Soil moisture and fungi affect seed survival in California grassland annual plants

Survival of seeds in the seed bank is important for the population dynamics of many plant species, yet the environmental factors that control seed survival at a landscape level remain poorly understood. These factors may include soil moisture, vegetation cover, soil type, and soil pathogens. Because many soil fungi respond to moisture and host species, fungi may mediate environmental drivers of seed survival. Here, I measure patterns of seed survival in California annual grassland plants across 15 species in three experiments. First, I surveyed seed survival for eight species at 18 grasslands and coastal sage scrub sites ranging across coastal and inland Santa Barbara County, California. Species differed in seed survival, and soil moisture and geographic location had the strongest influence on survival. Grasslands had higher survival than coastal sage scrub sites for some species. Second, I used a fungicide addition and exotic grass thatch removal experiment in the field to tease apart the relative impact of fungi, thatch, and their interaction in an invaded grassland. Seed survival was lower in the winter (wet season) than in the summer (dry season), but fungicide improved winter survival. Seed survival varied between species but did not depend on thatch. Third, I manipulated water and fungicide in the laboratory to directly examine the relationship between water, fungi, and survival. Seed survival declined from dry to single watered to continuously watered treatments. Fungicide slightly improved seed survival when seeds were watered once but not continually. Together, these experiments demonstrate an important role of soil moisture, potentially mediated by fungal pathogens, in driving seed survival.     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

Population age structure and recent

Mid to late 20th century expansion of Dracophyllum scrub into tussock grassland on subantarctic Campbell Island has been attributed to the collective effects of global warming, cessation of farming in 1931, and continued grazing by feral sheep. To determine the importance of these, we dated the timing of scrub expansion by aging 241 Dracophyllum plants in 17 plots chosen to sample the range of environments this shrub/ small tree occupies on Campbell Island. Three plots, in lowland, well-drained, locations were dominated by large, old Dracophyllum that had established between 1846 and 1940. Virtually all shrubs in the remaining plots had established after 1940, with peaks in 1970 and 1985. The pattern of establishment does not coincide with any marked change in the temperature regime, although a prolonged period of relatively dry winters (c. 1970–1990) coincides with a late surge of regeneration on very wet sites. The removal of feral sheep from different parts of the island at different times is also unrelated to the pattern of establishment. In contrast, Dracophyllum spread follows farm abandonment in 1931 when regular burning ceased, suggesting that Dracophyllum is invading sites from which it was excluded by fire. However, the earliest reports from the mid 19th century indicate that tussock grassland was previously the dominant vegetation cover on the island, with limited Dracophyllum scrub. It is possible that the reduction of the dense tussock grasslands by fire and grazing in late 19th–early 20th century opened a regeneration window for Dracophyllum scrub to spread once burning ceased.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

Impacts of invasive annuals on soil carbon and nitrogen storage in southern California depend on the identity of the invader

Non‐native plant invasions can alter nutrient cycling processes and contribute to global climate change. In southern California, California sage scrub (hereafter sage scrub), a native shrub‐dominated habitat type in lowland areas, has decreased to <10% of its original distribution. Postdisturbance type‐conversion to non‐native annual grassland, and increasingly to mustard‐dominated invasive forbland, is a key contributor to sage scrub loss. To better understand how type‐conversion by common invasive annuals impacts carbon (C) and nitrogen (N) storage in surface soils, we examined how the identity of the invader (non‐native grasses, Bromus spp.; and non‐native forbs, Brassica nigra), microbial concentrations, and soil properties interact to influence soil nutrient storage in adjacent native and invasive habitat types at nine sites along a coast to inland gradient. We found that the impact of type‐conversion on nutrient storage was contingent upon the invasive plant type. Sage scrub soils stored more C and N than non‐native grasslands, whereas non‐native forblands had nutrient storage similar to or higher than sage scrub. We calculate that >940 t C km^?2 and >60 t N km^?2 are lost when sage scrub converts to grass‐dominated habitat, demonstrating that grass invasions are significant regional contributors to greenhouse gas emissions. We found that sites with greater total C and N storage were associated with high cation exchange capacities and bacterial concentrations. Non‐native grassland habitat type was a predictor of lower total C, and soil pH, which was greatest in invasive habitats, was a predictor of lower total N. We demonstrate that modeling regional nutrient storage requires accurate classification of habitat type and fine‐scale quantification of cation exchange capacity, pH, and bacterial abundance. Our results provide evidence that efforts to restore and conserve sage scrub enhance nutrient storage, a key ecosystem service reducing atmospheric CO₂ concentrations.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

