degree to which soil disturbance and endozoochorous seed dispersal drive plant invasions. To test if soil disturbance promotes plant invasion, we conducted an exclosure experiment in which we recorded emergence, establishment and biomass of seedlings of seven non‐native plant species planted in no‐rooting, boar‐rooting and artificial rooting patches in Patagonia, Argentina. To examine the role of boar in dispersing seeds we germinated viable seeds from 181 boar droppings and compared this collection to the soil seed bank by collecting a soil sample adjacent to each dropping. We found that both establishment and biomass of non‐native seedlings in boar‐rooting patches were double those in no‐rooting patches. Values in artificial rooting patches were intermediate between those in boar‐rooting and no‐rooting treatments. By contrast, we found that the proportion of non‐native seedlings in the soil samples was double that in the droppings, and over 80% of the germinated seeds were native species in both samples. Lastly, an effect size test showed that soil disturbance by wild boar rather than endozoochorous dispersal facilitates plant invasions. These results have implications for both the native and introduced ranges of wild boar, where rooting disturbance may facilitate community composition shifts.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

Comparative responses of model C3 and C4 plants to drought in low and elevated CO2

Interactive effects of CO₂ and water availability have been predicted to alter the competitive relationships between C3 and C4 species over geological and contemporary time scales. We tested the effects of drought and CO₂ partial pressures (pCO₂) ranging from values of the Pleistocene to those predicted for the future on the physiology and growth of model C3 and C4 species. We grew co-occurring Abutilon theophrasti (C3) and Amaranthus retroflexus (C4) in monoculture at 18 (Pleistocene), 27 (preindustrial), 35 (current), and 70 (future) Pa CO₂ under conditions of high light and nutrient availability. After 27 days of growth, water was withheld from randomly chosen plants of each species until visible wilting occurred. Under well-watered conditions, low pCO₂ that occurred during the Pleistocene was highly limiting to C3 photosynthesis and growth, and C3 plants showed increased photosynthesis and growth with increasing pCO₂ between the Pleistocene and future CO₂ values. Well-watered C4 plants exhibited increased photosynthesis in response to increasing pCO₂, but total mass and leaf area were unaffected by pCO₂. In response to drought, C3 plants dropped a large amount of leaf area and maintained relatively high leaf water potential in remaining leaves, whereas C4 plants retained greater leaf area, but at a lower leaf water potential. Furthermore, drought-treated C3 plants grown at 18 Pa CO₂ retained relatively greater leaf area than C3 plants grown at higher pCO₂ and exhibited a delay in the reduction of stomatal conductance that may have occurred in response to severe carbon limitations. The C4 plants grown at 70 Pa CO₂ showed lower relative reductions in net photosynthesis by the end of the drought compared to plants at lower pCO₂, indicating that CO₂ enrichment may alleviate drought effects in C4 plants. At the Pleistocene pCO₂, C3 and C4 plants showed similar relative recovery from drought for leaf area and biomass production, whereas C4 plants showed higher recovery than C3 plants at current and elevated pCO₂. Based on these model systems, we conclude that C3 species may not have been at a disadvantage relative to C4 species in response to low CO₂ and severe drought during the Pleistocene. Furthermore, C4 species may have an advantage over C3 species in response to increasing atmospheric CO₂ and more frequent and severe droughts.     

Morning vs afternoon sun patches in experimental forest gaps: consequences of temporal incongruency of resources to birch regeneration

We investigated whether the timing of high light availability as sun patches within forest gaps, independent of total or peak photosynthetic photon flux (PPF), influences the physiology and growth of four coexisting birch species (Betula alleghaniensis, B. lenta, B. papyrifera, and B. populifolia). Birch seedlings were grown for two years along either the east or west sides of experimental gap structures and at two moisture levels. Seedlings positioned in the west received sun patches earlier in the day than those in the east, and environmental conditions for carbon gain were generally more favorable during the earlier sunpatches in the west; air and leaf temperatures were lower, and relative humidity higher, relative to conditions during sun patches in the cats, simulating patterns observed in natural forest gaps. Seedlings positioned along the west edges of gaps fixed more carbon earlier in the day than those in the east, and in many cases, peak net photosynthetic rates were greater for west positioned seedlings. In year two, leaf-level integrated daily carbon gain was greater for west- than eastpositioned plants, and for the most pioneer species, B. populifolia, differences between west and east seedlings were greatest at lower soil moisture levels. Despite some small effects on leaf gas exchange, the timing of high light availability, and its temporal congruence with other factors critical to carbon gain, had no significant effects on first or second year seedling biomass. The responses of birch seedlings to controlled variations in the timing of high light availability were generally much smaller than birch seedling responses to variations in other components of daily light regimes such as total integrated and peak PPF.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

