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Background and aimsThe evolution of selfing from outcrossing may be the most common transition in plant reproductive systems and is associated with a variety of ecological circumstances and life history strategies. The most widely discussed explanation for these associations is the reproductive assurance hypothesis - the proposition that selfing is favoured because it increases female fitness when outcross pollen receipt is limited. Here an alternative explanation, the time limitation hypothesis, is addressed, one scenario of which proposes that selfing may evolve as a correlated response to selection for a faster life cycle in seasonally deteriorating environments.MethodsArtificial selection for faster maturation (early flowering) or for low herkogamy was performed on Clarkia unguiculata (Onagraceae), a largely outcrossing species whose closest relative, C. exilis, has evolved higher levels of autogamous selfing. Direct responses to selection and correlated evolutionary changes in these traits were measured under greenhouse conditions. Direct responses to selection on early flowering and correlated evolutionary changes in the node of the first flower, herkogamy, dichogamy, gas exchange rates and water use efficiency (WUE) were measured under field conditions.Key resultsLines selected for early flowering and for low herkogamy showed consistent, statistically significant responses to direct selection. However, there was little or no evidence of correlated evolutionary changes in flowering date, floral traits, gas exchange rates or WUE.ConclusionsThese results suggest that the maturation rate and mating system have evolved independently in Clarkia and that the time limitation hypothesis does not explain the repeated evolution of selfing in this genus, at least through its indirect selection scenario. They also suggest that the life history and physiological components of drought escape are not genetically correlated in Clarkia, and that differences in gas exchange physiology between C. unguiculata and C. exilis have evolved independently of differences in mating system and life history.

Background and Aims Mating systems of plants are diverse and evolutionarily labile. Abiotic environmental factors, such as seasonal drought, may impose selection on physiological traits that could lead to transitions in mating system if physiological traits are genetically correlated with traits that influence mating system. Within Clarkia, self-fertilizing taxa have higher photosynthetic rates, earlier flowering phenology, faster individual floral development and more compressed flowering periods than their outcrossing sister taxa, potentially reducing the selfing taxa's exposure to drought. In theory, this contrast in trait combinations between sister taxa could have arisen via correlated evolution due to pleiotropy or genetic linkage. Alternatively, each trait may evolve independently as part of a life history that is adaptive in seasonally dry environments. Methods To evaluate these hypotheses, we examined relationships between photosynthetic rates (adjusted for plant height and leaf node position) and outcrossing rates (estimated by allozyme variation in progeny arrays) during two consecutive years in multiple wild populations of two mixed-mating Clarkia taxa, each of which is sister to a derived selfing taxon. If the negative association between photosynthetic rate and outcrossing previously observed between sister taxa reflects correlated evolution due to a strong negative genetic correlation between these traits, then a similarly negative relationship would be observed within populations of each taxon. By contrast, if the combination of elevated photosynthetic rates and reduced outcrossing evolved independently within taxa, we predicted no consistent relationship between photosynthetic rate and outcrossing rate. Key Results We found no significant difference in outcrossing rates within populations between groups of plants with high versus low photosynthetic rates. Conclusions Overall, these results provide support for the hypothesis that the joint divergence in photosynthetic rate and mating system observed between Clarkia sister taxa is the result of independent evolutionary transitions.

In habitats where resource availability declines during the growing season, selection may favor early-flowering individuals. Under such ephemerally favorable conditions, late-blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation (PL) of seed set and outcrossing rates between early- and late-flowering individuals in two mixed-mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008-2010, we hand-pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open-pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open-pollinated flowers. Second (2009), we conducted an anther-removal experiment to estimate self-pollen deposition. Seed set in Clarkia unguiculata was not pollen-limited. Clarkia xantiana ssp. xantiana was pollen-limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator-dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 - and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules.