Measures of resilience: the response of coastal sage scrub to fire

Measures of four components of resilience are developed and used to quantify the response of coastal sage scrub to fire in southern California: (1) elasticity (rate of recovery following disturbance), (2) amplitude (threshold of disturbance beyond which recovery to the original state no longer occurs), (3) malleability (extent of alteration of the new stable-state from the original) and (4) damping (extent and duration of oscillation in an ecosystem parameter following disturbance). Vegetation and soil properties measured before fire, and for the first 5–6 yr after fire on four coastal (Venturan association) and four inland (Riversidian association) sites of coastal sage were used to follow changes. In addition, results from a simulation model of post-fire succession in Venturan coastal sage scrub (the FINICS model of Malanson) were used to examine resilience behavior over a 200 yr period. Resilience behavior of coastal sage scrub is critically influenced by the presence of a competitive mix of inherently strongly and weakly resprouting species. Sites dominated by weak resprouters exhibit lower elasticity and less damping of year-to-year fluctuations in composition in the early post-fire years. Sites with a mixture of weak and strong resprouters have a lower threshold of disturbance (amplitude) before species extirpation occurs, a result intensified by a higher frequency of disturbance. Malleability is also greater in these systems under higher disturbance frequency.Nomenclature follows. P.A. Munz, 1974. A Flora of Southern California. Univ. California Press.     

Restoration of Florida scrub vegetation in an old field through 23 years after planting

Florida scrub, a fire‐maintained, xeromorphic shrubland, hosts many rare and declining species, but most scrub habitat has been lost to development and agriculture. Formerly cultivated sites offer an opportunity to restore Florida scrub. In this 23‐year study, we test whether scrub species could be restored to an old field and whether scrub vegetation composition and structure could be reestablished over time. Scrub oaks were planted in a section of an old field in 1992 after the site was cleared and treated with herbicide. Additional oaks and other scrub species were planted in 1993. We determined survival and growth annually of a marked sample of scrub species. We sampled vegetation cover in two height strata annually on 10 permanent line‐intercept transects in the old grove and on 20 transects in adjacent, intact scrub. Initial survival of Quercus geminata exceeded that of Q. chapmanii or Q. myrtifolia. Cover of scrub oaks greater than 0.5 m increased from 1.3% in 1992 to 65.2% in 2010. However, the vegetation structure of large Q. geminata did not resemble that of native scrub. Adaptive management led to cutting large oaks in the fall of 2010, and burning the site in early 2011. Cutting and burning appeared to stimulate sprouting and clonal spread of scrub oaks. Ordination analysis indicated directional change related to increasing scrub oak cover and time since planting but the old field still differed from intact scrub. Vegetation has developed toward scrub composition and structure but exotic grasses persist.     

Patterns of floristic richness in vegetation communities of California: regional scale analysis with multi-temporal NDVI

Aim Applying water‐energy dynamics and heterogeneity theory to explain species richness via remote sensing could allow for the regional characterization and monitoring of vegetation community assemblages and their environment. We assess the relationship of multi‐temporal normalized difference vegetation index (NDVI) to plant species richness in vegetation communities. Location California, USA. Methods Sub‐regions containing species inventories for chaparral, coastal sage scrub, foothill woodland, and yellow pine forest communities were intersected with a vegetation community map and an AVHRR NDVI time series for 1990, 1991, 1992, 1995 and 1996. Principal components analysis reduced the AVHRR data to three variables representing the sum and temporal trajectories of NDVI within each community. A fourth variable representing heterogeneity was tested using the standard deviation of the first component. Quadratic forms of these variables were also tested. Species richness was analysed by stepwise regression. Results Chaparral, coastal sage scrub, and yellow pine forest had the best relationships between species richness and NDVI. Richness of chaparral was related to NDVI heterogeneity and spring greenness (r² varied between 0.26 and 0.62 depending on year of NDVI data). Richness of coastal sage scrub was nonlinearly related to annual NDVI and heterogeneity (r² 0.63–0.81), with peak richness at intermediate values. Foothill woodland richness was related to heterogeneity in a monotonic curvilinear fashion (r² 0.28–0.35). Yellow pine forest richness was negatively related to spring greenness and positively related to heterogeneity (r² 0.40–0.46). Main Conclusions While NDVI's relationship to species richness varied, the selection of NDVI variables was generally consistent across years and indicated that spatial variability in NDVI may reflect important patterns in water‐energy use that affect plant species richness. The principal component axis that should correspond closely with annual mean NPP showed a less prominent role. We conclude that plant species richness for coarse vegetation associations can be characterized and monitored at a regional scale and over long periods of time using relatively coarse resolution NDVI data.     