灌丛化对黄土高原草地植物群落结构和地上生物量的影响

中国北方草地普遍出现灌丛化现象,灌丛化改变植物群落结构、植物多样性和生产力,直接影响着草地生态保护与可持续利用.该研究以黄土高原灌丛化草地为研究对象,通过植被调查,分析比较不同坡向的灌丛斑块与禾草斑块植物群落结构(物种组成、优势种及物种多样性)和地上生物量的差异.结果发现:(1)灌丛化草地不同坡向对物种多样性及地上生物...     

Festuca campestris density and defoliation regulate abundance of the rhizomatous grass Poa pratensis in a fallow field

Invasion by the rhizomatous grass Kentucky bluegrass (Poa pratensis) is a global phenomenon, including into foothills rough fescue (Festuca campestris) grasslands of southwestern Alberta, Canada. In order to better understand the competitive relationships between these species, we conducted a fallow field study where rough fescue bunchgrass tussocks were transplanted at one of three planting densities (15, 30, or 45 cm spacing), and then subject to various treatments in a factorial design, including one‐time intensive summer defoliation and seeding of bluegrass into adjacent bare soil. Rough fescue plants exhibited marked intraspecific competition, as high planting densities increased tussock mortality, while decreasing plant tiller counts and relative inflorescence production, together with plant and tiller‐specific mass. However, high densities of the bunchgrass also reduced the cover and biomass of encroaching bluegrass, coincidental with reduced resource (soil moisture and light) availability in mid‐summer. Although summer defoliation increased rough fescue tiller counts, this disturbance reduced plant and tiller mass, and also increased Kentucky bluegrass. We conclude that while high densities of nondefoliated stands of rough fescue may increase resistance to bluegrass encroachment, a reduction in either fescue plant density or vigor via defoliation can increase the risk of bluegrass invasion within northern temperate grassland.     

Effects of Initial Site Treatments on Early Growth and Three-Year Survival of Idaho Fescue

Prairies in the Pacific Northwest have been actively restored for over a decade. Competition from non‐native woody and herbaceous species has been presumed to be a major cause for the failure of restoration projects. In this research, plugs of the native prairie bunchgrass, Festuca idahoensis Elmer var. roemeri (Pavlick), were grown from seed in a nursery and transplanted into a grassland site dominated by non‐native pasture grasses. The growth of the plants was followed for three years, and biomass of all volunteer plants was measured. Before planting, five treatments were applied to the plots: removal of vegetation by burning, removal of vegetation by an herbicide‐and‐till procedure, soil impoverishment by removal of organic matter, fertilizer application, and compost mulch application. Initial growth of Idaho fescue plugs was greatest with fertilizer and compost mulch. Plants grown in mulched plots were also able to photosynthesize later into the dry summer season. After the first year, plots initially fertilized or composted had the lowest survival rate of Idaho fescue. Impoverished and herbicide‐and‐till plots had the greatest 3‐year survival. Mulched plots supported the greatest weed growth after three years. Stressful environments give a competitive advantage to Idaho fescue in prairie restoration projects. As weedy species increase, growth and survival of Idaho fescue decreases.     

Germination biology and seedling growth of Clusia hilariana Schltdl., a dominant CAM-tree of drought-prone sandy coastal plains

The spatial and temporal fluctuations of water availability can be an obstacle for recruitment of many species in the restinga and might restrict seed germination and seedling growth in specific regeneration safe-sites. Clusia hilariana is one of the most dominant species of Restinga de Jurubatiba. This species has a high proportion of seedling establishment occurring inside the tanks of soil bromeliads underneath vegetation patches. Given the thin seed coats, the fast germination time and seed dispersal of C. hilariana during the dry season, we hypothesized that their major regeneration niche (the tanks of soil bromeliads) is related to susceptibility of seed germination and also seedling growth to low water availability. To test this hypothesis, seeds were germinated under decreasing water potentials using PEG 6000 solutions and seedlings were grown under varying water regimes. The percentage of seed germination progressively decreased at lower water potentials. After 38 days in ?1.0 MPa no seeds germinated. However, approximately 90% of seeds germinated when transferred to Ψ = 0 MPa. The relative growth rates of seedlings of C. hilariana did not differ between water treatments. Thus, the major regeneration niche of C. hilariana is not a consequence of a high sensitivity of seeds and seedlings to water shortage. Nonetheless, C. hilariana showed an array of seed and seedling traits that may help to overcome establishment constraints of the harsh environment of restingas.     