Premise of the studyOne proposed function of long styles is to intensify selection among male gametophytes relative to short styles. If so, given sufficient competition, longer styles will have higher rates of pollen tube attrition (failure to reach the style base) within the style than shorter ones. Alternatively, style length may influence pollen receipt, which itself may affect attrition rates.MethodsWe tested these predictions by collecting senescing styles from wild populations of two insect-pollinated Clarkia species. We examined the number of pollen grains adhering to the stigma, length of styles, and rates of attrition from the stigma surface to the stigma-style junction (SSJ), from the SSJ to the style base, and from the stigma surface to the style base. Multivariate analyses estimated the independent effects of pollen grains per stigma, the number of pollen tubes at the SSJ, and style length on attrition.Key resultsStyle length was generally positively correlated with pollen receipt, and the number of pollen grains per stigma was positively correlated with all three attrition rates. In neither species was any attrition rate affected by style length independent of the number of pollen grains per stigma.ConclusionsPollen attrition was mediated by style length, but the function of style length was primarily to increase the number of germinating pollen grains, which affected attrition rates either through stigma clogging or pollen-pollen interactions. Style length may have a direct effect on pollen receipt due to the stigma's position relative to pollinator body parts, but traits correlated with style length may also directly affect pollen receipt.

Premise of the studyOne explanation for the evolution of selfing, the drought escape hypothesis, proposes that self-fertilization may evolve under conditions of intensifying seasonal drought as part of a suite of traits that enable plants to accelerate the completion of their life cycle, thereby escaping late-season drought. Here, we test two fundamental assumptions of this hypothesis in Clarkia xantiana: (1) that a seasonal decline in precipitation causes an increase in drought stress and (2) that this results in changes in physiological performance, reflecting these deteriorating conditions.MethodsWe examined seasonal and interannual variation in abiotic environmental conditions (estimated by ambient temperature, relative humidity, predawn leaf water potentials, and carbon isotope ratios) and physiological traits (photosynthesis, conductance, transpiration, instantaneous water-use efficiency, ascorbate peroxidase and glutathione reductase activities, quantum yield of photosystem II, PSII potential efficiency) in field populations of C. xantiana in 2009 and 2010.Key resultsIn both years, plants experienced intensifying drought across the growing season. Gas exchange rates decreased over the growing season and were lower in 2009 (a relatively dry year) than in 2010, suggesting that the temporal changes from early to late spring were directly linked to the deteriorating environmental conditions.ConclusionsSeasonal declines in transpiration rate may have increased survival by protecting plants from desiccation. Concomitant declines in photosynthetic rate likely reduced the availability of resources for seed production late in the season. Thus, the physiological patterns observed are consistent with the conditions required for the drought escape hypothesis.

Temporal variation in stress might signify changes in an animal's internal or external environment, while spatial variation in stress might signify variation in the quality of the habitats that individual animals experience. Habitat-induced variations in stress might be easiest to detect in highly territorial animals, and especially in species that do not take advantage of common strategies for modulating habitat-induced stress, such as migration (escape in space) or hibernation (escape in time). Spatial and temporal variation in response to potential stressors has received little study in wild animals, especially at scales appropriate for relating stress to specific habitat characteristics. Here, we use the American pika (Ochotona princeps), a territorial small mammal, to investigate stress response within and among territories. For individually territorial animals such as pikas, differences in habitat quality should lead to differences in stress exhibited by territory owners. We indexed stress using stress-associated hormone metabolites in feces collected non-invasively from pika territories every 2 weeks from June to September 2018. We hypothesized that differences in territory quality would lead to spatial differences in mean stress and that seasonal variation in physiology or the physical environment would lead to synchronous variation across territories through time. We used linear mixed-effects models to explore spatiotemporal variation in stress using fixed effects of day-of-year and broad habitat characteristics (elevation, aspect, site), as well as local variation in habitat characteristics hypothesized to affect territory quality for this saxicolous species (talus depth, clast size, available forage types). We found that temporal variation within territories was greater than spatial variation among territories, suggesting that shared seasonal stressors are more influential than differences in individual habitat quality. This approach could be used in other wildlife studies to refine our understanding of habitat quality and its effect on individual stress levels as a driver of population decline. [ABSTRACT FROM AUTHOR]