Climate warming drives local extinction: Evidence from observation and experimentation.

Despite increasing concern about elevated extinction risk as global temperatures rise, it is difficult to confirm causal links between climate change and extinction. By coupling 25 years of in situ climate manipulation with experimental seed introductions and both historical and current plant surveys, we identify causal, mechanistic links between climate change and the local extinction of a widespread mountain plant (Androsace septentrionalis). Climate warming causes precipitous declines in population size by reducing fecundity and survival across multiple life stages. Climate warming also purges belowground seed banks, limiting the potential for the future recovery of at-risk populations under ameliorated conditions. Bolstered by previous reports of plant community shifts in this experiment and in other habitats, our findings not only support the hypothesis that climate change can drive local extinction but also foreshadow potentially widespread species losses in subalpine meadows as climate warming continues.

Pollinator-mediated competition influences selection for flower-color displacement in sympatric monkeyflowers.

PREMISE OF STUDY: When coflowering plant species share pollinators, pollinator-mediated competition may favor divergent floral characters associated with pollinator attraction. One potential outcome of this process is that sympatric populations will display increased divergence in floral traits compared with allopatric populations. We developed a new system to study the pattern and process of character displacement. In the central Sierra Nevada of California, USA, Mimulus bicolor is a spring wildflower with two flower-color morphs, one of which resembles coflowering M. guttatus. METHODS: We documented a fine-scale geographic pattern of character displacement in sympatric and allopatric patches and, using experimental arrays, measured seed set in M. bicolor color morphs in the presence versus absence of M. guttatus. KEY RESULTS: In sympatric arrays yellow, guttatus-like M. bicolor morphs had lower relative fitness (0.35 ± 0.05) and reduced conspecific pollen deposition compared with the distinct alternative morph, whereas in allopatric arrays yellow, guttatus-like morphs were occasionally strongly favored. CONCLUSIONS: Pollinator-mediated competition with M. guttatus is consistent with ecological character displacement in M. bicolor and likely contributes to a geographic pattern of character displacement.

Propagule limitation, disparate habitat quality, and variation in phenotypic selection at a local species range boundary.

Adaptation to novel conditions beyond current range boundaries requires the presence of suitable sites within dispersal range, but may be impeded when emigrants encounter poor habitat and sharply different selection pressures. We investigated fine-scale spatial heterogeneity in ecological dynamics and selection at a local population boundary of the annual plant Gilia tricolor. In two years, we planted G. tricolor seeds in core habitat, margin habitat at the edge of the local range, and exterior habitat in order to measure spatial and temporal variation in habitat quality, opportunity for selection, and selection on phenotypic traits. We found a striking decline in average habitat quality with distance from the population core, yet some migrant seeds were successful in suitable, unoccupied microsites at and beyond the range boundary. Total and direct selection on four out of five measured phenotypic traits varied across habitat zones, as well as between years. Moreover, the margin habitat often exerted unique selection pressures that were not intermediate between core and exterior habitats. This study reveals that a combination of ecological and evolutionary forces, including propagule limitation, variation in habitat quality and spatial heterogeneity in phenotypic selection may reduce opportunities for adaptive range expansion, even across a very local population boundary.

A role for indirect facilitation in maintaining diversity in a guild of African acacia ants.

Determining how competing species coexist is essential to understanding patterns of biodiversity. Indirect facilitation, in which a competitively dominant species exerts a positive effect on one competitor by more strongly suppressing a third, shared competitor, is a potentially potent yet understudied mechanism for competitive coexistence. Here we provide evidence for indirect facilitation in a guild of four African Acacia ant species that compete for nesting space on the host plant Acacia drepanolobium, showing that a competitively dominant acacia ant species indirectly creates establishment opportunities for the most subordinate species that may help to maintain diversity. Using long-term observational data and field experiments, we demonstrate that the competitively dominant ant species outcompetes two competitively intermediate species, while tolerating colonies of the subordinate competitor; this creates opportunities for local colonization and establishment of colonies of the subordinate species within the dominant species' territories. Host plants occupied by this subordinate species are then more likely to be colonized by the intermediate species, which in turn are more likely to be displaced by the dominant species. This process has the potential to generate a cyclical succession of ant species on host trees, contributing to stable coexistence within this highly competitive community.

