The morphology and post-hatching development of the skull of Bolitoglossa nicefori (Caudata: Plethodontidae): developmental implications of recapitulation and repatterning

The cranial morphology of the direct-developing salamander Bolitoglossa nicefori and its post-hatching development are described and compared with that of other urodeles. Four stages of cranial development are defined on the basis of conspicuous events that occur during post-hatching ontogeny. The adult skull morphology of B. nicefori is similar to that of other plethodontids; however, some regions show interspecific variation. The post-hatching ontogeny of the skull and the stage of ossification observed in the hatchlings of B. nicefori show two important ontogenetic features: (1) a mosaic of early larval, metamorphic and post-metamorphic skull features in hatchlings, and (2) absence of characteristic larval elements in skull and hyoid apparatus. The distinctive stage of ossification in the hatchlings of B. nicefori could be caused by heterochronic changes in the ossification sequence, compared to the ontogeny of metamorphic salamanders. The possible heterochronic changes and the absence of larval traits are perhaps due to ontogenetic repatterning, yet without an obvious impact on the adult skull morphology (absence of morphological novelties). This might indicate a compartmentalized development. Further studies should be performed in order to establish the possible occurrence of recapitulatory patterns or ontogenetic repatterning in the skull morphogenesis of B. nicefori during its embryonic development.     

Ontogenetic plasticity of anatomical and ecophysiological traits and their correlations in Iris pumila plants grown in contrasting light conditions

To better understand what directs and limits the evolution of phenotype, constraints in the realization of the optimal phenotype need to be addressed. That includes estimations of variability of adaptively important traits as well as their correlation structures, but also evaluation of how they are affected by relevant environmental conditions and development phases. The aims of this study were to analyze phenotypic plasticity, genetic variability and correlation structures of important Iris pumila leaf traits in different light environments and ontogenetic phases, and estimate its evolutionary potential. Stomatal density, specific leaf area, total chlorophyll concentration and chlorophyll a/b ratio were analyzed on I. pumila full‐sib families in the seedling phase and on the same plants after 3 years of growth in contrasting light conditions typical for ontogenetic stage in question. There was a significant phenotypic plasticity in both ontogenetic stages, but significant genetic variability was detected only for chlorophyll concentrations. Correlations of the same trait between different stages were weak due to changes in environmental conditions and difference in ontogenetic reaction norms of different genotypes. Ontogenetic variability of correlation structures was detected, where correlations and integration were higher in seedlings compared with adult plants 3 years later. Correlations were affected by environmental conditions, with integration being higher in the lower light conditions, but correlations between phases being stronger in the higher light treatment. These findings demonstrated that the analyzed traits can be selected and can mostly evolve independently in different environments and ontogenetic stages, with low genetic variability as a potentially main constraint.     

Development and plasticity of endangered shrub Lindera melissifolia (Lauraceae) seedlings under contrasting light regimes

Lindera melissifolia (Walt.) Blume seedlings were raised in a growth chamber to determine the effects of light availability on shoot growth pattern, and basic leaf and stem growth. Lindera melissifolia seedlings exhibited a sympodial shoot growth pattern for 3 months following emergence from the soil medium, but this pattern was characterized by a reduction in leaf blade area approximately 30 days after emergence, followed by increases in leaf blade area. Seedlings receiving low light were 76% taller than seedlings receiving high light. Seedlings receiving low light also had larger leaf blade dimensions, blade area, seedling leaf area, and greater mass. Seedlings raised in high light had a greater proportional distribution of biomass in the roots, suggesting possible water stress from greater vapor pressure deficits. Furthermore, these seedlings displayed sharp angles of blade inclination and blade folding – acclimation that reduces exposure to light and subsequent higher leaf temperatures in open environments. These differences in morphological response to light resulted in high phenotypic variability in L. melissifolia seedlings. Lindera melissifolia seedling development showed a brief period of phenotypic plasticity, followed by ontogenetic plasticity. The short period of phenotypic plasticity may, however, have profound ecological implications for the conservation and recovery of this federally endangered shrub. Further experimentation should take into account the development of ontogenetic standards for comparisons of plant traits in addition to temporal standards.     

