1,3 Dipolar Cycloaddition of Münchnones: Factors behind the Regioselectivity.

The 1,3 dipolar cycloaddition reactions of münchnones and alkenes provide an expedite synthetic way to substituted pyrroles, an exceedingly important structural motif in the pharmaceutical and material science fields of research. The factors governing their regioselectivity rationalization are not well understood. Using several approaches, we investigate a set of 14 reactions (featuring two münchnones, 12 different alkenes, and two alkynes). The Natural Bond Theory and the Non-Covalent Interaction Index analyses of the noncovalent interaction energies fail to predict the experimental major regioisomer. Employing global cDFT descriptors or local ones such as the Fukui function and dual descriptor yields similarly inaccurate predictions. Only the local softness pairing, within Pearson's Hard and Soft Acids and Bases principle, constitutes a reliable predictor for the major reaction product. By taking into account an estimator for the steric effects, the correct regioisomer is predicted. Steric effects play a major role in driving the regioselectivity, as was corroborated by energy decomposition analysis of the transition states.

Disturbance and the (surprising?) role of ecosystem engineering in explaining spatial patterns of non-native plant establishment.

Different conceptions of disturbance differ in the degree to which they appeal to mechanisms that are general and equivalent, or species-, functional group-, or interaction-specific. Some concepts of disturbance, for example, predict that soil disturbances and herbivory have identical impacts on species richness via identical mechanisms (reduction in biomass and in competition). An alternative hypothesis is that the specific traits of disturbance agents (small mammals) and plants differentially affect the richness or abundance of different plant groups. We tested these hypotheses on a degu (Octodon degus) colony in central Chile. We ask whether native and non-native forbs respond differently to degu bioturbation on runways versus herbivory on grazing lawns. We ask whether this can explain the increase in non-native plants on degu colonies. We found that biopedturbation did not explain the locations of non-native plants. We did not find direct evidence of grazing increasing non-native herbs either, but a grazing effect appears to be mediated by grass, which is the dominant cover. Further, we provide supplementary evidence to support our interpretation that a key mechanism of non-native spread is the formation of dry soil conditions on grazing lawns. Thus, ecosystem engineering (alteration of soil qualities) may be an outcome of disturbances, in which each interacts with specific plant traits, to create the observed pattern of non-native spread in the colony. Based on these results, we propose to extend Jentsch and White (Ecology, 100, 2019, e02734) concept of combined pulse/ disturbance events to the long-term process duality of ecosystem engineering/ disturbance.

COSORE: A community database for continuous soil respiration and other soil-atmosphere greenhouse gas flux data.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil-to-atmosphere CO2 flux, commonly though imprecisely termed soil respiration (RS ), is one of the largest carbon fluxes in the Earth system. An increasing number of high-frequency RS measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open-source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long-term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured RS , the database design accommodates other soil-atmosphere measurements (e.g. ecosystem respiration, chamber-measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package.

Freezing and water availability structure the evolutionary diversity of trees across the Americas.

The historical course of evolutionary diversification shapes the current distribution of biodiversity, but the main forces constraining diversification are still a subject of debate. We unveil the evolutionary structure of tree species assemblages across the Americas to assess whether an inability to move or an inability to evolve is the predominant constraint in plant diversification and biogeography. We find a fundamental divide in tree lineage composition between tropical and extratropical environments, defined by the absence versus presence of freezing temperatures. Within the Neotropics, we uncover a further evolutionary split between moist and dry forests. Our results demonstrate that American tree lineages tend to retain their ancestral environmental relationships and that phylogenetic niche conservatism is the primary force structuring the distribution of tree biodiversity. Our study establishes the pervasive importance of niche conservatism to community assembly even at intercontinental scales.

Soil invertebrate diversity loss and functional changes in temperate forest soils replaced by exotic pine plantations.

