Kevin J. Rice

SPATIAL HETEROGENEITY EXPLAINS THE SCALE DEPENDENCE OF THE NATIVE–EXOTIC DIVERSITY RELATIONSHIP

While small-scale studies show that more diverse native communities are less invasible by exotics, studies at large spatial scales often find positive correlations between native and exotic diversity. This large-scale pattern is thought to arise because landscapes with favorable conditions for native species also have favorable conditions for exotic species. From theory, we proposed an alternative hypothesis: the positive relationship at large scales is driven by spatial heterogeneity in species composition, which is driven by spatial heterogeneity in the environment. Landscapes with more spatial heterogeneity in the environment can sustain more native and more exotic species, leading to a positive correlation of native and exotic diversity at large scales. In a nested data set for grassland plants, we detected negative relationships between native and exotic diversity at small spatial scales and positive relationships at large spatial scales. Supporting our hypothesis, the positive relationships between native and exotic diversity at large scales were driven by positive relationships between native and exotic beta diversity. Further, both native and exotic diversity were positively correlated with spatial heterogeneity in abiotic conditions (variance of soil depth, soil nitrogen, and aspect) but were uncorrelated with average abiotic conditions, supporting the spatial-heterogeneity hypothesis but not the favorable-conditions

Effects of competition on resource availability and growth of a California bunchgrass

In California, little is known about the sensitivity of native bunchgrasses to competition or to changes in resource availability. We investigated the effect of nonnative annual vegetation on resource availability and growth of a native bunchgrass, Nassella pulchra, in a pair of factorial field experiments that incorporated effects of both interspecific and intraspecific competition as well as variation in soil depth. Plots of differing target densities and neighborhoods were used to assess changes in aboveground (light) and be- lowground (water) resource availability over multiple seasons in two sites with differing soil depth. N. pulchra grown without interspecific competitors grew larger and produced more culms at all planting densities compared to plants in plots with interspecific com- petitors. Intraspecific competition significantly influenced growth only in the absence of interspecific competition. Reproductive effort, as measured by flowering culm production, was more sensitive than vegetative growth to both forms of competition. Light availability and variability at the soil surface was greatly reduced by the nonnative annual neighborhood. As expected, soil moisture was rapidly depleted by annuals to 30 cm in all plots. In deep- soil plots, soil moisture was reduced at 60-150 cm depths only when annual vegetation was removed, and depletion was correlated with N. pulchra basal area. This result suggests that the interspecific neighborhood reduced root growth in N. pulchra and its subsequent ability to use deep moisture. Within Californias inland grasslands, nonnative annual veg- etation has changed seasonal patterns of resource availability. We conclude that (1) increased competition for light during the spring, when growth of annuals is most rapid, suppresses growth and reproduction of N. pulchra; (2) by suppressing bunchgrass growth, annual grasses reduce access to belowground resources by competitive interference; and (3) the loss of perennial grasses in California grasslands and the general dominance by nonnative annual species results in the relative underutilization of deep soil resources. These con- clusions suggest that the dominance of California grasslands by nonnative annual vegetation has shifted the primary limiting resource from soil moisture to light and the timing of resource limitation from summer to winter and spring.

Ecological genetics of Bromus tectorum. I : A hierarchical analysis of phenotypic variation

SummaryAn understanding of how genetic differentiation and phenotypic plasticity may interact to promote the spread of an introduced species requires information on the hierarchical distribution of genetic variation within the species in its new range. For example, a lack of genetic variation within marginal populations of an introduced species may slow its rate of spread into new habitats. In a glasshouse study, we examined the phenotypic variation among populations, among families, within families, and the homogeneity of variances within families for morphological and life history characters of an alien, self-pollinating annual grass, Bromus tectorum. The populations of B. tectorum studied were collected from both large, central populations in steppe habitats and small, peripheral populations in forest sites along a broad temperature-moisture gradient in western North America. Most variation in average flowering time was due to differences among seed source populations; among family variation contributed less than 20% to the total variance for flowering time. Populations from arid steppe habitats were the earliest flowering while the population from the most mesic forest habitat was last to flower. In contrast, the within-family variance was a major contributor to the total variance for plant dry weight, seed number per plant, total seed weight, and individual seed weight. The amount of total variation explained by among-family differences ranged between 18% for average seed weight to 30% for total plant dry weight. There was no consistent difference in within-population genetic variability between large, central populations in steppe habitats and smaller, potentially more isolated populations in forest habitats. Significant heterogeneity in within-family variance in some of the source populations suggest that families differ in the capacity for phenotypic response to environmental variation. Considered independently from source population, there was no consistent trend linking a particular trait to increased heterogeneity of within-family variances

LOCAL ADAPTATION, PATTERNS OF SELECTION, AND GENE FLOW IN THE CALIFORNIAN SERPENTINE SUNFLOWER (HELIANTHUS EXILIS)