A comparative analysis of leaf shape of wheat, barley and maize using an empirical shape model

Background and Aims

The phenotypes of grasses show differences depending on growth conditions and ontogenetic stage. Understanding these responses and finding suitable mathematical formalizations are an essential part of the development of plant and crop models. Usually, a marked change in architecture between juvenile and adult plants is observed, where dimension and shape of leaves are likely to change. In this paper, the plasticity of leaf shape is analysed according to growth conditions and ontogeny.

Methods

Leaf shape of Triticum aestivum, Hordeum vulgare and Zea mays cultivars grown under varying conditions was measured using digital image processing. An empirical leaf shape model was fitted to measured shape data of single leaves. Obtained values of model parameters were used to analyse the patterns in leaf shape.

Key Results

The model was able to delineate leaf shape of all studied species. The model error was small. Differences in leaf shape between juvenile and adult leaves in T. aestivum and H. vulgare were observed. Varying growth conditions impacted leaf dimensions but did not impact leaf shape of the respective species.

Conclusions

Leaf shape of the studied T. aestivum and H. vulgare cultivars was remarkably stable for a comparable ontogenetic stage (leaf rank), but differed between stages. Along with other aspects of grass architecture, leaf shape changed during the transition from juvenile to adult growth phase. Model-based analysis of leaf shape is a method to investigate these differences. Presented results can be integrated into architectural models of plant development to delineate leaf shape for different species, cultivars and environmental conditions.     

Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

Are ontogenetic shifts in foliar structure and resource acquisition spatially conditioned in tank‐bromeliads?

The phenotypic plasticity of plants has been explored as a function of either ontogeny (apparent plasticity) or environment (adaptive plasticity), although few studies have analyzed these factors together. In the present study, we take advantage of the dispersal of Aechmea mertensii bromeliads by Camponotus femoratus or Pachycondyla goeldii ants in shaded and sunny environments, respectively, to quantify ontogenetic changes in morphological, foliar, and functional traits, and to analyze ontogenetic and ant species effects on 14 traits. Most of the morphological (plant height, number of leaves), foliar (leaf thickness, leaf mass area, total water content, trichome density), and functional (leaf δ¹³C) traits differed as a function of ontogeny. Conversely, only leaf δ¹⁵N showed an adaptive phenotypic plasticity. On the other hand, plant width, tank width, longest leaf length, stomatal density, and leaf C concentration showed an adaptation to local environment with ontogeny. The exception was leaf N concentration, which showed no trend at all. Aechmea mertensii did not show an abrupt morphological modification such as in heteroblastic bromeliads, although it was characterized by strong, size‐related functional modifications for CO₂ acquisition. The adaptive phenotypic variation found between the two ant species indicates the spatially conditioned plasticity of A. mertensii in the context of insect‐assisted dispersal. However, ant‐mediated effects on phenotypic plasticity in A. mertensii are not obvious because ant species and light environment are confounding variables. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 299–312.     

Ontogenetic changes in crown architecture and leaf arrangement: effects on light capture efficiency in three tree species differing in leaf longevity

Pronounced strategy shifts along ontogeny have been observed in several tree species, mainly because of the trend to maximize growth during the seedling stage, which constitutes the most vulnerable part of the tree’s life cycle. Our aim here was to analyze the ontogenetic changes in crown characteristics and light capture patterns in three Quercus species: the evergreens Quercus ilex and Quercus suber and the deciduous Quercus faginea co-occurring in a Mediterranean open woodland. The seedlings were distributed in the large clearings among the adults and received full sunlight. We constructed three-dimensional models of the aerial parts of seedlings and mature trees of the three species, using the YplantQMC program. Large differences between growth stages were observed for all variables. The seedlings exhibited smaller branch sizes and crown densities than those observed in the adult trees. Leaf angles to horizontal also tended to increase during ontogeny, whereas leaf dispersion and the observed distances between leaves tended to decrease. The amount of photosynthetic radiation absorbed per unit leaf area throughout the growing season was lower in adult specimens than in young specimens. Changes in absorption efficiency during ontogeny were more intense for the species with longer leaf life span at maturity. We conclude that more intense ontogenetic shifts in species with longer leaf life span reflect the priority change from the maximization of short-term productivity at the seedling stage to maximizing leaf longevity during the adult stage.     

