Early Growth and Physiological Acclimation to Shade and Water Restriction of Seven Sclerophyllous Species of the Mediterranean Forests of Central Chile.

The success of using active restoration in Mediterranean-type climate zones mostly depends on an appropriate matching of plant species and specific management prescriptions upon establishment. In this study, we assessed the early growth and short-term physiological acclimation of seven common species found in the sclerophyllous forests in central Chile to water restriction and shading. We established a nursery experiment that included three treatments (T0: sun-exposed and water-restricted, T1: sun-exposed and fully irrigated, and T2: shaded and fully irrigated) and seven tree species differing in their shade and drought tolerance (Quillaja saponaria Molina, Aristotelia chilensis (Mol.) Stuntz, Peumus boldus Molina, Lithraea caustica (Mol.) Hook. and Arn, Luma apiculata (DC.) Burret, Colliguaja odorifera Molina, and Escallonia pulverulenta (Ruiz and Prav.) Pers). We measured the increment in seedling height and different leaf morpho-physiological traits during two months in the dry season. Based on the measured traits, none of the species took advantage of the higher water availability in T1 relative to T0, but most of the species responded to the shade in T2, regardless of their shade or drought tolerance. Height increments due to shade varied from 0% in P. boldus to 203% in L. apiculata. Overall, all the species responded similarly to the treatments in specific leaf area, chlorophyll content index, photosynthetic rate, stomatal conductance, and intrinsic water use efficiency. This suggests that the species exhibited similar acclimation patterns of these parameters to shade and drought, even regarding the variation in midday xylem water potential found in the water-restricted treatment T0 (from -1.5 MPa in P. boldus to -3.1 MPa in E. pulverulenta). In this study, shading had a higher positive effect on the seedling performance of sclerophyllous species than watering, which at operational level highlights the need for investing in tree shelters when using these species in restoration programs.

Morpho-physiological and anatomical responses of two Urochloa hybrids under shade: Dataset article.

Silvopastoral systems are an important strategy for sustainable livestock production. However, to expand their implementation, it is crucial to identify and develop forage materials that maintain good production and quality while being tolerant to shade conditions as well as other biotic and abiotic stresses. A field trial was conducted to evaluate the morpho-anatomical and agronomic responses of two Urochloa hybrids (Camello and Talisman) under two light conditions: shade (28 % light intensity) and full exposure. The trial followed a randomised complete block design with split-plot arrangement, where each treatment corresponded to a plot with three replications. Morphological and anatomical parameters were recorded in three technical replicates of each replication. Histological leaf sections were analysed for the percentage of adaxial epidermis, abaxial epidermis, vascular tissue, colourless parenchyma, Kranz sheath, bulliform cells, sclerenchyma, and chlorenchyma. Measurements in leaf included relative chlorophyll concentration, leaf area, leaf length, and leaf width. Evaluations in plant included height and number of tillers. Agronomic parameters such as plant cover and dry biomass were recorded for each plot. Additionally, six leaf imprints were made on the leaf undersides to observe stomatal morphology, and their length was recorded. Furthermore, plants from each treatment were grown in soil-filled tubes within the same plots. Root system photographs were taken, and in three replications per treatment, root length, root diameter, root volume, root surface area, and the depth at which 95 % of roots were concentrated (D95) were determined. These data can be utilised by the scientific community and breeders to conduct analyses and meta-analyses to identify shade tolerance mechanisms and develop genetic materials tolerant to changing climatic conditions while being optimal for use in silvopastoral systems.

Trait coordination at leaf level explains the resistance to excess light stress in shade-tolerant tropical tree species.

Plant functioning in response to the environment is an important issue for vegetation reassembly of degraded lands because of both low and excessive sunlight influence the performance of young plants. However, how shade-tolerant tree species deal with excess of light energy remains unclear due to the contrasting results among studies and the subjective classifications of species shade tolerance. From a quantitative classification of shade tolerance of 12 tropical tree species planted in the field under contrasting light conditions, we hypothesized that shade-tolerant species are capable of effective long-term acclimation to high-light conditions. Plant size and survival of species were measured at the beginning and 38 months after planting. We also measured functional traits associated with carbon economy and non-photochemical energy dissipation. Under high-light condition, more light-demanding species showed higher growth rates associated with higher values of functional traits that improve the CO2 assimilation capacity. By contrast, more shade-tolerant species showed higher survival that was associated with higher values of qN, leaf K contents and leaf thickness. The concomitant increase in these traits, as well as their greater plasticity to light in WUEi, Fv/Fm and qN, confers on these species strong photoprotection to avoid high-light stress and to persist under open field conditions. The results challenge the common assumption that only fast-growing and light-demanding tree species are suitable for vegetation reassembly in full sunlight conditions and to improve the environmental conditions for other species.

Germination of small tropical seeds has distinct light quality and temperature requirements, depending on microhabitat.

The coexistence of plant species in tropical rainforests is related to specific abiotic resources, varying according to the occurrence microhabitat of each species. Light quality is the main abiotic factor influencing germination of small seeds; however, studies often do not discriminate its effect from that of light irradiance. This study compared specific requirements for seed germination of ten small-seeded species, with restricted occurrence in only one of three contrasting microhabitats: forest understorey, edge of clearings and open areas. Laboratory experiments were carried out to test temperature regime (constant or fluctuating), light quality (R:FR) and light irradiance (PAR), which reproduce high and low conditions commonly found in the microhabitats. Seed germination of all species occurred between 20 and 30 °C, only seeds of open area species were able to germinate at 35 °C and no species required alternating temperatures to germinate. Irrespective of species and microhabitat, a decrease in the R:FR reduced the germination percentage; however, there were differences in the capacity to germinate at low R:FR. The values of R:FR50% were higher for open area and edge species (0.441-0.345) than for understorey species (0.181-0.109), with few exceptions. For all species and most of the tests, germination was not influenced by PAR. Light quality is the most important light signal for germination of small seeds; irradiance has little effect. Our results suggest two distinct patterns of germination for small-seeded species: open area and edge species are light-demanding and require high R:FR to germinate, while understorey species are shade-tolerant and germinate at low R:FR. These differences are responsible for distinct microhabitat occurrence and help to explain the coexistence of species in tropical forests.

