Ecological stress memory in wood architecture of two Neotropical hickory species from central-eastern Mexico.

BACKGROUND: Drought periods are major evolutionary triggers of wood anatomical adaptive variation in Lower Tropical Montane Cloud Forests tree species. We tested the influence of historical drought events on the effects of ecological stress memory on latewood width and xylem vessel traits in two relict hickory species (Carya palmeri and Carya myristiciformis) from central-eastern Mexico. We hypothesized that latewood width would decrease during historical drought years, establishing correlations between growth and water stress conditions, and that moisture deficit during past tree growth between successive drought events, would impact on wood anatomical features. We analyzed latewood anatomical traits that developed during historical drought and pre- and post-drought years in both species. RESULTS: We found that repeated periods of hydric stress left climatic signatures for annual latewood growth and xylem vessel traits that are essential for hydric adaptation in tropical montane hickory species. CONCLUSIONS: Our results demonstrate the existence of cause‒effect relationships in wood anatomical architecture and highlight the ecological stress memory linked with historical drought events. Thus, combined time-series analysis of latewood width and xylem vessel traits is a powerful tool for understanding the ecological behavior of hickory species.

A late Cretaceous fagalean inflorescence preserved in amber from New Jersey.

PREMISE OF THE STUDY: An inflorescence with three pistillate flowers in amber from the early Upper Cretaceous (Turonian, ~90-94 million years ago) of central New Jersey represents the oldest known flowers with features present in an early stem complex of the Fagales. The inflorescence has characteristics of Nothofagaceae, but also has strikingly distinct characters that suggest it is intermediate between Nothofagus and other Fagales. This intermediacy is consistent with its northern hemisphere distribution. METHODS: We investigated this new fossil by comparing it with extant and fossil members of the Fagales using light microscopy and nano-computed tomography. In addition, for exploring its relationships, we mapped the morphological characters onto a widely accepted molecular-based tree of modern basal Fagales using standard methods of character optimization. KEY RESULTS: The phylogenetic position of the fossil inflorescence can be unequivocally determined by the presence of unique features, singly and in combination, that are found only in "basal" members of Fagales. The fossil adds critical information on the features of the early stem Fagales, evolution of the cupule in Nothofagaceae and Fagaceae, and a reasonable biogeographic hypothesis for the differentiation of southern (e.g., Nothofagaceae) and northern hemisphere Fagales. CONCLUSIONS: This new fossil provides insight into the early evolution of Fagales and suggests that early stem Fagales that had not yet differentiated into modern families were present in the Late Cretaceous of North America. Based on available evidence, the fossil is best interpreted as an early stem member of the Fagales, with features that suggest a transition from a more generalized Nothofagus-like fagalean ancestor with some unique presumably plesiomorphic features. The presence of an enlarged perianth and flexuous styles also suggests the possibility of insect pollination, which has been lost in all Fagales with the exception of some members of subfamily Castaneoideae in Fagaceae sensu stricto (which otherwise are very different from this fossil). The poorly developed, bract-like cupule valves of the fossil can be interpreted as primitive (i.e., incipient) or as reduced from more developed cupules that are found in most modern Fagaceae and Nothofagaceae.

Xylem adjusts to maintain efficiency across a steep precipitation gradient in two coexisting generalist species.

Background and Aims: Trees adjust the configuration of their conductive system in response to changes in water availability, maximizing efficiency in wet environments and increasing safety in dry habitats. However, evidence of this general trend is not conclusive. Generalist species growing across broad climatic gradients provide an ideal framework to assess intra-specific xylem adjustments under contrasting environmental conditions. Our aims were to compare the response of xylem traits to variations in precipitation of two co-occurring generalist tree species, and to assess climate control on xylem trait variability and co-ordination. Methods: We evaluated xylem traits of Embothrium coccineum (Proteaceae, evergreen) and Nothofagus antarctica (Nothofagaceae, deciduous) in three areas across an abrupt precipitation gradient, from 500 to 2500 mm, in southern Chile. We measured wood density, vessel lumen area and density, percentage of conductive area and vessel grouping, and estimated the hydraulic function from anatomical measurements in 60 individuals per species. Key Results: Both species shared a common pattern of response along the precipitation gradient, with an increase in vessel density with dryness, but without changes in estimated hydraulic conductivity. Xylem traits in E. coccineum were more variable and more responsive to the climate gradient, decreasing vessel lumen area and increasing wood density, whereas vessel grouping showed contrasting patterns between species. Additionally, the analysis of trait co-ordination at the individual level revealed a tighter co-ordination among xylem traits in E. coccineum. Conclusions: Estimated xylem efficiency was maintained in combination with different levels of expected xylem safety within species. Reduction in vessel lumen area was compensated through large increases in vessel density, thus breaking the trade-off between xylem efficiency and safety. Otherwise, the existence of alternative internal adjustments in coexisting species to face similar climatic constraints might increase resilience of temperate forests against unpredictable changes in climatic conditions.

Insights into intraspecific wood density variation and its relationship to growth, height and elevation in a treeline species.

The wood economics spectrum provides a general framework for interspecific trait-trait coordination across wide environmental gradients. Whether global patterns are mirrored within species constitutes a poorly explored subject. In this study, I first determined whether wood density co-varies together with elevation, tree growth and height at the within-species level. Second, I determined the variation of wood density in different stem parts (trunk, branch and twigs). In situ trunk sapwood, trunk heartwood, branch and twig densities, in addition to stem growth rates and tree height were determined in adult trees of Nothofagus pumilio at four elevations in five locations spanning 18° of latitude. Mixed effects models were fitted to test relationships among variables. The variation in wood density reported in this study was narrow (ca. 0.4-0.6 g cm-3 ) relative to global density variation (ca. 0.3-1.0 g cm-3 ). There was no significant relationship between stem growth rates and wood density. Furthermore, the elevation gradient did not alter the wood density of any stem part. Trunk sapwood density was negatively related to tree height. Twig density was higher than branch and trunk densities. Trunk heartwood density was always significantly higher than sapwood density. Negative across-species trends found in the growth-wood density relationship may not emerge as the aggregate of parallel intraspecific patterns. Actually, trees with contrasting growth rates show similar wood density values. Tree height, which is tightly related to elevation, showed a negative relationship with sapwood density.

Does water availability regulate biomass partitioning between trunk and branches?

The extent to which a vertical trunk is differentiated from its branches is a key trait for the architecture of trees and may affect interspecific relationships. In this study, we analysed the effect of soil water availability on biomass partitioning for Nothofagus pumilio by means of a nursery experiment. Juvenile trees were subject to three irrigation conditions: no irrigation, intermediate irrigation and high irrigation. Irrigation conditions emulated the mean precipitation of the most representative environments inhabited by N. pumilio. Changes in soil water availability modified the biomass partitioning patterns of trees. In comparison to the other two conditions, high irrigation caused: (i) a higher ratio of biomass partitioning to stems than roots; (ii) more trunk growth in relation to its branches; and (iii) more photosynthetic organs relative to the aboveground biomass. Trunk size relative to that of its most recent branches was not increased by water availability. Water availability may play a significant role in the capacity of N. pumilio for space occupation due to the effects on axis differentiation.