Environmental and Ontogenetic Effects on Intraspecific Trait Variation of a Macrophyte Species across Five Ecological Scales

Although functional trait variability is increasingly used in community ecology, the scale- and size-dependent aspects of trait variation are usually disregarded. Here we quantified the spatial structure of shoot height, branch length, root/shoot ratio and leaf number in a macrophyte species Potamogeton maackianus, and then disentangled the environmental and ontogenetic effects on these traits. Using a hierarchical nested design, we measured the four traits from 681 individuals across five ecological scales: lake, transect, depth stratus, quadrat and individual. A notable high trait variation (coefficient variation: 48–112%) was observed within species. These traits differed in the spatial structure, depending on environmental factors of different scales. Shoot height and branch length were most responsive to lake, transect and depth stratus scales, while root/shoot ratio and leaf number to quadrat and individual scales. The trait variations caused by environment are nearly three times higher than that caused by ontogeny, with ontogenetic variance ranging from 21% (leaf number) to 33% (branch length) of total variance. Remarkably, these traits showed non-negligible ontogenetic variation (0–60%) in each ecological scale, and significant shifts in allometric trajectories at lake and depth stratus scales. Our results highlight that environmental filtering processes can sort individuals within species with traits values adaptive to environmental changes and ontogenetic variation of functional traits was non-negligible across the five ecological scales.     

Ontogenetic transition in leaf traits: a new cost associated with the increase in leaf longevity

Aims Recent work has identified a worldwide 'economics' spectrum of correlated leaf traits that mainly reflects the compromises between maximizing leaf longevity and short-term productivity. However, during the early stages of tree growth different species tend to exhibit a common strategy, because competition for soil water and nutrients forces the maximization of short-term productivity owing to the need for rapid growth during the most vulnerable part of the tree's life cycle. Accordingly, our aim here was to compare the variations that occur during ontogeny in the different leaf traits (morphology and leaf chemical composition) of several coexisting Mediterranean woody species differing in their leaf life spans and to test our hypothesis that tree species with a long leaf life span should exhibit larger shifts in leaf characteristics along ontogeny.Methods Six Mediterranean tree species differing in leaf life span, selected from three plots located in central-western Spain, were studied during three growth stages: seedlings, juveniles and mature trees. Leaf life span, leaf morphology (leaf area, dry weight, thickness and mass per unit area) and chemical composition (N and fibre concentrations) were measured in all six species. The magnitude of the ontogenetic changes in the different traits was estimated and related to the mean leaf longevity of the different species.Important findings Along ontogeny, strong changes were observed in all variables analysed. The early growth stages showed lower leaf thickness, leaf thickness and mass per unit area and N, cellulose and hemicellulose concentrations than mature trees, but a higher lignin content. However, these changes were especially marked in species with a longer leaf life span at maturity. Interspecific differences in leaf life span, leaf morphology and chemical composition were stronger at the mature stage than at the seedling stage. We conclude that greater plasticity and more intense strategy shifts along ontogeny are necessarily associated with long leaf life span. Our results thus provide a new aspect that should be incorporated into the analysis of the costs and benefits associated with the different strategies related to leaf persistence displayed by the different species. Accordingly, the intensity of the alterations in leaf traits among different growth stages should be added to the suite of traits that change along the leaf economics spectrum.     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Herbivore attack in Casearia nitida influenced by plant ontogenetic variation in foliage quality and plant architecture

Traits influencing plant quality as food and/or shelter for herbivores may change during plant ontogeny, and as a consequence, influence the amount of herbivory that plants receive as they develop. In this study, differences in herbivore density and herbivory were evaluated for two ontogenetic stages of the tropical tree Casearia nitida. To assess plant ontogenetic differences in foliage quality as food for herbivores, nutritional and defensive traits were evaluated in saplings and reproductive trees. Predatory arthropods were quantified and the foraging preferences of a parasitoid wasp of the genus Zacremnops were assessed. In addition, survival rates of lepidopteran herbivores (Geometridae) were evaluated experimentally. Herbivore density was three times higher and herbivory was 66% greater in saplings than in reproductive trees. Accordingly, concentrations of total foliar phenolics were higher in reproductive trees than in saplings, whereas leaf toughness, water and nitrogen concentration did not vary between ontogenetic stages. Survival rates of lepidopteran larvae exposed to natural enemies were equivalent in reproductive trees and saplings. Given the greater herbivore density on saplings, equal survival rates implied a greater foraging effort of predators on reproductive trees. Furthermore, observed foraging of parasitoid wasps was restricted to reproductive trees. I propose that herbivore density, and as a consequence, leaf damage were lower in reproductive trees than in saplings due to both traits influencing food quality, and architectural or unmeasured indirect defensive traits influencing foraging preference of natural enemies of herbivores.     