Effects of shade on guinea grass genotypes Megathyrsus maximus (Poales: Poaceae) / Efectos de la sombra sobre genotipos del zacate guinea Megathyrsus maximus (Poales: Poaceae)

Introduction: Light stress is an important factor limiting the biomass yield while combining forage production with crops or forestry. Guinea grass is a widely adapted perennial fodder grass. The species exhibits high degree of variation for morphology, adaptation and biomass yield. Objective: Since there is a need in identifying shade adapted forage grasses for the expanding area under agroforestry/silvipastures, the present investigation took the task of understanding how the morphologically distinct genotypes of guinea grass respond under different shaded intensities. Methods: In the present study, forty-four genotypes related with the shade response were studied in varying shading conditions (pure sunlight, 25, 50 and 75 % shade) created artificially. Results: Based on green and dry matter yields ranking, the genotype IG 01-98 performed the best followed by genotypes IG 01-92, IG 97-5, IG 97-6 and IG 01-89 in decreasing order. Particularly, IG 01-93 was identified as the best performing under 50 % shading conditions. On the other hand, most of the top ranking genotypes performed well both under open and up to 50 % of shade. Morphologically, these genotypes were taller and possessed longer and broader leaves. Under shaded conditions (over 50 %), leaf length and width showed an increasing trend compared to open conditions. Also, chlorophyll content increased with shading intensity. Conclusions: Most of the genotypes collected from the southern Indian humid tropical environment with early flowering nature were tolerant to shade. Differential genotypic response was observed for biomass yield and yield attributes under shade. The study established appreciable variability for shade tolerance among genotypes.

Introducción: El estrés leve es un factor importante que limita el rendimiento de la biomasa al tiempo que combina laproducción forrajera con los cultivos o la silvicultura. El zacate guinea es una planta de forraje perenne ampliamente adaptada. La especie presenta un alto grado de variación en su morfología, adaptación y rendimiento de biomasa. Objetivo: Como existe la necesidad de la identificación de pastos forrajeros adaptados a la sombra para el área de expansión bajo agroforestería /silvicultura, la presente investigación se realizó para entender cómo los genotipos morfológicamente distintos al zacate guinea responden a diferentes intensidades de sombra. Métodos: Se estudiaron 44 genotipos y su respuesta a los niveles de sombra: luz solar pura y 25, 50 y 75 % de sombra (creados artificialmente). Resultados: Basado en la clasificación de rendimientos demateria verde y seca, el genotipo IG 01-98 fue el mejor, seguido de los genotipos IG 01-92, IG 97-5, IG 97-6 e IG 01-89 en orden decreciente. Por su parte, el IG 01-93 fue identificado como el de mayor rendimiento bajo el 50 % de condición de sombra. La mayoría de los genotipos de primer nivel tuvieron un buen desempeño, tanto bajo sombra abierta como al 50 %. Morfológicamente, estos genotipos fueron más altos y poseían hojas más largas y más anchas. En condiciones de sombra, de más del 50 %, la longitud y ancho de la hoja mostraron una tendencia creciente en comparación con la condición abierta. El contenido de clorofila aumentó con la intensidad de la sombra. Conclusiones: La mayoría de los genotipos recolectados al sur de la India en un ambiente tropical húmedo y tienen una floración temprana, fueron tolerantes a la sombra. Se observó una respuesta genotípica diferencial para los atributos de rendimiento y rendimiento de biomasa bajo sombra. El estudio mostró la existencia de una variabilidad apreciable para la tolerancia a la sombra entre los genotipos.

Shade tolerance within the context of the successional process in tropical rain forests

Shade tolerance (the capacity to survive and grow over long periods under shade) is a key component of plant fitness and the foundation of current theories of forest succession in tropical rain forests. It serves as a paradigm to understand the optimal allocation of limited resources under dynamic light regimes. I analyze how tropical rain forest succession influences the expression of ecophysiological mechanisms leading to shade tolerance, and identify future areas that will increase our understanding of the ecological and evolutionary consequences of this phenomenon. Shade tolerance is a multivariate, continuous functional trait reflecting the growth-mortality trade-off of investing resources under limited light vs. exploiting high light conditions. I propose the life cycle successional trajectory model of Gómez-Pompa & Vázquez-Yanes as an integrative tool to understand tropical rain forest succession. This model shows how species distribute along the successional environmental gradient based on their degree of shade tolerance and represents a more integrative paradigm to understand the interface between different aspects of species diversity (ontogenetic variation and functional diversity) throughout succession. It proposes that different trait combinations determining shade tolerance are expressed at different stages of the life cycle, which affects how and when plants enter the successional trajectory. Models explaining the expression of shade tolerance (resource availability, carbon gain, CSR, resource competition) are based on whole-plant economics and are not mutually exclusive. The analysis of shade tolerance is biased towards tree seedlings in the understory of mature forests. Other life stages (juvenile and adult trees), life forms, and microhabitats throughout the forest profile are almost always excluded from these analyses. More integrative explanations based on the distribution of functional traits among species, ontogenetic stages, and the nature of the environmental gradient are being developed based on long-term data and chronosequence comparisons. In summary, shade-tolerance is a complex phenomenon, is determined by multiple characters that change ontogenetically over space and time and entails considerable plasticity. Current methods do not account for this plasticity. Understanding the nature of shade tolerance and its functional basis is critical to comprehending plant performance and improving the management, restoration and conservation of tropical rain forests given the combined threats of global warming and habitat loss.