Fire history and patterns of Venturan subassociations of Californian coastal sage scrub

Californian coastal sage scrub has floristically distinct subassociations with sharp boundaries in the Santa Monica Mountains. This mesoscale biogeographic pattern has been variously attributed to the timing and pattern of fire and to differences in the moisture availability on sites. An examination of the actual fire history of sites reveals that recent fire events are unlikely to have caused the observed patterns. Sites with similar fire histories are not as similar in vegetation as sites with different fire histories but similar aspect. Single short fire intervals do not result in dissimilar communities; fires are unlikely to recur with the same boundaries as these two subassociations. A simulation of shrub response to fire interval and intensity indicates that long-term fire trends are also unlikely to have caused the distinction or the sharp boundaries of the two subassociations. Both subassociations include species with life history characteristics adapted to short and to long fire intervals. The subassociations exist without regard to fire history.     

Influence of grazing and fire on breeding birds and perennial plants in Cyprus scrub and forest systems

Grazing and browsing by sheep and goats has been an important anthropogenic influence on ecosystems in the Mediterranean region for centuries. This influence has changed significantly in recent decades, with a general shift from range grazing to the penning of animals. The intermediate disturbance hypothesis (IDH) proposes that perturbation - including anthropogenic disturbance - is the norm for Mediterranean ecosystems, and that higher species diversity is found under conditions of continuous, moderate disturbance. Here we test the IDH as it relates to grazing of scrub and open forest habitats in Cyprus, while also testing for the effects of fire. We carried out surveys of breeding birds and vegetation at 48 study sites in scrub and open woodland across Cyprus. We estimated relative grazing pressure (past and present) and fire history at these sites, and tested for associations between these factors, breeding bird species richness and perennial vegetation diversity. We found moderate current grazing reduces density of lower and middle level scrub, and a higher level of diversity of perennial vegetation at moderately over intensively grazed sites. We found that moderate historical grazing pressure had a positive influence on richness of breeding bird species in lower scrub habitat such as phrygana, but that this effect was weaker the taller and denser the habitat. By contrast, current grazing pressure had a negative effect on species richness in lower scrub, but species richness in grazed habitats was higher in denser, taller scrub habitat such as garrigue and maquis. Our study suggests grazing plays an important role in maintaining habitat heterogeneity but the impact on avian species richness depends on the density and height of the vegetation.     

Recruitment of Quercus agrifolia in central California: the importance of shrub-dominated patches

Abstract. Many perennial plants strongly enhance the survival of seedlings of other species. We studied patterns of long-term recruitment of Quercus agrifolia (Coastal live oak) associated with shrub-dominated communities by counting Q. agrifolia recruits on a time sequence of historical aerial photographs and comparing recruitment among mapped patches of coastal sage scrub, chaparral, and grassland in an 1120-ha landscape. Because we could not identify new recruits in existing woodlands with aerial photographs, we studied the recruitment of Q. agrifolia in this vegetation type indirectly by comparing population size structures and the spatial relationships between shrubs and recruits among woodlands that varied in understory community type. At the landscape scale, recruitment was higher in coastal sage scrub vegetation than predicted by the extent of its coverage, commensurate with the spatial coverage of chaparral, and very low in grassland. Recruitment within woodland communities also varied considerably. In woodland communities on sheltered, north-oriented topography with understories dominated by shrubs, there were large numbers of small Q. agrifolia, and recruits were not significantly spatially associated with shrubs within plots. In woodlands with herbaceous understories there were few individuals in the small size classes, and recruits were strongly spatially associated with shrubs within plots. Woodlands with shrub-dominated understories have population structures that appear to be stable, but woodlands with herbaceous understories exhibit size structures associated with declining populations. Quercus recruitment into shrub-dominated patches corresponds with previous documentation of facilitative relationships between shrubs and oak seedlings, and suggests the occurrence of an unusual form of patch dynamics in these landscapes.     

Restoring Sage‐grouse nesting habitat through removal of early successional conifer

Conifer woodlands have expanded into sagebrush (Artemisia spp.) ecosystems and degrade habitat for sagebrush obligate species such as the Greater Sage‐grouse (Centrocercus urophasianus). Conifer management is increasing despite a lack of empirical evidence assessing outcomes to grouse and their habitat. Although assessments of vegetation recovery after conifer removal are common, comparisons of successional trends with habitat guidelines or actual data on habitat used by sage‐grouse is lacking. We assessed impacts of conifer encroachment on vegetation characteristics known to be important for sage‐grouse nesting. Using a controlled repeated measures design, we then evaluated vegetation changes for 3 years after conifer removal. We compared these results to data from 356 local sage‐grouse nests, rangewide nesting habitat estimates, and published habitat guidelines. We measured negative effects of conifer cover on many characteristics important for sage‐grouse nesting habitat including percent cover of forbs, grasses, and shrubs, and species richness of forbs and shrubs. In untreated habitat, herbaceous vegetation cover was slightly below the cover at local nest sites, while shrub cover and sagebrush cover were well below cover at the nest sites. Following conifer removal, we measured increases in herbaceous vegetation, primarily grasses, and sagebrush height. Our results indicate that conifer abundance can decrease habitat suitability for nesting sage‐grouse. Additionally, conifer removal can improve habitat suitability for nesting sage‐grouse within 3 years, and trajectories indicate that the habitat may continue to improve in the near future.     