Asymmetric and frequency-dependent pollinator-mediated interactions may influence competitive displacement in two vernal pool plants.

A plant species immigrating into a community may experience a rarity disadvantage due to competition for the services of pollinators. These negative reproductive interactions have the potential to lead to competitive displacement or exclusion of a species from a site. In this study, we used one- and two-species arrays of potted plants to test for density and frequency dependence in pollinator-mediated and above-ground intraspecific and interspecific competition between two species of Limnanthes that have overlapping ranges, but rarely occur in close sympatry. There were asymmetric competitive effects; the species responded differently to their frequency within 16-plant replacement series arrays. Limnanthes douglasii rosea experienced stronger reductions in lifetime and per-flower fertility, likely due to pollinator-mediated competition with Limnanthes alba. This effect may be linked to asymmetrical competition through heterospecific pollen transfer. This study demonstrates that pollinator-mediated competition may discourage establishment of L. d. rosea in sites already occupied by its congener.

Complex consequences of herbivory and interplant cues in three annual plants.

Information exchange (or signaling) between plants following herbivore damage has recently been shown to affect plant responses to herbivory in relatively simple natural systems. In a large, manipulative field study using three annual plant species (Achyrachaena mollis, Lupinus nanus, and Sinapis arvensis), we tested whether experimental damage to a neighboring conspecific affected a plant's lifetime fitness and interactions with herbivores. By manipulating relatedness between plants, we assessed whether genetic relatedness of neighboring individuals influenced the outcome of having a damaged neighbor. Additionally, in laboratory feeding assays, we assessed whether damage to a neighboring plant specifically affected palatability to a generalist herbivore and, for S. arvensis, a specialist herbivore. Our study suggested a high level of contingency in the outcomes of plant signaling. For example, in the field, damaging a neighbor resulted in greater herbivory to A. mollis, but only when the damaged neighbor was a close relative. Similarly, in laboratory trials, the palatability of S. arvensis to a generalist herbivore increased after the plant was exposed to a damaged neighbor, while palatability to a specialist herbivore decreased. Across all species, damage to a neighbor resulted in decreased lifetime fitness, but only if neighbors were closely related. These results suggest that the outcomes of plant signaling within multi-species neighborhoods may be far more context-specific than has been previously shown. In particular, our study shows that herbivore interactions and signaling between plants are contingent on the genetic relationship between neighboring plants. Many factors affect the outcomes of plant signaling, and studies that clarify these factors will be necessary in order to assess the role of plant information exchange about herbivory in natural systems.

Mutualism and adaptive divergence: co-invasion of a heterogeneous grassland by an exotic legume-rhizobium symbiosis.

Species interactions play a critical role in biological invasions. For example, exotic plant and microbe mutualists can facilitate each other's spread as they co-invade novel ranges. Environmental context may influence the effect of mutualisms on invasions in heterogeneous environments, however these effects are poorly understood. We examined the mutualism between the legume, Medicago polymorpha, and the rhizobium, Ensifer medicae, which have both invaded California grasslands. Many of these invaded grasslands are composed of a patchwork of harsh serpentine and relatively benign non-serpentine soils. We grew legume genotypes collected from serpentine or non-serpentine soil in both types of soil in combination with rhizobium genotypes from serpentine or non-serpentine soils and in the absence of rhizobia. Legumes invested more strongly in the mutualism in the home soil type and trends in fitness suggested that this ecotypic divergence was adaptive. Serpentine legumes had greater allocation to symbiotic root nodules in serpentine soil than did non-serpentine legumes and non-serpentine legumes had greater allocation to nodules in non-serpentine soil than did serpentine legumes. Therefore, this invasive legume has undergone the rapid evolution of divergence for soil-specific investment in the mutualism. Contrary to theoretical expectations, the mutualism was less beneficial for legumes grown on the stressful serpentine soil than on the non-serpentine soil, possibly due to the inhibitory effects of serpentine on the benefits derived from the interaction. The soil-specific ability to allocate to a robust microbial mutualism may be a critical, and previously overlooked, adaptation for plants adapting to heterogeneous environments during invasion.

Stroke, regional anesthesia in the sitting position, and hypotension: a review of 4169 ambulatory surgery patients.

