Morphology, germination, and geographic distribution of Pentaclethra macroloba (Fabaceae): a hyperdominant Amazonian tree / Morfología, germinación y distribución geográfica de Pentaclethra macroloba (Fabaceae): árbol amazónico hiperdominante

Introduction: Pentaclethra macroloba is a hyperdominant tree in the Amazon estuary of great socioeconomic importance for the region because the oil from its seeds is a powerful herbal medicine. Objective: We aimed to characterize the morphological structure, the morphological adaptations in response to the daily flooding of the Amazon estuary and the biogeographic area of P. macroloba. Methods: Detailed description of the external morphology, from germination to the adult tree, was obtained from individuals located in floodplain forest, Northeast Amazonia. The occupation area and the geographical extension of P. macroloba were determined from point of geographical coordinates of botanical samples deposited in the digital collections of Mobot and SpeciesLink. Results: Adult individuals have adapted structures in response to daily flooding, such as: adventitious roots to increase respiratory efficiency and lenticels in the trunk, serving as a connection point for oxygen transport between the root and the aerial part. Dried fruit favours the activation of the explosive dehiscence mechanism, allowing the seed to be expelled long distance. Deltoid shape of the seed allows water fluctuation and more efficient dispersal. Seedling is phanerocotylar hypogeal and with one pair of reserve cotyledons that provide the seedling an extra source of energy to escape the flood. Germination rate was 78 % and the speed was 0.2 seeds.day-1. Circular buffer method revealed the presence of 123 subpopulations of P. macroloba distributed in a radius of 5 699 943 km² across the Neotropical region. Conclusions: Much of the morphological structures of P. macroloba are adaptive and evolutionary responses to the periodically flooded environment of the Amazon estuary, showing that these environments select the trees, best adapted, to inhabit the flood. P. macroloba has a wide geographical area denotes the plasticity of adapting to different environments, which may justify its monodominance in some regions.

Introducción: Pentaclethra macroloba es un árbol hiperdominante en el estuario del Amazonas, de gran importancia socioeconómica para la región, pues el aceite de sus semillas es un poderoso medicamento natural. Objetivo: Nuestro objetivo fue caracterizar la estructura morfológica, las adaptaciones morfológicas en respuesta a las inundaciones diarias del estuario del Amazonas y el área biogeográfica de P. macroloba. Métodos: Una descripción detallada de la morfología externa, desde la germinación hasta el árbol adulto, se obtuvo de individuos ubicados en el bosque periódicamente inundado, al noreste de la Amazonia. El área de ocupación y la extensión geográfica de P. macroloba se determinaron a partir del punto de coordenadas geográficas de muestras botánicas depositadas en las colecciones digitales de Mobot y SpeciesLink. Resultados: Los individuos adultos tienen estructuras adaptadas en respuesta a las inundaciones diarias, tales como: raíces adventicias para aumentar la eficiencia respiratoria y lenticelas en el tronco, que sirven como punto de conexión para el transporte de oxígeno entre la raíz y la parte aérea. Los frutos secos favorecen la activación del mecanismo de dehiscencia explosiva, permitiendo que la semilla sea expulsada a larga distancia. La forma deltoidea de la semilla permite la fluctuación en el agua y una dispersión más eficiente. La plántula es hipogel fanerocotiledones y con un par de cotiledones de reserva que proporcionan a la plántula una fuente extra de energía para escapar de la inundación. La tasa de germinación fue del 78 % y la velocidad fue de 0.2 semillas día-1. El método Circular buffer reveló la presencia de 123 subpoblaciones de P. macroloba distribuidas en un radio de 5 699 943 km² a lo largo de la región neotropical. Conclusiones: Gran parte de las estructuras morfológicas de P. macroloba son respuestas adaptativas y evolutivas al ambiente periódicamente inundado del estuario del Amazonas, lo que demuestra que estos ambientes actúan como filtro ambiental seleccionado las especies mejor adaptadas al medio. Su amplia área geográfica denota la plasticidad de adaptarse a diferentes ambientes, lo que puede justificar su monodominancia en algunas regiones.

Regulatory effects of root pruning on leaf nutrients, photosynthesis, and growth of trees in a closed-canopy poplar plantation.