Abstract The traditional view of the species as the fundamental unit of evolution has been challenged by observations that in heterogeneous environments, gene flow may be too restricted to overcome the effects of local selection. Whether a species evolves as a cohesive unit depends critically on the dynamic balance between homogenizing gene flow among populations and potentially disruptive local adaptation. To examine this evolutionary balance between “global” gene flow and local selection, we studied northern Californian populations of Helianthus exilis, the serpentine sunflower, within a mosaic of contrasting serpentine and nonserpentine areas that differ considerably in soil chemistry and water availability. Local adaptation to riparian and serpentine habitats was studied in Helianthus exilis along with an analysis of gene flow patterns among populations within these habitats. Local adaptation was assessed in H. exilis during 2002 and 2003 using reciprocal transplant experiments at multiple locations within serpentine and riparian habitats. Effects of competition and germination date on the expression of local adaptation were also examined within the reciprocal transplant experiments. Local adaptation was detected in both years at the local site level and at the level of habitat. The analysis of the transplanted populations indicated that the patterns of selection differed considerably between riparian and serpentine sites. Differential survivorship occurred in serpentine habitats, whereas selection on reproductive output predominated in riparian habitats. Local adaptation was expressed only in the absence of competition. Local adaptation in terms of survivorship was most strongly expressed in treatments with delayed seed germination. Microsatellite markers were used to quantify population genetic parameters and examine the patterns of gene flow among sampled populations. Analysis of molecular markers revealed a system of population patches that freely exchange genes with each other. Strong selection seems to maintain ecotypic variation within this endemic sunflower species, while extensive gene flow among populations prevents local speciation between serpentine and riparian ecotypes.

Ecological genetics of Bromus tectorum III. The demography of reciprocally sown populations

SummaryBy incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.

LOCAL ADAPTATION IN TWO SUBSPECIES OF AN ANNUAL PLANT : IMPLICATIONS FOR MIGRATION AND GENE FLOW

Plant populations often adapt to local environmental conditions. Here we demonstrate local adaptation in two subspecies of the California native annual Gilia capitata using standard reciprocal transplant techniques in two sites (coastal and inland) over three consecutive years. Subspecies performance in each site was measured in four ways: probability of seedling emergence, early vegetative size (length of longest leaf), probability of flowering, and total number of inflorescences produced per plant. Analysis of three of the four variables demonstrated local adaptation through site‐by‐subspecies interactions in which natives outperformed immigrants. The disparity between natives and immigrants in their probability of emergence and probability of flowering was greater at the coastal site than at the inland site. Treated in isolation, these two fitness components suggest that migration from the coast to the inland site may be less restricted by selection than migration in the opposite direction. Two measurements of individual size (leaf length and number of inflorescences), suggest (though not strongly) that immigrants may be subject to weaker selection at the coastal site than at the inland site. A standard cohort life table is used to compare replacement rates (R0) for each subspecies at each site. Comparisons of R0s suggest that immigrants are under a severe demographic disadvantage at the coastal site, but only a small disadvantage at the inland site. The results point out the importance of integrating over several fitness components when documenting the magnitude of local adaptation.

INTRASPECIFIC AND DIFFUSE COMPETITION: THE RESPONSE OF NASSELLA PULCHRA IN A CALIFORNIA GRASSLAND

In inland California grasslands, the high densities of alien annual species have altered the growing environment for native perennial grasses. Using variable-density plots, we measured the influence of intraspecific competition (conspecifics only) and diffuse competition (mixed-composition neighborhoods that include conspecifics) on growth and survival of Nassella pulchra, purple needlegrass. We assessed the effects of intraspecific and diffuse competition in weeded plots and unweeded plots, respectively, across a density gradient of N. pulchra plants (16-356 plants/M2). We used summer fire and spring sheep grazing to reduce diffuse competition in unweeded plots. The potential effect of rooting volume on competitive interactions was explored by establishing plots on two sites of different soil depth. Diffuse competition had an overriding influence on N. pulchra growth in all treatments. Intraspecific competitive effects were apparent only in the absence of diffuse competition. The effects of grazing and soil depth on growth were only short-lived interactions with the burning treatment. Burning was a longer-lived interaction, but only in weeded plots. Plant mortality was significantly increased by diffuse competition. Overall, N. pulchra survival was greatest in weeded plots, in grazed plots, and in deeper soil plots. The growth of N. pulchra individuals was negatively affected by alien annual species in all treatment combinations. Our data indicate that recruitment of N. pulchra within inland California grasslands is reduced by the adverse environment created by high densities of alien annual species. Successful attempts to increase populations of N. pulchra through management of the grassland community must involve significant modification of the biotic environment.

