Coastal Prairie Research Articles

Vertisol Crack Extent Associated with Gilgai and Soil Moisture in the Texas Gulf Coast Prairie

Long‐term observations of in situ crack formation and closure in shrink‐swell soils are rare, but important to understanding hydrology in shrink‐swell soils. To analyze spatial and temporal variability of crack development in a Vertisol with gilgai, soil cracks were measured on a 100‐m2 area of Laewest clay (fine, smectitic, hyperthermic Typic Hapludert) with native tallgrass vegetation on 42 dates from 1989 to 1998. Our objectives were to (i) report the distribution of Vertisol cracking across gilgai microtopography; (ii) estimate crack depth as a function of crack width considering gilgai; (iii) investigate the relationship of surface cracking and soil moisture considering gilgai and hysteresis. All surface cracks were mapped on scaled diagrams with width categorized, and some crack depth measured. Gravimetric soil moisture corresponding to crack measurements was measured on 18 dates, and on an additional 32 dates without crack measurements. Drying, wetting, and uniform soil moisture conditions were classified from the difference in soil moisture from 10‐ to 25‐cm depths. Microtopography was quantified using a digital elevation model. Results showed that crack area density was greatest on microhighs and microslopes, though microlows had the largest cracking potential. The linear correlation between crack depth and width was moderately strong (r2 = 0.5), and not affected significantly by gilgai and hysteresis. However, taking hysteresis into account improved the linear regression models of crack area density versus soil moisture (up to r2 = 0.69) on both microhighs and microlows. Antecedent soil moisture seemed to impact in situ crack area density. Further field studies are recommended.

Factors Influencing Selection of Road Culverts as Winter Roost Sites by Western Burrowing Owls

The conversion of coastal prairie to farmland in southern Texas has drastically reduced the number of available animal burrows, thereby forcing western Burrowing Owls (Athene cunicularia hypugaea) wintering in southern Texas to use nontraditional roost sites such as roadside culverts. We studied factors influencing the selection of road culverts as roost sites by Burrowing Owls by comparing characteristics of 34 occupied and 100 unoccupied culverts. All occupied culverts were in agricultural habitat. Culverts with small diameters (≤16 cm) and those with an east—west orientation were occupied by Burrowing Owls in greater proportions than were culverts with larger diameters or different orientations. Occupied culverts were also associated with absence of grass, absence of woody vegetation, and presence of crop stubble. Our results provide guidelines for making drainage culverts more attractive to Burrowing Owls, but use of roadside culverts by Burrowing Owls may expose the owls to an increased risk of mortality from vehicle collisions. To avoid this dilemma, our guidelines for culverts could also be adapted as criteria for installation of artificial burrows in habitats suitable for wintering Burrowing Owls.

Nutrient Encounter by a Generalist Herbivore in a Heterogeneous Landscape

Most landscapes are heterogeneous in vegetation composition and thus, in the distribution of nutrients required by herbivores for growth and reproduction. Hispid cotton rats (Sigmodon hispdus) inhabit Texas coastal prairies which are characterized by habitat patches dominated by dicots, monocots or both [mixed] plant types. Hispid cotton rats must obtain nutrients necessary for reproduction by ingesting both dicots and monocots, and reproductive females concentrate their activity in mixed patches. Mixed patches may be selected because they have high nutrient contents or because presence of both monocot and dicot food plants lowers cost of foraging movements. Because hispid cotton rats select a nutritionally complete diet and may detect protein by odor and taste cues, we hypothesized that nutrient concentrations may cue position of foraging paths. We used nutrient maps to measure amount of protein, P and Ca encountered during foraging from radiotracked individuals. We compared these values to those obtained from randomly generated foraging paths. Actual and random paths did not differ in amount of protein, P or Ca encountered. Differences in nutient accumulation between seasons were explained by seasonal differences in availability. We conclude that foraging paths do not respond to nutrient content of plants, but that reproductive females likely occupy mixed habitat patches because presence of both dicot and monocot food plants decreases time and energy spent foraging.