Predatory Potential of Euseius alatus (Phytoseiidae) on Different Life Stages of Oligonychus ilicis (Tetranychidae) on Coffee Leaves Under Laboratory Conditions

This study evaluated the predatory capacity of Euseius alatus (DeLeon) as a biological control agent of the pest mite Oligonychus ilicis (McGregor) on coffee leaves under laboratory conditions, using arenas containing 25 O. ilicis per coffee (Coffea arabica) leaf to one specimen of each stage of the predator mite. The functional response and oviposition rate of adult females of E. alatus were evaluated on coffee leaf arenas and offered from 1 to 125 immature stages of O. ilicis per arena. The number of preys killed and the number of eggs laid by the predator were evaluated every 24 h during 8 days. The preys consumed were daily replaced. Male and female adults of E. alatus were the most efficient in killing all developmental stages of O. ilicis. Larvae and nymphs of O. ilicis were the most consumed by all stages of the predatory mite. The functional response and oviposition rates of E. alatus increased as the prey density increased, with a positive and highly significant correlation. Regression analysis suggested a type II functional response, with a maximum predation of 22 O. ilicis/arena and a maximum oviposition rate of 1.7 eggs/day at a density of 70 O. ilicis/arena.     

Intraspecific Trait Variation Driven by Plasticity and Ontogeny in Hypochaeris radicata

The importance of intraspecific variation in plant functional traits for structuring communities and driving ecosystem processes is increasingly recognized, but mechanisms governing this variation are less studied. Variation could be due to adaptation to local conditions, plasticity in observed traits, or ontogeny. We investigated 1) whether abiotic stress caused individuals, maternal lines, and populations to exhibit trait convergence, 2) whether trait variation was primarily due to ecotypic differences or trait plasticity, and 3) whether traits varied with ontogeny. We sampled three populations of Hypochaeris radicata that differed significantly in rosette diameter and specific leaf area (SLA). We grew nine maternal lines from each population (27 lines total) under three greenhouse conditions: ambient conditions (control), 50% drought, or 80% shade. Plant diameter and relative chlorophyll content were measured throughout the experiment, and leaf shape, root∶shoot ratio, and SLA were measured after five weeks. We used hierarchical mixed-models and variance component analysis to quantify differences in treatment effects and the contributions of population of origin and maternal line to observed variation. Observed variation in plant traits was driven primarily by plasticity. Shade significantly influenced all measured traits. Plant diameter was the only trait that had a sizable proportion of trait variation (30%) explained by population of origin. There were significant ontogenetic differences for both plant diameter and relative chlorophyll content. When subjected to abiotic stress in the form of light or water limitation, Hypochaeris radicata exhibited significant trait variability. This variation was due primarily to trait plasticity, rather than to adaptation to local conditions, and also differed with ontogeny.     

ONTOGENETIC NICHE SHIFT IN THE BRACHIOPOD TEREBRATALIA TRANSVERSA: RELATIONSHIP BETWEEN THE LOSS OF ROTATION ABILITY AND ALLOMETRIC GROWTH

