Fruit wings accelerate germination in Anacyclus clavatus.

PREMISE: The lateral membranous expansions of fruits, commonly referred to as wings, have long been theorized to serve only dispersal functions. Alternatively, because winged fruits typically have earlier seed germination than unwinged fruits, we hypothesized that wings could increase the contact surface with water, ultimately triggering earlier germination. METHODS: We investigated this alternative hypothesis by exploring the potential role of fruit wings on germination in the heterocarpic species Anacyclus clavatus (Desf.) Pers. (Asteraceae), which produces both winged and unwinged fruits. First, we measured the speed and degree of water absorption in winged and unwinged fruits. Second, we investigated the effects of wings on germination performance, by either reducing wing size or by preventing water absorption by sealing wings with wax. Next, we tested the influence of water availability on the germination performance of winged and unwinged fruits by reducing the water potential. RESULTS: Winged fruits absorbed more water at a faster rate than unwinged fruits. The sealing of wings delayed germination, whereas germination time was not significantly altered by wing cutting. The restriction of water availability by decreasing water potential significantly delayed seed germination of unwinged fruits, whereas winged fruits remained unaffected. CONCLUSIONS: Altogether, our results support the effect of wings on germination and cast doubt on the unique role of wings in dispersal. Whether or not wings contribute to dispersal, we propose that they also improve seed germination and seedling establishment by facilitating water absorption after the release from their mother plants.

Fire-scarred fossil tree from the Late Triassic shows a pre-fire drought signal.

Exploring features of wood anatomy associated with fire scars found on fossil tree trunks is likely to increase our knowledge of the environmental and ecological processes that occurred in ancient forests and of the role of fire as an evolutionary force. In Petrified Forest National Park, Arizona, where Late Triassic fossil trees are exposed, we found 13 examples of fossil logs with external features resembling modern fire scars. One specimen with the unambiguous external features of a fire scar was collected for analysis of its fossilized wood. A light-colored band composed of compressed and distorted tracheids was associated with the scarring event. Cell lumen diameter and cell wall thickness in the pre-scarring fossilized wood show a response similar to that described in modern trees experiencing drought conditions. Tracheids in the post-scarring wood are initially smaller, and then become larger than average following a recovery period, as is often observed in modern conifers following fire. The responses in external morphology and wood anatomy to drought and fire were similar to those of some modern trees and support the view that some forests may have experienced conditions favoring the evolution of fire-adapted traits for more than 200 million years.

Within-individual phenotypic plasticity in flowers fosters pollination niche shift.

Phenotypic plasticity, the ability of a genotype of producing different phenotypes when exposed to different environments, may impact ecological interactions. We study here how within-individual plasticity in Moricandia arvensis flowers modifies its pollination niche. During spring, this plant produces large, cross-shaped, UV-reflecting lilac flowers attracting mostly long-tongued large bees. However, unlike most co-occurring species, M. arvensis keeps flowering during the hot, dry summer due to its plasticity in key vegetative traits. Changes in temperature and photoperiod in summer trigger changes in gene expression and the production of small, rounded, UV-absorbing white flowers that attract a different assemblage of generalist pollinators. This shift in pollination niche potentially allows successful reproduction in harsh conditions, facilitating M. arvensis to face anthropogenic perturbations and climate change.

Low growth resilience to drought is related to future mortality risk in trees.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

A synthesis of radial growth patterns preceding tree mortality.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Secondary Growth and Carbohydrate Storage Patterns Differ between Sexes in Juniperus thurifera.

Differences in reproductive costs between male and female plants have been shown to foster sex-related variability in growth and C-storage patterns. The extent to which differential secondary growth in dioecious trees is associated with changes in stem carbohydrate storage patterns, however, has not been fully assessed. We explored the long-term radial growth and the seasonal variation of non-structural carbohydrate (NSC) content in sapwood of 40 males and 40 females Juniperus thurifera trees at two sites. NSC content was analyzed bimonthly for 1 year, and tree-ring width was measured for the 1931-2010 period. Sex-related differences in secondary growth and carbohydrate storage were site-dependent. Under less restrictive environmental conditions females grew more and stored more non-soluble sugars than males. Our results reinforce that sex-related differences in growth and resource storage may be a consequence of local adaptation to environmental conditions. Seasonal variation in soluble sugars concentration was opposite to cambial activity, with minima seen during periods of maximal secondary growth, and did not differ between the sexes or sites. Trees with higher stem NSC levels at critical periods showed higher radial growth, suggesting a common mechanism irrespective of site or sex. Sex-related patterns of secondary growth were linked to differences in non-soluble sugars content indicating sex-specific strategies of long-term performance.