Discovery of a large clonal patch of a social amoeba: implications for social evolution

Studies of genetic population structures of clonally reproducing macro-organisms have revealed large areas where only one clone is found. These areas, referred to as clonal patches, have not been shown to occur in free-living microbes until now. In free-living microbes, high genetic diversity at local scales is usually maintained by high rates of dispersal. We report, however, a highly dense, 12-m clonal patch of the social amoeba Dictyostelium discoideum in a cattle pasture located in a Texas Gulf Coast prairie. We confirm the presence of only one clone by the analysis of 65 samples and amplification of 10 polymorphic microsatellite loci. Samplings of additional cattle pastures nearby showed higher clonal diversity, but with a density of D. discoideum isolates lower than in the clonal patch. These findings show that high rates of microbial dispersal do not always produce genetic diversity at local scales, contrary to the findings of previous studies. The existence of clonal patches may be particularly important for microbial social evolution.

Effect of Terrestrial Mollusc Herbivory on Holocarpha macradenia (Asteraceae) Seedlings in California Coastal Prairie under Different Clipping Regimes

Herbivory is generally overlooked as a factor influencing the viability of endangered plant populations in western North American grasslands. I conducted experiments to determine the effects of terrestrial mollusc herbivory for survival of a federally endangered, annual forb, Holocarpha macradenia (Santa Cruz tarplant), in California's coastal prairie. In addition, I utilized litter removal and clipping regimes from a continuing study of site-specific responses to evaluate the relationship between herbivory rates from terrestrial molluscs and disturbance regimes. I planted seedlings of H. macradenia into copper bordered quadrats that excluded molluscs and into non-copper bordered controls at two sites in California's coastal prairie. Three treatment regimes of vegetation clipping and litter removal were utilized. Herbivory damage was assessed using an ordinal scale every two weeks throughout the rainy season. Molluscs were also trapped and identified. Herbivory amounts were significantly less when molluscs were excluded from plots for all disturbance regimes; however, there was no significant difference in herbivory levels between the disturbance regimes. A significant amount of invertebrate herbivory on H. macradenia was by the non-native slug, Deroceras reticulatum (grey garden slug), and this herbivory significantly affected the survivability of H. macradenia seedlings in California coastal prairie.

METAPOPULATION DYNAMICS OVERRIDE LOCAL LIMITS ON LONG‐TERM PARASITE PERSISTENCE

A simple null model, particularly germane to small and vulnerable organisms such as parasites, is that local conditions set a stage upon which larger-scale dynamics play out. Soil moisture strongly influences survival of entomopathogenic nematodes (EPN), which in turn drive trophic cascades by protecting vegetation from root-feeding herbivores. In this study, we examine the mechanisms responsible for patchy occurrence of an entomopathogenic nematode, Heterorhabditis marelatus, in a California coastal prairie. One hypothesis proposes that biotic factors such as competition and natural enemies could regulate occurrence of EPN populations. We found that fungi and other enemies of EPN, although locally potent, did not explain the patterns of incidence across sites. Abiotic factors also have strong effects on EPN persistence, especially for vulnerable free-living stages. Thus, we tested the hypothesis that patchy occurrence of EPN on a large landscape was driven by differences in soil moisture. Our research uses long-term data on nematode incidence in combination with a landscape-level experiment to demonstrate the lack of a correlation between soil moisture and long-term persistence. A year-long experiment showed EPN mortality was weakly correlated with soil moisture among our study sites. Thirteen years of data, however, showed that colonization rates were highly correlated with long-term persistence. Sites with highest long-term persistence experienced the highest rates of rhizosphere colonization, extinction, and turnover. As a result, we concluded that metapopulation dynamics override limitations set by local and short-term abiotic conditions to determine long-term persistence in this parasite-driven trophic cascade.