The effects of fire on ant communities in north-western Patagonia: the importance of habitat structure and regional context

Abstract. We investigated the effects of recent fires on the native ant communities in two habitats of north-west Patagonia that differ in vegetation structural complexity. Using bait traps, we sampled ants in replicated scrub and steppe areas including paired burned and unburned sites. Fires significantly reduced plant cover and ant diversity only in scrub sites. The drop in diversity was due to (a) a reduction in the abundance of rare species associated with woody vegetation, and (b) an increase in the abundance of the dominant species, which thrive in more xeric microclimatic conditions. Consequently, ant assemblage structure of burned scrub approaches that of steppe sites. Our findings suggest that the effects of disturbances on ant assemblages depends both on habitat characteristics, which in turn determine the extent of the changes induced by the disturbance, and on the regional context of the ant fauna, which in turn determines the ability of the ants to deal with the post-disturbance conditions.     

Phytophthora Root Rot: Assessing the potential threat to Australia's oldest national park

Summary   Royal National Park, Australia's oldest national park, is a significant reserve for conservation of the flora and fauna that are characteristic of the Hawkesbury Sandstone in New South Wales. Since at least 1974, Phytophthora Root Rot (caused by Phytophthora cinnamomi ) has been known to occur in the Park, but there is no knowledge of the extent of infestation or the potential impacts of the disease within the Park. This study investigated the distribution of the pathogen within Royal National Park at two scales: a systematic survey by vegetation type, and a targeted survey of populations of Waratah and Spear Grass-tree. These two species are known to be susceptible to Phytophthora Root Rot and are therefore potential indicators of the impact of the pathogen on vegetation in Royal National Park. Phytophthora cinnamomi was recovered from all vegetation types sampled but most commonly in heathland/open scrub vegetation. The pathogen was easily recovered from sites containing Spear Grass-tree, but was not isolated from any sites containing Waratah. Because of the widespread distribution of P. cinnamomi , we conclude that hygiene measures will be of little use to prevent the spread of the pathogen within Royal National Park. Monitoring of the occurrence and spread of disease symptoms in plants and applying phosphite to protect susceptible rare or threatened flora may be the most appropriate management options.     

Alternative causes of edge-abundance relationships in birds and small mammals of California coastal sage scrub

Changes in the distribution and abundance of bird and small mammal species at urban-wildland edges can be caused by different factors. Edges can affect populations directly if animals respond behaviorally to the edge itself or if proximity to edge directly affects demographic vital rates (an "ecotonal" effect). Alternatively, urban edges can indirectly affect populations if edges alter the characteristics of the adjacent wildland vegetation, which in turn prompts a response to the altered habitat (a "matrix" or "habitat" effect). We studied edge effects of birds and small mammals in southern Californian coastal sage scrub, and assessed whether edge effects were attributable to direct behavioral responses to edges or to animal responses to changes in habitat at edges. Vegetation species composition and structure varied with distance from edge, but the differences varied among study sites. Because vegetation characteristics were correlated with distance from edge, responses to habitat were explored by using independently-derived models of habitat associations to calibrate vegetation measurements to the habitat affinities of each animal species. Of sixteen species examined, five bird and one small mammal species responded to edge independently of habitat features, and thus habitat restoration at edges is expected to be an ineffective conservation measure for these species. Two additional species of birds and one small mammal responded to habitat gradients that coincided with distance from edge, such that the effect of edge on these species was expressed via potentially reversible habitat degradation.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Population Structure and Growth Patterns of Opuntia echios var. gigantea along an Elevational Gradient in the Galípagos Islands1

Cacti growing in forests potentially experience growth limitation due to reduced light availability. To test this hypothesis, we studied the population structure of Opuntia echios var. gigantea at 15 sites on the south side of Isla Santa Cruz, Galípagos Islands, Ecuador. Populations were located in communities ranging from arid scrub at low elevations to closed‐canopy tropical dry forest at higher elevations. Ordination confirmed the existence of a strong elevation‐vegetation gradient. Opuntia abundance peaked at lower elevations (ca 30 m), with lower densities in closed‐canopy sites. For populations in scrub vegetation, density declined fairly regularly with plant height. Populations in forested sites had few plants of intermediate height, suggesting periodic recruitment. Scrub populations had random dispersion, while those in forests were aggregated. The change in spatial pattern may be related to a change in primary reproductive mode from asexual propagation via fallen fruits to propagation via fallen cladodes. Height was significantly correlated with stem diameter. Intercepts of these relationships increased toward higher elevations, probably in response to the increasing height of the surrounding canopy.