BACKGROUND AND OBJECTIVE: Intraoperative hypotension is used to reduce surgical bleeding. Case reports of stroke after general anesthesia in the sitting position led us to collect data (patient demographics, medical risk factors for stroke, intraoperative hemodynamics) about the incidence of stroke after surgery in the sitting position. METHODS: This study reviewed 4169 (3000 retrospective, 1169 prospective) ambulatory shoulder surgeries in the sitting position. For the prospective cohort, patients were queried postoperatively regarding stroke, with corroboration from 4 databases (anesthesia department quality assurance, hospital case management, state-reportable events, and hospital information system diagnostic code databases). For the retrospective cohort, rate of stroke was determined via the same 4 databases. RESULTS: No patient had a stroke (95% confidence interval, 0%-0.07%). Risk factors for perioperative stroke were present in 40% of patients. Brachial plexus nerve block with intravenous sedation was used for 95.7% (retrospective) and 99.8% (prospective) of the cohorts. Many patients (47%) experienced intraoperative hypotension by at least one definition: 40% (retrospective) and 30% (prospective) had at least a 30% decrease in mean arterial pressure; 27% (retrospective) and 24% (prospective) had a mean pressure less than 66 mm Hg; and 13% (retrospective) and 12% (prospective) had a systolic blood pressure of less than 90 mm Hg. CONCLUSIONS: No strokes were observed in 4169 patients. The estimated upper limit of the 95% confidence interval for stroke after regional anesthesia for shoulder surgery in the seated position is 0.07%, despite frequent incidence of hypotension.

The high cost of mutualism: effects of four species of East African ant symbionts on their myrmecophyte host tree.

Three recent meta-analyses of protective plant-ant mutualisms report a surprisingly weak relationship between herbivore protection and measured demographic benefits to ant-plants, suggesting high tolerance for herbivory, substantial costs of ant-mediated defense, and/or benefits that are realized episodically rather than continuously. Experimental manipulations of protective ant-plant associations typically last for less than a year, yet virtually all specialized myrmecophytes are long-lived perennials for which the costs and benefits of maintaining ant symbionts could accrue at different rates over the host's lifetime. To complement long-term monitoring studies, we experimentally excluded each of four ant symbionts from their long-lived myrmecophyte host trees (Acacia drepanolobium) for 4.5 years. Ant species varied in their effectiveness against herbivores and in their effects on intermediate-term growth and reproduction, but the level of herbivore protection provided was a poor predictor of the net impact they had on host trees. Removal of the three Crematogaster species resulted in cumulative gains in host tree growth and/or reproduction over the course of the experiment, despite the fact that two of those species significantly reduce chronic herbivore damage. In contrast, although T. penzigi is a relatively poor defender, the low cost of maintaining this ant symbiont apparently eliminated negative impacts on overall tree growth and reproduction, resulting in enhanced allocation to new branch growth by the final census. Acacia drepanolobium is evidently highly tolerant of herbivory by insects and small browsers, and the costs of maintaining Crematogaster colonies exceeded the benefits received during the study. No experimental trees were killed by elephants, but elephant damage was uniquely associated with reduced tree growth, and at least one ant species (C. mimosae) strongly deterred elephant browsing. We hypothesize that rare but catastrophic damage by elephants may be more important than chronic herbivory in maintaining the costly myrmecophyte habit in this system.

Fitness variation and local distribution limits in an annual plant population.

Understanding how genetic variation shapes species' distributions involves examining how variation is distributed across a species' range as well as how it responds to underlying environmental heterogeneity. We examined patterns of fitness variation across the local distribution of an annual composite (Lasthenia fremontii) spanning a small-scale inundation gradient in a California vernal pool wetland. Using seeds collected from the center and edge of a population, paternal half-sib families were generated and transplanted back to the center and edge of the original population. All transplants were adapted to the conditions at the center of the population. The effect of the environment on the opportunity for selection depended on the model of selection assumed. Under a model of hard selection, variance in absolute fitness was lower among transplants at the edge of the population than at the center. Under a model of soft selection, the variance in relative fitness was similar between center and edge microhabitats. Given that this population is likely well-mixed, differences in habitat quality between center and edge microhabitats will likely cause selection at the center of the population to dominate the evolutionary trajectory of this population.