The global expansion of tree plantations is often claimed to have positive effects for mitigating global warming, preventing soil erosion, and reducing biodiversity loss. However, questions remain unanswered about the impacts of plantations on belowground diversity and soil properties. Here, we examine how forestry plantations of exotic trees affect critical soil functions and the composition of invertebrate assemblages, by comparing invertebrate diversity and soil physico-chemical properties between non-native Pinus radiata plantations, and nearby native forests in a region of extensive plantation activity in south-central Chile. We quantified differences in diversity, abundance, and community composition of soil invertebrates, as well as fundamental soil properties such as soil water content, water infiltration, nutrient status, and pH. We show that in this landscape mosaic of native forest and plantations, both soil invertebrate communities and physical soil properties differed significantly between systems, despite similar soil origins and topographies. We found a significant loss of soil carbon and a major reduction in taxonomic and functional diversity of soil invertebrates in pine plantation sites. Soil biotic and abiotic characteristics of plantations differed significantly from native forests in plantation-dominated south-central Chile, with profound consequences for ecosystem processes and resilience to future climate change.

Coupling of microbial nitrogen transformations and climate in sclerophyll forest soils from the Mediterranean Region of central Chile.

The Mediterranean region of central Chile is experiencing extensive "mega-droughts" with detrimental effects for the environment and economy of the region. In the northern hemisphere, nitrogen (N) limitation of Mediterranean ecosystems has been explained by the decoupling between N inputs and plant uptake during the dormant season. In central Chile, soils have often been considered N-rich in comparison to other Mediterranean ecosystems of the world, yet the impacts of expected intensification of seasonal drought remain unknown. In this work, we seek to disentangle patterns of microbial N transformations and their seasonal coupling with climate in the Chilean sclerophyll forest-type. We aim to assess how water limitation affects microbial N transformations, thus addressing the impact of ongoing regional climate trends on soil N status. We studied four stands of the sclerophyll forest-type in Chile. Field measurements in surface soils showed a 67% decline of free-living diazotrophic activity (DA) and 59% decrease of net N mineralization rates during the summer rainless and dormant season, accompanied by a stimulation of in-situ denitrification rates to values 70% higher than in wetter winter. Higher rates of both free-living DA and net N mineralization found during spring, provided evidence for strong coupling of these two processes during the growing season. Overall, the experimental addition of water in the field to litter samples almost doubled DA but had no effect on denitrification rates. We conclude that coupling of microbial mediated soil N transformations during the wetter growing season explains the N enrichment of sclerophyll forest soils. Expected increases in the length and intensity of the dry period, according to climate change models, reflected in the current mega-droughts may drastically reduce biological N fixation and net N mineralization, increasing at the same time denitrification rates, thereby potentially reducing long-term soil N capital.

Assessing the influence of life form and life cycle on the response of desert plants to past climate change: Genetic diversity patterns of an herbaceous lineage of Nolana along western South America.

PREMISE OF THE STUDY: Plant responses to past climate change could have been shaped by life-history traits. Here we explore the influence of life form on the response of xerophytic plants to Quaternary climate fluctuations, through a comparison of genetic patterns of codistributed herbaceous and shrubby lineages of the genus Nolana. METHODS: We reconstructed the phylogeographic history of a herbaceous lineage of three species of Nolana distributed from a northern arid zone (30°S) to a southern wet-temperate (42°S) zone, by sequencing two cpDNA regions. Results were compared with similar data published earlier for a congeneric, codistributed shrubby lineage. KEY RESULTS: We detected significant genetic differentiation among populations. Divergence of all haplotypes occurred during the Pleistocene, between 245 and 62 kyr ago. For both the shrubby and herbaceous lineages, the greatest haplotype diversity was found in their northern range. However, herbs also retained some diversity at higher latitude. Herbaceous populations were less genetically structured and less differentiated than shrubby ones. CONCLUSION: Genetic evidence revealed that both lineages of Nolana survived climate change through the Quaternary, experiencing population collapses and recoveries. Phylogeographic histories present similarities between the two lineages, but also marked differences that can be explained by their differences in life form and life cycle. While the shrubby lineage followed the classical pattern of postglacial expansion toward higher latitudes, species in the herbaceous lineage showed evidence of long-lasting persistence at the southern edge of their current range, suggesting for the first time multiple glacial refugia for a xerophytic plant in southern South America.

Relationship between soil nutrients and mycorrhizal associations of two Bipinnula species (Orchidaceae) from central Chile.

