Topography influences diurnal and seasonal microclimate fluctuations in hilly terrain environments of coastal California.

Understanding the topographic basis for microclimatic variation remains fundamental to predicting the site level effects of warming air temperatures. Quantifying diurnal fluctuation and seasonal extremes in relation to topography offers insight into the potential relationship between site level conditions and changes in regional climate. The present study investigated an annual understory temperature regime for 50 sites distributed across a topographically diverse area (>12 km2) comprised of mixed evergreen-deciduous woodland vegetation typical of California coastal ranges. We investigated the effect of topography and tree cover on site-to-site variation in near-surface temperatures using a combination of multiple linear regression and multivariate techniques. Sites in topographically depressed areas (e.g., valley bottoms) exhibited larger seasonal and diurnal variation. Elevation (at 10 m resolution) was found to be the primary driver of daily and seasonal variations, in addition to hillslope position, canopy cover and northness. The elevation effect on seasonal mean temperatures was inverted, reflecting large-scale cold-air pooling in the study region, with elevated minimum and mean temperature at higher elevations. Additionally, several of our sites showed considerable buffering (dampened diurnal and seasonal temperature fluctuations) compared to average regional conditions measured at an on-site weather station. Results from this study help inform efforts to extrapolate temperature records across large landscapes and have the potential to improve our ecological understanding of fine-scale seasonal climate variation in coastal range environments.

Climate change, tree demography, and thermophilization in western US forests.

Climate change is driving widespread changes in ecological communities. Warming temperatures often shift community composition toward more heat-tolerant taxa. The factors influencing the rate of this "thermophilization" process remain unclear. Using 10-y census data from an extensive forest plot network, we show that mature tree communities of the western United States have undergone thermophilization. The mean magnitude of climate warming over the 10-y study interval was 0.32 °C, whereas the mean magnitude of thermophilization was 0.039 °C. Differential tree mortality was the strongest demographic driver of thermophilization, rather than growth or recruitment. Thermophilization rates are associated with recent changes in temperature and hydrologic variables, as well as topography and disturbance, with insect damage showing the strongest standardized effect on thermophilization rates. On average, thermophilization occurred more rapidly on cool, north-facing hillslopes. Our results demonstrate that warming temperatures are outpacing the composition of western US forest tree communities, and that climate change may erode biodiversity patterns structured by topographic variation.

Global wind patterns shape genetic differentiation, asymmetric gene flow, and genetic diversity in trees.

Wind disperses the pollen and seeds of many plants, but little is known about whether and how it shapes large-scale landscape genetic patterns. We address this question by a synthesis and reanalysis of genetic data from more than 1,900 populations of 97 tree and shrub species around the world, using a newly developed framework for modeling long-term landscape connectivity by wind currents. We show that wind shapes three independent aspects of landscape genetics in plants with wind pollination or seed dispersal: populations linked by stronger winds are more genetically similar, populations linked by directionally imbalanced winds exhibit asymmetric gene flow ratios, and downwind populations have higher genetic diversity. For each of these distinct hypotheses, partial correlations between the respective wind and genetic metrics (controlling for distance and climate) are positive for a significant majority of wind-dispersed or wind-pollinated genetic data sets and increase significantly across functional groups expected to be increasingly influenced by wind. Together, these results indicate that the geography of both wind strength and wind direction play important roles in shaping large-scale genetic patterns across the world's forests. These findings have implications for various aspects of basic plant ecology and evolution, as well as the response of biodiversity to future global change.

Evolutionary relationships between drought-related traits and climate shape large hydraulic safety margins in western North American oaks.

Quantitative knowledge of xylem physical tolerance limits to dehydration is essential to understanding plant drought tolerance but is lacking in many long-vessel angiosperms. We examine the hypothesis that a fundamental association between sustained xylem water transport and downstream tissue function should select for xylem that avoids embolism in long-vessel trees by quantifying xylem capacity to withstand air entry of western North American oaks (Quercus spp.). Optical visualization showed that 50% of embolism occurs at water potentials below -2.7 MPa in all 19 species, and -6.6 MPa in the most resistant species. By mapping the evolution of xylem vulnerability to embolism onto a fossil-dated phylogeny of the western North American oaks, we found large differences between clades (sections) while closely related species within each clade vary little in their capacity to withstand air entry. Phylogenetic conservatism in xylem physical tolerance, together with a significant correlation between species distributions along rainfall gradients and their dehydration tolerance, suggests that closely related species occupy similar climatic niches and that species' geographic ranges may have shifted along aridity gradients in accordance with their physical tolerance. Such trends, coupled with evolutionary associations between capacity to withstand xylem embolism and other hydraulic-related traits, yield wide margins of safety against embolism in oaks from diverse habitats. Evolved responses of the vascular system to aridity support the embolism avoidance hypothesis and reveal the importance of quantifying plant capacity to withstand xylem embolism for understanding function and biogeography of some of the Northern Hemisphere's most ecologically and economically important plants.