Synergy of multiple partners, including freeloaders, increases host fitness in a multispecies mutualism.

Understanding cooperation is a central challenge in biology, because natural selection should favor "free-loaders" that reap benefits without reciprocating. For interspecific cooperation (mutualism), most approaches to this paradox focus on costs and benefits of individual partners and the strategies mutualists use to associate with beneficial partners. However, natural selection acts on lifetime fitness, and most mutualists, particularly longer-lived species interacting with shorter-lived partners (e.g., corals and zooxanthellae, tropical trees and mycorrhizae) interact with multiple partner species throughout ontogeny. Determining how multiple partnerships might interactively affect lifetime fitness is a crucial unexplored link in understanding the evolution and maintenance of cooperation. The tropical tree Acacia drepanolobium associates with four symbiotic ant species whose short-term individual effects range from mutualistic to parasitic. Using a long-term dataset, we show that tree fitness is enhanced by partnering sequentially with sets of different ant symbionts over the ontogeny of a tree. These sets include a "sterilization parasite" that prevents reproduction and another that reduces tree survivorship. Trees associating with partner sets that include these "parasites" enhance lifetime fitness by trading off survivorship and fecundity at different life stages. Our results demonstrate the importance of evaluating mutualism within a community context and suggest that lifespan inequalities among mutualists may help cooperation persist in the face of exploitation.

Characterizing selection on phenotypic plasticity in response to natural environmental heterogeneity.

Adaptive genetic differentiation and adaptive phenotypic plasticity can increase the fitness of plant lineages in heterogeneous environments. We examine the relative importance of genetic differentiation and plasticity in determining the fitness of the annual plant, Erodium cicutarium, in a serpentine grassland in California. Previous work demonstrated that the serpentine sites within this mosaic display stronger dispersal-scale heterogeneity than nonserpentine sites. We conducted a reciprocal transplant experiment among six sites to characterize selection on plasticity expressed by 180 full-sibling families in response to natural environmental heterogeneity across these sites. Multivariate axes of environmental variation were constructed using a principal components analysis of soil chemistry data collected at every experimental block. Simple linear regressions were used to characterize the intercept, and slope (linear and curvilinear) of reaction norms for each full-sibling family in response to each axis of environmental variation. Multiple linear regression analyses revealed significant selection on trait means and slopes of reaction norms. Multivariate analyses of variance demonstrated genetic differentiation between serpentine and nonserpentine lineages in the expression of plasticity in response to three of the five axes of environmental variation considered. In all but one case, serpentine genotypes expressed a stronger adaptive plastic response than nonserpentine genotypes.

Understanding the consequences of seed dispersal in a heterogeneous environment.

Plant distributions are in part determined by environmental heterogeneity on both large (landscape) and small (several meters) spatial scales. Plant populations can respond to environmental heterogeneity via genetic differentiation between large distinct patches, and via phenotypic plasticity in response to heterogeneity occurring at small scales relative to dispersal distance. As a result, the level of environmental heterogeneity experienced across generations, as determined by seed dispersal distance, may itself be under selection. Selection could act to increase or decrease seed dispersal distance, depending on patterns of heterogeneity in environmental quality with distance from a maternal home site. Serpentine soils, which impose harsh and variable abiotic stress on non-adapted plants, have been partially invaded by Erodium cicutarium in northern California, USA. Using nearby grassland sites characterized as either serpentine or non-serpentine, we collected seeds from dense patches of E. cicutarium on both soil types in spring 2004 and subsequently dispersed those seeds to one of four distances from their maternal home site (0, 0.5, 1, or 10 m). We examined distance-dependent patterns of variation in offspring lifetime fitness, conspecific density, soil availability, soil water content, and aboveground grass and forb biomass. ANOVA revealed a distinct fitness peak when seeds were dispersed 0.5 m from their maternal home site on serpentine patches. In non-serpentine patches, fitness was reduced only for seeds placed back into the maternal home site. Conspecific density was uniformly high within 1 m of a maternal home site on both soils, whereas soil water content and grass biomass were significantly heterogeneous among dispersal distances only on serpentine soils. Structural equation modeling and multigroup analysis revealed significantly stronger direct and indirect effects linking abiotic and biotic variation to offspring performance on serpentine soils than on non-serpentine soils, indicating the potential for soil-specific selection on seed dispersal distance in this invasive species.