A plantation of 5-year-old poplar Populus × euramericana cv. 'Neva' was used to study the regulatory effects of root pruning on nutrients, photosynthetic characteristics, and water-use efficiency (WUE) of leaves and growth rates of diameter at breast height (DBH; 1.3 m), tree height, and volume. Six root-pruning treatments were conducted with different combinations of intensity (at a distance of six, eight or ten times DBH from the trunk) and orientation (on two or four sides of the trees). Results showed that the N, P, K, photosynthetic rate, transpiration rate, and stomatal conductance of leaves were all significantly decreased by root pruning over the initial period following root pruning (30 days), but increased in the subsequent investigations. The values of the above indexes peaked in 8-2 treatment (i.e., eight times DBH distance on two sides). The leaf WUE in 8-2 treatment, and average growth rates of DBH, tree height and volume, were the highest among all treatments within 3 years of root pruning. The results indicated that the root pruning based on the appropriate selection of intensity and orientation had significant positive effects on leaf nutrients, photosynthesis, and growth of trees in a closed-canopy poplar plantation.

The water relations and xylem attributes of albino redwood shoots (Sequioa sempervirens (D. Don.) Endl.).

Plants that lack chlorophyll are rare and typically restricted to holoparasites that obtain their carbon, water and mineral resources from a host plant. Although not parasites in the traditional sense, albino foliage, such as the sprouts that sometimes develop from redwood tree trunks, are comparable in function. They occur sporadically, and can reach the size of shrubs and in rare cases, trees. Albino redwoods are interesting because in addition to their reduced carbon resources, the absence of chloroplasts may impede proper stomatal function, and both aspects may have upstream consequences on water transport and xylem quality. We examined the water relations, water transport and xylem anatomical attributes of albino redwoods and show that similar to achlorophyllous and parasitic plants, albino redwoods have notably higher stomatal conductance than green sprouts. Given that stem xylem tracheid size as well as water transport efficiency are nearly equivalent in both albino and green individuals, we attribute the increased leaf water loss in albino sprouts to lower leaf to xylem area ratios, which favour improved hydration relative to green sprouts. The stems of albino redwoods were more vulnerable to drought-induced embolism than green stems, and this was consistent with the albino's weaker tracheids, as characterized by wall thickness to lumen diameter measures. Our results are both complementary and consistent with previous research on achlorophyllous plants, and suggest that the loss of stomatal control and photosynthetic capacity results in substantial vascular and anatomical adjustments.

Phenological variation of leaf functional traits within species.

A basic assumption of the trait-based approach in plant ecology is that differences in functional trait values are greater between species than within species. We questioned this assumption by assessing (1) the relative extent of inter- and intraspecific leaf trait variation throughout a complete growing season (phenological variation) in a group of deciduous and evergreen woody species, and (2) whether species rankings based on leaf traits were maintained across the growing season. We analysed leaf mass per area (LMA) and leaf nutrient concentrations (C, N, P), including the C:N and N:P ratios. Intraspecific trait variation (ITV) due to phenology was significantly greater than interspecific variation for leaf N concentration on a mass basis (Nm; 68.90 %) and for the leaf C:N ratio (60.60 %), whereas interspecific variation was significantly higher than ITV for LMA (62.30 %) and for leaf C concentration on a mass (Cm) and area (Ca) basis (Cm 70.40 %; Ca 65.30 %). ITV was particularly low for LMA (<20 %). Species rankings were highly modified by phenology for a number of leaf traits (Pm, N:P ratio) but were relatively well conserved throughout the growing season for others (LMA, Nm). Patterns of ITV across the growing season differed significantly between deciduous and evergreen species for all traits except leaf P but did not vary between native and exotic species. Overall, our results show that intraspecific phenological variation in leaf traits may be similar to or greater than interspecific variation and that temporal patterns of ITV vary considerably among traits and species, especially for leaf nutrient concentrations, factors which can potentially affect quantitative interspecific relationships.

Tree-Ring Dating of the Reshui-1 Tomb in Dulan County, Qinghai Province, North-West China.