Range dynamics mediated by compensatory life stage responses to experimental climate manipulations.

The expectations of polar or upslope distributional shifts of species ranges in response to warming climate conditions have been recently questioned. Diverse responses of different life stages to changing temperature and moisture regimes may alter these predicted range dynamics. Furthermore, the climate driver(s) influencing demographic rates, and the contribution of each demographic rate to population growth rate (λ), may shift across a species range. We investigated these demographic effects by experimentally manipulating climate and measuring responses of λ in nine populations spanning the elevation range of an alpine plant (Ivesia lycopodioides). Populations exhibited stable growth rates (λ ~ 1) under naturally wet conditions and declining rates (λ < 1) under naturally dry conditions. However, opposing vital rate responses to experimental heating and watering lead to negligible or negative effects on population stability. These findings indicate that life stage-specific responses to changing climate can disrupt the current relationships between population stability and climate across species ranges.

Natural selection maintains species despite frequent hybridization in the desert shrub Encelia.

Natural selection is an important driver of genetic and phenotypic differentiation between species. For species in which potential gene flow is high but realized gene flow is low, adaptation via natural selection may be a particularly important force maintaining species. For a recent radiation of New World desert shrubs (Encelia: Asteraceae), we use fine-scale geographic sampling and population genomics to determine patterns of gene flow across two hybrid zones formed between two independent pairs of species with parapatric distributions. After finding evidence for extremely strong selection at both hybrid zones, we use a combination of field experiments, high-resolution imaging, and physiological measurements to determine the ecological basis for selection at one of the hybrid zones. Our results identify multiple ecological mechanisms of selection (drought, salinity, herbivory, and burial) that together are sufficient to maintain species boundaries despite high rates of hybridization. Given that multiple pairs of Encelia species hybridize at ecologically divergent parapatric boundaries, such mechanisms may maintain species boundaries throughout Encelia.

Topographic heterogeneity lengthens the duration of pollinator resources.

The availability of sufficient and diverse resources across time is important for maintenance of biodiversity and ecosystem functioning. In this study, we examine the potential for variation in environmental conditions across topographic gradients to extend floral resource timing. Flowering time on a landscape may vary across topography due to differences in abiotic factors, species turnover, or genotypic differences. However, the extent to which this variation in phenology affects overall flowering duration on a landscape, and the components of diversity that influence flowering duration, are unexplored. We investigate whether differences in flowering time due to topography yield an overall extension in duration of flowering resources in a northern California grassland. We recorded flowering time of pollinator resource species across four successive spring growing seasons (2015-2018) on paired north and south aspects. Flowering time differences were evaluated both at the community level and within species present on both paired aspects. The role of plasticity was examined in an experimental case study using genotypes of Lasthenia gracilis. We found that aspect is a strong determinant of phenology, with earlier flowering on warmer south-facing slopes. Aspect differences resulted in complementarity in timing of flowering resources across sites, as aspects that started flowering earlier also ended earlier. Complementarity between north and south aspects served to extend the flowering time of pollinator resources by an average of 4-8 days (8%-15%), depending on the year. This extension can be attributed to both within-species responses to aspect differences and species turnover. Flowering of L. gracilis genotypes was distinct across aspects, demonstrating that plasticity can drive the extension of flowering duration. Our findings indicate that heterogeneous topography can extend overall flowering time of pollinator resources, which may support pollinator biodiversity. Extension was most pronounced at the community level, which incorporates species turnover as well as plastic and genotypic differences within species.

Weather underground: Subsurface hydrologic processes mediate tree vulnerability to extreme climatic drought.

