Stephen D. Davis

Mortality of resprouting chaparral shrubs after a fire and during a record drought: physiological mechanisms and demographic consequences.

We examined postfire regeneration of chaparral shrubs during an intense drought. This study focused on the demography and physiology of shrub species that resprout from a basal lignotuber following fire. We found significant levels of resprout mortality when intense drought occurred in the year following fire during the period of shrub recovery. Three of the seven sampled resprouting species had the greatest or near greatest levels of mortality ever recorded when compared to previous studies. Most shrub mortality occurred during the drought after individuals had resprouted (i.e. individuals survived fire, resprouted and then subsequently died). Physiological measurements of species with high mortality suggested that resprout stems were highly embolized and xylem hydraulic conductivities were close to zero during the peak of the drought. In addition, lignotubers of two of the three species experiencing high mortality were depleted of starch. Population densities of most shrub species declined after the drought compared with their prefire levels, with the exception of one drought tolerant obligate seeding species. Resprouting shrub species may deplete their carbohydrate reserves during the resprouting process, making them particularly vulnerable to drought because of the need to transpire water to acquire the CO2 that is used to supply energy to a large respiring root system. Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.

Towards understanding resprouting at the global scale.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Chaparral Shrub Hydraulic Traits, Size, and Life History Types Relate to Species Mortality during California's Historic Drought of 2014.

Chaparral is the most abundant vegetation type in California and current climate change models predict more frequent and severe droughts that could impact plant community structure. Understanding the factors related to species-specific drought mortality is essential to predict such changes. We predicted that life history type, hydraulic traits, and plant size would be related to the ability of species to survive drought. We evaluated the impact of these factors in a mature chaparral stand during the drought of 2014, which has been reported as the most severe in California in the last 1,200 years. We measured tissue water potential, native xylem specific conductivity, leaf specific conductivity, percentage loss in conductivity, and chlorophyll fluorescence for 11 species in February 2014, which was exceptionally dry following protracted drought. Mortality among the 11 dominant species ranged from 0 to 93%. Total stand density was reduced 63.4% and relative dominance of species shifted after the drought. Mortality was negatively correlated with water potential, native xylem specific conductivity, and chlorophyll fluorescence, but not with percent loss in hydraulic conductivity and leaf specific conductivity. The model that best explained mortality included species and plant size as main factors and indicated that larger plants had greater survival for 2 of the species. In general, species with greater resistance to water-stress induced cavitation showed greater mortality levels. Despite adult resprouters typically being more vulnerable to cavitation, results suggest that their more extensive root systems enable them to better access soil moisture and avoid harmful levels of dehydration. These results are consistent with the hypothesis that short-term high intensity droughts have the strongest effect on mature plants of shallow-rooted dehydration tolerant species, whereas deep-rooted dehydration avoiding species fare better in the short-term. Severe droughts can drive changes in chaparral structure as a result of the differential mortality among species.

Exotic deer diminish post-fire resilience of native shrub communities on Santa Catalina Island, southern California

Browsing by exotic mule deer on Santa Catalina Island (SCI) off the coast of southern California may diminish the post-fire resilience of native shrublands. To assess this, deer exclosures were established following a wildfire to monitor post-fire recovery of three dominant, native shrub species (Heteromeles arbutifolia, Rhus integrifolia, and Rhamnus pirifolia). Post-fire resprout growth, mortality, and tissue water status as well as pre- and post-fire shrub density and cover were measured inside and outside of deer exclosures. We found that deer browsing significantly limited post-fire resprout growth and led to increased mortality of resprouting H. arbutifolia shrubs (88 % mortality outside compared to 11 % inside exclosures). Post-fire resprouts maintained favorable water status during the study despite drought conditions, indicating that water stress was not a proximate cause of resprout mortality. Deer browsing resulted in a >93 % reduction in canopy coverage of dominant shrub species. The dramatic reduction of native shrubs at this site may create opportunities for displacement by exotic species, resulting in eventual vegetation-type conversion. The observed link between intense browsing and post-fire shrub mortality provides much needed information concerning the environmental impact of exotic deer on SCI and illustrates the interaction between exotic herbivores and fire on an island system.