La tolerancia a la sombra (la capacidad de sobrevivir y crecer durante largos períodos bajo sombra profunda) es un componente clave del valor adaptativo de la planta y la base de las teorías actuales de la sucesión forestal de la selva tropical. Sirve como un paradigma para entender la asignación óptima de recursos limitados bajo regímenes dinámicos de luz. En esta revisión analizo cómo la sucesión de los bosques tropicales lluviosos influye en la expresión de los mecanismos ecofisiológicos que conducen a la tolerancia a la sombra, e identifico áreas futuras que pueden aumentar nuestra comprensión de las consecuencias ecológicas y evolutivas de este fenómeno. La tolerancia a la sombra es un rasgo funcional continuo y multivariable que refleja el balance de invertir recursos bajo condiciones de luz limitada versus crecer más rápidamente en condiciones de luz intensa. Propongo el modelo de ciclo de vida a lo largo de la trayectoria de sucesión de Gómez-Pompa y Vázquez-Yanes como una herramienta integradora para entender la sucesión de la selva tropical. Este modelo muestra cómo las especies se distribuyen a lo largo del gradiente ambiental en función de su grado de tolerancia a la sombra, y representa un paradigma más integrador para comprender la interacción entre los diferentes componentes de la diversidad de especies (diversidad taxonómica y funcional y variación ontogenética) a lo largo de la sucesión. El modelo propone que las diferentes combinaciones de caracteres funcionales que determinan la tolerancia a la sombra se expresan en diferentes etapas del ciclo de vida, y afectan cómo y cuándo las plantas ingresan en el proceso de sucesión. Los modelos que explican la expresión de tolerancia a la sombra (disponibilidad de recursos, ganancia de carbono, CSR, competencia de recursos) se basan en la economía de toda la planta y no son mutuamente excluyentes. Se están desarrollando explicaciones más integradoras basadas en la distribución de caracteres funcionales entre especies, etapas ontogenéticas, y micrositios, mediante el uso de estudios de cronosecuencia y metadatos colectados a largo plazo. El análisis de la tolerancia a la sombra está sesgado hacia las plántulas de árboles y el sotobosque. Otras formas de vida y microhábitats dentro del perfil del bosque están casi excluidas de estos análisis. En resumen, la tolerancia a la sombra es un fenómeno complejo, está determinada por múltiples caracteres funcionales que cambian ontogenéticamente en el espacio y el tiempo, e implica una considerable plasticidad. Los métodos actuales no toman en cuenta esta plasticidad. Comprender la naturaleza de la tolerancia a la sombra y su base funcional es fundamental para entender el crecimiento de la planta y mejorar la gestión, restauración, y conservación de los bosques tropicales, los cuales enfrentan las amenazas combinadas del calentamiento global y la pérdida de hábitat.

Life history traits influence the strength of distance- and density-dependence at different life stages of two Amazonian palms.

Background and Aims: Natural enemies are known to be important in regulating plant populations and contributing to species coexistence (Janzen-Connell effects). The strength of Janzen-Connell effects (both distance- and density-effects) varies across species, but the life history traits that may mediate such a variation are not well understood. This study examined Janzen-Connell effects across the life stages (seed through adult stages) of two sympatric palm species with distinct phenologies and shade tolerances, two traits that may mediate the strength and timing of Janzen-Connell effects. Methods: Populations of two common palm species, Attalea phalerata and Astrocaryum murumuru , were studied in Manu National Park, Peru. Seed predation experiments were conducted to assess Janzen-Connell effects at the seed stage. In the post-seed stages, spatial point pattern analyses of the distributions of individuals and biomass were used to infer the strength of distance- and density-effects. Key Results: Seed predation was both negative distance- and density-dependent consistent with the Janzen-Connell effects. However, only seedling recruitment for asynchronously fruiting Attalea phalerata was depressed near adults while recruitment remained high for synchronously fruiting Astrocaryum murumuru , consistent with weak distance-effects. Negative density-effects were strong in the early stages for shade-intolerant Attalea phalerata but weak or absent in shade-tolerant Astrocaryum murumuru. Conclusions: Distance- and density-effects varied among the life stages of the two palm species in a manner that corresponded to their contrasting phenology and shade tolerance. Generalizing such connections across many species would provide a route to understanding how trait-mediated Janzen-Connell effects scale up to whole communities of species.

Shade tolerance and herbivory are associated with RGR of tree species via different functional traits.