Drought extent and severity have increased and are predicted to continue to increase in many parts of the world. Understanding tree vulnerability to drought at both individual and species levels is key to ongoing forest management and preparation for future transitions in community composition. The influence of subsurface hydrologic processes is particularly important in water-limited ecosystems, and is an under-studied aspect of tree drought vulnerability. With California's 2013-2016 extraordinary drought as a natural experiment, we studied four co-occurring woodland tree species, blue oak (Quercus douglasii), valley oak (Quercus lobata), gray pine (Pinus sabiniana), and California juniper (Juniperus californica), examining drought vulnerability as a function of climate, lithology and hydrology using regional aerial dieback surveys and site-scale field surveys. We found that in addition to climatic drought severity (i.e., rainfall), subsurface processes explained variation in drought vulnerability within and across species at both scales. Regionally for blue oak, severity of dieback was related to the bedrock lithology, with higher mortality on igneous and metamorphic substrates, and to regional reductions in groundwater. At the site scale, access to deep subsurface water, evidenced by stem water stable isotope composition, was related to canopy condition across all species. Along hillslope gradients, channel locations supported similar environments in terms of water stress across a wide climatic gradient, indicating that subsurface hydrology mediates species' experience of drought, and that areas associated with persistent access to subsurface hydrologic resources may provide important refugia at species' xeric range edges. Despite this persistent overall influence of the subsurface environment, individual species showed markedly different response patterns. We argue that hydrologic niche segregation can be a useful lens through which to interpret these differences in vulnerability to climatic drought and climate change.

Plant hydraulic traits reveal islands as refugia from worsening drought.

Relatively mesic environments within arid regions may be important conservation targets as 'climate change refugia' for species persistence in the face of worsening drought conditions. Semi-arid southern California and the relatively mesic environments of California's Channel Islands provide a model system for examining drought responses of plants in potential climate change refugia. Most methods for detecting refugia are focused on 'exposure' of organisms to certain abiotic conditions, which fail to assess how local adaptation or acclimation of plant traits (i.e. 'sensitivity') contribute to or offset the benefits of reduced exposure. Here, we use a comparative plant hydraulics approach to characterize the vulnerability of plants to drought, providing a framework for identifying the locations and trait patterns that underlie functioning climate change refugia. Seasonal water relations, xylem hydraulic traits and remotely sensed vegetation indices of matched island and mainland field sites were used to compare the response of native plants from contrasting island and mainland sites to hotter droughts in the early 21st century. Island plants experienced more favorable water relations and resilience to recent drought. However, island plants displayed low plasticity/adaptation of hydraulic traits to local conditions, which indicates that relatively conserved traits of island plants underlie greater hydraulic safety and localized buffering from regional drought conditions. Our results provide an explanation for how California's Channel Islands function as a regional climate refugia during past and current climate change and demonstrate a physiology-based approach for detecting potential climate change refugia in other systems.

Multiple axes of ecological vulnerability to climate change.

Observed ecological responses to climate change are highly individualistic across species and locations, and understanding the drivers of this variability is essential for management and conservation efforts. While it is clear that differences in exposure, sensitivity, and adaptive capacity all contribute to heterogeneity in climate change vulnerability, predicting these features at macroecological scales remains a critical challenge. We explore multiple drivers of heterogeneous vulnerability across the distributions of 96 vegetation types of the ecologically diverse western US, using data on observed climate trends from 1948 to 2014 to highlight emerging patterns of change. We ask three novel questions about factors potentially shaping vulnerability across the region: (a) How does sensitivity to different climate variables vary geographically and across vegetation classes? (b) How do multivariate climate exposure patterns interact with these sensitivities to shape vulnerability patterns? (c) How different are these vulnerability patterns according to three widely implemented vulnerability paradigms-niche novelty (decline in modeled suitability), temporal novelty (standardized anomaly), and spatial novelty (inbound climate velocity)-each of which uses a distinct frame of reference to quantify climate departure? We propose that considering these three novelty paradigms in combination could help improve our understanding and prediction of heterogeneous climate change responses, and we discuss the distinct climate adaptation strategies connected with different combinations of high and low novelty across the three metrics. Our results reveal a diverse mosaic of climate change vulnerability signatures across the region's plant communities. Each of the above factors contributes strongly to this heterogeneity: climate variable sensitivity exhibits clear patterns across vegetation types, multivariate climate change data reveal highly diverse exposure signatures across locations, and the three novelty paradigms diverge widely in their climate change vulnerability predictions. Together, these results shed light on potential drivers of individualistic climate change responses and may help to inform effective management strategies.