BACKGROUND AND AIMS: Mycorrhizal associations are influenced by abiotic and biotic factors, including climate, soil conditions and the identity of host plants. However, the effect of environmental conditions on orchid mycorrhizal associations remains poorly understood. The present study examined how differences in soil nutrient availability are related to the diversity and composition of mycorrhizal fungi associated with two terrestrial orchid species from central Chile. METHODS: For 12 populations of Bipinnula fimbriata and B. plumosa, OTU (operational taxonomic unit) richness, phylogenetic diversity and community composition of mycorrhizal fungi in root samples were estimated using internal transcribed spacer (ITS) sequences. Then, these mycorrhizal diversity variables were related to soil nutrients and host species using generalized linear models and non-metric multidimensional scaling. KEY RESULTS: Variation in OTU composition of mycorrhizal fungi among sites was explained mainly by orchid host species. Fungi in Tulasnellaceae and Ceratobasidiaceae were isolated from both orchid species, but the former were more frequent in B. fimbriata and the latter in B. plumosa. Soil nutrients and orchid host species had significant effects on OTU richness and phylogenetic diversity. Mycorrhizal diversity decreased in habitats with higher N in both species and increased with P availability only in B. fimbriata CONCLUSIONS: The results suggest that soil nutrient availability modulates orchid mycorrhizal associations and provide support for the hypothesis that specialization is favoured by higher soil nutrient availability. Inter-specific differences in mycorrhizal composition can arise due to a geographical pattern of distribution of orchid mycorrhizal fungi, host preferences for fungal partners or differential performance of mycorrhizal fungi under different nutrient availabilities. Further experiments are needed to evaluate these hypotheses.

Functional traits variation explains the distribution of Aextoxicon punctatum (Aextoxicaceae) in pronounced moisture gradients within fog-dependent forest fragments.

Climate change and fragmentation are major threats to world forests. Understanding how functional traits related to drought tolerance change across small-scale, pronounced moisture gradients in fragmented forests is important to predict species' responses to these threats. In the case of Aextoxicon punctatum, a dominant canopy tree in fog-dependent rain forest patches in semiarid Chile, we explored how the magnitude, variability and correlation patterns of leaf and xylem vessel traits and hydraulic conductivity varied across soil moisture (SM) gradients established within and among forest patches of different size, which are associated with differences in tree establishment and mortality patterns. Leaf traits varied across soil-moisture gradients produced by fog interception. Trees growing at drier leeward edges showed higher leaf mass per area, trichome and stomatal density than trees from the wetter core and windward zones. In contrast, xylem vessel traits (vessels diameter and density) did not vary producing loss of hydraulic conductivity at drier leeward edges. We also detected higher levels of phenotypic integration and variability at leeward edges. The ability of A. punctatum to modify leaf traits in response to differences in SM availability established over short distances (<500 m) facilitates its persistence in contrasting microhabitats within forest patches. However, xylem anatomy showed limited plasticity, which increases cavitation risk at leeward edges. Greater patch fragmentation, together with fluctuations in irradiance and SM in small patches, could result in higher risk of drought-related tree mortality, with profound impacts on hydrological balances at the ecosystem scale.

Understanding litter decomposition in semiarid ecosystems: linking leaf traits, UV exposure and rainfall variability.

Differences in litter quality, microbial activity or abiotic conditions cannot fully account for the variability in decomposition rates observed in semiarid ecosystems. Here we tested the role of variation in litter quality, water supply, and UV radiation as drivers of litter decomposition in arid lands. And show that carry-over effects of litter photodegradation during dry periods can regulate decomposition during subsequent wet periods. We present data from a two-phase experiment, where we first exposed litter from a drought-deciduous and an evergreen shrub to natural UV levels during five, rainless summer months and, subsequently, in the laboratory, we assessed the carry-over effects of photodegradation on biomass loss under different irrigation treatments representing the observed range of local rainfall variation among years (15-240 mm). Photodegradation of litter in the field produced average carbon losses of 12%, but deciduous Proustia pungens lost >25%, while evergreen Porlieria chilensis less than 5%. Natural exposure to UV significantly reduced carbon-to-nitrogen and lignin:N ratios in Proustia litter but not in Porlieria. During the subsequent wet phase, remaining litter biomass was lower in Proustia than in Porlieria. Indeed UV exposure increased litter decomposition of Proustia under low and medium rainfall treatments, whereas no carry-over effects were detected under high rainfall treatment. Consequently, for deciduous Proustia carry-over effects of UV exposure were negligible under high irrigation. Litter decomposition of the evergreen Porlieria depended solely on levels of rainfall that promote microbial decomposers. Our two-phase experiment revealed that both the carry-over effects of photodegradation and litter quality, modulated by inter-annual variability in rainfall, can explain the marked differences in decomposition rates and the frequent decoupling between rainfall and litter decomposition observed in semiarid ecosystems.