Abstract: Many articulated brachiopods experience marked life habit variations during ontogeny because they experience their fluid environment at successively higher Reynolds numbers, and they can change the configuration of their inhalant and exhalant flows as body size increases. We show that the extant brachiopod Terebratalia transversa undergoes a substantial ontogenetic change in reorientation governed by rotation around the pedicle. T. transversa′s reorientation angle (maximum ability to rotate on the pedicle) decreases during ontogeny, from 180 degrees in juveniles to 10–20 degrees in individuals exceeding 5 mm, to complete cessation of rotation in individuals larger than 10 mm. Rotation ability is substantially reduced after T. transversa achieves the adult lophophore configuration and preferred orientation with respect to ambient water currents at a length of 2.5–5 mm. We hypothesize that the rotation angle of T. transversa is determined mainly by the position of ventral and dorsal points of attachment of dorsal pedicle muscles relative to the pedicle. T. transversa shows a close correlation between the ontogenetic change in reorientation angle and ontogeny of morphological traits that are related to points of attachment of dorsal pedicle muscles, although other morphological features can also limit rotation in the adult stage. The major morphological change in cardinalia shape and the observed reduction of rotation affect individuals 2.5–10 mm in length. The position of ventral insertions of dorsal pedicle muscles remains constant, but contraction of dorsal pedicle muscles is functionally handicapped because dorsal insertions shift away from the valve midline, rise above the dorsal valve floor, and become limited by a wide cardinal process early in ontogeny (<5 mm). The rate of increase of cardinal process width and of distance between dorsal pedicle muscle scars substantially decreases in the subadult stage (5–10 mm), and most of the cardinalia shell traits grow nearly isometrically in the adult stage (>10 mm). T. transversa attains smaller shell length in crevices than on exposed substrates. The proportion of small‐sized individuals and population density is lower on exposed substrates than in crevices, indicating higher juvenile mortality on substrates prone to grazing and physical disturbance. The loss of reorientation ability can be a consequence of morphological changes that strengthen substrate attachment and maximize protection against biotic or physical disturbance (1) by minimizing torques around the pedicle axis and/or (2) by shifting energy investments into attachment strength at the expense of the cost involved in reorientation.     

FLORAL ONTOGENY IN CYCLAMEN PERSICUM ‘F-1 ROSEMUNDE ROSE’ (PRIMULACEAE)

Floral ontogeny was examined in Cyclamen persicum ‘F-1 Rosemunde Rose’ using a combination of light and scanning electron microscopy. The leaf plastochron index (LPI), earlier calculated for leaf elongation, was used to determine the length of each stage of floral development. LPI will provide a useful tool for selecting flowers of a given stage from large plant populations or from plants where flowers are small or inaccessible during early ontogenetic stages. Most features of floral development are similar to those previously described for other primulaceous genera. The petal-stamen relationship, however, is unusual; stamens arise through periclinal cell divisions in the adaxial surface layers of common petal-stamen primordia. Anatomical evidence suggests that the placenta is formed both by appendicular initials which give rise to the ovules and ventral carpellary bundles, and receptacular cells which form some, if not all, of the central axis.     

Morphological Variations within the Ontogeny of Deinonychus antirrhopus (Theropoda,Dromaeosauridae)

This research resulted from the determination that MCZ 8791 is a specimen of Deinonychus antirrhopus between one and two years of age and that the morphological variations within particular growth stages of this taxon have yet to be described. The primary goal of the research is to identify ontogenetic variations in this taxon. Histological analyses determined that the Deinonychus specimens AMNH 3015 and MOR 1178 were adults. Comparisons are made between MCZ 8791 and these adult specimens. The holotype, YPM 5205, and the other associated specimens of this taxon within the YPM collection are similar in size and morphology to AMNH 3015. Further comparisons were made with the three partial specimens OMNH 50268, MCZ 4371, and MOR 1182. Although these specimens represent only a partial ontogenetic series, a number of morphological variations can be described. One secondary goal of this research is to compare the known pattern of variable, informative, ontogenetic characters in MCZ 8791 to a similar pattern of morphological characters in the sub-adult dromaeosaurid specimen Bambiraptor feinbergorum, AMNH FR: 30556. If the characters that have been determined to represent variable juvenile morphology in the ontogeny of Deinonychus are exhibited in Bambiraptor, this study will begin the process of determining whether a similar, conservative, ontogenetic pattern exists throughout the rest of Dromaeosauridae. If defensible, it may reduce the number of sympatric taxa within this clade. The other secondary goal relates to the forelimb function. The approximate body size, forelimb length, wrist development, and the presence of a more prominent olecranon on the ulna of MCZ 8791 support the hypothesis that juveniles of this taxon possessed some form of flight capability.     