Relative growth rate (RGR) plays an important role in plant adaptation to the light environment through the growth potential/survival trade-off. RGR is a complex trait with physiological and biomass allocation components. It has been argued that herbivory may influence the evolution of plant strategies to cope with the light environment, but little is known about the relation between susceptibility to herbivores and growth-related functional traits. Here, we examined in 11 evergreen tree species from a temperate rainforest the association between growth-related functional traits and (i) species' shade-tolerance, and (ii) herbivory rate in the field. We aimed at elucidating the differential linkage of shade and herbivory with RGR via growth-related functional traits. We found that RGR was associated negatively with shade-tolerance and positively with herbivory rate. However, herbivory rate and shade-tolerance were not significantly related. RGR was determined mainly by photosynthetic rate (Amax ) and specific leaf area (SLA). Results suggest that shade tolerance and herbivore resistance do not covary with the same functional traits. Whereas shade-tolerance was strongly related to Amax and to a lesser extent to leaf mass ratio (LMR) and dark respiration (Rd ), herbivory rate was closely related to allocation traits (SLA and LMR) and slightly associated with protein content. The effects of low light on RGR would be mediated by Amax , while the effects of herbivory on RGR would be mediated by SLA. Our findings suggest that shade and herbivores may differentially contribute to shape RGR of tree species through their effects on different resource-uptake functional traits.

Seedling growth and survival of five tree species in secondary forests and adjacent pastures in the montane rain forests of Southern Costa Rica / Crecimiento y supervivencia de plántulas de cinco especies de árboles en bosques secundarios y pastizales adyacentes en un bosque lluvioso montano del sur de Costa Rica

Abstract:Successful forest restoration in tropical environments is limited by the paucity of studies on the initial establishment and early survival requirements of seedlings of most native tropical tree species under disturbed conditions. Here, we evaluated the initial growth responses and the regeneration potential of seedlings of five tree species native to Costa Rica (Hasseltia floribunda, Inga densiflora, Persea americana, Tapirira mexicana and Trichilia pittieri). Seedlings were grown in secondary forests and adjacent open pastures under contrasting conditions of light availability. We quantified seedling growth, survival and herbivory from August 2010 to August 2011 on a monthly basis, and measured differences in leaf mass per area (LMA) at the end of the experiment. We found significant variation in growth responses between the understory of secondary forests and pastures. Growth in height was highest in pastures across all species, with I. densiflora, P. americana and T. mexicana showing the most striking differences. In contrast, H. floribunda and T. pittieri did not show differences in diameter growth between environments. Except for T. mexicana, herbivory increased throughout the experiment in all the species. Herbivory increased faster in pastures for H. floribunda and T. pittieri and showed higher rates in the forest understory for I. densiflora and P. americana. Seedling survival showed significant differences for all species across environments. Survival of H. floribunda and I. densiflora was higher in secondary forests, whereas the other species showed higher survival in pastures. LMA showed higher values in the pastures across all species, showing that individuals exposed to high light conditions had great photosynthetic rate and great leaf construction capacity. Due to their rapid growth and high survival, I. densiflora and T. mexicana showed great potential to restore abandoned pastures and secondary forests. Increasing our knowledge on the response of seedlings under disturbed conditions in tropical ecosystems is critical for improving the restoration of altered environments by matching the ecological amplitude of native species with specific environmental conditions. Rev. Biol. Trop. 64 (4): 1565-1583. Epub 2016 December 01.

Resumen:La restauración exitosa del bosque en ambientes tropicales está limitada por la carencia de estudios sobre los requisitos de hábitat, el establecimiento inicial, y la sobrevivencia de las plántulas de la mayoría de las especies de árboles tropicales nativos. En este estudio, evaluamos las respuestas en el crecimiento inicial y el potencial de regeneración de las plántulas de cinco especies de árboles nativos de Costa Rica (Hasseltia floribunda, Inga densiflora, Persea americana, Tapirira mexicana y Trichilia pittieri). Sembramos las plántulas en bosques secundarios y en pastizales abiertos adyacentes con condiciones de luz contrastantes. Cuantificamos el crecimiento, la supervivencia y el daño por herbivoría de las plántulas de forma mensual entre agosto 2010 y agosto 2011, y medimos las diferencias en el la masa foliar por área de la hoja al final del experimento. Existió una variación significativa en el crecimiento de las plántulas expuestas a la sombra en los bosques secundarios y aquellas que crecieron en pastizales. El crecimiento en altura fue mayor en pastizales en todas las especies; I. densiflora, P. americana y T. mexicana mostraron la mayor diferencia entre ambientes. En con- traste, H. floribunda y T. pittieri no mostraron diferencias en el crecimiento del diámetro entre ambientes. Con la excepción de T. mexicana, todas las especies mostraron un aumento en herbivoría durante el estudio. En los pastizales, el incremento de la herbivoría fue más rápido en H. floribunda y T. pittieri, mientras que para I. densiflora y P. americana fue mayor en el bosque. La sobrevivencia de plántulas fue diferente entre ambientes para todas las especies. La sobrevivencia de H. floribunda e I. densiflora fue mayor en el bosque secundario, mientras que en las demás especies hubo mayor sobrevivencia de plántulas en los pastizales. La masa foliar por área fue mayor en las plántulas que crecieron en los pastizales que en bosque secundario para todas las especies, lo que sugiere que los individuos que crecieron en condiciones de alta incidencia de luz tuvieron mayor tasa fotosintética y mayor capacidad de construcción de la hoja. Debido al rápido crecimiento y alta supervivencia I. densiflora y T. mexicana tienen gran potencial para la restauración de bosques secundarios y pastizales abandonados. Generar información sobre la respuesta de plántulas creciendo en bosques secundarios y pastizales en regeneración es crítico para mejorar los procesos de restauración de ambientes alterados. La restauración de estos bosques depende de la correspondencia entre las condiciones específicas de hábitat y la plasticidad ecológica de las especies nativas.

Carbon dynamics of Acer pseudoplatanus seedlings under drought and complete darkness.