Cryptic seedling herbivory by nocturnal introduced generalists impacts survival, performance of native and exotic plants.

Although much of the theory on the success of invasive species has been geared at escape from specialist enemies, the impact of introduced generalist invertebrate herbivores on both native and introduced plant species has been underappreciated. The role of nocturnal invertebrate herbivores in structuring plant communities has been examined extensively in Europe, but less so in North America. Many nocturnal generalists (slugs, snails, and earwigs) have been introduced to North America, and 96% of herbivores found during a night census at our California Central Valley site were introduced generalists. We explored the role of these herbivores in the distribution, survivorship, and growth of 12 native and introduced plant species from six families. We predicted that introduced species sharing an evolutionary history with these generalists might be less vulnerable than native plant species. We quantified plant and herbivore abundances within our heterogeneous site and also established herbivore removal experiments in 160 plots spanning the gamut of microhabitats. As 18 collaborators, we checked 2000 seedling sites every day for three weeks to assess nocturnal seedling predation. Laboratory feeding trials allowed us to quantify the palatability of plant species to the two dominant nocturnal herbivores at the site (slugs and earwigs) and allowed us to account for herbivore microhabitat preferences when analyzing attack rates on seedlings. The relationship between local slug abundance and percent cover of five common plant taxa at the field site was significantly negatively associated with the mean palatability of these taxa to slugs in laboratory trials. Moreover, seedling mortality of 12 species in open-field plots was positively correlated with mean palatability of these taxa to both slugs and earwigs in laboratory trials. Counter to expectations, seedlings of native species were neither more vulnerable nor more palatable to nocturnal generalists than those of introduced species. Growth comparison of plants within and outside herbivore exclosures also revealed no differences between native and introduced plant species, despite large impacts of herbivores on growth. Cryptic nocturnal predation on seedlings was common and had large effects on plant establishment at our site. Without intensive monitoring, such predation could easily be misconstrued as poor seedling emergence.

Factors driving distribution limits in an annual plant community.

Studies examining plant distribution patterns across environmental gradients have generally focused on perennial-dominated systems, and we know relatively little about the processes structuring annual communities. Here, the ecological factors determining local distribution patterns of five dominant annual species distributed across micro-topographic gradients in ephemeral California wetlands are examined. Over two growing seasons in three vernal pools, patterns of inundation and above-ground biomass were characterized across the microtopographic gradient, population boundaries for five dominant species were documented and a reciprocal transplant experiment and neighbor removal treatment were conducted to test the relative effects of within-pool elevation, competition and seed dispersal on plant performance. Despite large differences in inundation time between growing seasons, above-ground biomass and the elevation of population boundaries remained consistent. The predicted 'optimal' depth for each species shifted between years, but competition and recruitment limitation restricted species' abilities to track these conditions. The distributions of the focal taxa are primarily driven by differential responses to environmental conditions associated with different microtopographic positions along pool inundation gradients, and are reinforced by competition and dispersal constraints. The relative importance of competition, other environmental factors and dispersal patterns appear to contrast with results obtained in systems dominated by perennial plants.

One-year experience with day-of-surgery pregnancy testing before elective orthopedic procedures.

BACKGROUND: Elective surgery is generally postponed in pregnancy. A policy of testing for urine human chorionic gonadotropin (hCG) in all women of childbearing age on the day of surgery was initiated at an elective orthopedic surgery facility. This is a retrospective report of our 1 yr experience and the associated costs. METHODS: Records were reviewed from January 3, 2005, through January 2, 2006, to determine the number of urine hCG tests performed, and the disposition of all patients with a positive result. Costs were calculated using the charges for testing supplies and laboratory labor. RESULTS: During the first year of policy implementation, 2588 of 2595 women of childbearing age underwent urine hCG testing. Five patients had a positive result, and surgery was cancelled. Of these five, three were previously unrecognized pregnancies, one an unrecognized asymptomatic ectopic pregnancy, and one a false-positive result in a perimenopausal woman. Three other women had a "weak positive" urine result, followed by a negative serum hCG. Surgery proceeded in all three. The calculated cost was $5.03 per urine test, and $3273 for each true positive result. CONCLUSIONS: A policy of routinely performing urine hCG pregnancy tests in women of childbearing age on the day of surgery was effective in detecting unrecognized pregnancy. This resulted in a postponement of elective surgery in all cases. Of 2588 women tested, one had surgery postponed because of a false-positive result. The cost of $3273 per true positive test must be compared with the benefit.