A comparative study of dissolved organic carbon transport and stabilization in California forest and grassland soils

For soil carbon to be effectively sequestered beyond a timescale of a few decades, this carbon must become incorporated into passive reservoirs or greater depths, yet the actual mechanisms by which this occurs is at best poorly known. In this study, we quantified the magnitude of dissolved organic carbon (DOC) leaching and subsequent retention in soils of a coniferous forest and a coastal prairie ecosystem. Despite small annual losses of DOC relative to respiratory losses, DOC leaching plays a significant role in transporting C from surface horizons and stabilizing it within the mineral soil. We found that DOC movement into the mineral soil constitutes 22% of the annual C inputs below 40 cm in a coniferous forest, whereas only 2% of the C inputs below 20 cm in a prairie soil could be accounted for by this process. In line with these C input estimates, we calculated advective transport velocities of 1.05 and 0.45 mm year−1 for the forested and prairie sites, respectively. Radiocarbon measurements of field-collected DOC interpreted with a basic transport-turnover model indicated that DOC which was transported and subsequently absorbed had a mean residence time of 90–150 years. Given these residence times, the process of DOC movement and retention is responsible for 20% of the total mineral soil C stock to 1 m in the forest soil and 9% in the prairie soil. These results provide quantitative data confirming differences in C cycles in forests and grasslands, and suggest the need for incorporating a better mechanistic understanding of soil C transport, storage and turnover processes into both local and regional C cycle models.

Dissolved organic carbon chemistry and dynamics in contrasting forest and grassland soils

In this study, we examined changes in isotopic (13C and 14C) and spectroscopic (UV and 13C NMR) properties of dissolved organic carbon (DOC) in relation to soil organic matter (SOM) to elucidate the sources and sinks of DOC as water percolates through the soils of two contrasting upland coastal California ecosystems—a redwood forest and a coastal prairie. Despite differences in the distribution of C stocks and litter chemistry at these two sites, we found similar shifts in DOC chemistry with soil depth. DOC concentrations dropped rapidly with increasing depth, with an accompanying decrease in the C:N ratio, an increase in the δ13C value and an decrease in specific UV adsorption. In the grassland soil, Δ14C values declined from current atmospheric values (+70‰) in the surface horizon to −75‰ at 100 cm. In the redwood soil, the Δ14C value of 111‰ in O horizon leachates was indicative of OM with a residence time of 8–10 yrs, with a decrease in Δ14C values to −80‰ at 100 cm. Solid-state CP/MAS 13C NMR spectra were generally most similar to highly humified OM, with a general decrease in the relative abundance of aromatic compounds and an increase in the alkyl C/O-alkyl C ratio with increasing depth. All of these trends are consistent with the shifts in SOM properties with increasing depth, which are interpreted to mean a shift from fresh plant material to older, highly altered OM. In this Mediterranean climate, we found distinct seasonal shifts in the quantity and composition of DOC found in soil solution during the winter rainy period that was also consistent with a shift from recent labile substrates to older, highly altered OM. These results fit in with a growing body of literature suggesting that the source of much of the DOC within mineral soils is the local soil OM, and the 14C data, in particular, indicate that DOC at depth is not simply the fraction of surficial leachates that have not been adsorbed or decomposed. Rather, exchange reactions with a portion of the more stabilized SOM pool exert the strongest control on both the concentration and composition of DOC found in these soils.

Wood Decomposition Following a Perennial Lupine Die-Off: A 3-Year Litterbag Study

Woody debris is a conspicuous feature of many ecosystems and can be a large pool of stored carbon and nutrients. In the California coastal prairie, yellow bush lupines (Lupinus arboreus) experience mass die-offs, producing large quantities of woody detritus. Live lupines are fed upon by the stem-boring caterpillars of the ghost moth, Hepialus californicus, and outbreaks of ghost moths are one factor contributing to lupine die-offs. A common detritivore, the terrestrial isopod Porcellio scaber, frequently inhabits ghost moth tunnels in lupine wood. We used a litterbag experiment to test the hypothesis that H. californicus increases decomposition of woody lupine detritus by facilitating its use by P. scaber. Isopod access to wood was crossed with simulated ghost moth boring to measure the independent and interactive effects of these two arthropods on total mass loss, as well as on carbon, nitrogen, and lignin dynamics. Isopods initially colonized litterbags but were not more abundant on L. arboreus logs that had simulated ghost moth boring than on logs without boring. They were rare in litterbags collected at 12 months or later and had no effect on wood decomposition. Simulated ghost moth boring increased wood decomposition (P = 0.0021), from 50.5 to 55.1% mass loss after 3 years. This effect was likely due to increased surface area for microbial utilization of the wood. Lupine wood had an initial lignin content of 14.70 ± 0.67%, but lignin did not appear to decompose during the 3 years of this study, and by the end of the experiment accounted for 32.6 ± 1.12% of the remaining wood. Neither ghost moth boring nor isopod access affected lignin loss. Lupine wood from a die-off in 2002 was estimated to have contained three times more nitrogen per unit area than the yearly input of annual grass litter. The slow decomposition of lupine wood, however, restricts the rate at which nitrogen is released into the soil and results in the storage of carbon and nutrients in lupine wood for several years following such die-offs.