Carbon (C) storage is considered a key component to plant survival under drought and shade, although the combined effects of these factors on survival remain poorly understood. We investigated how drought and shade alter the C dynamics and survival of tree seedlings, and whether drought limits the access to or usage of stored C. We experimentally applied two levels of soil humidity (well-watered versus drought, the latter induced by dry-down) and light availability (light versus complete darkness) on 1-year-old seedlings of Acer pseudoplatanus L. for 3 months. We quantified the survival, biomass, growth rate and non-structural carbohydrates (NSC) of seedlings at their time of death or at the end of the experiment for those that survived. We found that the soil dried out faster when drought was combined with light than when it was combined with complete darkness. Seedlings subjected to both drought and light showed reduced growth and reached 100% mortality earlier than any other treatment, with the highest NSC concentrations at the time of death. Seedlings exposed to both drought and complete darkness died significantly earlier than seedlings exposed to complete darkness only, but had similar NSC concentrations at time of their death, suggesting that drought accelerated the use of stored C under complete darkness. Complete darkness significantly reduced seedling growth and whole-plant NSC concentrations regardless of soil humidity, while root NSC concentrations were significantly more reduced when complete darkness was combined with drought conditions. Thus, the C dynamics in A. pseudoplatanus seedlings under complete darkness was not hindered by drought, i.e., the access and use of stored C was not limited by drought. The contrasting growth and C storage responses driven by drought under light versus complete darkness are consistent with a key role of the drought progression in the C dynamics of trees.

Interactions between plant size and canopy openness influence vital rates and life-history tradeoffs in two neotropical understory herbs.

UNLABELLED: • PREMISE OF THE STUDY: For tropical forest understory plants, the ability to grow, survive, and reproduce is limited by the availability of light. The extent to which reproduction incurs a survival or growth cost may change with light availability, plant size, and adaptation to shade, and may vary among similar species.• METHODS: We estimated size-specific rates of growth, survival, and reproduction (vital rates), for two neotropical understory herbs (order Zingiberales) in a premontane tropical rainforest in Costa Rica. During three annual censuses we monitored 1278 plants, measuring leaf area, number of inflorescences, and canopy openness. We fit regression models of all vital rates and evaluated them over a range of light levels. The best fitting models were selected using Akaike's Information Criterion.• KEY RESULTS: All vital rates were significantly influenced by size in both species, but not always by light. Increasing light resulted in higher growth and a higher probability of reproduction in both species, but lower survival in one species. Both species grew at small sizes but shrank at larger sizes. The size at which shrinkage began differed among species and light environments. Vital rates of large individuals were more sensitive to changes in light than small individuals.• CONCLUSIONS: Increasing light does not always positively influence vital rates; the extent to which light affects vital rates depends on plant size. Differences among species in their abilities to thrive under different light conditions and thus occupy distinct niches may contribute to the maintenance of species diversity.

Leaf Morphological Plasticity of Tree Species from Two Developmental Stages in Araucaria Forest

This study compared the morphological and anatomical variations of the leaves of four shade-tolerant tree species Allophylus edulis (St.-Hil.) Radlk (Sapindaceae), Casearia sylvestris Sw. (Salicaceae), Cupania vernalis Cambess. (Sapindaceae) and Luehea divaricata Mart. (Malvaceae) from a fragment of Araucaria forest in two developmental stages. Morphological and anatomical traits, such as leaf and tissue thickness, leaf area, leaf dry mass, specific leaf area, leaf density and stomata density were measured from 30 leaves of each developmental stage. The phenotypic plasticity index was also calculated for each quantitative trait. The results showed that the four species presented higher mean values ​​for specific leaf area and spongy/palisade parenchyma ratio at young stage, and higher mean values ​​for stomata density, total and palisade parenchyma thickness in the adult stage. The plasticity index demonstrated that L. divricata presented highest plasticity for both the morphological and anatomical traits while A. edulis displayed the lowest plasticity index. The results of this study indicated that the leaves of these species exhibited distinct morphological traits at each stage of development to cope with acting environmental factors.

Influence of light conditions on the allometry and growth of the understory palm Geonoma undata subsp. edulis (Arecaceae) of neotropical cloud forests.

PREMISE OF THE STUDY: Knowledge on the growth responses of understory palms to changing light conditions within neotropical cloud forests is limited. The low light regime of these environments, in addition to persistent cloudiness, low ambient temperatures, and slow nutrient cycles, imposes significant constraints on biomass accumulation. Here, we evaluate how changes in the understory light conditions influenced the allometry and growth of G. undata subsp. edulis in two cloud forests in Costa Rica. METHODS: We examined the structural relationships between stem diameter, stem height, and crown area in reproductive and nonreproductive individuals. We related the variation in stem growth and crown area with allometry, leaf production and longevity, and light conditions that we measured using hemispherical photographs over 1 year. KEY RESULTS: The allometric and growth pattern of G. undata subsp. edulis was characterized by its investment in crown area, which was strongly and positively related to increments in palm height and reproduction. Growth, measured as the increase in crown area and stem height, was not explained by the variation in the light regime spanning 1 year. However, reproductive individuals were generally taller, more slender, and had larger leaf areas than nonreproductive individuals. CONCLUSIONS: Our results demonstrated that stem growth responses were mostly controlled by initial crown size rather than by temporal differences in the understory light regimes of cloud forests. These results suggest that cloud forest understory palms have a limited capacity to respond to light changes and rely mostly on an opportunistic strategy for biomass accumulation and reproduction.