Climate-change refugia: biodiversity in the slow lane.

Climate-change adaptation focuses on conducting and translating research to minimize the dire impacts of anthropogenic climate change, including threats to biodiversity and human welfare. One adaptation strategy is to focus conservation on climate-change refugia (that is, areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and sociocultural resources). In this Special Issue, recent methodological and conceptual advances in refugia science will be highlighted. Advances in this emerging subdiscipline are improving scientific understanding and conservation in the face of climate change by considering scale and ecosystem dynamics, and looking beyond climate exposure to sensitivity and adaptive capacity. We propose considering refugia in the context of a multifaceted, long-term, network-based approach, as temporal and spatial gradients of ecological persistence that can act as "slow lanes" rather than areas of stasis. After years of discussion confined primarily to the scientific literature, researchers and resource managers are now working together to put refugia conservation into practice.

Range edges in heterogeneous landscapes: Integrating geographic scale and climate complexity into range dynamics.

The impacts of climate change have re-energized interest in understanding the role of climate in setting species geographic range edges. Despite the strong focus on species' distributions in ecology and evolution, defining a species range edge is theoretically and empirically difficult. The challenge of determining a range edge and its relationship to climate is in part driven by the nested nature of geography and the multidimensionality of climate, which together generate complex patterns of both climate and biotic distributions across landscapes. Because range-limiting processes occur in both geographic and climate space, the relationship between these two spaces plays a critical role in setting range limits. With both conceptual and empirical support, we argue that three factors-climate heterogeneity, collinearity among climate variables, and spatial scale-interact to shape the spatial structure of range edges along climate gradients, and we discuss several ways that these factors influence the stability of species range edges with a changing climate. We demonstrate that geographic and climate edges are often not concordant across species ranges. Furthermore, high climate heterogeneity and low climate collinearity across landscapes increase the spectrum of possible relationships between geographic and climatic space, suggesting that geographic range edges and climatic niche limits correspond less frequently than we may expect. More empirical explorations of how the complexity of real landscapes shapes the ecological and evolutionary processes that determine species range edges will advance the development of range limit theory and its applications to biodiversity conservation in the context of changing climate.

Species Selection Regime and Phylogenetic Tree Shape.

Species selection, the effect of heritable traits in generating between-lineage diversification rate differences, provides a valuable conceptual framework for understanding the relationship between traits, diversification, and phylogenetic tree shape. An important challenge, however, is that the nature of real diversification landscapes-curves or surfaces which describe the propensity of species-level lineages to diversify as a function of one or more traits-remains poorly understood. Here, we present a novel, time-stratified extension of the QuaSSE model in which speciation/extinction rate is specified as a static or temporally shifting Gaussian or skewed-Gaussian function of the diversification trait. We then use simulations to show that the generally imbalanced nature of real phylogenetic trees, as well as their generally greater than expected frequency of deep branching events, are typical outcomes when diversification is treated as a dynamic, trait-dependent process. Focusing on four basic models (Gaussian-speciation with and without background extinction; skewed-speciation; Gaussian-extinction), we also show that particular features of the species selection regime produce distinct tree shape signatures and that, consequently, a combination of tree shape metrics has the potential to reveal the species selection regime under which a particular lineage diversified. We evaluate this idea empirically by comparing the phylogenetic trees of plant lineages diversifying within climatically and geologically stable environments of the Greater Cape Floristic Region, with those of lineages diversifying in environments that have experienced major change through the Late Miocene-Pliocene. Consistent with our expectations, the trees of lineages diversifying in a dynamic context are less balanced, show a greater concentration of branching events close to the present, and display stronger diversification rate-trait correlations. We suggest that species selection plays an important role in shaping phylogenetic trees but recognize the need for an explicit probabilistic framework within which to assess the likelihoods of alternative diversification scenarios as explanations of a particular tree shape. [Cape flora; diversification landscape; environmental change; gamma statistic; species selection; time-stratified QuaSSE model; trait-dependent diversification; tree imbalance.].