Mechanisms of Habitat Selection by the Hispid Cotton Rat (Sigmodon hispidus)

The Texas coastal prairie is composed of habitat patches characterized by monocots, dicots, or a mixture of both plant types. Radiotelemetry revealed that reproductive female hispid cotton rats (Sigmodon hispidus) preferred mixed habitats and avoided dicot habitats, whereas males did not show a preference. Such habitat selection can be attained by shorter distances moved or increased turning (i.e., area-restricted search). Reproductive females, but not males, moved shorter distances in mixed habitats. However, turns relative to straight-ahead movements for females, but not males, were fewer in mixed habitats than in monocot or dicot habitats suggesting directed foraging rather than area-restricted search. To obtain necessary amounts of nutrients, especially carbohydrates, protein, phosphorus, and calcium, reproductive females ingest both monocots and dicots. Directed movement may facilitate foraging for dicots because these food items occur in clumps in mixed habitats. Because their nutritional needs are less than those of females, males do not exhibit the same patterns of habitat selection. Differential patch occupancy was not explained by overhead plant cover, but more bare ground in dicot habitats may explain avoidance of these habitat types because of increased predation risk.

Seasonal and Spatial Controls over Nutrient Cycling in a Pacific Northwest Prairie

West Coast prairies in the US are an endangered ecosystem, and effective conservation will require an understanding of how changing climate will impact nutrient cycling and availability. We examined how seasonal patterns and micro-heterogeneity in edaphic conditions (% moisture, total organic carbon, % clay, pH, and inorganic nitrogen and phosphorus) control carbon, nitrogen, and phosphorus cycling in an upland prairie in western Oregon, USA. Across the prairie, we collected soils seasonally and measured microbial respiration, net nitrogen mineralization, net nitrification, and phosphorus availability under field conditions and under experimentally varied temperature and moisture treatments. The response variables differed in the degree of temperature and moisture limitation within seasons and how these factors varied across sampling sites. In general, we found that microbial respiration was limited by low soil moisture year-round and by low temperatures in the winter. Net nitrogen mineralization and net nitrification were never limited by temperature, but both were limited by excessive soil moisture in winter, and net nitrification was also inhibited by low soil moisture in the summer. Factors that enhanced microbial respiration tended to decrease soil phosphorus availability. Edaphic factors explained 76% of the seasonal and spatial variation in microbial respiration, 35% of the variation in phosphorus availability, and 29% of the variation in net nitrification. Much of the variation in net nitrogen mineralization remained unexplained (R 2 = 0.19). This study, for the first time, demonstrates the complex seasonal controls over nutrient cycling in a Pacific Northwest prairie.

Isotopic variability in large carbonate nodules in Vertisols: Implications for climate and ecosystem assessments

Vertisols occurring in the Coastal Prairie of Texas preserve distinctive patterns of carbon isotopic values with depth for both soil organic matter as well as pedogenic carbonate. These isotopic values may be used to reconstruct past climate and ecosystems (C3 versus C4 vegetation). Some soils contain large carbonate nodules (> 2 cm diameter) that exhibit δ 13C isotopic gradients of up to 2–3‰ across the nodule and have an internal structure that resembles concentric growth rings. These isotopic gradients are used to potentially track relative nodule movement in soil profiles. Some nodules possibly move within and even across microenvironments displaced by several centimeters. The isotopic gradients of the nodules may also track climate and ecosystem changes associated with relative changes in soil depth caused by soil movement. In order to make accurate climate and ecosystem interpretations, soil organic matter and pedogenic carbonate should be demonstrated to have formed in isotopic equilibrium at their respective soil depth.