Intraspecific functional trait variability across different spatial scales: a case study of two dominant trees in Korean pine broadleaved forest

Intraspecific functional trait variability plays an important role in the response of plants to environmental changes. However, it is still unclear how the variability differs across three nested spatial scales (individual, plot, and site) and which determinants (climatic, soil, and ontogenetic variables) shape the trait variability. Along a latitudinal gradient in Korean pine broadleaved forest of northeast China, we quantified the extent of intraspecific variability of four functional traits in two dominant trees Pinus koraiensis and Fraxinus mandshurica at eight sites, including specific leaf area, leaf dry matter content (morphological traits) and leaf nitrogen content, leaf phosphorus content (physiological traits). Results showed a large trait variation within and between species (coefficient variation: 6.07–23.3%). The leaf physiological traits of F. mandshurica and morphological traits of P. koraiensis were more responsive at site scale, while the morphological traits of F. mandshurica and physiological traits of P. koraiensis were more responsive at individual scale. In addition, abiotic and biotic factors explaining functional trait variation differ markedly between the two tree species, with physiological trait of F. mandshurica being more associated with climate and soil, while traits variability in P. koraiensis was not affected by climate, soil, and ontogeny, except for leaf phosphorus content. Overall, we can predict that the physiological traits of broadleaved species tend to be more sensitive to environmental changes, while pines are more sensitive to competition. It is critical to determine which spatial scale and trait type should be taken into account in predictive models of vegetation dynamics.     

Sun-shade adaptability of the red mangrove,Rhizophora mangle (Rhizophoraceae): Changes through ontogeny at several levels of biological organization

Rhizophora mangle L., the predominant neotropical mangrove species, occupies a gradient from low intertidal swamp margins with high insolation, to shaded sites at highest high water. Across a light gradient, R. mangle shows properties of both “light-demanding” and “shade-tolerant” species, and defies designation according to existing successional paradigms for rain forest trees. The mode and magnitude of its adaptability to light also change through ontogeny as it grows into the canopy. We characterized and compared phenotypic flexibility of R. mangle seedlings, saplings, and tree modules across changing light environments, from the level of leaf anatomy and photosynthesis, through stem and whole-plant architecture. We also examined growth and mortality differences among sun and shade populations of seedlings over 3 yr. Sun and shade seedling populations diverged in terms of four of six leaf anatomy traits (relative thickness of tissue layers and stomatal density), as well as leaf size and shape, specific leaf area (SLA), leaf internode distances, disparity in blade–petiole angles, canopy spread: height ratios, standing leaf numbers, summer (July) photosynthetic light curve shapes, and growth rates. Saplings showed significant sun/shade differences in fewer characters: leaf thickness, SLA, leaf overlap, disparity in bladepetiole angles, standing leaf numbers, stem volume and branching angle (first-order branches only), and summer photosynthesis. In trees, leaf anatomy was insensitive to light environment, but leaf length, width, and SLA, disparities in bladepetiole angles, and summer maximal photosynthetic rates varied among sun and shade leaf populations. Seedling and sapling photosynthetic rates were significantly depressed in winter (December), while photosynthetic rates in tree leaves did not differ in winter and summer. Seasonal and ontogenetic changes in response to light environment are apparent at several levels of biological organization in R. mangle, within constraints of its architectural baiiplan. Such variation has implications for models of stand carbon gain, and suggest that response flexibility may change with plant age.     

Prickly Poppies Can Get Pricklier: Ontogenetic Patterns in the Induction of Physical Defense Traits

Plant ontogeny is a common source of variation in defense and herbivory. Yet, few studies have investigated how the induction of physical defense traits changes across plant ontogeny. Physical defense traits are costly to produce, and thus, it was predicted that induction as a cost-saving strategy would be particularly favorable for seedlings, leading to ontogenetic declines in the inducibility of these traits. We tested for induction of three different physical defense traits (prickles, latex and leaf toughness) in response to mechanical defoliation and jasmonic acid application using prickly poppies (Argemone glauca and A. mexicana, Papaveraceae) as a model system. Genetic variation in the induction of physical defenses was tested using maternal sib-ships sampled from multiple populations. Both species induced higher densities of laminar prickles, although the magnitude of induction was much higher in the endemic Hawaiian prickly poppy, A. glauca, than in the cosmopolitan A. mexicana. The magnitude of prickle induction was also higher in young compared to older juvenile plant stages in A. glauca, demonstrating a strong role of ontogeny. Neither latex exudation nor leaf toughness was induced in either species. Although significant genetic variation was detected within and among populations for constitutive expression of physical defense traits in Argemone, there was no evidence for genetic variation in the induction of these traits. This study provides the first evidence for the induction of physical defenses in prickly poppies, emphasizing how an ontogenetically explicit framework can reveal new insights into plant defense. Moreover, this study illustrates how sister species comparisons between island vs. continental plants can provide new insights into plant functional and evolutionary ecology, highlighting a fruitful area for future research on more species pairs.