No local adaptation in leaf or stem xylem vulnerability to embolism, but consistent vulnerability segmentation in a North American oak.

Vulnerability to embolism varies between con-generic species distributed along aridity gradients, yet little is known about intraspecific variation and its drivers. Even less is known about intraspecific variation in tissues other than stems, despite results suggesting that roots, stems and leaves can differ in vulnerability. We hypothesized that intraspecific variation in vulnerability in leaves and stems is adaptive and driven by aridity. We quantified leaf and stem vulnerability of Quercus douglasii using the optical technique. To assess contributions of genetic variation and phenotypic plasticity to within-species variation, we quantified the vulnerability of individuals growing in a common garden, but originating from populations along an aridity gradient, as well as individuals from the same wild populations. Intraspecific variation in water potential at which 50% of total embolism in a tissue is observed (P50 ) was explained mostly by differences between individuals (>66% of total variance) and tissues (16%). There was little between-population variation in leaf/stem P50 in the garden, which was not related to site of origin aridity. Unexpectedly, we observed a positive relationship between wild individual stem P50 and aridity. Although there is no local adaptation and only minor phenotypic plasticity in leaf/stem vulnerability in Q. douglasii, high levels of potentially heritable variation within populations or strong environmental selection could contribute to adaptive responses under future climate change.

Beyond isohydricity: The role of environmental variability in determining plant drought responses.

Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012-2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a "common garden" thought experiment, how existing trait-based or response-based isohydricity metrics can be confounded by these environmental variations, leading to Type-1 (false positive) and Type-2 (false negative) errors; and (d) advocate for the use of model-based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale.

Microclimate and demography interact to shape stable population dynamics across the range of an alpine plant.

Heterogeneous terrain in montane systems results in a decoupling of climatic gradients. Population dynamics across species' ranges in these heterogeneous landscapes are shaped by relationships between demographic rates and these interwoven climate gradients. Linking demography and climate variables across species' ranges refines our understanding of the underlying mechanisms of species' current and future ranges. We explored the importance of multiple microclimatic gradients in shaping individual demographic rates and population growth rates in 16 populations across the elevational distribution of an alpine plant (Ivesia lycopodioides var. scandularis). Using integral projection modeling, we ask how each rate varies across three microclimate gradients: accumulated degree-days, growing-season soil moisture, and days of snow cover. Range-wide variation in demographic rates was best explained by the combined influence of multiple microclimatic variables. Different pairs of demographic rates exhibited both similar and inverse responses to the same microclimatic gradient, and the microclimatic effects often varied with plant size. These responses resulted in range-wide projected population persistence, with no declining populations at either elevational range edge or at the extremes of the microclimate gradients. The complex relationships between topography, microclimate and demography suggest that populations across a species' range may have unique demographic pathways to stable population dynamics.

Facets of phylodiversity: evolutionary diversification, divergence and survival as conservation targets.

Biodiversity is often described as having multiple facets, including species richness, functional diversity and phylogenetic diversity. In this paper, we argue that phylogenetic diversity itself has three distinct facets-lineage diversification, character divergence and survival time-that can be quantified using distinct branch length metrics on an evolutionary tree. Each dimension is related to different processes of macroevolution, has different spatial patterns and is tied to distinct goals for conserving biodiversity and protecting its future resilience and evolutionary potential. We compared the landscapes identified as top conservation priorities by each of these three metrics in a conservation gap analysis for California, a world biodiversity hotspot, using herbarium data on the biogeography and evolutionary relationships of more than 5000 native plant species. Our analysis incorporated a novel continuous metric of current land protection status, fine-scale data on landscape intactness and an optimization algorithm used to identify complementary priority sites containing concentrations of taxa that are evolutionarily unique, vulnerable due to small range size and/or poorly protected across their ranges. Top conservation priorities included pockets of coastal and northern California that ranked highly for all three phylodiversity dimensions and for species richness, as well as sites uniquely identified by each metric whose value may depend on whether properties such as genetic divergence, high net diversification or independent survival experience are most desirable in an Anthropocene flora.This article is part of the theme issue 'Biological collections for understanding biodiversity in the Anthropocene'.

The ecohydrological context of drought and classification of plant responses.