Holcus lanatus invasion slows decomposition through its interaction with a macroinvertebrate detritivore, Porcellio scaber

Detritus based food webs may mediate the impacts of invasive species on ecosystem processes. Holcus lanatus (L.) is an invasive perennial grass that is rapidly spreading in the coastal prairie of California. We used litterbags to determine if H. lanatus altered the rate of litter decomposition through its interaction with the dominant macroinvertebrate detritivore at our study site, the isopod Porcellio scaber (Latreille). Over the course of a year, H. lanatus litter loss was 15% less than annual grasses, a difference that was directly attributable to P. scaber. Although there was no effect of isopods on litter loss during the winter, when most decomposition occurred, isopods had substantial effects on litter loss of annual grasses during the summer. P. scaber had no effect on litter loss of H. lanatus litter. Our findings suggest that H. lanatus invasion slows decomposition in the coastal prairie because it is unusually refractory to an important detritivore. The decreased decomposition of H. lanatus litter, along with increased production during the growing season, lead to a doubling of the end-of-season litter biomass where H. lanatus had invaded.

Effects of nutrient loading and extreme rainfall events on coastal tallgrass prairies: invasion intensity, vegetation responses, and carbon and nitrogen distribution

AbstractSoil fertility and precipitation are major factors regulating transitions from grasslands to forests. Biotic regulation may influence the effects of these abiotic drivers. In this study, we examined the effects of extreme rainfall events, anthropogenic nutrient loading and insect herbivory on the ability of Chinese tallow tree (Sapium sebiferum) to invade coastal prairie to determine how these factors may influence woody invasion of a grassland. We manipulated soil fertility (NPK addition) and simulated variation in frequency of extreme rainfall events in a three growing season, full factorial field experiment. Adding water to or pumping water out of plots simulated increased and decreased rainfall frequencies. We added Sapium seeds and seedlings to each plot and manipulated insect herbivory on transplanted Sapium seedlings with insecticide. We measured soil moisture, Sapium performance, vegetation mass, and carbon and nitrogen in vegetation and soils (0–10 cm deep, 10–20 cm deep). Fertilization increased Sapium invasion intensity by increasing seedling survival, height growth and biomass. Insect damage was low and insect suppression had little effect in all conditions. Recruitment of Sapium from seed was very low and independent of treatments. Vegetation mass was increased by fertilization in both rainfall treatments but not in the ambient moisture treatment. The amount of carbon and nitrogen in plants was increased by fertilization, especially in modified moisture plots. Soil carbon and nitrogen were independent of all treatments. These results suggest that coastal tallgrass prairies are more likely to be impacted by nutrient loading, in terms of invasion severity and nutrient cycling, than by changes in the frequency of extreme rainfall events.