Tuyuhun and Tubo were two important states that thrived in north-western China during AD 311-900 in parallel with the Han Chinese dynasties of Sui and Tang periods. The Reshui Tomb Cluster located in Dulan County of the north-eastern Tibetan Plateau is an important cultural relic of the Tuyuhun-Tubo age. The official excavations of the Reshui tombs were regarded as top events in archaeology in the 1980s and 1990s in China. The Reshui-1 Tomb is the largest one among the tombs in the area. Since its excavation, there have been debates on whether the owner of the tomb belonged to the Tuyuhun or Tubo ethnicity. Therefore, accurately dating the Reshui-1 Tomb has a critical place in studying the Tubo and Tuyuhun histories. We collected 7 discs and 11 increment cores of Qilian juniper (Juniperus przewalskii Kom.) from the exposed and fallen beams of the roof of the Reshui-1Tomb. The lengths of the 16 tree-ring records are between 69 and 152 years. Based on a previously developed master dating chronology using Qilian juniper samples from the eastern Qaidam Basin, the calendar dates of the 16 specimens were determined by the COFECHA program and visual dating procedure. The average inter-series correlation among the dated sample series is 0.696, indicating good quality of cross-dating. The year of the outermost rings is AD 715 for the 7 discs and 4 out of the 9 increment cores. Moreover, the ring-width variations of the samples are consistent with the existing chronologies from the region. The presence of late-wood of AD 715 in the samples indicated that the Reshui-1 Tomb was completed in late AD 715 or early 716, which means that the Reshui-1 Tomb was finished in the Tubo age. This date provides direct evidence for archaeologists to determine the owner's ethnicity and identify of the Reshui-1 Tomb.

Timing and climate forcing of volcanic eruptions for the past 2,500 years.

Volcanic eruptions contribute to climate variability, but quantifying these contributions has been limited by inconsistencies in the timing of atmospheric volcanic aerosol loading determined from ice cores and subsequent cooling from climate proxies such as tree rings. Here we resolve these inconsistencies and show that large eruptions in the tropics and high latitudes were primary drivers of interannual-to-decadal temperature variability in the Northern Hemisphere during the past 2,500 years. Our results are based on new records of atmospheric aerosol loading developed from high-resolution, multi-parameter measurements from an array of Greenland and Antarctic ice cores as well as distinctive age markers to constrain chronologies. Overall, cooling was proportional to the magnitude of volcanic forcing and persisted for up to ten years after some of the largest eruptive episodes. Our revised timescale more firmly implicates volcanic eruptions as catalysts in the major sixth-century pandemics, famines, and socioeconomic disruptions in Eurasia and Mesoamerica while allowing multi-millennium quantification of climate response to volcanic forcing.

The length of the dry season may be associated with leaf scleromorphism in cerrado plants.

Despite limitations of low fertility and high acidity of the soils, the cerrado flora is the richest amongst savannas. Many cerrado woody species show sclerophyllous leaves, which might be related to the availability of water and nutrients in the soil. To better understand the function and structure of cerrado vegetation within its own variations, we compared two cerrado communities: one in its core region in central Brazil (Brasília, DF) and the other on its southern periphery (Itirapina, SP). We contrasted the length of the dry season, soil fertility rates, leaf concentrations of N, P, K, Ca and Mg and the specific leaf area (SLA) between these communities. The dry season was shorter on the periphery, where the soil was more fertile although more acidic. Plants from the periphery showed higher SLA and higher leaf concentrations of N, P, Ca and Mg. We propose that the higher SLA of plants from the periphery is related to the shorter dry season, which allows better conditions for nutrient uptake.

A 3,500-year tree-ring record of annual precipitation on the northeastern Tibetan Plateau.

An annually resolved and absolutely dated ring-width chronology spanning 4,500 y has been constructed using subfossil, archaeological, and living-tree juniper samples from the northeastern Tibetan Plateau. The chronology represents changing mean annual precipitation and is most reliable after 1500 B.C. Reconstructed precipitation for this period displays a trend toward more moist conditions: the last 10-, 25-, and 50-y periods all appear to be the wettest in at least three and a half millennia. Notable historical dry periods occurred in the 4th century BCE and in the second half of the 15th century CE. The driest individual year reconstructed (since 1500 B.C.) is 1048 B.C., whereas the wettest is 2010. Precipitation variability in this region appears not to be associated with inferred changes in Asian monsoon intensity during recent millennia. The chronology displays a statistical association with the multidecadal and longer-term variability of reconstructed mean Northern Hemisphere temperatures over the last two millennia. This suggests that any further large-scale warming might be associated with even greater moisture supply in this region.