Decadal trends in the pollinator assemblage of Eucryphia cordifolia in Chilean rainforests.

Long-term studies of plant-pollinator interactions are almost nonexistent in the scientific literature. The objective of the present study was to determine changes and trends in the pollinator assemblage of ulmo (Eucryphia cordifolia; Cunoniaceae), a canopy-emergent tree found in Chilean temperate rainforests. We assessed the temporal variability of the pollinator assemblage and identified possible modulators of the observed temporal shifts. We sampled insect visitors to the flowers of 16 individual trees of E. cordifolia during 10 consecutive flowering seasons (2000-2009), recording a total of 137 pollinator species with a mean number of species per year of 44. Only three pollinator species (2.2%) were recorded every year. Two bee species accounted for 50% of all insect visits to flowers. One bee species, Bombus dahlbomii (native), was dominant in one season, whereas Apis mellifera (exotic) dominated during the next season. These interannual shifts in population abundances presented first-order dynamics that were characterized by oscillations with a period of 2 years. Changes in the abundances of the dominant pollinators, as well as differences in temperature and precipitation during insect emergence and flowering, led to a nested temporal structure of pollinator composition. Furthermore, the abundances of less common pollinators were sensitive to the abundance of the dominant bee species and to monthly maximum temperatures and the average precipitation during spring and summer. Based on our results and those from other studies, we predict a decline in the numbers of Bombus dahlbomii and nondominant native pollinators in response to new exotic arrivals.

Ecosystem properties self-organize in response to a directional fog-vegetation interaction.

Feedbacks between vegetation and resource inputs can lead to the local, self-organization of ecosystem properties. In particular, feedbacks in response to directional resources (e.g., coastal fog, slope runoff) can create complex spatial patterns, such as vegetation banding. Although similar feedbacks are thought to be involved in the development of ecosystems, clear empirical examples are rare. We created a simple model of a fog-influenced, temperate rainforest in central Chile, which allows the comparison of natural banding patterns to simulations of various putative mechanisms. We show that only feedbacks between plants and fog were able to replicate the characteristic distributions of vegetation, soil water, and soil nutrients observed in field transects. Other processes, such as rainfall, were unable to match these diagnostic distributions. Furthermore, fog interception by windward trees leads to increased downwind mortality, leading to progressive extinction of the leeward edge. This pattern of ecosystem development and decay through self-organized processes illustrates, on a relatively small spatial and temporal scale, the patterns predicted for ecosystem evolution.

Increased drought impacts on temperate rainforests from southern South America: results of a process-based, dynamic forest model.

Increased droughts due to regional shifts in temperature and rainfall regimes are likely to affect forests in temperate regions in the coming decades. To assess their consequences for forest dynamics, we need predictive tools that couple hydrologic processes, soil moisture dynamics and plant productivity. Here, we developed and tested a dynamic forest model that predicts the hydrologic balance of North Patagonian rainforests on Chiloé Island, in temperate South America (42°S). The model incorporates the dynamic linkages between changing rainfall regimes, soil moisture and individual tree growth. Declining rainfall, as predicted for the study area, should mean up to 50% less summer rain by year 2100. We analysed forest responses to increased drought using the model proposed focusing on changes in evapotranspiration, soil moisture and forest structure (above-ground biomass and basal area). We compared the responses of a young stand (YS, ca. 60 years-old) and an old-growth forest (OG, >500 years-old) in the same area. Based on detailed field measurements of water fluxes, the model provides a reliable account of the hydrologic balance of these evergreen, broad-leaved rainforests. We found higher evapotranspiration in OG than YS under current climate. Increasing drought predicted for this century can reduce evapotranspiration by 15% in the OG compared to current values. Drier climate will alter forest structure, leading to decreases in above ground biomass by 27% of the current value in OG. The model presented here can be used to assess the potential impacts of climate change on forest hydrology and other threats of global change on future forests such as fragmentation, introduction of exotic tree species, and changes in fire regimes. Our study expands the applicability of forest dynamics models in remote and hitherto overlooked regions of the world, such as southern temperate rainforests.