Many recent studies on drought-induced vegetation mortality have explored how plant functional traits, and classifications of such traits along axes of, for example, isohydry-anisohydry, might contribute to predicting drought survival and recovery. As these studies proliferate, the consistency and predictive value of such classifications need to be carefully examined. Here, we outline the basis for a systematic classification of plant drought responses that accounts for both environmental conditions and functional traits. We use non-dimensional analysis to integrate plant traits and metrics of environmental variation into groups that can be associated with alternative drought stress pathways (hydraulic failure and carbon limitation), and demonstrate that these groupings predict physiological drought outcomes using both synthetic and measured data. In doing so, we aim to untangle some confounding effects of environment and trait variations that undermine current classification schemes, advocate for more careful treatment of the environmental context within which plants experience and respond to drought, and outline a pathway towards a general classification of drought vulnerability.

Spatial phylogenetics of the native California flora.

BACKGROUND: California is a world floristic biodiversity hotspot where the terms neo- and paleo-endemism were first applied. Using spatial phylogenetics, it is now possible to evaluate biodiversity from an evolutionary standpoint, including discovering significant areas of neo- and paleo-endemism, by combining spatial information from museum collections and DNA-based phylogenies. Here we used a distributional dataset of 1.39 million herbarium specimens, a phylogeny of 1083 operational taxonomic units (OTUs) and 9 genes, and a spatial randomization test to identify regions of significant phylogenetic diversity, relative phylogenetic diversity, and phylogenetic endemism (PE), as well as to conduct a categorical analysis of neo- and paleo-endemism (CANAPE). RESULTS: We found (1) extensive phylogenetic clustering in the South Coast Ranges, southern Great Valley, and deserts of California; (2) significant concentrations of short branches in the Mojave and Great Basin Deserts and the South Coast Ranges and long branches in the northern Great Valley, Sierra Nevada foothills, and the northwestern and southwestern parts of the state; (3) significant concentrations of paleo-endemism in Northwestern California, the northern Great Valley, and western Sonoran Desert, and neo-endemism in the White-Inyo Range, northern Mojave Desert, and southern Channel Islands. Multiple analyses were run to observe the effects on significance patterns of using different phylogenetic tree topologies (uncalibrated trees versus time-calibrated ultrametric trees) and using different representations of OTU ranges (herbarium specimen locations versus species distribution models). CONCLUSIONS: These analyses showed that examining the geographic distributions of branch lengths in a statistical framework adds a new dimension to California floristics that, in comparison with climatic data, helps to illuminate causes of endemism. In particular, the concentration of significant PE in more arid regions of California extends previous ideas about aridity as an evolutionary stimulus. The patterns seen are largely robust to phylogenetic uncertainty and time calibration but are sensitive to the use of occurrence data versus modeled ranges, indicating that special attention toward improving geographic distributional data should be top priority in the future for advancing understanding of spatial patterns of biodiversity.

Species richness and endemism in the native flora of California.

PREMISE OF THE STUDY: California's vascular flora is the most diverse and threatened in temperate North America. Previous studies of spatial patterns of Californian plant diversity have been limited by traditional metrics, non-uniform geographic units, and distributional data derived from floristic descriptions for only a subset of species. METHODS: We revisited patterns of sampling intensity, species richness, and relative endemism in California based on equal-area spatial units, the full vascular flora, and specimen-based distributional data. We estimated richness, weighted endemism (inverse range-weighting of species), and corrected weighted endemism (weighted endemism corrected for richness), and performed a randomization test for significantly high endemism. KEY RESULTS: Possible biases in herbarium data do not obscure patterns of high richness and endemism at the spatial resolution studied. High species richness was sometimes associated with significantly high endemism (e.g., Klamath Ranges) but often not. In Stebbins and Major's (1965) main endemism hotspot, Southwestern California, species richness is high across much of the Peninsular and Transverse ranges but significantly high endemism is mostly localized to the Santa Rosa and San Bernardino mountains. In contrast, species richness is low in the Channel Islands, where endemism is significantly high, as also found for much of the Death Valley region. CONCLUSIONS: Measures of taxonomic richness, even with greater weighting of range-restricted taxa, are insufficient for identifying areas of significantly high endemism that warrant conservation attention. Differences between our findings and those in previous studies appear to mostly reflect the source and scale of distributional data, and recent analytical refinements.