Removal of nutrient limitations in forest gaps enhances growth rate and resistance to cavitation in subtropical canopy tree species differing in shade tolerance.

A 4-year fertilization experiment with nitrogen (N) and phosphorus (P) was carried out in natural gaps of a subtropical forest in northeastern Argentina. Saplings of six dominant canopy species differing in shade tolerance were grown in five control and five N + P fertilized gaps. Hydraulic architectural traits such as wood density, the leaf area to sapwood area ratio (LA : SA), vulnerability to cavitation (P50) and specific and leaf-specific hydraulic conductivity were measured, as well as the relative growth rate, specific leaf area (SLA) and percentage of leaf damage by insect herbivores. Plant growth rates and resistance to drought-induced embolisms increased when nutrient limitations were removed. On average, the P50 of control plants was -1.1 MPa, while the P50 of fertilized plants was -1.6 MPa. Wood density and LA : SA decreased with N + P additions. A trade-off between vulnerability to cavitation and efficiency of water transport was not observed. The relative growth rate was positively related to the total leaf surface area per plant and negatively related to LA : SA, while P50 was positively related to SLA across species and treatments. Plants with higher growth rates and higher total leaf area in fertilized plots were able to avoid hydraulic dysfunction by becoming less vulnerable to cavitation (more negative P50). Two high-light-requiring species exhibited relatively low growth rates due to heavy herbivore damage. Contrary to expectations, shade-tolerant plants with relatively high resistance to hydraulic dysfunction and reduced herbivory damage were able to grow faster. These results suggest that during the initial phase of sapling establishment in gaps, species that were less vulnerable to cavitation and exhibited reduced herbivory damage had faster realized growth rates than less shade-tolerant species with higher potential growth rates. Finally, functional relationships between hydraulic traits and growth rate across species and treatments were maintained regardless of soil nutrient status.

Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree.

BACKGROUND AND AIMS: Sex allocation has been studied mainly in small herbaceous plants but much less in monoecious wind-pollinated trees. The aim of this study was to explore changes in gender segregation and sex allocation by Pinus halepensis, a Mediterranean lowland pine tree, within tree crowns and between trees differing in their size or crown shape. METHODS: The production of new male and female cones and sex allocation of biomass, nitrogen and phosphorus were studied. The relationship between branch location, its reproductive status and proxies of branch vigour was also studied. KEY RESULTS: Small trees produced only female cones, but, as trees grew, they produced both male and female cones. Female cones were produced mainly in the upper part of the crown, and male cones in its middle and lower parts. Lateral branch density was correlated with the number of male but not female cones; lateral branches were more dense in large than in small trees and even denser in hemispherical trees. Apical branches grew faster, were thicker and their phosphorus concentration was higher than in lateral shoots. Nitrogen concentration was higher in cone-bearing apical branches than in apical vegetative branches and in lateral branches with or without cones. Allocation to male relative to female function increased with tree size as predicted by sex allocation theory. CONCLUSIONS: The adaptive values of sex allocation and gender segregation patterns in P. halepensis, in relation to its unique life history, are demonstrated and discussed. Small trees produce only female cones that have a higher probability of being pollinated than the probability of male cones pollinating; the female-first strategy enhances population spread. Hemispherical old trees are loaded with serotinous cones that supply enough seeds for post-fire germination; thus, allocation to males is more beneficial than to females.

Architectural plasticity in young Eucalyptus marginata on restored bauxite mines and adjacent natural forest in south-western Australia.