Does plant architectural complexity increase with increasing habitat complexity?: a test with a pioneer shrub in the Brazilian Cerrado / A complexidade arquitetural de plantas aumenta com a complexidade do habitat?: um teste com um arbusto pioneiro de Cerrado

Understanding variation in plant traits in heterogeneous habitats is important to predict responses to changing environments, but trait-environment associations are poorly known along ecological gradients. We tested the hypothesis that plant architectural complexity increases with habitat complexity along a soil fertility gradient in a Cerrado (Neotropical savanna) area in southeastern Brazil. Plant architecture and productivity (estimated as the total number of healthy infructescences) of Miconia albicans (SW.) Triana were examined in three types of vegetation which together form a natural gradient of increasing soil fertility, tree density and canopy cover: grasslands (campo sujo, CS), shrublands (cerrado sensu strico, CE) and woodlands (cerradão, CD). As expected, plants growing at the CS were shorter and had a lower branching pattern, whereas plants at the CD were the tallest. Unexpectedly, however, CD plants did not show higher architectural complexity compared to CE plants. Higher architectural similarity between CE and CD plants compared to similarity between CS and CE plants suggests reduced expression of functional architectural traits under shade. Plants growing at the CE produced more quaternary shoots, leading to a larger number of infructescences. This higher plant productivity in CE indicates that trait variation in ecological gradients is more complex than previously thought. Nematode-induced galls accounted for fruit destruction in 76.5% infructescences across physiognomies, but percentage of attack was poorly related to architectural variables. Our data suggest shade-induced limitation in M. albicans architecture, and point to complex phenotypic variation in heterogeneous habitats in Neotropical savannas.(AU)

O entendimento da variação dos atributos de plantas em ambientes heterogêneos é importante para prever respostas às mudanças climáticas; entretanto, a resposta de atributos das plantas às mudanças ambientais é pouco conhecida ao longo de gradientes ecológicos. Testou-se a hipótese de que a complexidade arquitetural de Miconia albicans (SW.) Triana aumenta com a complexidade ambiental, ao longo de um gradiente de fertilidade de solo em um Cerrado no sudeste brasileiro. A arquitetura e a produtividade (estimada por meio do número total de infrutescências sadias) foram examinadas em três tipos de vegetação, que formam um gradiente de fertilidade de solo, de densidade de árvores e de cobertura vegetal: campo sujo (CS), cerrado sensu stricto (CE) e cerradão (CD). Plantas crescendo no CS foram menores e apresentaram menor complexidade arquitetural, enquanto plantas no CD foram maiores. No entanto, de forma inesperada, plantas no CD não demonstraram maior complexidade arquitetural quando comparadas às plantas de CE, e a maior similaridade arquitetural entre plantas de CE e CD, quando comparadas com plantas de CE e CS, sugere que a expressão de atributos funcionais da arquitetura é limitada sob condições de sombra. Plantas de CE produziram mais ramos quaternários e maior número de infrutescências, sendo que a maior produtividade destas plantas indica que a variação de atributos em gradientes ecológicos é mais complexa do que se pensava anteriormente. Galhas de nematóides destruíram 76,5% das infrutescências nas vegetações, mas a porcentagem de ataque não esteve fortemente correlacionada com as variáveis arquiteturais. Os resultados do presente trabalho sugerem limitação na expressão da arquitetura da planta induzida pela sombra e apontam para uma complexa variação fenotípica na colonização de ambientes heterogêneos de savanas neotropicais.(AU)

Does plant architectural complexity increase with increasing habitat complexity? A test with a pioneer shrub in the Brazilian Cerrado / A complexidade arquitetural de plantas aumenta com a complexidade do habitat? Um teste com um arbusto pioneiro de Cerrado

Understanding variation in plant traits in heterogeneous habitats is important to predict responses to changing environments, but trait-environment associations are poorly known along ecological gradients. We tested the hypothesis that plant architectural complexity increases with habitat complexity along a soil fertility gradient in a Cerrado (Neotropical savanna) area in southeastern Brazil. Plant architecture and productivity (estimated as the total number of healthy infructescences) of Miconia albicans (SW.) Triana were examined in three types of vegetation which together form a natural gradient of increasing soil fertility, tree density and canopy cover: grasslands (campo sujo, CS), shrublands (cerrado sensu strico, CE) and woodlands (cerradão, CD). As expected, plants growing at the CS were shorter and had a lower branching pattern, whereas plants at the CD were the tallest. Unexpectedly, however, CD plants did not show higher architectural complexity compared to CE plants. Higher architectural similarity between CE and CD plants compared to similarity between CS and CE plants suggests reduced expression of functional architectural traits under shade. Plants growing at the CE produced more quaternary shoots, leading to a larger number of infructescences. This higher plant productivity in CE indicates that trait variation in ecological gradients is more complex than previously thought. Nematode-induced galls accounted for fruit destruction in 76.5% infructescences across physiognomies, but percentage of attack was poorly related to architectural variables. Our data suggest shade-induced limitation in M. albicans architecture, and point to complex phenotypic variation in heterogeneous habitats in Neotropical savannas.