An Experimental Test of Resistance to Cheatgrass Invasion: Limiting Resources at Different Life Stages

Variable densities of an invasive species may represent variation in invasion resistance, due to variation in resource availability. This study determined whether low- and high-density cheatgrass (Bromus tectorum L.) patches within a shadscale-bunchgrass community of western Utah, USA, can be explained by variation in resource availability. It also explored the possible role of seed limitation and enemy pressure on invasion patterns. Two parallel field experiments were conducted:(1) increasing resources within low-density cheatgrass patches and, conversely (2) reducing resources within high-density cheatgrass patches. Treatments were applied at three life stages separately and across all stages. In low-density cheatgrass patches (assumed to represent high resistance), a disturbance that reduced soil compaction had the strongest positive effect, significantly increasing biomass by 250% and density by 104% in comparison to the control. The second strongest effect was reducing neighbors (native grasses), which significantly increased cheatgrass biomass and density. These results indicate that resources are present in low-density cheatgrass patches, but they are unavailable without disturbance and/or are exploited by competitors, and hence represent resistance to invasion. In high-density cheatgrass patches (assumed to represent low resistance), nitrogen availability was important in maintaining cheatgrass densities. Reducing nitrogen (via sucrose addition) significantly decreased density (by 37%) but not biomass. Life stages of cheatgrass were differentially affected by these resource manipulations. In addition, herbivore (primarily grasshoppers) and pathogen (head smut) pressures were documented to affect cheatgrass density, but did not explain resistance patterns. Instead, we found that differential resource availability explains the observed variation in cheatgrass density, and variation in natural resistance.     

Lack of recruitment in Lavandula stoechas subsp. pedunculata: a case of safe-site limitation

Lavandula stoechas subsp. pedunculata regeneration depends exclusively on the establishment of new individuals. Seed availability and seedling emergence and survival are therefore critical life stages and processes for species regeneration. In this study, seedling emergence and survival was monitored for two years in the scrub, both in clearings and adjacent to adult plants, and the surrounding perennial grassland, at 1, 3 and 5 m from the scrub. Soil seed bank spatial distribution was also studied for one year in the same two habitats, using the same sampling design. Soil seed availability in the scrub is high regardless of the distance from the adult individuals. On the contrary, the adjacent grassland shows a drastic fall in seed density, and almost no seedlings were observed there. In the scrub, seedling density was negatively related to distance from the three nearest adult plants in the clearings, and positively related to adult plant size beneath the adult Lavandula plants. There was also a negative relationship between seedling density and the percentage of bare soil. Only one seedling survived the first drought period, with no detection of effects of either position with respect to adult individuals or seedling density. We hypothesized that the study populations suffer a lack of appropriate safe sites within the scrubland while in the adjacent perennial grassland, observed low seed availability was added to safe-site limitation. That results in a lack of successful seedling establishments and a poor expansion potential of Lavandula scrublands, whose edges remain static in the short and medium term. As found in other Mediterranean scrubland, recruitment may only occur in years with particularly favourable weather, under disturbance regimes that increase seedling survival probability or when external dispersal agents increased seed availability in adequate places for Lavandula establishment.     

Plant responses to selective grazing by bison: interactions between light,herbivory and water stress

Two abundant tallgrass prairie forb species, Ambrosia psilostachya and Vernonia baldwinii, are commonly found intact in patches where the grasses have been selectively grazed by bison. Microclimatic patterns and physiological responses of these forbs were measured in grazed and ungrazed patches. These experiments demonstrated that bison herbivory indirectly enhanced water availability and productivity of forbs growing in grazed patches. This was due primarily to the reduction in transpiring grass leaf area in grazed patches and an increase in light availability. In grazed patches, incident light at forb mid-canopy height was 53% greater than ungrazed sites at midseason and soil temperatures were always warmer (e.g., 10°C at 5 cm), perhaps enabling forbs to initiate growth earlier in the spring. Enhanced leaf xylem pressure potential and stomatal conductance in plants in grazed areas were most evident when water availability was low (i.e., late in the growing season and over short-term dry periods characteristic of the tallgrass prairie environment). Relative to individuals in ungrazed areas, end-of-season biomass of A. psilostachya was 40% greater and reproductive biomass and head number of V. baldwinii was 45% and 40% greater, respectively, in plants in grazed patches. A favorable growing environment maintained in grazed patches during periods of water limitation enhances carbon gain in forbs leading to increased biomass and potential fitness.     