The aboveground architecture of Eucalyptus marginata (Jarrah) was investigated in chronosequences of young trees (2.5, 5 and 10 m height) growing in a seasonally dry climate in a natural forest environment with intact soils, and on adjacent restored bauxite mine sites on soils with highly modified A and B horizons above an intact C horizon. Compared to forest trees, trees on restored sites were much younger and faster growing, with straighter, more clearly defined main stems and deeper, narrower crowns containing a greater number of branches that were longer, thinner and more vertically angled. Trees on restored sites also had a higher fraction of biomass in leaves than forest trees, as indicated by 20-25% thicker leaves, 30-70% greater leaf area, 10-30% greater leaf area to sapwood area ratios and 5-30% lesser branch Huber values. Differences in crown architecture and biomass distribution were consistent with putatively greater soil-water, nutrient and light availability on restored sites. Our results demonstrate that under the same climatic conditions, E. marginata displays a high degree of plasticity of aboveground architecture in response to the net effects of resource availability and soil environment. These differences in architecture are likely to have functional consequences in relation to tree hydraulics and growth that, on larger scales, is likely to affect the water and carbon balances of restored forest ecosystems. This study highlights substrate as a significant determinant of tree architecture in water-limited environments. It further suggests that the architecture of young trees on restored sites may need to change again if they are to survive likely longer-term changes in resource availability.

Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree.

The Caricaceae is a small family of tropical trees and herbs in which most species are dioecious. In the present study, we extend our previous work on dioecy in the Caricaceae, characterising the morphological variation in sexual expression in flowers of the dioecious tree Jacaratia mexicana. We found that, in J. mexicana, female plants produce only pistillate flowers, while male plants are sexually variable and can bear three different types of flowers: staminate, pistillate and perfect. To characterise the distinct types of flowers, we measured 26 morphological variables. Our results indicate that: (i) pistillate flowers from male trees carry healthy-looking ovules and are morphologically similar, although smaller than, pistillate flowers on female plants; (ii) staminate flowers have a rudimentary, non-functional pistil and are the only flowers capable of producing nectar; and (iii) perfect flowers produce healthy-looking ovules and pollen, but have smaller ovaries than pistillate flowers and fewer anthers than staminate flowers, and do not produce nectar. The restriction of sexual variation to male trees is consistent with the evolutionary path of dioecy from hermaphrodite ancestors through the initial invasion of male-sterile plants and a subsequent gradual reduction in female fertility in cosexual individuals (gynodioecy pathway), but further work is needed to confirm this hypothesis.

Adaptation of fine roots to annual fertilization and irrigation in a 13-year-old Pinus pinaster stand.

Effects of fertilization and irrigation on fine roots and fungal hyphae were studied in 13-year-old maritime pine (Pinus pinaster Aït. in Soland), 7 years after the initiation of the treatments. The fertilization trials consisted of a phosphorus treatment, a complete fertilizer treatment (N, P, K, Ca and Mg), and an unfertilized treatment (control). Fertilizers were applied annually and were adjusted according to foliar target values. Two irrigation regimes (no irrigation and irrigation of a set amount each day) were applied from May to October. Root samples to depths of 120 cm were collected in summer of 2005, and the biomass of small roots (diameter 2-20 mm) and fine roots (diameter

Chlorpyrifos immersion to eliminate third instars of Japanese beetle (Coleoptera: Scarabaeidae) in balled and burlapped trees and subsequent treatment effects on red maple.

This study examined chlorpyrifos immersion of balled and burlapped (B&B) nursery trees for elimination of third instars of Japanese beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae), and for phytotoxicity on red maple, Acer rubrum L. Trees were harvested as 45- and 60-cm-diameter B&B and immersed in chlorpyrifos at U.S. Domestic Japanese Beetle Harmonization Plan rate (0.24 kg active ingredient [AI/100 liters) or lower rates of 0.015, 0.03, 0.06, and 0.12 kg (AI)/100 liters. The 0.03, 0.06, and 0.24 kg (AI) rates provided 100% control of Japanese beetle grubs in both 45- and 60-cm B&B. The 0.015 and 0.12 kg (AI) chlorpyrifos rates were 100% effective in three tests. However, in another test, 0.015 and 0.12 kg (AI) chlorpyrifos treatments had four (93% control) and one (98% control) grubs recovered, respectively. Root ball soils consisted of loam, silt loam, or clay loam texture classifications. Trunk diameter and internode growth of red maple harvested as 45-cm B&B decreased linearly with increasing chlorpyrifos dip rate during the first year, but effects were unapparent in the second year. Chlorpyrifos rates had no measurable impact on growth of red maples harvested as 60-cm B&B. No visual phytotoxicity symptoms were detected for chlorpyrifos rate or root ball size treatments. In conclusion, results support lowering the U.S. Domestic Japanese Beetle Harmonization Plan chlorpyrifos dip rate for category 2 states to at least 0.03 kg (AI) for B&B diameters < or =60 cm. Chlorpyrifos rates < 0.24 kg (AI) will lower cost, reduce worker exposure, and lessen potential environmental contamination.