Pre-dispersal predation effect on seed packaging strategies and seed viability.

An increased understanding of intraspecific seed packaging (i.e. seed size/number strategy) variation across different environments may improve current knowledge of the ecological forces that drive seed evolution in plants. In particular, pre-dispersal seed predation may influence seed packaging strategies, triggering a reduction of the resources allocated to undamaged seeds within the preyed fruits. Assessing plant reactions to pre-dispersal seed predation is crucial to a better understanding of predation effects, but the response of plants to arthropod attacks remains unexplored. We have assessed the effect of cone predation on the size and viability of undamaged seeds in populations of Juniperus thurifera with contrasting seed packaging strategies, namely, North African populations with single-large-seeded cones and South European populations with multi-small-seeded cones. Our results show that the incidence of predation was lower on the single-large-seeded African cones than on the multi-small-seeded European ones. Seeds from non-preyed cones were also larger and had a higher germination success than uneaten seeds from preyed cones, but only in populations with multi-seeded cones and in cones attacked by Trisetacus sp., suggesting a differential plastic response to predation. It is possible that pre-dispersal seed predation has been a strong selective pressure in European populations with high cone predation rates, being a process which maintains multi-small-seeded cones and empty seeds as a strategy to save some seeds from predation. Conversely, pre-dispersal predation might not have a strong effect in the African populations with single-large-seeded cones characterized by seed germination and filling rates higher than those in the European populations. Our results indicate that differences in pre-dispersal seed predators and predation levels may affect both selection on and intraspecific variation in seed packaging.

Different growth sensitivity to climate of the conifer Juniperus thurifera on both sides of the Mediterranean Sea.

Mediterranean plants cope with cold wet winters and dry hot summers, with a drought gradient from northwest to southeast. Limiting climatic conditions have become more pronounced in the last decades due to the warming trend and rainfall decrease. Juniperus thurifera L., a long-lived conifer tree endemic to the western Mediterranean region, has a disjunct distribution in Europe and Africa, making it a suitable species to study sensitivity to climate in both sides of the Mediterranean Basin. Tree-ring width chronologies were built for three J. thurifera stands at Spain (Europe) and three in Morocco (Africa) and correlated with monthly temperature and precipitation. The temporal stability of climate-growth relationships was assessed using moving correlations; the drought effect on growth was calculated using the monthly standardized precipitation-evapotranspiration index (SPEI) at different temporal scales. In the wettest stands, increasing spring temperature and summer precipitation enhanced growth, while in the driest stands, growth was enhanced by higher spring precipitation and lower summer temperature. The climate-growth correlations shifted during the twentieth century, especially since the 1970s. Particularly noticeable is the recent negative correlation with previous autumn and winter precipitation in the wettest stands of J. thurifera, probably related with an effect of cloud cover or flooding on carbon storage depletion for next year growth. The driest stands were affected by drought at long time scales, while the wettest stands respond to drought at short time scales. This reveals a different strategy to cope with drought conditions, with populations from drier sites able to cope with short periods of water deficit.

Sex-specific, age-dependent sensitivity of tree-ring growth to climate in the dioecious tree Juniperus thurifera.

Tree features may modulate the sensitivity of radial growth to climate, leading to a nonuniform response. Age-related increases in climatic sensitivity have been observed repeatedly. Sex-related climatic sensitivity is also possible because of the long-term differential reproductive cost between the sexes. This study analysed the simultaneous effects of age and sex on the sensitivity of tree-ring growth to climate. Ring widths were measured from 50 female and 50 male Juniperus thurifera trees, 50-350 yr old, growing under a Mediterranean continental climate. Response functions were calculated based on tree-ring chronologies and monthly climatic records. Climatic sensitivity decreased with increasing age. Young trees (50-100 yr) were the most climatically sensitive to June-July precipitation, which affected growth positively. We found a significant interaction between age and sex in the climatic response of J. thurifera, with young females the most sensitive to summer water stress. Our results suggest that age-dependent climatic sensitivity can be determined by site-specific limiting environmental conditions and species-specific architectural and physiological adjustments during ontogeny. This study supports that the different ontogenetic stages of J. thurifera differ in their root structural traits and that sex-related sensitivity to summer drought may be attributable to less efficient water use by females.