Isolation of Microsatellite Markers in a Chaparral Species Endemic to Southern California, Ceanothus megacarpus (Rhamnaceae)

Premise of the study: Microsatellite (simple sequence repeat [SSR]) markers were developed for Ceanothus megacarpus, a chaparral species endemic to coastal southern California, to investigate potential processes (e.g., fragmentation, genetic drift, and interspecific hybridization) responsible for the genetic structure within and among populations distributed throughout mainland and island populations. Methods and Results: Four SSR-enriched libraries were used to develop and optimize 10 primer sets of microsatellite loci containing either di-, tri-, or tetranucleotide repeats. Levels of variation at these loci were assessed for two populations of C. megacarpus. Observed heterozygosity ranged from 0.250 to 0.885, and number of alleles ranged between four and 21 per locus. Eight to nine loci also successfully amplified in three other species of Ceanothus. Conclusions: These markers should prove useful for evaluating the influence of recent and historical processes on genetic variation in C. megacarpus and related species.

An individual-based model of chaparral vegetation response to frequent wildfires

The Santa Monica Mountains are home to many species of chaparral shrubs that provide vegetative cover and whose deep roots contribute to the stability of the steep slopes. Recently, native chaparral have been threatened by an unprecedented drought and frequent wildfires in Southern California. Besides the damage from the wildfires themselves, there is the potential for subsequent structural losses due to erosion and landslides. In this paper, we develop a mathematical model that predicts the impact of drought and frequent wildfires on chaparral plant community structure. We begin by classifying chaparral into two life history types based on their response to wildfires. Nonsprouters are completely killed by a fire, but their seeds germinate in response to fire cues. Facultative sprouters survive by resprouting but also rely on seed germination for post-fire recovery. The individual-based model presented here simulates the growth, seed dispersal, and resprouting behavior of individual shrubs across two life history types as they compete for space and resources in a rectangular domain. The model also incorporates varying annual rainfall and fire frequency as well as the competition between plants for scarce resources. The parameters were fit using seedling and resprout survivorship data as well as point quarter sampling data from 1986 to 2014 at a biological preserve within the natural landscape of the Malibu campus of Pepperdine University. The simulations from our model reproduce the change in plant community structure at our study site which includes the local extinction of the nonsprouter Ceanothus megacarpus due to shortened fire return intervals. Our simulations predict that a combination of extreme drought and frequent wildfires will drastically reduce the overall density of chaparral, increasing the likelihood of invasion by highly flammable exotic grasses. The simulations further predict that the majority of surviving shrubs will be facultative sprouting species such as Malosma laurina.

Seasonal changes in tissue-water relations for eight species of ferns during historic drought in California

PREMISE OF THE STUDY California experienced severe drought between 2012 and 2016. During this period, we compared seasonal changes in tissue-water relations among eight fern species in the Santa Monica Mountains of southern California to elucidate differential mechanisms of drought survival and physiological performance during extreme water deficits. METHODS We monitored seasonal changes in water potential (Ψmd) and dark-adapted chlorophyll fluorescence (Fv/Fm), assessed tissue-water relations including osmotic potential at saturation and the turgor loss point (Ψπ, sat and Ψπ, tlp), and measured, for two evergreen species, xylem-specific and leaf-specific hydraulic conductivity (Ks and Kl) and vulnerability of stem xylem to water stress-induced embolism (water potential at 50% loss hydraulic conductivity, Ψ50). KEY RESULTS Species grew in either riparian or chaparral understory. The five chaparral species had a wider range of seasonal water potentials, root depths, and frond phenological traits, including one evergreen, two summer-deciduous, and two desiccation-tolerant (resurrection) species. Evergreen species were especially diverse, with an evergreen riparian species maintaining seasonal water potentials above -1.3 MPa, while an evergreen chaparral species had seasonal water potentials below -8 MPa. In those two species the Ψ50 values were -2.5 MPa and -4.3 MPa, respectively. CONCLUSIONS Observed differences in physiological performance among eight fern species reflected niche partitioning in water utilization and habitat preference associated with distinct phenological traits. We predict differential survival among fern species as future drought events in California intensify, with desiccation-tolerant resurrection ferns being the most resistant.