WHEN THERE IS NO ESCAPE: THE EFFECTS OF NATURAL ENEMIES ON NATIVE, INVASIVE, AND NONINVASIVE PLANTS

An important question in the study of biological invasions is the degree to which successful invasion can be explained by release from control by natural enemies. Natural enemies dominate explanations of two alternate phenomena: that most introduced plants fail to establish viable populations (biotic resistance hypothesis) and that some introduced plants become noxious invaders (natural enemies hypothesis). We used a suite of 18 phylogenetically related native and nonnative clovers (Trifolium and Medicago) and the foliar pathogens and invertebrate herbivores that attack them to answer two questions. Do native species suffer greater attack by natural enemies relative to introduced species at the same site? Are some introduced species excluded from native plant communities because they are susceptible to local natural enemies? We address these questions using three lines of evidence: (1) the frequency of attack and composition of fungal pathogens and herbivores for each clover species in four years of common garden experiments, as well as susceptibility to inoculation with a common pathogen; (2) the degree of leaf damage suffered by each species in common garden experiments; and (3) fitness effects estimated using correlative approaches and pathogen removal experiments. Introduced species showed no evidence of escape from pathogens, being equivalent to native species as a group in terms of infection levels, susceptibility, disease prevalence, disease severity (with more severe damage on introduced species in one year), the influence of disease on mortality, and the effect of fungicide treatment on mortality and biomass. In contrast, invertebrate herbivores caused more damage on native species in two years, although the influence of herbivore attack on mortality did not differ between native and introduced species. Within introduced species, the predictions of the biotic resistance hypothesis were not supported: the most invasive species showed greater infection, greater prevalence and severity of disease, greater prevalence of herbivory, and greater effects of fungicide on biomass and were indistinguishable from noninvasive introduced species in all other respects. Therefore, although herbivores preferred native over introduced species, escape from pest pressure cannot be used to explain why some introduced clovers are common invaders in coastal prairie while others are not.

The role of soil resources in an exotic tree invasion in Texas coastal prairie

Summary Variation in resource requirements among plant species may cause changes in community composition when resource supply rates vary. Because exotic plants may differ in their requirements compared with native plants, changes in resource levels or ratios may change their invasive potential. The concentration hypothesis, nutrient balance hypothesis and resource ratio hypothesis make different predictions regarding the effects of nutrient additions on plant communities and the success of invaders. We investigated these hypotheses using Chinese tallow tree (Sapium sebiferum), which is an aggressive invader of grasslands in the southern USA. Soil nitrogen, phosphorus and potassium were manipulated in a full factorial field experiment in a Texas coastal prairie. Survival of Sapium seedlings over three growing seasons was independent of all three types of nutrient additions, alone or in any combination. Adding nitrogen or potassium increased the mass and height of Sapium seedlings. Adding phosphorus increased the mass and height of Sapium seedlings only when nitrogen was also added. These results support the nutrient balance hypothesis, which assumes that higher tissue concentrations of a nutrient reflect a greater ability of a plant species to take up and/or conserve that nutrient. The above‐ground biomass of background prairie vegetation was independent of the addition of each type of nutrient alone or in any combination. However, the functional composition of the community shifted from graminoids to forbs with the addition of nitrogen and from graminoids to woody plants with the addition of phosphorus. These results support the concentration hypothesis, which assumes that higher tissue concentrations of a nutrient reflect a greater requirement for that nutrient. The contrasting responses of native woody vegetation and Sapium to phosphorus addition suggest that Sapium may have unusually low demands for phosphorus relative to nitrogen for a woody plant in this community. These results show that potassium or nitrogen addition, perhaps in combination with phosphorus addition, has the potential to accelerate the invasion of Sapium into Texas coastal prairies. Because anthropogenic nutrient additions are common in grasslands worldwide, this may help to explain the frequent success of woody species invading grasslands.

HABITAT RELATIONSHIPS OF BIRDS OVERWINTERING IN A MANAGED COASTAL PRAIRIE

Abstract Grassland birds are considered to be rapidly declining in North America. Management approaches for grassland birds frequently rely on prescribed burning to maintain habitat in suitable condition. We evaluated the relationships among years since burn, vegetation structure, and overwintering grassland bird abundance in coastal prairie. Le Conte's Sparrows (Ammodramus leconteii) were most common in areas that had: (1) been burned within the previous 2 years, (2) medium density herbaceous vegetation, and (3) sparse shrub densities. Savannah Sparrows (Passerculus sandwichensis) were associated with areas: (1) burned within 1 year, (2) with sparse herbaceous vegetation, and (3) with sparse shrub densities. Sedge Wrens (Cistothorus platensis) were most common in areas that had: (1) burned greater than 2 years prior and (2) dense herbaceous vegetation. Swamp Sparrows (Melospiza georgiana): (1) were most common in areas of dense shrubs, (2) not related to time since burnings, and (3) demonstrated no relat...