Diverging drought-tolerance strategies explain tree species distribution along a fog-dependent moisture gradient in a temperate rain forest.

The study of functional traits and physiological mechanisms determining species' drought tolerance is important for the prediction of their responses to climatic change. Fog-dependent forest patches in semiarid regions are a good study system with which to gain an understanding of species' responses to increasing aridity and patch fragmentation. Here we measured leaf and hydraulic traits for three dominant species with contrasting distributions within patches in relict, fog-dependent forests in semiarid Chile. In addition, we assessed pressure-volume curve parameters in trees growing at a dry leeward edge and wet patch core. We predicted species would display contrasting suites of traits according to local water availability: from one end favoring water conservation and reducing cavitation risk, and from the opposite end favoring photosynthetic and hydraulic efficiency. Consistent with our hypothesis, we identified a continuum of water use strategies explaining species distribution along a small-scale moisture gradient. Drimys winteri, a tree restricted to the humid core, showed traits allowing efficient water transport and high carbon gain; in contrast, Myrceugenia correifolia, a tree that occurs in the drier patch edges, exhibited traits promoting water conservation and lower gas exchange rates, as well low water potential at turgor loss point. The most widespread species, Aextoxicon punctatum, showed intermediate trait values. Osmotic compensatory mechanism was detected in M. correifolia, but not in A. punctatum. We show that partitioning of the pronounced soil moisture gradients from patch cores to leeward edges among tree species is driven by differential drought tolerance. Such differences indicate that trees have contrasting abilities to cope with future reductions in soil moisture.

Extreme climatic events change the dynamics and invasibility of semi-arid annual plant communities.

Extreme climatic events represent disturbances that change the availability of resources. We studied their effects on annual plant assemblages in a semi-arid ecosystem in north-central Chile. We analysed 130 years of precipitation data using generalised extreme-value distribution to determine extreme events, and multivariate techniques to analyse 20 years of plant cover data of 34 native and 11 exotic species. Extreme drought resets the dynamics of the system and renders it susceptible to invasion. On the other hand, by favouring native annuals, moderately wet events change species composition and allow the community to be resilient to extreme drought. The probability of extreme drought has doubled over the last 50 years. Therefore, investigations on the interaction of climate change and biological invasions are relevant to determine the potential for future effects on the dynamics of semi-arid annual plant communities.

Bromeliad growth and stoichiometry: responses to atmospheric nutrient supply in fog-dependent ecosystems of the hyper-arid Atacama Desert, Chile.

Carbon, nitrogen, and phosphorus (C, N, P) stoichiometry influences the growth of plants and nutrient cycling within ecosystems. Indeed, elemental ratios are used as an index for functional differences between plants and their responses to natural or anthropogenic variations in nutrient supply. We investigated the variation in growth and elemental content of the rootless terrestrial bromeliad Tillandsia landbeckii, which obtains its moisture, and likely its nutrients, from coastal fogs in the Atacama Desert. We assessed (1) how fog nutrient supply influences plant growth and stoichiometry and (2) the response of plant growth and stoichiometry to variations in nutrient supply by using reciprocal transplants. We hypothesized that T. landbeckii should exhibit physiological and biochemical plastic responses commensurate with nutrient supply from atmospheric deposition. In the case of the Atacama Desert, nutrient supply from fog is variable over space and time, which suggests a relatively high variation in the growth and elemental content of atmospheric bromeliads. We found that the nutrient content of T. landbeckii showed high spatio-temporal variability, driven partially by fog nutrient deposition but also by plant growth rates. Reciprocal transplant experiments showed that transplanted individuals converged to similar nutrient content, growth rates, and leaf production of resident plants at each site, reflecting local nutrient availability. Although plant nutrient content did not exactly match the relative supply of N and P, our results suggest that atmospheric nutrient supply is a dominant driver of plant growth and stoichiometry. In fact, our results indicate that N uptake by T. landbeckii plants depends more on N supplied by fog, whereas P uptake is mainly regulated by within-plant nutrient demand for growth. Overall, these findings indicate that variation in fog nutrient supply exerts a strong control over growth and nutrient dynamics of atmospheric plants, which are ubiquitous across fog-dominated ecosystems.