O entendimento da variação dos atributos de plantas em ambientes heterogêneos é importante para prever respostas às mudanças climáticas; entretanto, a resposta de atributos das plantas às mudanças ambientais é pouco conhecida ao longo de gradientes ecológicos. Testou-se a hipótese de que a complexidade arquitetural de Miconia albicans (SW.) Triana aumenta com a complexidade ambiental, ao longo de um gradiente de fertilidade de solo em um Cerrado no sudeste brasileiro. A arquitetura e a produtividade (estimada por meio do número total de infrutescências sadias) foram examinadas em três tipos de vegetação, que formam um gradiente de fertilidade de solo, de densidade de árvores e de cobertura vegetal: campo sujo (CS), cerrado sensu stricto (CE) e cerradão (CD). Plantas crescendo no CS foram menores e apresentaram menor complexidade arquitetural, enquanto plantas no CD foram maiores. No entanto, de forma inesperada, plantas no CD não demonstraram maior complexidade arquitetural quando comparadas às plantas de CE, e a maior similaridade arquitetural entre plantas de CE e CD, quando comparadas com plantas de CE e CS, sugere que a expressão de atributos funcionais da arquitetura é limitada sob condições de sombra. Plantas de CE produziram mais ramos quaternários e maior número de infrutescências, sendo que a maior produtividade destas plantas indica que a variação de atributos em gradientes ecológicos é mais complexa do que se pensava anteriormente. Galhas de nematóides destruíram 76,5% das infrutescências nas vegetações, mas a porcentagem de ataque não esteve fortemente correlacionada com as variáveis arquiteturais. Os resultados do presente trabalho sugerem limitação na expressão da arquitetura da planta induzida pela sombra e apontam para uma complexa variação fenotípica na colonização de ambientes heterogêneos de savanas neotropicais.

Composição bromatológica e produtividade do capim-andropógonem diferentes idades de rebrota em sistema silvipastoril / Bromatological composition and productivity of Andropogon grass at different ages of resprout in silvopastoral system

Avaliou-se a composição e produtividade do capim-andropógon (Andropogon gayanus Kunth.) em diferentes idades de rebrota, associado às espécies arbóreas pau-darco (Tabebuia serratifolia) e jatobá (Hymenaea courbaril) no período chuvoso, em Teresina, Piauí. Adotou-se o delineamento de blocos ao acaso, em esquema fatorial 3 x 2, com parcelas subdivididas, sendo as parcelas os ambientes (sombreamento com jatobá e pau-darco e área aberta) e posições (nascente e poente), e as subparcelas as idades (35, 49 e 63 dias), com cinco repetições. Houve interação ambiente x idade para MS na planta, FDN e FDA na planta e nas folhas e NIDN nas folhas. A PB nas folhas à sombra foi superior (9,65 ± 0,94%) em relação à área aberta (9,16 ± 1,41%). Houve efeito linear decrescente (p < 0,01) da idade sobre PB na planta e folhas. A partir dos 47,7 e 48,8 dias houve redução na relação folha/colmo do capim-andropógon em pau-darco e jatobá, respectivamente. É viável o cultivo da gramínea em sistema silvipastoril, pela estabilidade na composição de MS e FDA e nas proporções de NIDN e NIDA na planta e nas folhas, bem como para PB na planta e FDN nas folhas, com maior teor de PB nas folhas da gramínea. Não houve diferença nos teores de FDN na gramínea em função da idade entre os ambientes.(AU)

The study evaluated composition and productivity of Andropogon grass at different ages of resprout, associated with the species Tabebuia serratifolia and Hymenaea courbaril, during the rainy period, in Teresina, Piauí. A random blocks design was adopted, in a 3 x 2 factorial scheme, with subdivided parcels, being the parcels the environments (shading with Tabebuia serratifolia, Hymenaea courbaril and open area) and position (rising and setting), and subparcels the ages (35, 49 and 63 days), with five repetitions. There was interaction between the environment and age for DM in the plant, NDF and ADF in the plant and leaves, and NDIN in leaves. The CP in leaves in the shade was higher (9.65 ± 0.94%) compared to the open area (9.16 ± 1.41%). There was a linear effect (p < 0.01) of the age on CP in the plant and leaves. From days 47.7 and 48.8 there was a reduction in the leaf/steam ratio of Andropogon grass under T. serratifolia and H. courbaril, respectively. The culture of the grass in silvopastoral systems is viable, due to the stability in the composition of DM and ADF in the ratios of NDIN and ADIN in the plant and leaves, as well as for CP in plant and NDF in leaves, with higher CP levels in leaves of the grass. There was no difference in the levels of NDF in the grass in function of the age among environments.(AU)

Composição bromatológica e produtividade do capim-andropógonem diferentes idades de rebrota em sistema silvipastoril / Bromatological composition and productivity of Andropogon grass at different ages of resprout in silvopastoral system

Avaliou-se a composição e produtividade do capim-andropógon (Andropogon gayanus Kunth.) em diferentes idades de rebrota, associado às espécies arbóreas pau-d’arco (Tabebuia serratifolia) e jatobá (Hymenaea courbaril) no período chuvoso, em Teresina, Piauí. Adotou-se o delineamento de blocos ao acaso, em esquema fatorial 3 x 2, com parcelas subdivididas, sendo as parcelas os ambientes (sombreamento com jatobá e pau-d’arco e área aberta) e posições (nascente e poente), e as subparcelas as idades (35, 49 e 63 dias), com cinco repetições. Houve interação ambiente x idade para MS na planta, FDN e FDA na planta e nas folhas e NIDN nas folhas. A PB nas folhas à sombra foi superior (9,65 ± 0,94%) em relação à área aberta (9,16 ± 1,41%). Houve efeito linear decrescente (p < 0,01) da idade sobre PB na planta e folhas. A partir dos 47,7 e 48,8 dias houve redução na relação folha/colmo do capim-andropógon em pau-d’arco e jatobá, respectivamente. É viável o cultivo da gramínea em sistema silvipastoril, pela estabilidade na composição de MS e FDA e nas proporções de NIDN e NIDA na planta e nas folhas, bem como para PB na planta e FDN nas folhas, com maior teor de PB nas folhas da gramínea. Não houve diferença nos teores de FDN na gramínea em função da idade entre os ambientes.