Nutrient availability constrains the hydraulic architecture and water relations of savannah trees.

Leaf and whole plant-level functional traits were studied in five dominant woody savannah species from Central Brazil (Cerrado) to determine whether reduction of nutrient limitations in oligotrophic Cerrado soils affects carbon allocation, water relations and hydraulic architecture. Four treatments were used: control, N additions, P additions and N plus P additions. Fertilizers were applied twice yearly, from October 1998 to March 2004. Sixty-three months after the first nutrient addition, the total leaf area increment was significantly greater across all species in the N- and the N + P-fertilized plots than in the control and in the P-fertilized plots. Nitrogen fertilization significantly altered several components of hydraulic architecture: specific conductivity of terminal stems increased with N additions, whereas leaf-specific conductivity and wood density decreased in most cases. Average daily sap flow per individual was consistently higher with N and N + P additions compared to the control, but its relative increase was not as great as that of leaf area. Long-term additions of N and N + P caused midday PsiL to decline significantly by a mean of 0.6 MPa across all species because N-induced relative reductions in soil-to-leaf hydraulic conductance were greater than those of stomatal conductance and transpiration on a leaf area basis. Phosphorus-fertilized trees did not exhibit significant changes in midday PsiL. Analysis of xylem vulnerability curves indicated that N-fertilized trees were significantly less vulnerable to embolism than trees in control and P-fertilized plots. Thus, N-induced decreases in midday PsiL appeared to be almost entirely compensated by increases in resistance to embolism. Leaf tissue water relations characteristics also changed as a result of N-induced declines in minimum PsiL: osmotic potential at full turgor decreased and symplastic solute content on a dry matter basis increased linearly with declining midday PsiL across species and treatments. Despite being adapted to chronic nutrient limitations, Cerrado woody species apparently have the capacity to exploit increases in nutrient availability by allocating resources to maximize carbon gain and enhance growth. The cost of increased allocation to leaf area relative to water transport capacity involved increased total water loss per plant and a decrease in minimum leaf water potentials. However, the risk of increased embolism and turgor loss was relatively low as xylem vulnerability to embolism and leaf osmotic characteristics changed in parallel with changes in plant water status induced by N fertilization.

[Regional and extra-local pollen in tundra pollen samples].

Patterns of pollen spectra formation in the tundra zone of Eurasia were considered. Changes in total pollen concentration were traced in subfossil pollen samples of the tundra zone. The data on subfossil pollen spectra were used to evaluate the proportion between local and regional plus extra-local components of tundra pollen samples as well as the changes in concentration of pollen of Scots and Siberian stone pines as well as of tree and shrub birches. The diameter of dwarf birch pollen was determined in different tundra subzones of Western Siberia. The role of extra-local and regional pollen was considered for all vegetation subzones of tundra.

Fine roots, arbuscular mycorrhizal hyphae and soil nutrients in four neotropical rain forests: patterns across large geographic distances.

* It is commonly hypothesized that stand-level fine root biomass increases as soil fertility decreases both within and among tropical forests, but few data exist to test this prediction across broad geographic scales. This study investigated the relationships among fine roots, arbuscular mycorrhizal (AM) fungi and soil nutrients in four lowland, neotropical rainforests. * Within each forest, samples were collected from plots that differed in fertility and above-ground biomass, and fine roots, AM hyphae and total soil nutrients were measured. * Among sites, total fine root mass varied by a factor of three, from 237+/-19 g m-2 in Costa Rica to 800+/-116 g m-2 in Brazil (0-40 cm depth). Both root mass and length were negatively correlated to soil nitrogen and phosphorus, but AM hyphae were not related to nutrients, root properties or above-ground biomass. * These results suggest that understanding how soil fertility affects fine roots is an additional factor that may improve the representation of root functions in global biogeochemical models or biome-wide averages of root properties in tropical forests.