Microsatellite markers for the relict tree Aextoxicon punctatum: the only species in the Chilean endemic family Aextoxicaceae.

PREMISE OF THE STUDY: We screened 10 microsatellite loci for the dioecious, rainforest tree Aextoxicon punctatum, a species belonging to a monotypic family and genus, endemic to southwestern South America (30-43°S). METHODS AND RESULTS: Polymorphisms were evaluated in 108 adult trees from four populations, including the northern and southern extremes of the geographic range of Aextoxicon in Chile. All 10 microsatellites revealed polymorphic variation. A total of 69, 57, 59, and 69 alleles were found in 40 (Fray Jorge), 19 (Santa Ines), 21 (Quebrada del Tigre), and 28 (Guabun) individual trees, respectively. The mean expected heterozygosity per population ranged from 0.70 to 0.72. CONCLUSIONS: These polymorphic microsatellites will be useful in assessing the genetic structure and conservation status of Aextoxicon throughout its historically fragmented geographic range. Parentage analysis will provide additional insights into the key historical and contemporary processes that have mediated population differentiation in this species.

A network analysis of plant-pollinator interactions in temperate rain forests of Chiloé Island, Chile.

This study characterizes the structure of a plant-pollinator network in a temperate rain forest of Chiloé Island, southern Chile, where woody species are strongly dependent on biotic pollinators, and analyzes its robustness to the loss of participating species. Degree distribution, nestedness, and expected species persistence were evaluated. In addition, we assessed the roles of predefined subsets of plants (classified by life forms) and pollinators (grouped by taxonomic orders) in the network's structure and dynamics. For this, we simulated the complete removal of each plant and pollinator subset and analyzed the resultant connectivity patterns, as well as the expected long-term species losses by running a stochastic model. Finally, we evaluated the sensitivity of the network structure to the loss of single species in order to identify potential targets for conservation. Our results show that the plant-pollinator network of this Chilean temperate rain forest exhibits a nested structure of interactions, with a degree distribution best described by a power law model. Model simulations revealed the importance of trees and hymenopterans as pivotal groups that maintain the core structure of the pollination network and guarantee overall species persistence. The hymenopterans Bombus dahlbomii and Diphaglossa gayi, the shrubs Tepualia stipularis and Ugni molinae, the vines Mitraria coccinea and Asteranthera ovata, and the entire set of tree species exerted a disproportionately large influence on the preservation of network structure and should be considered as focal species for conservation programs given current threats from selective logging and habitat loss.

Evolution of autonomous selfing accompanies increased specialization in the pollination system of Schizanthus (Solanaceae).

The co-occurrence of elaborate flowers visited by specific groups of pollinators and capacity for autonomous selfing in the same plant species has puzzled evolutionary biologists since the time of Charles Darwin. To examine whether autonomous selfing and floral specialization evolved in association, we quantified the autofertility level (AFI) in nine Schizanthus species characterized by a wide range of pollination specialization, revealing AFI values of 0.02 to complete selfing. An independent contrasts analysis conducted on AFIs and number of functional pollinator groups showed that autonomous selfing evolved from an ancestral outcrossing system as plants became increasingly specialized (r = -0.82). To assess whether autonomous selfing together with specialization acts as a reproductive assurance mechanism, we estimated spatial and interannual variation in fruit set due to pollinator failure in two closely related high Andean Schizanthus species differing in their specialization levels. Variation in pollinator failure rate was more pronounced and autonomous selfing increased fruit production over biotically assisted pollination in the more specialized species. Our study suggests that specialized pollination deems species more vulnerable to pollinator fluctuation thus promoting the evolution of delayed autonomous selfing.