Effect of Seed Source, Topsoil Removal, and Plant Neighbor Removal on Restoring California Coastal Prairies

Abstract Grasslands are hot spots of biodiversity but are now widely threatened by changes in patterns of disturbances, such as grazing and fire regimes, exotic species invasions, and cultivation. The goal of this experiment was to find the most appropriate combination of treatments to reintroduce Danthonia californica, a formerly dominant perennial bunchgrass, into degraded California coastal prairies. Danthonia californica was sown from seed and transplanted at two sites and at two grazing intensities (grazed/ungrazed) in a multifactorial experiment testing the effects of (1) local versus nonlocal seed sources; (2) topsoil removal; and (3) reduction of plant neighbors. Seed emergence was very low, suggesting that transplanting may be a better option to reintroduce D. californica. Although transplants grown from nonlocal seeds survived better initially at both sites, transplants from local seeds had higher survival after 1.5 year at one site. This suggests that short‐term plant establishment studies may be misleading. Topsoil removal greatly enhanced transplant survival, and neighbor removal primarily increased transplant growth. Our results suggest that removing topsoil prior to transplanting seedlings grown from local seeds is the most promising method to reintroduce D. californica. However, the benefits of removing topsoil to provide safe sites for plant establishment should be weighed carefully against potential negative effects on the native seed bank and microbial communities on a site‐specific basis.

Seed banks in plant conservation: Case study of Santa Cruz tarplant restoration

Although conservation and restoration practitioners have focused on maximizing aboveground population size and seed set of rare plants, a clear understanding of seed bank dynamics is crucial to managing these species. Santa Cruz tarplant ( Holocarpha macradenia) is a threatened annual forb restricted to coastal prairie habitats in central California. Holocarpha produces disk achenes germinating within a year of production and ray achenes forming a persistent seed bank. We constructed both deterministic and stochastic demographic models for a restored Holocarpha population, using demographic rates measured separately for unmanipulated plants and plants growing in plots where vegetation was clipped. The deterministic models indicated that regardless of germination from the seed bank, the population would decline without clipping or similar treatments that enhance survival and reproductive output. Deterministic models showed only a slight positive effect of increased ray seed germination rates on population growth, which would need to be balanced against a potential loss of buffering against environmental variation as the dormant seed bank was reduced. Our stochastic simulations suggested that extinction risk for Holocarpha populations would be minimized by intermediate levels of ray seed germination. Thus, managers should focus on improving the performance of aboveground plants before considering actions to stimulate germination, since the former will yield a greater increase in deterministic population growth and not sacrifice any buffering effect of the seed bank. This case study emphasizes the importance of considering dormant seeds and seed banks in designing successful restoration and management strategies for plant species at risk of extinction.

Mechanisms of resistance to invasion in a California grassland: the roles of competitor identity, resource availability, and environmental gradients

Resistance to the invasion of exotic plants may sometimes result from the strong effects of a relatively small number of resident species. Understanding the mechanisms by which such species resist invasion could provide important insights for the management of invaded ecosystems. Furthermore, the individualistic responses of community members to resource availability and environmental gradients could drive spatial variation in resistance at the local to landscape scales. We tested the resistance of monoculture plots of three native perennial grasses from the California coastal prairie to the invasion of the European perennial grass Holcus lanatus. We also used a watering treatment that increased early summer water availability and a natural elevational gradient in resource availability and soil texture to evaluate how resident identity interacted with abiotic resistance to affect Holcus establishment. Two native species, Festuca rubra and Calamagrostis nutkaensis, exhibited strong resistance, correlated with their negative effects on light availability. A third native grass, Bromus carinatus var. maritimus, had either no effect or a weakly facultative effect on Holcus performance relative to bare plots. Water addition did not alter the resistance of these species, but the elevation gradient did. Holcus invasion increased with improving abiotic conditions towards the slope bottom in bare and Bromus plots, but invasion decreased towards the bottom in Calamagrostis plots, where better conditions favored competitive residents. These results support the idea that resistance to invasion can sometimes be due to a subset of native species, and that the resistance provided by even a single species is likely to vary across the landscape. Identifying the mechanisms by which species resist invasion could facilitate the selection of management strategies that at best increase, or at worst do not decrease, natural resistance.