The study evaluated composition and productivity of Andropogon grass at different ages of resprout, associated with the species Tabebuia serratifolia and Hymenaea courbaril, during the rainy period, in Teresina, Piauí. A random blocks design was adopted, in a 3 x 2 factorial scheme, with subdivided parcels, being the parcels the environments (shading with Tabebuia serratifolia, Hymenaea courbaril and open area) and position (rising and setting), and subparcels the ages (35, 49 and 63 days), with five repetitions. There was interaction between the environment and age for DM in the plant, NDF and ADF in the plant and leaves, and NDIN in leaves. The CP in leaves in the shade was higher (9.65 ± 0.94%) compared to the open area (9.16 ± 1.41%). There was a linear effect (p < 0.01) of the age on CP in the plant and leaves. From days 47.7 and 48.8 there was a reduction in the leaf/steam ratio of Andropogon grass under T. serratifolia and H. courbaril, respectively. The culture of the grass in silvopastoral systems is viable, due to the stability in the composition of DM and ADF in the ratios of NDIN and ADIN in the plant and leaves, as well as for CP in plant and NDF in leaves, with higher CP levels in leaves of the grass. There was no difference in the levels of NDF in the grass in function of the age among environments.

Interspecific variation in seed size and safe sites in a temperate rain forest.

• The safe site concept could have utility in community ecology if predictive relationships between plant traits and safe site characteristics could be identified. Here we examine the proposal that the nature and relative abundance of safe sites are systematically related to seed size, in an assemblage of 17 woody species in a temperate forest. • We compared the degree of association of seedlings of each species with elevated microsites, and examined life-history correlates of this variation. • Seed size explained 45% of interspecific variation in percentage of seedlings growing on logs and other elevated substrates. Neither specific leaf area nor an index of light requirements gave significant increases in explanatory power. Phylogenetically independent contrasts gave a similar, but weaker, relationship with seed size. Most small-seeded species were overrepresented on elevated microsites, whereas two large-seeded taxa were underrepresented on these substrates. Safe sites of small-seeded species were therefore more spatially restricted than those of large-seeded taxa, as elevated surfaces occupied only 8% of the forest floor. • Safe site differentiation may help to explain the wide range of seed size present within many communities, as well as species coexistence in forests.

Germination requirements and seedling shade tolerance are not correlated in a Chilean temperate rain forest.

• In a comparative study of 26 Chilean temperate woody species, the following questions were addressed. Are germination requirements correlated with seedling light requirements? Can germination requirements be used to delimit pioneer and nonpioneer guilds? • To assess dependence of germination on canopy gaps, germination percentages were compared experimentally in gap and understorey environments. Seedling light requirements were quantified by determining the mean light environment naturally occupied by juveniles of each species. • Germination was significantly higher in the gap than in the understorey in > 30% of the species, although only one germinated exclusively in the gap. In the understorey, germination was higher in 19% of species, whereas 50% were indifferent to germination environment. Several light-demanding species germinated abundantly in the understorey. Both cross-species analysis and phylogenetically independent contrasts showed that seedling light requirements were not correlated with responsiveness of germination to the gap environment. • Results suggest that germination requirements and seedling light requirements are not coupled, and that requirement of canopy gaps for germination cannot be used to delimit pioneer and nonpioneer guilds in a Chilean temperate rain forest.

Relationships of leaf dark respiration with light environment and tissue nitrogen content in juveniles of 11 cold-temperate tree species.

It has been argued that plants adapted to low light should have lower carbon losses via dark respiration (Rd) than those not so adapted, and similarly, all species would be expected to down-regulate Rd in deep shade, because the associated advantages of high metabolic potential cannot be realized in such habitats. In order to test these hypotheses, and to explore the determinants of intraspecific variation in respiration rates, we measured Rd, leaf mass per unit area (LMA), and nitrogen content of mature foliage in juveniles of 11 cold-temperate tree species (angiosperms and conifers), growing in diverse light environments in forest understories in northern Minnesota. Among the seven angiosperm species, respiration on mass, area, and nitrogen bases showed significant negative overall relationships with shade tolerance level. Mass-based respiration rates (Rd mass) of angiosperms as a group showed a significant positive overall relationship with an index of light availability (percentage canopy openness, %CO). Rd mass of most conifers also showed evidence of acclimation of Rd mass to light availability. LMA of all species also increased with increasing %CO, but this response was generally much stronger in angiosperms than in conifers. As a result, the response of area-based respiration (Rd area) to %CO was dominated by ΔRd mass for conifers, and by ΔLMA for most angiosperms, i.e., functional types differed in the components of acclimation of Rd area to light availability. Among the seven angiosperm species, the relationships of leaf N on a mass basis (N mass) with %CO were modulated by shade tolerance: negative slopes in shade-tolerant species may be related to the steep increases in LMA of these taxa along gradients of increasing light intensity, and associated dilution of N-rich, metabolically active tissue by increasing investment in leaf structural components. Although N mass was therefore an unreliable predictor of variation in Rd mass along light gradients, respiration per unit leaf N (Rd/N) was significantly positively correlated with %CO for most species. This probably reflects variation in the proportion of leaf N allocated to protein and/or the influence of leaf carbohydrate status on Rd. Species shade tolerance differences were not significantly correlated with the magnitude of either ΔRd mass or ΔRd area, indicating that variation in acclimation potential of Rd is much less important than inherent differences in this trait. Acclimation of Rd mass to light availability appears to be a generalized feature of juvenile trees, and the important ecological trade-off is likely between high metabolic capacity in high light and low respiratory losses in low light.