A new approach to preanesthetic site verification after 2 cases of wrong site peripheral nerve blocks.

OBJECTIVE: We discuss the importance of a "preanesthetic site verification" and highlight 2 significant modifications to a policy developed at our institution in 2003. CASE REPORT: The report describes 2 cases of wrong site peripheral nerve blocks that initiated protocol amendments to address shortcomings of the original policy. Two specific limitations were identified to improve upon. First, the practitioner must not overlook the site verification as it is essential prior to every block. Second, time delays between the verification and block performance should be minimized. CONCLUSIONS: The "preanesthetic site verification" is an integral part of preventing wrong site block and surgery. To ensure that it is carried out before every peripheral nerve block, a unique multidisciplinary approach was adopted in which the block needles were removed from anesthesia carts and transferred to a separate container in the area of the circulating nurse. The anesthesiologist must now request a block needle from the circulating nurse immediately prior to block performance and confirm the site at that time. This safety process emulates the presurgical site verification that takes place before a scalpel is passed to a surgeon. Furthermore, the circulating nurse must remain at the bedside until block initiation to make sure that delays between site verification and block performance do not impinge on correct site placement.

Breakdown of an ant-plant mutualism follows the loss of large herbivores from an African savanna.

Mutualisms are key components of biodiversity and ecosystem function, yet the forces maintaining them are poorly understood. We investigated the effects of removing large mammals on an ant-Acacia mutualism in an African savanna. Ten years of large-herbivore exclusion reduced the nectar and housing provided by plants to ants, increasing antagonistic behavior by a mutualistic ant associate and shifting competitive dominance within the plant-ant community from this nectar-dependent mutualist to an antagonistic species that does not depend on plant rewards. Trees occupied by this antagonist suffered increased attack by stem-boring beetles, grew more slowly, and experienced doubled mortality relative to trees occupied by the mutualistic ant. These results show that large mammals maintain cooperation within a widespread symbiosis and suggest complex cascading effects of megafaunal extinction.

Collinsia sparsiflora in serpentine and nonserpentine habitats: using F2 hybrids to detect the potential role of selection in ecotypic differentiation.

Here we document phenotypic differences between serpentine and nonserpentine ecotypes of Collinsia sparsiflora, as well as patterns of selection in these contrasting soil habitats. We transplanted the two parental ecotypes and experimental F2 hybrids into six field sites, and collected morphological, phenological and fitness data on emergent plants. To focus on edaphically mediated selection, rather than on pollinator-mediated selection, we used pollinator-exclusion cages. Transplanted parentals of the two ecotypes showed genetic differentiation for floral traits, but not for phenological traits or cotyledon size. For the F2 hybrids growing on serpentine soils, there was significant directional selection on cotyledon size, flower size and flower shape. However, the pattern of selection did not differ significantly between serpentine and nonserpentine sites. Overall, we did not see evidence for divergent selection on the two soil types. We conclude that differences in floral traits between the ecotypes do not reflect adaptation to physical conditions associated with soil type, and that there are unmeasured traits that must be contributing to ecotypic differentiation.

Ecological factors limiting the distribution of Gilia tricolor in a California grassland mosaic.

Recent evolutionary models of range limits emphasize the importance of ecological and demographic factors operating at species' margins. This study aims to establish the ecological context driving population boundaries in Gilia tricolor, a native California annual restricted to distinct habitat patches in the coastal range of California. A transplant experiment in one hillside G. tricolor population examined the roles of competition and soil chemistry as well as litter and biomass accumulation in setting local population boundaries. Results indicate that boundaries are maintained primarily by inhibition of seedling emergence by vegetation and litter, and that upslope and downslope population boundaries are heterogeneous in litter biomass and transplant performance. Consistent emergence inhibition in undisturbed, peripheral sites maintains limits to the distribution of G. tricolor in this population. Fine-scaled ecological heterogeneity and heterogeneous boundary conditions likely play important roles in limiting adaptation and subsequent range expansion at population boundaries in G. tricolor.