TESTING LIFE HISTORY CORRELATES OF INVASIVENESS USING CONGENERIC PLANT SPECIES

We used three congeneric annual thistles, which vary in their ability to invade California (USA) annual grasslands, to test whether invasiveness is related to differences in life history traits. We hypothesized that populations of these summer-flowering Centaurea species must pass through a demographic gauntlet of survival and reproduction in order to persist and that the most invasive species (C. solstitialis) might possess unique life history characteristics. Using the idea of a demographic gauntlet as a conceptual framework, we compared each congener in terms of (1) seed germination and seedling establishment, (2) survival of rosettes subjected to competition from annual grasses, (3) subsequent growth and flowering in adult plants, and (4) variation in breeding system. Grazing and soil disturbance is thought to affect Centaurea establishment, growth, and reproduction, so we also explored differences among congeners in their response to clipping and to different sizes of soil disturbance. We found minimal differences among congeners in either seed germination responses or seedling establishment and survival. In contrast, differential growth responses of congeners to different sizes of canopy gaps led to large differences in adult size and fecundity. Canopy-gap size and clipping affected the fecundity of each species, but the most invasive species (C. solstitialis) was unique in its strong positive response to combinations of clipping and canopy gaps. In addition, the phenology of C. solstitialis allows this species to extend its growing season into the summer—a time when competition from winter annual vegetation for soil water is minimal. Surprisingly, C. solstitialis was highly self-incompatible while the less invasive species were highly self-compatible. Our results suggest that the invasiveness of C. solstitialis arises, in part, from its combined ability to persist in competition with annual grasses and its plastic growth and reproductive responses to open, disturbed habitat patches. Corresponding Editor: D. P. C. Peters.

Patterns of tree dieback in Queensland, Australia: the importance of drought stress and the role of resistance to cavitation

During the extreme 1992–1997 El Niño drought event, widespread stem mortality, or tree “dieback”, of both mature and juvenile eucalypts occurred within the tropical savannas of northeast Australia. Most of the dieback occurred in individuals of the ironbark species complex (Eucalyptus crebra – E. xanthoclada) while individuals of the bloodwood species Corymbia erythrophloia, exhibited significantly less stem mortality. Indicative of greater water stress, predawn and midday xylem water potentials of ironbark adults and saplings were significantly more negative than predawn values of bloodwoods. The very negative xylem water potentials in ironbarks suggest that stem mortality in both adult and juvenile ironbarks results from drought-induced embolism and that ironbarks perhaps have a shallower and less extensive root system than bloodwoods. Although predawn and midday water potentials for ironbark adults and saplings were similar, a census of mature and juvenile ironbark trees indicated that mortality was higher in adult trees. Cavitation vulnerability curves indicated that ironbark saplings may be better buffered against cavitation than adult trees. If they possess smaller root systems, saplings are more likely than adults to experience low xylem water potentials, even in non-drought years. Xylem conduits produced in adult trees during periods of normal rainfall, although perhaps more efficient in water conduction, may be more vulnerable to cavitation during infrequent severe droughts.

Competitive effects of grassland annuals on soil water and blue oak (Quercus douglasii) seedlings

Four California annual grassland species were used to examine the hypothesis that different plant species have equivalent competitive effects. We investigated the effects of the annuals on soil water availability and the growth responses of blue oak (Quercus douglasii Hook & Arn.) to neighbor—induced water depletion. Neighborhoods of annuals were composed of species from California annual grassland with differing phenology and morphology that were hypothesized to show non—equivalent competitive effects on both a per—individual— and a per—unit—tissue basis. Three introduced species of winter annuals were sown at each of three densities (10, 30, or 100 seeds/dm2) around a single oak acorn. The grass Bromus diandrus Roth. and the forb Erodium botrys Cav. flower in early spring, while the grass B. mollis L. flowers slightly later. A native summer annual forb, Hemizonia luzulaefolia DC., was sown at only the intermediate density. Plants were grown outdoors in 15 cm diameter X 1 m deep tubes filled with soil from an oak woodland site. Identity and density of annual species had independent and interactive effects on the water resource level. Soil water potential was negatively density—dependent, remaining highest when oaks were grown without competitors. At the intermediate density soil water potentials decreased in the following order: B. mollis (slowest), B. diandrus, E. botrys, and H. luzulaefolia (most rapid). This order followed the pattern of root length developed in the neighborhoods, which was species— and density—dependent. The relationship between sowing density and neighborhood dry mass also differed among the annual species. Correlations between root length and soil water potential were stronger for the forbs than for the grasses. A given root length of the summer annual, H. Luzulaefolia, depleted soil water to a greater extent than did the same root length of any species. Oak seedling growth, gas exchange rates, and survivorship were negatively related to the rate and extent of soil water depletion. Additionally, the relationship between annual root length and oak seedling height or root length differed by neighborhood species. Thus, neighborhood species were non—equivalent in competitive effect on both a per—individual and a per—unit—root—length basis. Phenologically different species may represent different functional groups of competitors.