Foraging decisions underlying restricted space use: effects of fire and forage maturation on large herbivore nutrient uptake

Recent models suggest that herbivores optimize nutrient intake by selecting patches of low to intermediate vegetation biomass. We assessed the application of this hypothesis to plains bison (Bison bison) in an experimental grassland managed with fire by estimating daily rates of nutrient intake in relation to grass biomass and by measuring patch selection in experimental watersheds in which grass biomass was manipulated by prescribed burning. Digestible crude protein content of grass declined linearly with increasing biomass, and the mean digestible protein content relative to grass biomass was greater in burned watersheds than watersheds not burned that spring (intercept; F_1,251 = 50.57, P < 0.0001). Linking these values to published functional response parameters, ad libitum protein intake, and protein expenditure parameters, Fryxell's (Am. Nat., 1991, 138 , 478) model predicted that the daily rate of protein intake should be highest when bison feed in grasslands with 400–600 kg/ha. In burned grassland sites, where bison spend most of their time, availability of grass biomass ranged between 40 and 3650 kg/ha, bison selected foraging areas of roughly 690 kg/ha, close to the value for protein intake maximization predicted by the model. The seasonal net protein intake predicted for large grazers in this study suggest feeding in burned grassland can be more beneficial for nutrient uptake relative to unburned grassland as long as grass regrowth is possible. Foraging site selection for grass patches of low to intermediate biomass help explain patterns of uniform space use reported previously for large grazers in fire‐prone systems.     

Carbon Addition as a Countermeasure Against Biological Invasion by Plants

Increased nitrogen availability is known to favor invasion by non-native plants into natural grasslands. This suggests that decreasing nitrogen availability might serve as a countermeasure against invasion. One way to at least temporarily decrease nitrogen availability to plants is to increase microbial nitrogen uptake by adding carbon to the soil, and sawdust is a carbon source whose low cost could make it a practical conservation tool. To test whether adding sawdust to soil can counter the tendency of nitrogen enrichment to promote invasions by non-native plants, we hand-tilled 1.5kg sawdust/m² into the upper soil of the bare, nitrogen-rich patches left by dead shrubs of the nitrogen-fixing shrub Lupinus arboreus in two nearby areas with contrasting levels of invasion in a coastal grassland in northern California. After two years, in both areas, patches with sawdust had 40% less biomass of non-native plants than patches without sawdust, whereas biomass of native plants was not affected by sawdust addition. The more negative effect of sawdust on non-native species was specifically due to an effect on non-native grasses; adding sawdust increased the frequency of both native and non-native forbs. Results suggest that adding carbon as sawdust to soil can help counter invasion of grassland by non-native plants when invasion is being promoted by increased nitrogen availability, especially when the major invasive species are grasses.     

Factors affecting the invasion success of Senecio inaequidens and S. pterophorus in Mediterranean plant communities

Question: Plant invasions result from complex interactions between species traits, community characteristics and environmental variations. We examined the effect of these interactions on the invasion potential of two invasive Senecio species, S. inaequidens and S. pterophorus, across three Mediterranean plant communities in a natural park. Location: Catalonia, NE Spain. Methods: We carried out two series of experimental seedling transplantations, in the spring and fall of 2003, in grassland, shrubland and Quercus ilex forest. Competition with neighbouring plants and water availability were manipulated. We evaluated the survival, growth and reproduction with respect to each treatment combination. Results: Any habitat can be colonised if disturbance occurs. In the absence of disturbance, shrubland enhanced the survival of seedlings. Competition with resident vegetation dramatically reduced survival in grassland and forest when establishment occurred in the spring. However, establishment in the fall promoted invasion in grassland and shrubland, even in the undisturbed treatment. Grassland allowed the highest growth and reproductive performance of both species while forest was the most resistant habitat to invasion. S. inaequidens had a higher growth rate and a shorter pre‐reproductive period than S. pterophorus. S. pterophorus produced more biomass and was more dependent on water availability than S. inaequidens. Conclusions: In the light of our results, we recommend surveying open shrublands and grasslands after periods of rainfall. Special attention should be paid to S. pterophorus, which is currently spreading. A preliminary assessment of the invasive‐ness of this plant is given in this study.